Posts Tagged ‘bark’

The ecology of bark thickness (2): another twist

December 3rd, 2016 No comments

Sometime ago we proposed that "at the global scale, a significant proportion of the variability in bark thickness is explained by the variability in fire regimes", and specifically predicted that frequent low intensity fires select for thick bark [1]. In addition, we suggested that differentiating between inner and outer bark thickness would help to better understand the functional role of bark, especially in non-fire prone ecosystems. The paucity of available data at a global scale limited an empirical demonstration of the proposed framework.

A recent paper has now provided evidence for the fire hypothesis of bark thickness at a global scale [2, 3]. Specifically, Rosell [2] regressed bark thickness against fire frequency and climate parameters and showed that the most sensitive part of the bark in relation to fire was the outer bark, while the inner bark was quite variable and slightly related to both fire and climate [2]. In the early paper [1] we also mentioned that little was known about the role of bark thickness in arid ecosystems. Recent research support the role of bark as a fire protection mechanisms in some arid ecosystems [4, 5].

To advance in the relationship between bark thickness and fire, it is necessarily to consider not only fire frequency, but also fire intensity, and to scale these fire characteristics with plant life-histories ([3], see figure below). This is because the relationship between fire regime and bark thickness is not expected to be simple and linear, but a bit more complex, including some threshold-type relationships (figure below).

Little by little we are improving our understanding on the role of bark as a fire-protection mechanism, and how fire regimes has shaped bark thickness in many ecosystems.


Figure: Bark thickness as a function of fire regime: flame height (an indicator of fire intensity) and mean fire return interval (fire frequency). Fire regime is scaled by the characteristics of the plant (height to the base of the crown and longevity, respectively). The shaded area represents the areas where thick bark is adaptive for fire protection, i.e., when return intervals are shorter than the lifespan of the plant and fires are of low intensity (flame height is shorter than the distance to the base of the crown, e.g., surface fires); the shade area is limited thresholds (values of 1 in the axes). The unshaded area represents the conditions where thick barks are not adaptive (thin bark is more likely), i.e., when fires are crown-fires or when the return interval is long (in relation to the longevity of the plant). From [3].


[1] Pausas, J.G. 2015. Bark thickness and fire regime. Functional Ecology 29:317-327. [doi | pdf | suppl. | blog]

[2] Rosell J.A. 2016. Bark thickness across the angiosperms: more than just fire. New Phytologist 211: 90–102

[3 ] Pausas J.G. 2017. Bark thickness and fire regime: another twist. New Phytologist 213: 13-15. [doi| pdf] <- New!

[4] Schubert, A. T., Nano, C. E. M., Clarke, P. J. & Lawes, M. J. 2016. Evidence for bark thickness as a fire-resistance trait from desert to savanna in fire-prone inland Australia. Plant Ecol. 217: 683-696.

[5] Cousins, S. R., Witkowski, E. T. F. & Pfab, M. F. 2016. Beating the blaze: Fire survival in the fan aloe (Kumara plicatilis), a succulent monocotyledonous tree endemic to the Cape fynbos, South Africa. Austral Ecol. 41:466-479.

Ecology and evolution in fire-prone ecosystems

February 28th, 2015 2 comments

During the last years I've been working in many topics related to fire ecology and plant evolution in ecosystems subject to recurrent fires (mainly mediterranean and savanna ecosystems). Because I believe knowledge should be spread around easily, I make my results available to the public in my web page (see publications list) and in this blog. However, having the cumulative list of paper published each year is not very convenient for people searching for a specific topic. For this reason, I'm rearranging most of my articles by topics as follows:

1. Fire history
2. Fire regime: climate & fuel
3. Fire traits (resprouting, postfire germination, serotiny, bark thickness, flammability, data & methods)
4. Fire & plant strategies (in Mediterranean ecosystems, in pines, in savannas, community assembly)
5. Fire & evolution
6. Some fire-adapted species (Pinus halepensis, Quercus suber, Ulex parviflorus)
7. Fire & vegetation modelling
8. Plant-animal interactions
9. Restoration & conservation

See: fire-ecology-evolution.html

Some papers may be repeated if they clearly fit in more than one topic; some papers, mainly old ones, do not fit well in any of these topics and have not been included (at least at the moment), they still can be found in the section of publications sorted by year. I'm still working on this rearrangement, so some modifications are possible; and any comment is welcome.
I hope this is useful for somebody!

Publications: by year | by topic | books


The ecology of bark thickness

December 1st, 2014 No comments

Bark is a vital and very visible part of woody plants, yet the functional and evolutionary ecology of the bark is still poorly understood. In a recent article I have studied one of the bark properties: bark thickness [1]. Bark thickness is very variable among woody plants and fire is a key factor selecting for a thick bark. This is because barks are very good heat insulators and under low intensity fires, small differences in bark thickness provides a great increase in the stem protection and survival. Consequently, at the global scale, an important proportion of the variability in bark thickness should be explained by the variability in fire regimes. In this paper I provide evidences supporting the role of fire regime in shaping bark thickness (in conjunction with other plant traits) on a global scale [1].

