Vapour pressure deficit modulates hydraulic function and structure of tropical rainforests under nonlimiting soil water supply.

Atmospheric conditions are expected to become warmer and drier in the future, but little is known about how evaporative demand influences forest structure and function independently from soil moisture availability, and how fast-response variables (such as canopy water potential and stomatal conductance) may mediate longer-term changes in forest structure and function in response to climate change. We used two tropical rainforest sites with different temperatures and vapour pressure deficits (VPD), but nonlimiting soil water supply, to assess the impact of evaporative demand on ecophysiological function and forest structure. Common species between sites allowed us to test the extent to which species composition, relative abundance and intraspecific variability contributed to site-level differences. The highest VPD site had lower midday canopy water potentials, canopy conductance (gc ), annual transpiration, forest stature, and biomass, while the transpiration rate was less sensitive to changes in VPD; it also had different height-diameter allometry (accounting for 51% of the difference in biomass between sites) and higher plot-level wood density. Our findings suggest that increases in VPD, even in the absence of soil water limitation, influence fast-response variables, such as canopy water potentials and gc , potentially leading to longer-term changes in forest stature resulting in reductions in biomass.

The role of big trees and abundant species in driving spatial patterns of species richness in an Australian tropical rainforest.

Big trees and abundant species dominate forest structure and composition. As a result, their spatial distribution and interactions with other species and individuals may contribute disproportionately to the emergence of spatial heterogeneity in richness patterns. We tested scale-dependent spatial patterning and species richness structures to understand the role of individual trees (big trees) and species (abundant species) in driving spatial richness patterns on a 25 ha plot in a diverse tropical forest of Australia. The individual species area relationship (ISAR) was used to assess species richness in neighborhoods ranging from 1 to 50 m radii around all big trees (≥70 cm dbh, n = 296) and all species with more than 100 individuals in the plot (n = 53). A crossed ISAR function was also used to compute species richness around big trees for trees of different size classes. Big individuals exert some spatial structuring on other big and mid-sized trees in local neighborhoods (up to 30 m and 16 m respectively), but not on small trees. While most abundant species were neutral with respect to richness patterns, we identified consistent species-specific signatures on spatial patterns of richness for 14 of the 53 species. Seven species consistently had higher than expected species richness in their neighborhood (species "accumulators"), and seven had lower than expected (species "repellers") across all spatial scales. Common traits of accumulators and repeller species suggest that niche partitioning along disturbance gradients is a primary mechanism driving spatial richness patterns, which is then manifested in large-scale spatial heterogeneity in species distributions across the plot.

Tropical tree mortality has increased with rising atmospheric water stress.

Evidence exists that tree mortality is accelerating in some regions of the tropics1,2, with profound consequences for the future of the tropical carbon sink and the global anthropogenic carbon budget left to limit peak global warming below 2 °C. However, the mechanisms that may be driving such mortality changes and whether particular species are especially vulnerable remain unclear3-8. Here we analyse a 49-year record of tree dynamics from 24 old-growth forest plots encompassing a broad climatic gradient across the Australian moist tropics and find that annual tree mortality risk has, on average, doubled across all plots and species over the last 35 years, indicating a potential halving in life expectancy and carbon residence time. Associated losses in biomass were not offset by gains from growth and recruitment. Plots in less moist local climates presented higher average mortality risk, but local mean climate did not predict the pace of temporal increase in mortality risk. Species varied in the trajectories of their mortality risk, with the highest average risk found nearer to the upper end of the atmospheric vapour pressure deficit niches of species. A long-term increase in vapour pressure deficit was evident across the region, suggesting that thresholds involving atmospheric water stress, driven by global warming, may be a primary cause of increasing tree mortality in moist tropical forests.

Forest system hydraulic conductance: partitioning tree and soil components.

