Biome States Research Articles

Long-term stability of temperate Australian wet forest-moorland mosaics despite recurrent fires associated with late Holocene climate change

ContextThe primary factors(s) responsible for the maintenance of Alternative biome states (ABS) in world forest biomes remains unclear and debatable, partly due to insufficient long-term ecological data from suitable ecosystem sites. The occurrence of moorland in southern and western Tasmanian wet temperate forest presents a suitable setting to test for ABS and understand the main stabilizing factors of ABS.ObjectivesWe use a palaeoecological approach to test for ABS and identify the degree of vegetation change and the effect of climate change and fire occurrence associated with ABS in southern Tasmania.MethodsSediment sequence from sink-hole lake in a forest and nearby pond in southern Tasmania were analysed for pollen and charcoal to reconstruct histories of forest, moorland and fire in the Tasmanian Wilderness World Heritage Area (TWWHA). Reconstructions were compared to palaeoclimate record.ResultsMoorland and forest in southern Tasmania have occupied the same habitat for at least the last 2600 years, and neither past climate change nor fire occurrence affected the stability of the vegetation mosaic. We suspect that localized environmental settings, such as topography and edaphic conditions are the primary stabilizing factors of the forest-moorland mosaics.ConclusionsThe observed stable vegetation mosaics in our study is contrary to the dominant ecological paradigm of landscape dynamics currently used to manage the TWWHA, and there is a need to refine the ecological basis of fire management in the area. Similar targeted palaeoecological studies are needed to fully understand the underlying factors responsible for the persistence of treeless vegetation in world forest biomes.

Caracterización de ecotonos entre biomas de bosque y biomas abiertos en un área protegida en Uruguay

Alternative Biome States theory (ABS) helps to explain the presence of open biomes (grasslands, shrublands) in environments sufficiently warm and humid for the development of forest biomes. In these environments it is possible to find areas dominated by open biomes that, according to this theory, respond mainly to fire and herbivory. The aims of this work are to identify whether there are different types of forest biome-open biome ecotones based on woody composition, describe them and explore their relationship with livestock management and environmental variables, using a protected area in Uruguay as a case study. We sampled woody vegetation and other environmental variables in transects (delimited in 200m) in 10 sites across four livestock management systems. We identified two types of forest biome-open biome ecotones, one dominated by shrubland and the other dominated by grassland. Our results agree with the ABS theory and suggest the mechanisms determining the woody composition and the structure of the ecotone vegetation are cattle and use of fire and/or rotary cutter. The changes in vegetation structure and woody richness were partly abrupt, which is consistent with what is proposed by the ABS theory. However, the composition also interacted with site factors such as rockiness and distance to the watercourse, suggesting that site-level environmental variables are acting synergistically with vegetation removal to determine vegetation composition. We found that these ecotones are sensitive to management and environmental variables; so, it is necessary to incorporate this landscape and site heterogeneity to articulate productive activity with nature conservation.

Alternative biome states challenge the modelling of species' niche shifts under climate change

Abstract It is common to characterise the species niche using climate and global species distribution maps. This is then used to predict changes in distribution under a warming climate. This approach assumes that climate is a major driver of species distribution and that each species responds individually (sensu Gleason) to climate. However, in many world landscapes, for a given climate, strikingly different vegetation types co‐occur: forests and non‐forests. These two alternative biome states are maintained by different feedback processes and have radically different species with contrasting shade and disturbance tolerance traits. We propose that to improve predictions of species distribution changes under a novel climate, we need to consider the presence or absence of forest shade, as species are likely to respond individually only within their forest or non‐forest biome, and not across biomes. Synthesis. By considering shade as a biotic filter in niche modelling, we are not only improving our predictive capacity, but we are also reconciling the two views of communities: both the individualistic (within biome) and the organismal (across biomes) views of the community concept become relevant and complementary.