Forest environments with very frequent (and low intensity) understory fires select for trees with thick bark at the base of the bole. In some savannas, trees do not have specially thick barks as they tend to growth quickly to escape the height affected by grass fires. Savannas living in poor soils may not be able to growth quickly and thus trees can only survive if they have a very thick bark in the whole plant (including in the thin branches). In Mediterranean ecosystems, fires are less frequent than in savannas, and there is time for the accumulation of fine woody biomass. Consequently, fires burns intensely (crown fires) and thus small differences in bark thickness do not increase stem survival; in such conditions, most species have relatively thin barks. In wet tropical forests, tree barks are very thin because fire are very rare and thus a thick bark is not advantageous. In very arid ecosystems, fuels are too sparse for fire spread, and thus the observed variability in bark thickness is related to other factors like a response to water stress. In conclusion, fire regimes can explain a large proportion of the variability of bark thickness at the global scale, and thus this trait varies across ecosystems in a predictable manner.


Figure: Examples of trees with thick bark: A. Myrcia bella (Myrtaceae, Brazil); B. Quercus suber (Fagaceae, Mediterranean Basin), in the cover of the book 'Cork oak Woodlands on the Edge' [2]; C: Eremanthus seidelii (Asteraceae, Brazil); and D: Enterolobium gummiferum (Fabaceae), small top branch. Photos from [1] and [2].


[1] Pausas, J.G. 2015. Bark thickness and fire regime. Functional Ecology   [doi | pdf | suppl.]

[2] Aronson J., Pereira J.S., Pausas J.G. (eds). 2009. Cork Oak Woodlands on the Edge: conservation, adaptive management, and restoration. Island Press, Washington DC. 315 pp. [The book]

Afrotropical and neotropical savannas are different

July 29th, 2013 No comments

Savannas are typically ecosystems dominated by grasses with a variable tree density (e.g., [1]). However, the savanna biome is very large, it occurs in different continents, and includes a large variability in the vegetation structure and composition. Fire and herbivory are the main disturbance factors shaping savannas worldwide and because the different climatic conditions and the different evolutionary histories among different savannas, fire and herbivory regimes also varies among savannas. Because plants are not adapted to fire and herbivory “per se” but to specific regimes of herbivory and fire [2], we expect different strategies to cope with these disturbances in different savannas. In this framework, we have recently compared savannas from Africa and from South America (afrotropical and neotropical savannas respectively) [3]: Afrotropical savannas have a dryer climate and are more intensely grazed than neotropical savannas, and thus the amount of available fuel is typically lower in afrotropical than in the neotropical savannas. Consequently fires tend to be more intense in neotropical savannas. In afrotropical conditions, young woody plants tend to grow quickly in height to soon locate the canopy above the flame zone before the next fire, and above the browsing height. Thus these plants tend to have a pole-like or lanky architecture (the lanky strategy). In contrast, in neotropical savannas where herbivory pressure is lower they require a thick corky bark for protection against relatively intense fires (the corky strategy) [3]. Despite the two fire escape strategies appear in both Africa and South America, we suggest that the lanky strategy is more adaptive in afrotropical savannas, while the corky strategy is more adaptive in neotropical savannas [3].

Figure: Diospyros hispida A.DC. (Ebenaceae), a South American example of a plant with the corky strategy. Although the trunk was fully burned one year earlier (dark branches and trunk), the bark protected the lateral buds which enabled epicormic resprouting and the formation of lateral resprouts (light grey branches). This photo was taken in Emas National Park (cerrado ecosystem, Brazil) at the beginning of the rainy season (2011) when this deciduous plant starts to produce new leaves (Photo: V.L. Dantas). For an example of the lanky strategy see [4].

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

[2] Keeley J.E., Pausas J.G., Rundel P.W., Bond W.J., Bradstock R.A. 2011. Fire as an evolutionary pressure shaping plant traits. Trends Plant Sci. 16(8): 406-411. [doi | trends | pdf]

[3] Dantas V. & Pausas J.G. 2013. The lanky and the corky: fire-escape strategies in savanna woody species Journal of Ecology 101: 1265-1272 [doi | pdf]

[4] Archibald, S. & Bond, W.J. 2003. Growing tall vs growing wide: tree architecture and allometry of Acacia karoo in forest, savanna, and arid environments. Oikos, 102: 3-14.