Soil-leaf hydraulic conductance determines canopy-atmosphere coupling in vegetation models, but it is typically derived from ex-situ measurements of stem segments and soil samples. Using a novel approach, we derive robust in-situ estimates for whole-tree conductance (ktree ), 'functional' soil conductance (ksoil ), and 'system' conductance (ksystem , water table to canopy), at two climatically different tropical rainforest sites. Hydraulic 'functional rooting depth', determined for each tree using profiles of soil water potential (Ψsoil ) and sap flux data, enabled a robust determination of ktree and ksoil . ktree was compared across species, size classes, seasons, height above nearest drainage (HAND), two field sites, and to alternative representations of ktree ; ksoil was analysed with respect to variations in site, season and HAND. ktree was lower and changed seasonally at the site with higher vapour pressure deficit (VPD) and rainfall; ktree differed little across species but scaled with tree circumference; rsoil (1/ksoil ) ranged from 0 in the wet season to 10× less than rtree (1/ktree ) in the dry season. VPD and not rainfall may influence plot-level k; leaf water potentials and sap flux can be used to determine ktree , ksoil and ksystem ; Ψsoil profiles can provide mechanistic insights into ecosystem-level water fluxes.

Tropical tree growth sensitivity to climate is driven by species intrinsic growth rate and leaf traits.

A better understanding of how climate affects growth in tree species is essential for improved predictions of forest dynamics under climate change. Long-term climate averages (mean climate) drive spatial variations in species' baseline growth rates, whereas deviations from these averages over time (anomalies) can create growth variation around the local baseline. However, the rarity of long-term tree census data spanning climatic gradients has so far limited our understanding of their respective role, especially in tropical systems. Furthermore, tree growth sensitivity to climate is likely to vary widely among species, and the ecological strategies underlying these differences remain poorly understood. Here, we utilize an exceptional dataset of 49 years of growth data for 509 tree species across 23 tropical rainforest plots along a climatic gradient to examine how multiannual tree growth responds to both climate means and anomalies, and how species' functional traits mediate these growth responses to climate. We show that anomalous increases in atmospheric evaporative demand and solar radiation consistently reduced tree growth. Drier forests and fast-growing species were more sensitive to water stress anomalies. In addition, species traits related to water use and photosynthesis partly explained differences in growth sensitivity to both climate means and anomalies. Our study demonstrates that both climate means and anomalies shape tree growth in tropical forests and that species traits can provide insights into understanding these demographic responses to climate change, offering a promising way forward to forecast tropical forest dynamics under different climate trajectories.

Releasing incompatible males drives strong suppression across populations of wild and Wolbachia-carrying Aedes aegypti in Australia.

Releasing sterile or incompatible male insects is a proven method of population management in agricultural systems with the potential to revolutionize mosquito control. Through a collaborative venture with the "Debug" Verily Life Sciences team, we assessed the incompatible insect technique (IIT) with the mosquito vector Aedes aegypti in northern Australia in a replicated treatment control field trial. Backcrossing a US strain of Ae. aegypti carrying Wolbachia wAlbB from Aedes albopictus with a local strain, we generated a wAlbB2-F4 strain incompatible with both the wild-type (no Wolbachia) and wMel-Wolbachia Ae. aegypti now extant in North Queensland. The wAlbB2-F4 strain was manually mass reared with males separated from females using Verily sex-sorting technologies to obtain no detectable female contamination in the field. With community consent, we delivered a total of three million IIT males into three isolated landscapes of over 200 houses each, releasing ∼50 males per house three times a week over 20 wk. Detecting initial overflooding ratios of between 5:1 and 10:1, strong population declines well beyond 80% were detected across all treatment landscapes when compared to controls. Monitoring through the following season to observe the ongoing effect saw one treatment landscape devoid of adult Ae. aegypti early in the season. A second landscape showed reduced adults, and the third recovered fully. These encouraging results in suppressing both wild-type and wMel-Ae. aegypti confirms the utility of bidirectional incompatibility in the field setting, show the IIT to be robust, and indicate that the removal of this arbovirus vector from human-occupied landscapes may be achievable.

The importance of large-diameter trees in the wet tropical rainforests of Australia.

