Posts Tagged ‘alternative states’

Microbes, herbivores, and wildfires

May 4th, 2020 No comments

Plants are the largest biomass component of most terrestrial ecosystems, and litter decomposition is considered the dominant process by which nutrients return to plants. In a recent paper [1] we show that in terrestrial ecosystems, there are three major pathways by which plant biomass is degraded into forms that release nutrients again available to plants: microbial decomposition; vertebrate herbivory; and wildfires. These processes act at different spatial and temporal scales, have different niches, and generates different ecological and evolutionary feedbacks. The three processes can occur in a given ecosystem (competing for the same resource, biomass), but the relative importance of each varies with the micro- and macro-environmental conditions (see Figure below).

Wildfires and herbivory are two powerful biomass consumers; they generate feedback processes that maintain vegetation at states of lower biomass than would be expected from the physical environmental conditions (alternative vegetation states [2]). In addition, wildfires and herbivory also select for light-loving species with a set of adaptive traits to persist under these consumers [3,4]. That is, both herbivory and fire can influence the mix and attributes of plant species, while the mix and attributes of plants also influence the fire and grazing regimes. These ecological and evolutionary feedbacks make fire and herbivory distinct from other abiotic disturbances such as cyclones, landslides, avalanches, volcanoes, or floods, where plants may respond – but the disturbance will not change in response to these plant responses. That is, wildfires, herbivory, and microbial decomposition can be viewed as biotic processes that structure our ecosystems and the biosphere, at different temporal and spatial scales.

This holistic view in which microbes, herbivores, and wildfires play a joint role in the functioning of ecosystems contributes to a better understanding of the diversity of mechanisms regulating the biosphere.

Figure: Plant biomass and vegetation structure in terrestrial ecosystems are determined by three feedback processes: vertebrate herbivory (H), microbial decomposition (D), and wildfires (W). These three processes also interact with each other (e.g., competition for biomass; but positive interactions also exist). Relative importance of each of the three ecosystem pathways varies in the environmental space (niche), here defined by the water availability and soil fertility. Illustration by Dharmaberen Studio. From [1].


[1] Pausas J.G. & Bond W.J. 2020. On the three major recycling pathways in terrestrial ecosystems. Trends Ecol. & Evol. [doi | pdf]

[2] Pausas J.G. & Bond W.J. 2020. Alternative biome states in terrestrial ecosystems. Trends Plant Sci. 25: 250-263. [doi | sciencedirect | cell | pdf]

[3] Keeley J.E., Bond W.J., Bradstock R.A., Pausas J.G. & Rundel P.W. 2012. Fire in Mediterranean Ecosystems: Ecology, Evolution and Management. Cambridge University Press. [the book]  

[4] Bond, W. J. 2019. Open Ecosystems: Ecology and Evolution Beyond the Forest Edge. Oxford University Press.


Alternative Biome States

January 8th, 2020 No comments

There is growing interest in the application of alternative stable state (ASS) theory to explain major vegetation patterns in tropical ecosystems [1] and beyond [2]. In a recent paper [3] we introduced the theory as applied to the puzzle of non-forested (open) biomes growing in climates that are warm and wet enough to support forests (alternative biome states, ABSs; Fig. 1). 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 (Table 1 below) for exploring forest/nonforest mosaics. Within climatic and edaphic constraints, consumers (fire and herbivores) can produce vastly different ecosystems from the climate potential and have done so for millions of years [4]. This framework explains not only tropical forest–savanna landscapes, but also other landscape mosaics across the globe (Fig. 2).

Fig. 1. Generalized feedback processes in fire-prone landscapes where open and closed biomes (e.g., a grassland and forest) are alternative stable states maintained by stabilizing feedbacks, while perturbations generate abrupt transitions among states (destabilizing factors). From: [3].

Fig. 2. Examples of multibiome landscape mosaics where closed forests alternate with open biomes (grasslands) that are maintained by mammal herbivory and fire. From: [3].

Table 1. Comparison of the three main dynamic processes assembling disturbance-prone communities and landscapes: classical (facilitation) succession, autosuccession, and ABS. From: [3].


[1] Dantas V.L., Hirota M., Oliveira R.S., Pausas J.G. 2016. Disturbance maintains alternative biome states. Ecol. Lett. 19: 12-19. [doi | wiley | pdf | suppl.]

[2] Pausas, J.G. 2015. Alternative fire-driven vegetation states. J. Veget. Sci. 26:4-6. [doi | pdf | suppl.]

[3] Pausas J.G. & Bond W.J. 2020. Alternative biome states in terrestrial ecosystems. Trends Plant Sci. [doi | sciencedirect| pdf]

[4] Pausas J.G. & Bond W.J. 2019. Humboldt and the reinvention of nature. J. Ecol. 107: 1031-1037. [doi | jecol blog | jgp blog | pdf]  

Scale mismatch in ecology

January 2nd, 2017 No comments

A recent paper suggested that fire-vegetation feedback processes may be unnecessary to explain tree cover patterns in tropical ecosystems and that climate-fire determinism is an alternative possibility [1]. This conclusion was based on the fact that it is possible to reproduce observed broad scale patterns in tropical regions (e.g., a trimodal frequency distribution of tree cover) using a simple model that does not explicitly incorporate fire-vegetation feedback processes. We argue that this reasoning is misleading because these two mechanisms (feedbacks vs fire-climate control) operate at different spatial and temporal scales [2]. It is not possible to evaluate the role of a process acting at fine scales (e.g., fire-vegetation feedbacks) using a model designed for reproducing regional-scale pattern; i.e., there is a mismatch between the scale of the question and the scale of the approach for addressing the question. While the distribution of forest and savannas are partially determined by climate, the most parsimonious explanation for their environmental overlaps (as alternative states) is the existence of feedback processes [3,4], as has been shown in many ecosystems, not only tropical ones [4]. Climate is unlikely to be an alternative to feedback processes; rather, climate and fire-vegetation feedbacks are complementary processes acting at different spatial and temporal scales [2].
Figure: Fire activity (based on remotely sensed data) for savannas and forests located in the range of environmental conditions where both occurs, for Africa and South America (Afrotropics and Neotropics, respectively). From [2,3].