Alternative biome states of African terrestrial vegetation and the potential drivers: A continental-scale study

The alternative stable state (ASS) theory provides a plausible framework to explain the spatial distribution of biomes and their dynamics. Existing studies to test the alternative biome states (ABSs) mainly focused on tree-dominated biomes. It is still uncertain whether ABSs are present in a wide range of terrestrial biomes. This study was to examine the ABSs in the terrestrial vegetated areas of Africa and the maintaining factors. The potential landscapes were reconstructed separately for forest, grassland and shrubland using the MODIS Vegetation Continuous Field data along the gradients of temperature, precipitation and aridity index (AI, the ratio of precipitation to potential evapotranspiration). The differences of soil organic carbon density (SOCD), fire count and grazing intensity were compared to test the feedback hypothesis to maintain the ABSs. The results showed that AI performed well in detecting the ABSs at the continental scale of Africa. Forest (at the wetter end) and shrubland (at the drier end) were well separated along the AI axis. Forest had three stable states (i.e. closed forest, woody savanna, and savanna) and shrubland had two stable states (i.e. closed shrubland and open shrubland). Grassland had two stable states (i.e. dense grassland and sparse grassland) distributing in a large AI range. The stable states that shared a specific AI range were regarded as the ABSs. Climate aridity greatly determined the distribution of the ABSs but the positive feedbacks between vegetation and SOCD, fire count, and livestock density played potential roles in driving the shifts between the ABSs. Our study indicated that the ABSs commonly existed in varied biomes (both tree-dominated and non-tree-dominated) in the African continent, which provided an enlarged picture of the ABSs of the terrestrial biomes. The findings contribute to a deeper understanding of large scale vegetation patterns and their dynamics and facilitate to macro management of the terrestrial ecosystems in facing the possible regime shifts of the biomes.

Floristic evidence for alternative biome states in tropical Africa.

The idea that tropical forest and savanna are alternative states is crucial to how we manage these biomes and predict their future under global change. Large-scale empirical evidence for alternative stable states is limited, however, and comes mostly from the multimodal distribution of structural aspects of vegetation. These approaches have been criticized, as structure alone cannot separate out wetter savannas from drier forests for example, and there are also technical challenges to mapping vegetation structure in unbiased ways. Here, we develop an alternative approach to delimit the climatic envelope of the two biomes in Africa using tree species lists gathered for a large number of forest and savanna sites distributed across the continent. Our analyses confirm extensive climatic overlap of forest and savanna, supporting the alternative stable states hypothesis for Africa, and this result is corroborated by paleoecological evidence. Further, we find the two biomes to have highly divergent tree species compositions and to represent alternative compositional states. This allowed us to classify tree species as forest vs. savanna specialists, with some generalist species that span both biomes. In conjunction with georeferenced herbarium records, we mapped the forest and savanna distributions across Africa and quantified their environmental limits, which are primarily related to precipitation and seasonality, with a secondary contribution of fire. These results are important for the ongoing efforts to restore African ecosystems, which depend on accurate biome maps to set appropriate targets for the restored states but also provide empirical evidence for broad-scale bistability.

Mohammed Armani.

Mohammed Armani.

Joint Phylogenetic Estimation of Geographic Movements and Biome Shifts during the Global Diversification of Viburnum.