Large trees are keystone structures in many terrestrial ecosystems. They contribute disproportionately to reproduction, recruitment and succession, and influence the structure, dynamics and diversity of forests. Recently, researchers have become concerned about evidence showing rapid declines in large, old trees in a range of ecosystems across the globe. We used ≥10 cm diameter at breast height (DBH) stem inventory data from 20, 0.5 ha forest plots spanning the wet tropical rainforest of Queensland, Australia to examine the contribution of large-diameter trees to above ground biomass (AGB), richness, dominance, mortality and recruitment. We show consistencies with tropical rainforest globally in that large-diameter trees (≥70 cm DBH) contribute much of the biomass (33%) from few trees (2.4% of stems ≥10 cm DBH) with the density of the largest trees explaining much of the variation (62%) in AGB across plots. Measurement of AGB in the largest 5% of trees allows plot biomass to be predicted with ~85% precision. In contrast to rainforest in Africa and America, we show that a high proportion of tree species are capable of reaching a large-diameter in Australian wet tropical rainforest resulting in weak biomass hyperdominance (~10% of species account for 50% of the biomass) leading to high potential resilience to regional disturbances and global environmental change. We show that the high AGB in Australian tropical forests is driven primarily by the high density of large trees coupled with contributions from high densities of medium size trees. Australian wet tropical rainforests are well positioned to maintain the current densities of large-diameter trees and high AGB into the future due to the species richness of large trees and a high density of replacement smaller trees.

BioTIME: A database of biodiversity time series for the Anthropocene.

MOTIVATION: The BioTIME database contains raw data on species identities and abundances in ecological assemblages through time. These data enable users to calculate temporal trends in biodiversity within and amongst assemblages using a broad range of metrics. BioTIME is being developed as a community-led open-source database of biodiversity time series. Our goal is to accelerate and facilitate quantitative analysis of temporal patterns of biodiversity in the Anthropocene. MAIN TYPES OF VARIABLES INCLUDED: The database contains 8,777,413 species abundance records, from assemblages consistently sampled for a minimum of 2 years, which need not necessarily be consecutive. In addition, the database contains metadata relating to sampling methodology and contextual information about each record. SPATIAL LOCATION AND GRAIN: BioTIME is a global database of 547,161 unique sampling locations spanning the marine, freshwater and terrestrial realms. Grain size varies across datasets from 0.0000000158 km2 (158 cm2) to 100 km2 (1,000,000,000,000 cm2). TIME PERIOD AND GRAIN: BioTIME records span from 1874 to 2016. The minimal temporal grain across all datasets in BioTIME is a year. MAJOR TAXA AND LEVEL OF MEASUREMENT: BioTIME includes data from 44,440 species across the plant and animal kingdoms, ranging from plants, plankton and terrestrial invertebrates to small and large vertebrates. SOFTWARE FORMAT: .csv and .SQL.

Contrasting effects of defaunation on aboveground carbon storage across the global tropics.

Defaunation is causing declines of large-seeded animal-dispersed trees in tropical forests worldwide, but whether and how these declines will affect carbon storage across this biome is unclear. Here we show, using a pan-tropical data set, that simulated declines of large-seeded animal-dispersed trees have contrasting effects on aboveground carbon stocks across Earth's tropical forests. In our simulations, African, American and South Asian forests, which have high proportions of animal-dispersed species, consistently show carbon losses (2-12%), but Southeast Asian and Australian forests, where there are more abiotically dispersed species, show little to no carbon losses or marginal gains (±1%). These patterns result primarily from changes in wood volume, and are underlain by consistent relationships in our empirical data (∼2,100 species), wherein, large-seeded animal-dispersed species are larger as adults than small-seeded animal-dispersed species, but are smaller than abiotically dispersed species. Thus, floristic differences and distinct dispersal mode-seed size-adult size combinations can drive contrasting regional responses to defaunation.

Testing the generality of above-ground biomass allometry across plant functional types at the continent scale.