[1] Good, P., Harper, A., Meesters, A., Robertson, E. & Betts, R. (2016) Are strong fire–vegetation feedbacks needed to explain the spatial distribution of tropical tree cover? Global Ecol. and Biogeogr. 25, 16-25.

[2] Pausas J.G. & Dantas V.L. 2017. Scale matters: Fire-vegetation feedbacks are needed to explain tropical tree cover at the local sacle. Global Ecol. and Biogeogr. [doiwiley | pdf]

[3] Dantas V.L., Hirota M., Oliveira R.S., Pausas J.G. 2016. Disturbance maintains alternative biome states. Ecology Letters 19: 12-19. [doi | wiley | pdf | suppl | blog]

[4] Pausas, J.G. 2015. Alternative fire-driven vegetation states. J. Veget. Sci. 26:4-6. [doi | pdf | suppl.] | blog]


Disturbance maintains alternative biome states

November 9th, 2015 No comments

It is becoming more and more evident that climate alone does not explain spatial and temporal patterns of the world vegetation, and that disturbance regimes explain an important part of the variability in vegetation and biome composition and distribution [1]. This has been suggested specially in tropical ecosystems where alternative vegetation states (e.g., forests and savannas) are possible for a given climatic conditions [2]. For instance, in dry years, surface fires may enter in forests and kill fire-sensitive trees and select for fire-resistant woody species with open crown architectures that generates well lit communities with a flammable grassy understory. Forest trees and savannas trees have a marked difference in bark thickness (thinner in the former) and thus a contrasted sensitivity to surface fires [3]. Thus, a switch to a forest state from a savanna depends on a sufficiently long fire interval or high resource availability allowing the outcompetition of shade-intolerant savanna trees and grasses (i.e. the inhibition of fires) by means of a closed canopy of forest trees. Similarly, herbivory can also exert a control on woody biomass and favour herbivory-resistant shrubs and grasses. However, empirical (field-based) evidence for alternative sates were very limited. In a recent paper [4] we used field data to show that, for a wide range of environmental conditions (in South America and Africa), 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 states in the Afrotropics, depending on the relative importance of fire and herbivory feedbacks. That is, we found evidence for a disturbance-driven bistability in the Neotropics and a disturbance-driven tristability in Afrotropics (figure below).


Fig. Top: Frequency distribution of basal area in afrotropical (tristability) and neotropical (bistability) ecosystems. Bottom: The discontinuous pattern of basal area along the resources gradient for both afrotropical and neotropical ecosystems (red: wooded grasslands; orange: savannas; green: forests). Note that there are regions of the gradient where two alternative vegetation types are possible; they are maintained by different disturbance regime (see [4]).

[1] Pausas, J.G. 2015. Alternative fire-driven vegetation states. J. Veget. Sci. 26: 4-6. [doi | pdf | suppl.]

[2] Dantas V., Batalha MA & Pausas JG. 2013. Fire drives functional thresholds on the savanna-forest transition. Ecology 94:2454-2463. [doi | pdf | appendix]

[3] Pausas, J.G. 2015. Bark thickness and fire regime. Funct. Ecol. 29: 317-327. [doi | pdf | suppl.]

[4] Dantas V.L., Hirota M., Oliveira R.S., Pausas J.G. 2016. Disturbance maintains alternative biome states. Ecology Letters 19:12-19 [doi | wiley | pdf |supp.– New!

[5] Update (a new relevant paper): Pausas J.G. & Dantas V.L. 2017. Scale matters: Fire-vegetation feedbacks are needed to explain tropical tree cover at the local sacle. Global Ecol. Biogeogr. 26: 395–399. [doi | pdf | post ]

Alternative fire-driven vegetation states

November 1st, 2014 No comments

One of the clearest pieces of evidence for the role of fire in shaping vegetation is the occurrence of alternative vegetation types maintained by different fire regimes in a given climate. The different flammability of alternative communities generates different fire feedback processes that maintain contrasted vegetation types with clear boundaries in a given environment; and fire exclusion blurs this structure. This has been well documented in tropical landscapes (e.g., [1]) that are often mosaics of two alternative stable states – savannas and forests – with distinct structures and functions and sharp boundaries. Currently, there is an increasing evidence that alternative fire-driven vegetation states do occur in other environments, including temperate forests ([2, 3] and figure below). That is, the existence of alternative fire-driven vegetation states may be more frequent than previously thought, although human activities may favour one of the states and mask the original bistability.


Figure: Factors determining the transition between two alternative vegetation states (fire sensitive forest and fire resilient shrubland) in a temperate landscape in Patagonia. Human factors (global warming, increased ignitions, and livestock grazing) favour transition to shrublands. From [2].

[1] Dantas V., Batalha MA & Pausas JG. 2013. Fire drives functional thresholds on the savanna-forest transition. Ecology 94:2454-2463.  [doi | pdf | appendix]

[2] Pausas, J.G. 2015. Alternative fire-driven vegetation states. Journal of Vegetation Science 26: 4-6 [doi | pdf | suppl.]

[3] Paritsis J., Veblen T.T. & Holz A. 2014. Positive fire feedbacks contribute to shifts from Nothofagus pumilio forests to fire-prone shrublands in Patagonia. J. Veget. Sci., 26.