17Drought threatens tropical rainforests over seasonal to decadal timescales [1][2][3][4] , but the drivers 18 of tree mortality following drought remain poorly understood 5,6 . It has been suggested that 19 reduced availability of non-structural carbohydrates (NSC) critically increases mortality risk 20 through insufficient carbon supply to metabolism ('carbon starvation') 7,8 . However little is 21 known about how NSC stores are affected by drought, especially over the long term, and 22 whether they are more important than hydraulic processes in determining drought-induced 23 mortality. Using data from the world's longest-running experimental drought study in tropical 24 rainforest (in the Brazilian Amazon), we test whether carbon starvation or deterioration of the 25 water-conducting pathways from soil to leaf trigger tree mortality. Biomass loss from 26 mortality in the experimentally-droughted forest increased substantially after >10 years of 27 reduced soil moisture availability. The mortality signal was dominated by the death of large 28 trees, which were at a much greater risk of hydraulic deterioration than smaller trees. 29However, we find no evidence that the droughted trees suffered carbon starvation, as their 30 NSC concentrations were similar to those of un-droughted trees, and growth rates did not 31 decline in either living or dying individuals. Our results indicate that hydraulics, rather than 32 carbon starvation, triggers tree death from drought in tropical rainforest. 34Drought-response observations from both field-scale experiments and natural droughts have 35 demonstrated increased mortality over the short-term (1-3 years), with notably higher 36 vulnerability for some taxa, and for larger trees 6,9,10 . After several years of drought, 37 recovering growth rates in smaller trees, dbh (diameter at breast height) <40 cm, and reduced 38 mortality have been recorded at different locations 6,11,12 . However, the long-term (>10 yr) 39 sensitivity of tropical forests to predicted prolonged and repeated water deficit [1][2][3] we synthesise these data to test whether long-term soil moisture deficit alters NSC storage 64 and use in tropical rainforest trees, and if this, or hydraulic processes, are most strongly 65 associated with increased mortality rates. 66By 2014, following 13 years of the TFE treatment, cumulative biomass loss through mortality 67 was 41.0±2.7% relative to pre-treatment values (Fig. 1a), and the rate of loss had increased 68 substantially since the previous reported value of 17.2±0.8%, after 7 years of TFE 6 . 69Accelerating biomass loss and failure to recover substantially, or to reach a new 70 equilibrium 13 , has led to a committed flux to the atmosphere from decomposing necromass of 71 101.9±19.1 Mg C ha -1 (Fig. 1a). This biomass loss has been driven by elevated mortality in 72 the largest trees (Fig. 1b), as previously observed over shorter timescales 6 , and has created a 73 canopy that has had a persistently lower average leaf area index during 2010-2014 74 (12.0±1...

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The allocation of the net primary productivity (NPP) of an ecosystem between canopy, woody tissue and fine roots is an important descriptor of the functioning of that ecosystem, and an important feature to correctly represent in terrestrial ecosystem models. Here, we collate and analyse a global dataset of NPP allocation in tropical forests, and compare this with the representation of NPP allocation in 13 terrestrial ecosystem models. On average, the data suggest an equal partitioning of allocation between all three main components (mean 34 + 6% canopy, 39 + 10% wood, 27 + 11% fine roots), but there is substantial site-to-site variation in allocation to woody tissue versus allocation to fine roots. Allocation to canopy (leaves, flowers and fruit) shows much less variance. The mean allocation of the ecosystem models is close to the mean of the data, but the spread is much greater, with several models reporting allocation partitioning outside of the spread of the data. Where all main components of NPP cannot be measured, litterfall is a good predictor of overall NPP (r 2 ¼ 0.83 for linear fit forced through origin), stem growth is a moderate predictor and fine root production a poor predictor. Across sites the major component of variation of allocation is a shifting allocation between wood and fine roots, with allocation to the canopy being a relatively invariant component of total NPP. This suggests the dominant allocation trade-off is a 'fine root versus wood' trade-off, as opposed to the expected 'root-shoot' trade-off; such a trade-off has recently been posited on theoretical grounds for old-growth forest stands. We conclude by discussing the systematic biases in estimates of allocation introduced by missing NPP components, including herbivory, large leaf litter and root exudates production. These biases have a moderate effect on overall carbon allocation estimates, but are smaller than the observed range in allocation values across sites.