Phylogeny, molecular sequences, fossils, biogeography, and biome occupancy are all lines of evidence that reflect the singular evolutionary history of a clade, but they are most often studied separately, by first inferring a fossil-dated molecular phylogeny, then mapping on ancestral ranges and biomes inferred from extant species. Here we jointly model the evolution of biogeographic ranges, biome affinities, and molecular sequences, while incorporating fossils to estimate a dated phylogeny for all of the 163 extant species of the woody plant clade Viburnum (Adoxaceae) that we currently recognize in our ongoing worldwide monographic treatment of the group. Our analyses indicate that while the major Viburnum lineages evolved in the Eocene, the majority of extant species originated since the Miocene. Viburnum radiated first in Asia, in warm, broad-leaved evergreen (lucidophyllous) forests. Within Asia, we infer several early shifts into more tropical forests, and multiple shifts into forests that experience prolonged freezing. From Asia, we infer two early movements into the New World. These two lineages probably first occupied warm temperate forests and adapted later to spreading cold climates. One of these lineages (Porphyrotinus) occupied cloud forests and moved south through the mountains of the Neotropics. Several other movements into North America took place more recently, facilitated by prior adaptations to freezing in the Old World. We also infer four disjunctions between Asia and Europe: the Tinus lineage is the oldest and probably occupied warm forests when it spread, whereas the other three were more recent and in cold-adapted lineages. These results variously contradict published accounts, especially the view that Viburnum radiated initially in cold forests and, accordingly, maintained vessel elements with scalariform perforations. We explored how the location and biome assignments of fossils affected our inference of ancestral areas and biome states. Our results are sensitive to, but not entirely dependent upon, the inclusion of fossil biome data. It will be critical to take advantage of all available lines of evidence to decipher events in the distant past. The joint estimation approach developed here provides cautious hope even when fossil evidence is limited. [Biogeography; biome; combined evidence; fossil pollen; phylogeny; Viburnum.].

African biomes are most sensitive to changes in CO<sub>2</sub> under recent and near-future CO<sub>2</sub> conditions

Abstract. Current rates of climate and atmospheric change are likely higher than during the last millions of years. Even higher rates of change are projected in CMIP5 climate model ensemble runs for some Representative Concentration Pathway (RCP) scenarios. The speed of ecological processes such as leaf physiology, demography or migration can differ from the speed of changes in environmental conditions. Such mismatches imply lags between the actual vegetation state and the vegetation state expected under prevailing environmental conditions. Here, we used a dynamic vegetation model, the adaptive Dynamic Global Vegetation Model (aDGVM), to study lags between actual and expected vegetation in Africa under a changing atmospheric CO2 mixing ratio. We hypothesized that lag size increases with a more rapidly changing CO2 mixing ratio as opposed to slower changes in CO2 and that disturbance by fire further increases lag size. Our model results confirm these hypotheses, revealing lags between vegetation state and environmental conditions and enhanced lags in fire-driven systems. Biome states, carbon stored in vegetation and tree cover in Africa are most sensitive to changes in CO2 under recent and near-future levels. When averaged across all biomes and simulations with and without fire, times to reach an equilibrium vegetation state increase from approximately 242 years for 200 ppm to 898 years for 1000 ppm. These results have important implications for vegetation modellers and for policy making. Lag effects imply that vegetation will undergo substantial changes in distribution patterns, structure and carbon sequestration even if emissions of fossils fuels and other greenhouse gasses are reduced and the climate system stabilizes. We conclude that modelers need to account for lag effects in models and in data used for model testing. Policy makers need to consider lagged responses and committed changes in the biosphere when developing adaptation and mitigation strategies.

Alternative Biome States in Terrestrial Ecosystems.

There is growing interest in the application of alternative stable state (ASS) theory to explain major vegetation patterns of the world. Here, we introduce the theory as applied to the puzzle of nonforested (open) biomes growing in climates that are warm and wet enough to support forests (alternative biome states, ABSs). Long thought to be the product of deforestation, diverse lines of evidence indicate that many open ecosystems are ancient. They have also been characterized as 'early successional' even where they persist for millennia. ABS is an alternative framework to that of climate determinism and succession for exploring forest/nonforest mosaics. This framework explains not only tropical forest-savanna landscapes, but also other landscape mosaics across the globe.

Estimating 20-year land-use change and derived CO2 emissions associated with crops, pasture and forestry in Brazil and each of its 27 states.

