The leaf economics spectrum1,2 and the global spectrum of plant forms and functions3 revealed fundamental axes of variation in plant traits, which represent different ecological strategies that are shaped by the evolutionary development of plant species2. Ecosystem functions depend on environmental conditions and the traits of species that comprise the ecological communities4. However, the axes of variation of ecosystem functions are largely unknown, which limits our understanding of how ecosystems respond as a whole to anthropogenic drivers, climate and environmental variability4,5. Here we derive a set of ecosystem functions6 from a dataset of surface gas exchange measurements across major terrestrial biomes. We find that most of the variability within ecosystem functions (71.8%) is captured by three key axes. The first axis reflects maximum ecosystem productivity and is mostly explained by vegetation structure. The second axis reflects ecosystem water-use strategies and is jointly explained by variation in vegetation height and climate. The third axis, which represents ecosystem carbon-use efficiency, features a gradient related to aridity, and is explained primarily by variation in vegetation structure. We show that two state-of-the-art land surface models reproduce the first and most important axis of ecosystem functions. However, the models tend to simulate more strongly correlated functions than those observed, which limits their ability to accurately predict the full range of responses to environmental changes in carbon, water and energy cycling in terrestrial ecosystems7,8.

Abstract. The dynamics of terrestrial ecosystems are shaped by the coupled cycles of carbon, nitrogen, and phosphorus, and these cycles are strongly dependent on the availability of water and energy. These interactions shape future terrestrial biosphere responses to global change. Here, we present a new terrestrial ecosystem model, QUINCY (QUantifying Interactions between terrestrial Nutrient CYcles and the climate system), which has been designed from scratch to allow for a seamless integration of the fully coupled carbon, nitrogen, and phosphorus cycles with each other and also with processes affecting the energy and water balances in terrestrial ecosystems. This new model includes (i) a representation of plant growth which separates source (e.g. photosynthesis) and sink (growth rate of individual tissues, constrained by temperature and the availability of water and nutrients) processes; (ii) the acclimation of many ecophysiological processes to meteorological conditions and/or nutrient availability; (iii) an explicit representation of vertical soil processes to separate litter and soil organic matter dynamics; (iv) a range of new diagnostics (leaf chlorophyll content; 13C, 14C, and 15N isotope tracers) to allow for a more in-depth model evaluation. In this paper, we present the model structure and provide an assessment of its performance against a range of observations from global-scale ecosystem monitoring networks. We demonstrate that QUINCY v1.0 is capable of simulating ecosystem dynamics across a wide climate gradient, as well as across different plant functional types. We further provide an assessment of the sensitivity of key model predictions to the model's parameterisation. This work lays the ground for future studies to test individual process hypotheses using the QUINCY v1.0 framework in the light of ecosystem manipulation observations, as well as global applications to investigate the large-scale consequences of nutrient-cycle interactions for projections of terrestrial biosphere dynamics.

Abstract. Phenology is essential to our understanding of biogeochemical cycles and the climate system. We develop a global mechanistic model of leaf phenology based on the hypothesis that phenology is a strategy for optimal carbon gain at the canopy level so that trees adjust leaf gains and losses in response to environmental factors such as light, temperature and soil moisture, to achieve maximum carbon assimilation. We fit this model to five years of satellite observations of leaf area index (LAI) using a Bayesian fitting algorithm. We show that our model is able to reproduce phenological patterns for all vegetation types and use it to explore variations in growing season length and the climate factors that limit leaf growth for different biomes. Phenology in wet tropical areas is limited by leaf age physiological constraints while at higher latitude leaf seasonality is limited by low temperature and light availability. Leaf growth in grassland regions is limited by water availability but often in combination with other factors. This model will advance the current understanding of phenology for ecosystem carbon models and our ability to predict future phenological behaviour.

Abstract. Seasonal and year-to-year variations in leaf cover imprint significant spatial and temporal variability on biogeochemical cycles, and affect land-surface properties related to climate. We develop a demographic model of leaf phenology based on the hypothesis that trees seek an optimal leaf area index (LAI) as a function of available light and soil water, and fit it to spaceborne observations of LAI over the Amazon basin, 2001Amazon basin, -2005. We find the model reproduces the spatial and temporal LAI distribution whilst also predicting geographic variation in leaf age from the basin centre (2.1 ± 0.2 years), through to the lowest values over the deciduous eastern and southern Amazon (6 ± 2 months). The model explains the observed increase in LAI during the dry season as a net addition of leaves in response to increased solar radiation. We anticipate our work to be a starting point from which to develop better descriptions of leaf phenology to incorporate into more sophisticated earth system models.