Leaf regrowth stage as a morpho-physiological indicator of Bromus valdivianus and Lolium perenne mixed pasture defoliation in New Zealand grazing system : a thesis presented in partial fulfilment of the requirements for the degree of Doctor of Philosophy in Plant Science at Massey University, Manawatū, New Zealand

Bromus valdivianus Phill. is a perennial grass species native to the South of Chile. Its better fitness for growing under low water availabilities compared with Lolium perenne L. has been assessed in field and glasshouse studies. However, Bromus valdivianus morpho-physiological attributes, such as root development under field conditions, and competitive ability are likely to manifest differently under contrasting defoliation criteria based on leaf regrowth stage. In addition, Lolium perenne and B. valdivianus growth can be complementary throughout the year under rainfed conditions. Therefore, the objective of the present thesis was to determine defoliation criteria based on leaf regrowth stage of mixed pastures (50/50% L. perenne and B. valdivianus) and consequences for species succession and functional traits development of the species. Four studies were developed in the thesis, two glasshouse and two field studies. The first glasshouse study evaluated the growth response and water-soluble carbohydrate (WSC) accumulation of B. valdivianus at three different defoliation frequencies. The results suggested that B. valdivianus increased herbage mass production and root biomass at low defoliation frequency (i.e., at 3.5–4.0 leaf regrowth stage). This was related to a higher WSC accumulation in the tiller base. In addition, under low soil water availabilities (20–25% of field capacity) WSC increased by ~20%, which indicated a drought resistant strategy of this species. The second study proved the growth enhancement (mainly root length and biomass) of B. valdivianus under competition with L. perenne compared with intraspecific competition when soil water shifted from high to low availabilities. In field studies, B. valdivianus mixed with L. perenne increased ~15% accumulated herbage mass compared to the species monocultures, which supported higher production during dry periods. This was due to the niche complementarity and asynchrony in herbage growth between the species, with B. valdivianus capable to maintain a steady tiller population throughout the year and root biomass accumulation at depth. Whereas L. perenne grew more than B. valdivianus under low level of oxygen in the soil and under optimal growth conditions and presented a higher nutritive value than B. valdivianus during winter, spring and autumn. The latter supports the partial grazing preference for L. perenne shown by sheep during part of the year, as measured in the second field study. This lower nutritive value (less energy) of B. valdivianus monocultures was overcame in the mixture, with a good overall value for high animal production. Regarding the mixture defoliation criterion, it was shown that it can be based on the optimal leaf regrowth stage of either species, as herbage mass production was similar between defoliation frequencies based on optimal leaf regrowth stage of either of the species. However, under defoliation based on optimal leaf regrowth stage of B. valdivianus, root biomass accumulation at depth increased ~45%. Overall, the results of the present thesis evidenced that Bromus valdivianus can successfully grow alongside L. perenne and, therefore, the mixture can increase forage production of New Zealand farming system in the event of climate change