Tropical savannas and dry forests

This is the author accepted manuscript. The final version is available from Elsevier via the DOI in this recordIn the tropics, research, conservation and public attention focus on rain forests, but this neglects that half of the global tropics have a seasonally dry climate. These regions are home to dry forests and savannas (Figures 1 and 2), and are the focus of this Primer. The attention given to rain forests is understandable. Their high species diversity, sheer stature and luxuriance thrill biologists today as much as they did the first explorers in the Age of Discovery. Although dry forest and savanna may make less of a first impression, they support a fascinating diversity of plant strategies to cope with stress and disturbance including fire, drought and herbivory. Savannas played a fundamental role in human evolution, and across Africa and India they support iconic megafauna. Pennington et al. introduce seasonally dry biomes in the tropics – savannahs and dry forests

Underground trees inhabit varied environmental extremes across the Afrotropics

This is the final version. Available on open access from Oxford University Press via the DOI in this recordAbstract Background and Aims Geoxyles, a distinctive feature of Afrotropical savannas and grasslands, survive recurrent disturbances by resprouting subshrub branches from large below-ground woody structures. Underground trees are a type of geoxyle that independently evolved within woody genera of at least 40 plant families in Africa. The environmental limits and determinants of underground tree biogeography are poorly understood, with the relative influence of frost and fire debated in particular. We aim to quantify variability in the niche of underground tree species relative to their taller, woody tree/shrub congeners. Methods Using occurrence records of four Afrotropical genera, Parinari (Chrysobalanaceae), Ozoroa (Anacardiaceae), Syzygium (Myrtaceae) and Lannea (Anacardiaceae), and environmental data of nine climate and disturbance variables, the biogeography and niche of underground trees are compared with their open and closed ecosystem congeners. Key Results Along multiple environmental gradients and in a multidimensional environmental space, underground trees inhabit significantly distinct and extreme environments relative to open and closed ecosystem congeners. Niche overlap is low among underground trees and their congeners, and also among underground trees of the four genera. Of the study taxa, Parinari underground trees inhabit hotter, drier and more seasonal environments where herbivory pressure is greatest. Ozoroa underground trees occupy relatively more fire-prone environments, while Syzygium underground trees sustain the highest frost frequency and occur in relatively wetter conditions with seasonal waterlogging. Lannea underground trees are associated with the lowest temperatures, highest precipitation, and varying exposure to disturbance. Conclusions While underground trees exhibit repeated convergent evolution, varied environments shape the ecology and biogeography of this iconic plant functional group. The multiplicity of extreme environments related to fire, frost, herbivory and waterlogging that different underground tree taxa occupy, and the distinctiveness of these environments, should be recognized in the management of African grassy ecosystems

Shade alters savanna grass layer structure and function along a gradient of canopy cover

This is the author accepted manuscript. The final version is available from Wiley via the DOI in this recordData availability statement: Data used for this study are available as supporting information.Aim: In savannas, a grass-dominated ground layer is key to ecosystem function via grass–fire feedbacks that maintain open ecosystems. With woody encroachment, tree density increases, thereby decreasing light in the ground layer and potentially altering ecosystem function. We investigated how light availability can filter individual grass species distributions and whether different functional traits are associated with response to a shade gradient in a landscape experiencing woody encroachment. Location: Savanna–forest mosaic in the Cerrado domain, southeastern Brazil. Methods: Along an encroachment gradient of increasing tree leaf area index (LAI) and shade, we determined how changing light availability alters grass diversity and ground layer structure relative to grass cover and grass functional traits (photosynthetic pathway, underground storage organs, bud protection and traits related to grass shape, size and leaf dimensions). Results: Increasing shade led to a decrease in grass cover and grass species richness, and also compositional and functional changes. We found that where tree LAI reached 1, grass cover was reduced by 50% and species richness by 30%. While C4 grass species abundances decreased with increasing shade, the opposite pattern was true for C3 grasses. There were only small differences in light preferences among C4 subtypes, with phosphoenolpyruvate carboxykinase (PCK) species tolerating slightly more shaded conditions. Persistence of some C4 species under more shaded conditions was possible, likely due to an ability to store starch reserves via underground storage organs. Conclusions: Woody encroachment changes diversity and structure of the grassy layer that is critical to the functioning of savanna ecosystems, highlighting the dependence of the diverse grass layer on open and sunny conditions. Our results suggest a threshold of tree cover close to LAI ≈ 1 as being critical to cerrado grassy layer conservation.National Science Foundation (NSF)São Paulo Research Foundation (FAPESP)National Council for Scientific and Technological DevelopmentCoordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES

Comment on “The extent of forest in dryland biomes”

This is the author accepted manuscript. The final version is available from American Association for the Advancement of Science via the DOI in this record.Bastin et al. (Reports, 12 May 2017, p. 635) infer forest as more globally extensive than previously estimated using tree cover data. However, their forest definition does not reflect ecosystem function or biotic composition. These structural and climatic definitions inflate forest estimates across the tropics and undermine conservation goals, leading to inappropriate management policies and practices in tropical grassy ecosystems

Fire History from Life-History: Determining the Fire Regime that a Plant Community Is Adapted Using Life-Histories

