Yerba maté (Ilex paraguariensis, Aquifoliaceae) is a subtropical, evergreen, dioecious, South American tree. In one preliminary study, it was observed that the functional strategy of yerba mate females, aiming to finish reproductive process, was increased transpiration relative to photosynthetic rates compared with males, on self-shaded leaves. We hypothesised that the long-term gas exchange response of males and females can evolve independently of phenological stage and cultivation type. In this spirit, the primary aim of the study was to analyse the physiological sexual dimorphism of this species, evaluating fluctuations of gas exchanges related to microclimate and phenological stages. A field study was conducted on adult plants of yerba maté cultivated in monoculture (MO) and in forest understorey (FUS), and measurements carried out in situ on microclimate and leaf gas exchange parameters. The photosynthetic photon flux density that was attained at leaf level in FUS was reduced roughly 10-fold compared with that in MO. Various leaf age populations were observed during a 2-year period at 2-month intervals and grouped into four categories: young, young-fully-expanded, fully-expanded and old. Young and youngfully-expanded leaves were the most active in photosynthesis. Leaves of female plants showed greater photosynthetic rate than those of male plants, which was expressed on all leaf age categories in MO, but only during vegetative stages previous to flowering and fruit ripening. The photosynthesis of young-fully-expanded leaves of females grown in FUS was superior to males but only during winter growth pause. The stomatal conductance differed in relation to cultivation type and leaf age but did not show the sexual differentiation. Physiological sexual dimorphism in yerba mate is shown to be plastic, responding to environmental conditions. The cost associated to the reproduction of yerba maté could be most easily met showing physiological differentiation of both sexes. A higher reproductive investment of females might be compensated for by exhibiting greater leaf photosynthesis than males that occurs in vegetative stages that precede flowering and fruit ripening. Sexual dimorphism and seasonal changes in Ilex paraguariensis M. Rakocevic et al. M. Rakocevic et al. Sexual dimorphism and seasonal changes in Ilex paraguariensis Ann Appl Biol 154 (2009) 291-301 ª

To assess differences in the lag-effect pattern in the relationship between yerba-mate biennial growth and environmental factors, a time-series analysis was performed. A generalized Poisson regression model was used to control time trends, temperature, growing degree days (GDD), rainfalls and night length (NL). It was hypothesized that the active growth and growth pauses in yerba-mate are controlled endogenously and modified by environment, and that genders would respond differently to environmental modifications. The patterns in the lag effect from the distributed-lag models were similar to those of time-series models with meteorological data means with lag = 0. GDD and NL were principal factors affecting biennial yerba-mate shoot elongation and the number of green leaves of females grown in monoculture, besides their significant effects on metamer emission and leaf area in males grown in monoculture. NL also had a significant influence on shoot elongation and leaf area of both genders grown in forest understorey (FUS), indicating that yerba-mate growth is synchronized by an internal clock sensitive to temperature adjustments. The morphological plasticity and the adaptation efforts of yerba-mate were more pronounced in monoculture than in FUS. Sexual dimorphism was expressed-males were more sensitive to environmental changes than females, especially in monoculture. Growth modifications were much more intense when plants were grown in a cultivation system that is less like yerba-mate natural habitat (monoculture) than in one resembling its natural habitat (FUS). Our data support the ecological specialization theory.

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