Land-use change (LUC) in Brazil has important implications on global climate change, ecosystem services and biodiversity, and agricultural expansion plays a critical role in this process. Concerns over these issues have led to the need for estimating the magnitude and impacts associated with that, which are increasingly reported in the environmental assessment of products. Currently, there is an extensive debate on which methods are more appropriate for estimating LUC and related emissions and regionalized estimates are lacking for Brazil, which is a world leader in agricultural production (e.g. food, fibres and bioenergy). We developed a method for estimating scenarios of past 20-year LUC and derived CO2 emission rates associated with 64 crops, pasture and forestry in Brazil as whole and in each of its 27 states, based on time-series statistics and in accordance with most used carbon-footprinting standards. The scenarios adopted provide a range between minimum and maximum rates of CO2 emissions from LUC according to different possibilities of land-use transitions, which can have large impacts in the results. Specificities of Brazil, like multiple cropping and highly heterogeneous carbon stocks, are also addressed. The highest CO2 emission rates are observed in the Amazon biome states and crops with the highest rates are those that have undergone expansion in this region. Some states and crops showing large agricultural areas have low emissions associated, especially in southern and eastern Brazil. Native carbon stocks and time of agricultural expansion are the most decisive factors to the patterns of emissions. Some implications on LUC estimation methods and standards and on agri-environmental policies are discussed.

Defining functional biomes and monitoring their change globally.

Biomes are important constructs for organizing understanding of how the worlds' major terrestrial ecosystems differ from one another and for monitoring change in these ecosystems. Yet existing biome classification schemes have been criticized for being overly subjective and for explicitly or implicitly invoking climate. We propose a new biome map and classification scheme that uses information on (i) an index of vegetation productivity, (ii) whether the minimum of vegetation activity is in the driest or coldest part of the year, and (iii) vegetation height. Although biomes produced on the basis of this classification show a strong spatial coherence, they show little congruence with existing biome classification schemes. Our biome map provides an alternative classification scheme for comparing the biogeochemical rates of terrestrial ecosystems. We use this new biome classification scheme to analyse the patterns of biome change observed over recent decades. Overall, 13% to 14% of analysed pixels shifted in biome state over the 30-year study period. A wide range of biome transitions were observed. For example, biomes with tall vegetation and minimum vegetation activity in the cold season shifted to higher productivity biome states. Biomes with short vegetation and low seasonality shifted to seasonally moisture-limited biome states. Our findings and method provide a new source of data for rigorously monitoring global vegetation change, analysing drivers of vegetation change and for benchmarking models of terrestrial ecosystem function.

Revising the biome concept for understanding and predicting global change impacts

AbstractBiomes are globally distributed, structurally and functionally similar vegetation units defined without reference to plant species composition. The boundaries between biomes are presumed to correspond with species turnover and changes in biogeochemical cycling. Determining the controls of biome distributions is thus critical for anticipating global change impacts. Historically, climate and soils have been understood to adequately explain the global distribution of biomes. Convergent evolution and environmental filtering are assumed to be pervasive, ultimately resulting in deterministic patterns of vegetation structure and function in relation to prevailing environmental conditions. Recent studies have highlighted significant problems with this view of biomes. Firstly, systematic structural and functional divergence within biomes has been identified when comparing environmentally similar, yet floristically distinct regions. Secondly, climatic determinism is being further undermined by evidence suggesting multiple stable biome states are possible for some combinations of climatic drivers. We argue that biomes remain useful and necessary constructs for organizing our knowledge of how ecosystems function and for predicting how they might respond to change. However, biome concepts should acknowledge the limits to predictability from environmental conditions alone and the influence of historical processes on modern vegetation patterns. We discuss how direct mapping of plant structure and function, the incorporation of insights into biome evolution and a new generation of vegetation models will lead to improvements in the concept of what biomes are, where they occur, and efforts to predict their distribution.

Disturbance maintains alternative biome states.