Accurate ground-based estimation of the carbon stored in terrestrial ecosystems is critical to quantifying the global carbon budget. Allometric models provide cost-effective methods for biomass prediction. But do such models vary with ecoregion or plant functional type? We compiled 15 054 measurements of individual tree or shrub biomass from across Australia to examine the generality of allometric models for above-ground biomass prediction. This provided a robust case study because Australia includes ecoregions ranging from arid shrublands to tropical rainforests, and has a rich history of biomass research, particularly in planted forests. Regardless of ecoregion, for five broad categories of plant functional type (shrubs; multistemmed trees; trees of the genus Eucalyptus and closely related genera; other trees of high wood density; and other trees of low wood density), relationships between biomass and stem diameter were generic. Simple power-law models explained 84-95% of the variation in biomass, with little improvement in model performance when other plant variables (height, bole wood density), or site characteristics (climate, age, management) were included. Predictions of stand-based biomass from allometric models of varying levels of generalization (species-specific, plant functional type) were validated using whole-plot harvest data from 17 contrasting stands (range: 9-356 Mg ha(-1) ). Losses in efficiency of prediction were <1% if generalized models were used in place of species-specific models. Furthermore, application of generalized multispecies models did not introduce significant bias in biomass prediction in 92% of the 53 species tested. Further, overall efficiency of stand-level biomass prediction was 99%, with a mean absolute prediction error of only 13%. Hence, for cost-effective prediction of biomass across a wide range of stands, we recommend use of generic allometric models based on plant functional types. Development of new species-specific models is only warranted when gains in accuracy of stand-based predictions are relatively high (e.g. high-value monocultures).

An estimate of the number of tropical tree species.

The high species richness of tropical forests has long been recognized, yet there remains substantial uncertainty regarding the actual number of tropical tree species. Using a pantropical tree inventory database from closed canopy forests, consisting of 657,630 trees belonging to 11,371 species, we use a fitted value of Fisher's alpha and an approximate pantropical stem total to estimate the minimum number of tropical forest tree species to fall between ∼ 40,000 and ∼ 53,000, i.e., at the high end of previous estimates. Contrary to common assumption, the Indo-Pacific region was found to be as species-rich as the Neotropics, with both regions having a minimum of ∼ 19,000-25,000 tree species. Continental Africa is relatively depauperate with a minimum of ∼ 4,500-6,000 tree species. Very few species are shared among the African, American, and the Indo-Pacific regions. We provide a methodological framework for estimating species richness in trees that may help refine species richness estimates of tree-dependent taxa.

Global variability in leaf respiration in relation to climate, plant functional types and leaf traits.

Leaf dark respiration (Rdark ) is an important yet poorly quantified component of the global carbon cycle. Given this, we analyzed a new global database of Rdark and associated leaf traits. Data for 899 species were compiled from 100 sites (from the Arctic to the tropics). Several woody and nonwoody plant functional types (PFTs) were represented. Mixed-effects models were used to disentangle sources of variation in Rdark . Area-based Rdark at the prevailing average daily growth temperature (T) of each site increased only twofold from the Arctic to the tropics, despite a 20°C increase in growing T (8-28°C). By contrast, Rdark at a standard T (25°C, Rdark (25) ) was threefold higher in the Arctic than in the tropics, and twofold higher at arid than at mesic sites. Species and PFTs at cold sites exhibited higher Rdark (25) at a given photosynthetic capacity (Vcmax (25) ) or leaf nitrogen concentration ([N]) than species at warmer sites. Rdark (25) values at any given Vcmax (25) or [N] were higher in herbs than in woody plants. The results highlight variation in Rdark among species and across global gradients in T and aridity. In addition to their ecological significance, the results provide a framework for improving representation of Rdark in terrestrial biosphere models (TBMs) and associated land-surface components of Earth system models (ESMs).

A broad approach to abrupt boundaries: looking beyond the boundary at soil attributes within and across tropical vegetation types.

