j.g. pausas' blog

This video provides a powerful message from grasses to the world, written and narrated by William Bond: The Untold Story of Grasses.

Grasses created an open sunlit world, rich in plants and animals.

When you support a tree planting project … Stop. Think. Are the trees restoring a forest? Or are they destroying an ancient grassland? What might be lost?

Further readings

• Bond WJ. 2019. Open Ecosystems: Ecology and Evolution Beyond the Forest Edge. Oxford University Press.
• Pausas JG. & Bond WJ. 2019. Humboldt and the reinvention of nature. J. Ecol. 107: 1031-1037. [doi | jecol blog | pdf]
• Pausas JG. & Bond WJ. 2020. Alternative biome states in terrestrial ecosystems. Trends Plant Sci. 25: 250-263. [doi | sciencedirect | cell | pdf]
• Afforestation is not a solution to mitigate CO₂ emissions [link]
• Mythbusting forests, by WJ Bond [link]
• https://openecosystems.co.za/

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.

References

[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.

The role of large herbivores in explaining broad-scale ecological pattern has probably been underestimated [1]. For instance, they are important in maintaining many landscapes and biomes across the world [2]. In a recent paper we show that the different history of megafauna abundance and extinctions in different continents has shaped the dominance of many plant traits [3]. Tropical Africa (paelotropics) and tropical South America (neotropics) are a clear example of contrasting megafauna history under similar environmental conditions. By comparing plant traits of woody species in different biomes (savannas and forests) and for the two different continents, we found that continent explain better the differences in plant traits than biome, climate, or soil, and that the differences between continents are consistent with the higher impact of large vertebrates in Africa than in South America. For instance, plants in African savannas tend to be more thorny and to have higher wood density, i.e., traits related to defense against megaherbivores. In contrast, South American savannas (Cerrado) harbor more species with underground bud banks (geoxyles [4]), and thicker protective barks, i.e., traits related to protect from wildfires [4,5].

Megafauna was certainly present in South America before the Holocene overkilling by humans, however, it is unlikely they live in the brazilian savannas (cerrado); their weak and vulnerable stems (low height growth, low wood density, and lack of spines), are unlikely to have evolved in regions with abundant browsers. We hypotheses that megafauna in South America was distributed in: 1) an open version of the current seasonally dry tropical forests (SDTF, e.g., Chaco) as the proportion of thorny species is similar to African savannas (and much higher than the cerrado); and 2) the subtropical grasslands, as they currently need to be maintained by humans due to the missing megaherbivores (landscape anachronism [1]).

Overall our results suggest that variation in plant traits in the tropics is unlikely to be fully understood without considering historical factors, especially the direct and indirect impacts of megafauna. Looking at plants and thinking on their megafauna history may provide novel insights for understanding vegetation patterns across the globe [1].

References

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

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

[3] Dantas V & Pausas JG. 2020. Megafauna biogeography explains plant functional trait variability in the tropics. Glob. Ecol. & Biogeogr. [doi | pdf | data:dryad]

[4] Pausas JG, Lamont BB, Paula S, Appezzato-da-Glória B & Fidelis A. 2018. Unearthing belowground bud banks in fire-prone ecosystems. New Phytol. 217: 1435–1448. [doi | pdf | suppl. | BBB database]

[5] Pausas JG. 2017. Bark thickness and fire regime: another twist. New Phytol. 213: 13-15. [doi | wiley | pdf]  &  Pausas, J.G. 2015. Bark thickness and fire regime. Funct. Ecol. 29:317-327. [doi | pdf | suppl.]

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).

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References

[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]  

Behind the paper “Humboldt and the reinvention of nature” [1]. Text simultaneously published here and in JEcologyBlog.com

It all started when I was reading an excellent book by Andrea Wulf entitled The Invention of Nature: Alexander von Humboldt’s New World. The book provides many details about Humboldt’s fascinating life and the wide-ranging influence he had on science and society. When reading the book, you can easily understand the unquestionable role Humboldt played in the history of ecology and biogeography. One of his many contributions to science was to set the basis for explaining how environmental factors affect species distribution; for example, he demonstrated that vegetation systematically varies across the world with climate and showed the ecological similarities between altitude and latitude.
 

Fig. 1. Vegetation of Chimborazo (Ecuador) by Humboldt and Bonpland (1807).

 
A question that came to my mind was not covered by the book; namely, to what extent does Humboldt’s view bias our vision of nature? This is relevant because many classical naturalists and ecologists, such as Henry David Thoreau, Charles Darwin, George Perkins Marsh, John Muir, Rachel Carlson, Frederic E. Clements, and Henry A. Gleason, were all inspired by Humboldt. By spreading a vision, they shaped what is today mainstream ecology and the environmental movement. The current emphasis on the role of climate and soil in many ecological and evolutionary studies, the emphasis on forests as the potential and most important vegetation, and the difficulties many researchers have accepting the ecological and evolutionary role of disturbances at broad scales, suggest that we are still largely viewing nature through the eyes of Humboldt.

After reading the book, I was lucky enough to be in London for a conference and met William Bond in a pub just next to Kew Gardens. We stayed hours talking about many things, mainly fire, grazing, alternative stable states, and all the wonders of ecosystems maintained by disturbances. Our conversation jumped from one country to the other, from one biome to the other, and from one continent to another. And when talking about the overwhelming role that many researchers attribute to the environment when explaining broad temporal and spatial vegetation patterns [2], we glimpsed the long shadow of Humboldt. That conversation in a quiet London pub was the seed of this paper (and of another to come). Talking with William is always enjoyable because of his enthusiasm, experience, and creativity.

Now that we are approaching the 250th anniversary of Humboldt’s birth, it is instructive to evaluate his legacy of climate and soil as primary factors explaining broad vegetation patterns. There is increasing evidence that many open, non-forested ecosystems (savannas, grasslands, and shrublands) cannot be predicted by climate and soil – and are ancient and diverse systems maintained by fire and/or vertebrate herbivory. Paleoecological and phylogenetic studies have shown the key role of fire and grazing at geological time scales (Fig. 2). In this paper [1], we propose moving beyond the legacy of Humboldt by embracing fire and large mammal herbivory as key factors in explaining the ecology and evolution of world vegetation. This implies understanding grasslands, savannas, and shrublands as ancient and diverse ecosystems that require conservation, including the processes that maintain them (grazing and wildfires).
 

Fig. 2. Changes of the drivers related to plant consumers (fire and herbivory), together with the evolution of different vegetation types, and some plant traits (serotiny and thick bark of pines, epicormic resprouting in eucalypts), along the evolutionary history of plants. Upper pointing triangles are peaks of O2 atmospheric concentration and fire activity; lower-pointing triangles are megafauna extinction events, also associated to fire activity peaks. Note that modern fire regimes are very recent, and at this scale they are almost a point. From [1].

Fig. 3. Alexander von Humboldt and Aimé Bonpland on the foot of Chimborazo, painting by Friedrich Georg Weitsch (1810)

 
References
[1] Pausas J.G. & Bond W.J. 2019. Humboldt and the reinvention of nature. J. Ecol. [doi | jecolblog | pdf]

[2] Pausas J.G. & Lamont B.B. 2018. Ecology and biogeography in 3D: the case of the Australian Proteaceae J. Biogeogr. 45: 1469-1477. [doi | pdf]