Understanding the mechanisms controlling the distribution of biomes remains a challenge. Although tropical biome distribution has traditionally been explained by climate and soil, contrasting vegetation types often occur as mosaics with sharp boundaries under very similar environmental conditions. While evidence suggests that these biomes are alternative states, empirical broad-scale support to this hypothesis is still lacking. Using community-level field data and a novel resource-niche overlap approach, we show that, for a wide range of environmental conditions, fire feedbacks maintain savannas and forests as alternative biome states in both the Neotropics and the Afrotropics. In addition, wooded grasslands and savannas occurred as alternative grassy states in the Afrotropics, depending on the relative importance of fire and herbivory feedbacks. These results are consistent with landscape scale evidence and suggest that disturbance is a general factor driving and maintaining alternative biome states and vegetation mosaics in the tropics.

Soil nutrients in an African forest/savanna mosaic: Drivers or driven?

This study set out to test three hypotheses explaining the existence of closed canopied versus open ecosystems in the same landscape and on the same parent rocks. Our data lent no support to the first hypothesis that forests are ancient and grasslands are anthropogenically derived through deforestation. Carbon isotope data suggests the opposite; both open and closed vegetation had C4 isotope signals in deeper soils, indicating that grasslands are the ancient system. The second hypothesis is that closed forests require nutrient rich soils and are replaced by open vegetation on nutrient poor soils. Despite open vegetation frequently havinglower nutrient concentrations than closed vegetation in our study area, nutrient stocks to 1m far exceeded threshold requirements to build a forest at all our sites. We suggest that vegetation patterns in our study area are a result of a third hypothesis, that forests and savannas are alternative stable states each subject to different ecological processes with positive feedbacks maintaining each state. Fire in open grassy vegetation is a key disturbance maintaining the boundaries between the two states. Frequent fires may deplete nutrients in the surface layers of the soil, thereby slowing tree growth and further maintaining the open state of the vegetation. However, our data on the vertical distribution of nutrients show strikingly similar nutrient patterns in the alternative biome states, with high nutrient concentrations in the open sites, suggesting that nutrient limitation is not an important feedback helping to maintain open ecosystems in our study area. Our results suggest that, rather than being drivers of vegetation patterns in our study area, soil nutrients are responding to dynamic vegetation changes caused by other factors.

Increasing atmospheric CO2 overrides the historical legacy of multiple stable biome states in Africa

The dominant vegetation over much of the global land surface is not predetermined by contemporary climate, but also influenced by past environmental conditions. This confounds attempts to predict current and future biome distributions, because even a perfect model would project multiple possible biomes without knowledge of the historical vegetation state. Here we compare the distribution of tree- and grass-dominated biomes across Africa simulated using a dynamic global vegetation model (DGVM). We explicitly evaluate where and under what conditions multiple stable biome states are possible for current and projected future climates. Our simulation results show that multiple stable biomes states are possible for vast areas of tropical and subtropical Africa under current conditions. Widespread loss of the potential for multiple stable biomes states is projected in the 21st Century, driven by increasing atmospheric CO2 . Many sites where currently both tree-dominated and grass-dominated biomes are possible become deterministically tree-dominated. Regions with multiple stable biome states are widespread and require consideration when attempting to predict future vegetation changes. Testing for behaviour characteristic of systems with multiple stable equilibria, such as hysteresis and dependence on historical conditions, and the resulting uncertainty in simulated vegetation, will lead to improved projections of global change impacts.

The Global Extent and Determinants of Savanna and Forest as Alternative Biome States

Theoretically, fire-tree cover feedbacks can maintain savanna and forest as alternative stable states. However, the global extent of fire-driven discontinuities in tree cover is unknown, especially accounting for seasonality and soils. We use tree cover, climate, fire, and soils data sets to show that tree cover is globally discontinuous. Climate influences tree cover globally but, at intermediate rainfall (1000 to 2500 millimeters) with mild seasonality (less than 7 months), tree cover is bimodal, and only fire differentiates between savanna and forest. These may be alternative states over large areas, including parts of Amazonia and the Congo. Changes in biome distributions, whether at the cost of savanna (due to fragmentation) or forest (due to climate), will be neither smooth nor easily reversible.