Most research on boundaries between vegetation types emphasizes the contrasts and similarities between conditions on either side of a boundary, but does not compare boundary to non-boundary vegetation. That is, most previous studies lack suitable controls, and may therefore overlook underlying aspects of landscape variability at a regional scale and underestimate the effects that the vegetation itself has on the soil. We compared 25 soil chemistry variables in rainforest, sclerophyll vegetation and across rainforest-sclerophyll boundaries in north-eastern Queensland, Australia. Like previous studies, we did find some contrasts in soil chemistry across vegetation boundaries. However we did not find greater variation in chemical parameters across boundary transects than in transects set in either rainforest or woodland. We also found that soil on both sides of the boundary is more similar to "rainforest soil" than to "woodland soil". Transects in wet sclerophyll forests with increasing degrees of rainforest invasion showed that as rainforest invades wet sclerophyll forest, the soil beneath wet sclerophyll forest becomes increasingly similar to rainforest soil. Our results have implications for understanding regional vegetation dynamics. Considering soil-vegetation feedbacks and the differences between soil at boundaries and in non-boundary sites may hold clues to some of the processes that occur across and between vegetation types in a wide range of ecosystems. Finally, we suggest that including appropriate controls should become standard practice for studies of vegetation boundaries and edge effects worldwide.

Predation of cassowary dispersed seeds: is the cassowary an effective disperser?

Post-dispersal predation is a potentially significant modifier of the distribution of recruiting plants and an often unmeasured determinant of the effectiveness of a frugivore's dispersal service. In the wet tropical forests of Australia and New Guinea, the cassowary provides a large volume, long distance dispersal service incorporating beneficial gut processing; however, the resultant clumped deposition might expose seeds to elevated mortality. We examined the contribution of post-dispersal seed predation to cassowary dispersal effectiveness by monitoring the fate of 11 species in southern cassowary (Casuarius casuarius johnsonii Linnaeus) droppings over a period of 1 year. Across all species, the rate of predation and removal was relatively slow. After 1 month, 70% of seeds remained intact and outwardly viable, while the number fell to 38% after 1 year. The proportion of seeds remaining intact in droppings varied considerably between species: soft-seeded and large-seeded species were more likely to escape removal and predation. Importantly, across all species, seeds in droppings were no more likely to be predated than those left undispersed under the parent tree. We speculate that seed predating and scatter-hoarding rodents are responsible for the vast majority of predation and removal from droppings and that the few seeds which undergo secondary dispersal survive to germination. Our findings reinforce the conclusion that the cassowary is an important seed disperser; however, dispersal effectiveness for particular plant species can be reduced by massive post-dispersal seed mortality.

Incorporating patterns of disperser behaviour into models of seed dispersal and its effects on estimated dispersal curves.

The processes determining where seeds fall relative to their parent plant influence the spatial structure and dynamics of plant populations and communities. For animal dispersed species the factors influencing seed shadows are poorly understood. In this paper we test the hypothesis that the daily temporal distribution of disperser behaviours, for example, foraging and movement, influences dispersal outcomes, in particular the shape and scale of dispersal curves. To do this, we describe frugivory and the dispersal curves produced by the southern cassowary, Casuarius casuarius, the only large-bodied disperser in Australia's rainforests. We found C. casuarius consumed fruits of 238 species and of all fleshy-fruit types. In feeding trials, seeds of 11 species were retained on average for 309 min (+/-256 SD). Sampling radio-telemetry data randomly, that is, assuming foraging occurs at random times during the day, gives an estimated average dispersal distance of 239 m (+/-207 SD) for seeds consumed by C. casuarius. Approximately 4% of seeds were dispersed further than 1,000 m. However, observation of wild birds indicated that foraging and movement occur more frequently early and late in the day. Seeds consumed early in the day were estimated to receive dispersal distances 1.4 times the 'random' average estimate, while afternoon consumed seeds received estimated mean dispersal distances of 0.46 times the 'random' estimate. Sampling movement data according to the daily distribution of C. casuarius foraging gives an estimated mean dispersal distance of 337 m (+/-194 SD). Most animals' behaviour has a non-random temporal distribution. Consequently such effects should be common and need to be incorporated into seed shadow estimation. Our results point to dispersal curves being an emergent property of the plant-disperser interaction rather than being a property of a plant or species.