Vein and stomatal traits in leaves of three co-occurring Quercus species differing in leaf life span

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ORIGINAL PAPER

Vein and stomatal traits in leaves of three co‑occurring Quercus species differing in leaf life span S. Mediavilla1   · I. Martín2 · A. Escudero1 Received: 3 December 2019 / Revised: 13 April 2020 / Accepted: 8 May 2020 © Springer-Verlag GmbH Germany, part of Springer Nature 2020

Abstract We analyzed vein and stomatal traits (vein density and vein volume per leaf area, stomatal density, stomatal pore length and pore index) and their relationships with other morphological traits [leaf area, leaf thickness and leaf mass per unit area (LMA)] of three co-occurring Mediterranean tree species with contrasting leaf habits [Quercus faginea Lam., Q. suber L. and Q. ilex L. subsp. ballota (Desf.) Samp.]. The results showed that leaf size, thickness and LMA varied among the species in parallel with the differences in leaf longevity. By contrast, the traits most related to water use showed inconsistent differences among the three species. Stomatal pore index was lowest in the species with intermediate leaf life span. The species with longest leaf longevity had highest vein density but minimum vein volume per area. Vein and stomatal traits also varied independently from other leaf traits within each species. The absence of association between LMA and vein volume suggests that a large LMA is the result of the accumulation of other tissues, and not necessarily veins. We concluded that in contrast with most economic leaf traits that tend to vary in parallel with leaf life span, traits related to water use varied inconsistently with leaf duration in the three species studied. Stomatal and vein traits, in addition, were not related to the maximum stomatal conductance of the different species. Keywords  Leaf venation · Leaf life span · LMA · Stomatal traits · Leaf economics spectrum · Quercus species

Introduction Leaves are the most important organs for plant growth and, as a consequence, leaf traits, which determine leaf performance, exhibit dramatic changes among different species and in response to environmental gradients (Roth-Nebelsick et al. 2001; Brodribb et al. 2010; Zhang et al. 2018). Amid the most important leaf traits, besides form and size, are vein patterns, which are responsible for at least two essential functions. Veins allow the transport of water for photosynthesis and transpiration, as well as nutrients, photosynthate Communicated by Judy Simon. * S. Mediavilla [email protected] 1



Área de Ecología, Facultad de Biología, Universidad de Salamanca, Campus Miguel de Unamuno, 37007 Salamanca, Spain



Dpto. de Botánica y Fisiología Vegetal, Facultad de Biología, Universidad de Salamanca, Campus Miguel de Unamuno, 37007 Salamanca, Spain

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and molecular signals to the rest of the plant (Brodribb et al. 2007, 2010; Sack et al. 2012; Sack and Scoffoni 2013). Additionally, leaf veins provide mechanical support, allowing the plant to display its leaves toward light, and play a key role in defensive functions against herbivores and environmental stressors (Katifori et al. 2010; Méndez-Alonzo et al. 2013