Bark harvesting and Cork oak vulnerability to fire

July 11th, 2012 No comments

Cork oak (Quercus suber) is a strong fire-resistant tree species thank to is very thick and insulating corky bark [1-4]. In fact it is the only European tree with the capacity to resprout from epicormic buds in the canopy after an intense crown-fire [1]. However, the bark of the cork oak is periodically harvested for cork production (mainly for bottle tops but also for other uses, [2]) and thus bark harvesting increases the vulnerability of the tree to fire. In a recent paper we quantified the response of cork oak (tree mortality, stem mortality, and crown recovery) after fire [5]. The results showed that fire vulnerability was higher for trees with thin bark (young or recently debarked individuals) and decreased with increasing bark thickness until cork was 3–4 cm thick. This bark thickness corresponds to the moment when exploited trees are debarked again, meaning that exploited trees are vulnerable to fire during a long period. Exploited trees were also more likely to be top-killed than never-debarked trees, even for the same bark thickness. Additionally, vulnerability to fire increased with burn severity and with tree diameter, and was higher in trees burned in early summer or located in drier south-facing aspects. All these aspects need to be considered when managing cork oak woodlands specially nowadays that fire activity is increased [6]. Increasing the length of the cork harvesting cycle would increase the time during which the trees have a thicker bark and are better protected against fire injury. Since cork is the main economical income from these forests, stopping bark exploitation might be unrealistic in most cases. However, in fire-prone areas where conservation and tourism are the main objectives, stopping bark explotation would likely be the most effective option to increase ecosystem resilience to fire. The valorisation of many other services provided by cork oak forests [7] could create economic incentives to decrease the bark-exploitation dependency of these systems in the future.

Foto: Cork oak  resprouting from epicormic buds (By F. Catry)


[1] Pausas, J.G. 1997. Resprouting of Quercus suber in NE Spain after fire. J. Veg. Sci. 8: 703-706. [doi | pdf]

[2] Aronson, J., J. S. Pereira, and J. G. Pausas (eds). 2009. Cork Oak Woodlands on the Edge: Ecology, Adaptive Management, and Restoration. Island Press, Washington, DC. [web of the book]

[3] Pausas J.G. 2009. Convergent evolution., 8/Nov/2009. [link]

[4] Pausas J.G. 2011. Bark thickness: a world record?, 3/Jan/201. [link]

[5] Catry F., Moreira F., Pausas J.G., Fernandes P.M., Rego F., Cardillo E. & Curt T. 2012. Cork Oak vulnerability to fire: the role of bark harvesting, tree characteristics and abiotic factors. PLoS ONE 7: e39810. [doi | pdf ]

[6] Pausas J.G. & Fernández-Muñoz S. 2012. Fire regime changes in the Western Mediterranean Basin: from fuel-limited to drought-driven fire regime. Climatic Change 110: 215-226. [doi | springer | pdf]

[7] Bugalho M.N., Caldeira M.C., Pereira J.S., Aronson J., & Pausas J.G. 2011. Mediterranean Cork oak savannas require human use to sustain biodiversity and ecosystem services. Frontiers in Ecology and the Environment 9: 278-286. [doi | pdf | blog]


Bark thickness: a world record?

January 3rd, 2011 3 comments

The thickness of the bark is a trait of paramount importance in trees living in ecosystems with frequent surface (understory) fires (e.g., some coniferous forests, savanna woodlands, etc.). This is because the bark is a good insulator protecting vital tissues from the heat of the fire. Having a bark few millimeter thicker provide an advantage in such fire-prone ecosystems. Thus there has been a selection for thick barks in surface fire ecosystems [1]. A prominent example of a tree with a very thick and insulating bark is the Cork oak (Quercus suber) that growth in the western part of the Mediterranean Basin [2]. In such species the thicker is the bark, the better is the response after fire [3, 4]. This bark is so thick and insulating that it is used not only as bottle tops, but also as insulating material in many industrial applications. However the Mediterranean Basin has been densely populated from long ago and it is very difficult (if possible) to find Cork oak woodlands in "natural" conditions, and thus it is not easy to know how thick the bark of Cork oak could attain in natural conditions. Most trees are frequently debarked for obtaining cork (frequencies ranging from every 9 to every 12 years, depending of the site conditions).

Few days ago I visited an ethnographic museum in Aggius (Sardinia) and found a piece of Cork oak bark of about 22 cm thick (see picture below), which is pretty thick. I only know of one record of a thicker bark: 27 cm in a 140 years-old Cork oak that was never debarked [5]. Do you know of any tree (of the same or another species) in the world with a thicker bark? Is Cork oak the world record on bark thickness?

Figure: Piece of bark from a Cork oak (Quercus suber), in the ethnographic museum of Aggius (Sardinia).


[1] Pausas J.G. 2009. Convergent evolution., 8/Nov/2009. [link]

[2] Aronson J., Pereira J.S., Pausas J.G. (eds). 2009. Cork Oak Woodlands on the Edge: conservation, adaptive management, and restoration. Island Press, Washington DC.  [link]

[3] Pausas, J.G. 1997. Resprouting of Quercus suber in NE Spain after fire. J. Veg. Sci. 8: 703-706. [doi pdf]

[4] Catry F.X., Rego F., Moreira F., Fernandes F.M., Pausas J.G. 2010. Post-fire tree mortality in mixed forests of central Portugal. Forest Ecology & Management 206: 1184-1192. [doi | pdf]

[5] Natividade J.V. 1950. Subericultura. Direçao Geral dos Serviços Florestais e Aquícolas Lisbon, Portugal.

Convergent evolution

November 8th, 2009 No comments

Images from two different tree species (A and B), from different Families (and different Orders), taken in different continents...


The thick bark offers protection to fire and thus these species are both adapted to live in fire-prone ecosystems [1].

Can you guess the species name of A and B?    [ Answer: A | B ]


[1] See also: The ecology of bark thickness | The ecology of bark thickness (2): another twist


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