Summary Tree mortality rates appear to be increasing in moist tropical forests (MTFs) with significant carbon cycle consequences. Here, we review the state of knowledge regarding MTF tree mortality, create a conceptual framework with testable hypotheses regarding the drivers, mechanisms and interactions that may underlie increasing MTF mortality rates, and identify the next steps for improved understanding and reduced prediction. Increasing mortality rates are associated with rising temperature and vapor pressure deficit, liana abundance, drought, wind events, fire and, possibly, CO2 fertilization‐induced increases in stand thinning or acceleration of trees reaching larger, more vulnerable heights. The majority of these mortality drivers may kill trees in part through carbon starvation and hydraulic failure. The relative importance of each driver is unknown. High species diversity may buffer MTFs against large‐scale mortality events, but recent and expected trends in mortality drivers give reason for concern regarding increasing mortality within MTFs. Models of tropical tree mortality are advancing the representation of hydraulics, carbon and demography, but require more empirical knowledge regarding the most common drivers and their subsequent mechanisms. We outline critical datasets and model developments required to test hypotheses regarding the underlying causes of increasing MTF mortality rates, and improve prediction of future mortality under climate change.

Large herbivores and carnivores (the megafauna) have been in a state of decline and extinction since the Late Pleistocene, both on land and more recently in the oceans. Much has been written on the timing and causes of these declines, but only recently has scientific attention focused on the consequences of these declines for ecosystem function. Here, we review progress in our understanding of how megafauna affect ecosystem physical and trophic structure, species composition, biogeochemistry, and climate, drawing on special features of PNAS and Ecography that have been published as a result of an international workshop on this topic held in Oxford in 2014. Insights emerging from this work have consequences for our understanding of changes in biosphere function since the Late Pleistocene and of the functioning of contemporary ecosystems, as well as offering a rationale and framework for scientifically informed restoration of megafaunal function where possible and appropriate.For hundreds of millions of years, an abundance of large animals, the megafauna, was a prominent feature of the land and oceans. However, in the last few tens of thousands of years-a blink of an eye on many evolutionary and biogeochemical timescales-something dramatic happened to Earth's ecology; megafauna largely disappeared from vast areas, rendered either actually or functionally extinct (1, 2). Only in small parts of the world do megafauna exist at diversities anything close to their previous state, and, in many of these remaining regions, they are in a state of functional decline through population depletion and range contraction. In the oceans, a similar process has occurred over the last few hundred years: although there has been little absolute extinction, there has been a dramatic decline in the abundance of whales and large fish through overharvesting (3). Both on land and in oceans, declines continue today (4-7).Homo sapiens evolved and dispersed in a world teeming with giant creatures. Our earliest art forms, such as the haunting and mesmerizing Late Pleistocene cave paintings of Lascaux and Altamira, show that megafauna had a profound impact on the psyche and spirituality of our ancestors. To humans past and modern, they indicate resources, danger, power, and charisma, but, beyond these impacts, such large animals have profound and distinct effects on the nature and functioning of the ecosystems they inhabit.Martin (8) first posited a major human role in past megafaunal disappearances, and, since then, much has been written on their patterns and causes and the relative importance of human effects, climate change, and other factors (8)(9)(10)(11)(12)(13)(14)(15). Only recently has work begun to address the environmental consequences of this dramatic transition from a megafaunal to a nonmegafaunal world on Earth's ecology, as manifested through vegetation cover (16), plant-animal interactions (17), ecosystem structure (16, 18), trophic interactions (7), fire regimes (19), biogeochemical cycling (20), and climate (21,22).In this pap...