Wildfire is a fundamental disturbance process in many ecological communities, and is critical in maintaining the structure of some plant communities. In the past century, changes in global land use practices have led to changes in fire regimes that have radically altered the composition of many plant communities. As the severe biodiversity impacts of inappropriate fire management regimes are recognized, attempts are being made to manage fires within a more ‘natural’ regime. In this aim, the focus has typically been on determining the fire regime to which the community has adapted. Here we take a subtly different approach and focus on the probability of a patch being burnt. We hypothesize that competing sympatric taxa from different plant functional groups are able to coexist due to the stochasticity of the fire regime, which creates opportunities in both time and space that are exploited differentially by each group. We exploit this situation to find the fire probability at which three sympatric grasses, from different functional groups, are able to co-exist. We do this by parameterizing a spatio-temporal simulation model with the life-history strategies of the three species and then search for the fire frequency and scale at which they are able to coexist when in competition. The simulation gives a clear result that these species only coexist across a very narrow range of fire probabilities centred at 0.2. Conversely, fire scale was found only to be important at very large scales. Our work demonstrates the efficacy of using competing sympatric species with different regeneration niches to determine the probability of fire in any given patch. Estimating this probability allows us to construct an expected historical distribution of fire return intervals for the community; a critical resource for managing fire-driven biodiversity in the face of a growing carbon economy and ongoing climate change

How contemporary bioclimatic and human controls change global fire regimes

Anthropogenically driven declines in tropical savannah burnt area have recently received attention due to their effect on trends in global burnt area. Large-scale trends in ecosystems where vegetation has adapted to infrequent fire, especially in cooler and wetter forested areas, are less well understood. Here, small changes in fire regimes can have a substantial impact on local biogeochemistry. To investigate trends in fire across a wide range of ecosystems, we used Bayesian inference to quantify four primary controls on burnt area: fuel continuity, fuel moisture, ignitions and anthropogenic suppression. We found that fuel continuity and moisture are the dominant limiting factors of burnt area globally. Suppression is most important in cropland areas, whereas savannahs and boreal forests are most sensitive to ignitions. We quantify fire regime shifts in areas with more than one, and often counteracting, trends in these controls. Forests are of particular concern, where we show average shifts in controls of 2.3–2.6% of their potential maximum per year, mainly driven by trends in fuel continuity and moisture. This study gives added importance to understanding long-term future changes in the controls on fire and the effect of fire trends on ecosystem function

The importance of interacting climate modes on Australia’s contribution to global carbon cycle extremes

The global carbon cycle is highly sensitive to climate-driven fluctuations of precipitation, especially in the Southern Hemisphere. This was clearly manifested by a 20% increase of the global terrestrial C sink in 2011 during the strongest sustained La Niña since 1917. However, inconsistencies exist between El Niño/La Niña (ENSO) cycles and precipitation in the historical record; for example, significant ENSO-precipitation correlations were present in only 31% of the last 100 years, and often absent in wet years. To resolve these inconsistencies, we used an advanced temporal scaling method for identifying interactions amongst three key climate modes (El Niño, the Indian Ocean dipole, and the southern annular mode). When these climate modes synchronised (1999-2012), drought and extreme precipitation were observed across Australia. The interaction amongst these climate modes, more than the effect of any single mode, was associated with large fluctuations in precipitation and productivity. The long-term exposure of vegetation to this arid environment has favoured a resilient flora capable of large fluctuations in photosynthetic productivity and explains why Australia was a major contributor not only to the 2011 global C sink anomaly but also to global reductions in photosynthetic C uptake during the previous decade of drought

Determinants of woody encroachment and cover in African savannas

Savanna ecosystems are an integral part of the African landscape and sustain the livelihoods of millions of people. Woody encroachment in savannas is a widespread phenomenon but its causes are widely debated. We review the extensive literature on woody encroachment to help improve understanding of the possible causes and to highlight where and how future scientific efforts to fully understand these causes should be focused. Rainfall is the most important determinant of maximum woody cover across Africa, but fire and herbivory interact to reduce woody cover below the maximum at many locations. We postulate that woody encroachment is most likely driven by CO2 enrichment and propose a two-system conceptual framework, whereby mechanisms of woody encroachment differ depending on whether the savanna is a wet or dry system. In dry savannas, the increased water-use efficiency in plants relaxes precipitation-driven constraints and increases woody growth. In wet savannas, the increase of carbon allocation to tree roots results in faster recovery rates after disturbance and a greater likelihood of reaching sexual maturity. Our proposed framework can be tested using a mixture of experimental and earth observational techniques. At a local level, changes in precipitation, burning regimes or herbivory could be driving woody encroachment, but are unlikely to be the explanation of this continent-wide phenomenon

Stable Carbon and Nitrogen Isotopes in a Peat Profile Are Influenced by Early Stage Diagenesis and Changes in Atmospheric CO2 and N Deposition

In this study, we test whether the δ13C and δ15N in a peat profile are, respectively, linked to the recent dilution of atmospheric δ13CO2 caused by increased fossil fuel combustion and changes in atmospheric δ15N deposition. We analysed bulk peat and Sphagnum fuscum branch C and N concentrations and bulk peat, S. fuscum branch and Andromeda polifolia leaf δ13C and δ15N from a 30-cm hummock-like peat profile from an Aapa mire in northern Finland. Statistically significant correlations were found between the dilution of atmospheric δ13CO2 and bulk peat δ13C, as well as between historically increasing wet N deposition and bulk peat δ15N. However, these correlations may be affected by early stage kinetic fractionation during decomposition and possibly other processes. We conclude that bulk peat stable carbon and nitrogen isotope ratios may reflect the dilution of atmospheric δ13CO2 and the changes in δ15N deposition, but probably also reflect the effects of early stage kinetic fractionation during diagenesis. This needs to be taken into account when interpreting palaeodata. There is a need for further studies of δ15N profiles in sufficiently old dated cores from sites with different rates of decomposition: These would facilitate more reliable separation of depositional δ15N from patterns caused by other processes