[1] We used leaf gas exchange, sap flow, and eddy covariance measurements to investigate whether high temperature substantially limits CO 2 uptake at the LBA-ECO (Large-scale Biosphere-Atmosphere) km-83 tropical forest site in Brazil. Leaf-level temperature-photosynthesis curves, and comparisons of whole-canopy net ecosystem CO 2 exchange (NEE) with air temperature, showed that CO 2 uptake declined sharply during warm periods. Observations of ambient leaf microclimate showed that leaves oscillate between two states: a cool, dimly lit stage and a hot, brightly illuminated stage where leaf temperatures are often greater than 35°C. The leaf-level rates of photosynthesis decreased when shaded leaves ($ambient air temperature and < 500 mmol m À2 s À1 ) were transferred into a prewarmed, brightly illuminated chamber (35°to 38°C and 1000 mmol m À2 s À1 ), coincident with increased leaf temperature, increased evaporative demand, and stomatal closure. The rates of whole-canopy CO 2 uptake calculated at 5-min intervals increased initially at the onset of sunny periods that followed extended cloudy periods, but then decreased as the sunlight continued, leaf temperature and evaporative demand increased, and canopy conductance decreased. The forest at km-83 appears to be close to a high temperature threshold, above which CO 2 uptake drops sharply. This sensitivity results in part from the covariance between leaf temperature and leaf illumination; the brightly illuminated leaves that contribute disproportionately to canopy photosynthesis are warmed to the point that leaf gas exchange is curtailed.

The past was a world of giants, with abundant whales in the sea and large animals roaming the land. However, that world came to an end following massive late-Quaternary megafauna extinctions on land and widespread population reductions in great whale populations over the past few centuries. These losses are likely to have had important consequences for broad-scale nutrient cycling, because recent literature suggests that large animals disproportionately drive nutrient movement. We estimate that the capacity of animals to move nutrients away from concentration patches has decreased to about 8% of the preextinction value on land and about 5% of historic values in oceans. For phosphorus (P), a key nutrient, upward movement in the ocean by marine mammals is about 23% of its former capacity (previously about 340 million kg of P per year). Movements by seabirds and anadromous fish provide important transfer of nutrients from the sea to land, totalling ∼150 million kg of P per year globally in the past, a transfer that has declined to less than 4% of this value as a result of the decimation of seabird colonies and anadromous fish populations. We propose that in the past, marine mammals, seabirds, anadromous fish, and terrestrial animals likely formed an interlinked system recycling nutrients from the ocean depths to the continental interiors, with marine mammals moving nutrients from the deep sea to surface waters, seabirds and anadromous fish moving nutrients from the ocean to land, and large animals moving nutrients away from hotspots into the continental interior. There were giants in the world in those days.Genesis 6:4, King James version T he past was a world of giants, with abundant whales in the oceans and terrestrial ecosystems teeming with large animals. However, most ecosystems lost their large animals, with around 150 mammal megafaunal (here, defined as ≥44 kg of body mass) species going extinct in the late Pleistocene and early Holocene (1, 2). These extinctions and range declines continued up through historical times and, in many cases, into the present (3). No global extinctions are known for any marine whales, but whale densities might have declined between 66% and 99% (4-6). Some of the largest species have experienced severe declines; for example, in the Southern Hemisphere, blue whales (Balaenoptera musculus) have been reduced to 1% of their historical numbers as a result of commercial whaling (4). Much effort has been devoted to determining the cause of the extinctions and declines, with less effort focusing on the ecological impacts of the extinctions. Here, we focus on the ecological impacts, with a specific focus on how nutrient dynamics may have changed on land following the lateQuaternary megafauna extinctions, and in the sea and air following historical hunting pressures.Most biogeochemists studying nutrient cycling focus on in situ production, such as weathering or biological nitrogen (N) fixation, largely ignoring lateral fluxes by animals because they are considered of secondary importance (...