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Posts Tagged ‘traits’

Pinus canariensis

May 7th, 2017 No comments

The last post was about Pinus brutia [1] from the Eastern Mediterranean basin. Another pine of the mediterranean group (Pinaster group) is Pinus canariensis, endemic of Canary Islands, in the north west of Africa (in the Atlantic). P. canariensis have a thick bark and resprouts vigorously from stem buds (epicormic resprouting) after crown fires. In addition, it produce serotinous cones, a clear adaptation to recruit after fire [2,3]. Very few other trees have strong adaptations to both survival and regeneration postfire; P. canariensis is among the best fire-adapted trees in the world, likely to survive very different fire regimes.

Pictures of Pinus canariensis (by JG Pausas except mid-right from NASA).
· Top-left: 5 years after a crown-fire (La Palma, Canary Is.).
· Mid-left: plantation 2 years after fire (Vall d’Ebo, Alicante, eastern Spain; planted in the 50s).
· Bottom-left: Contrasted response of Pinus halepensis (left; fire-killed serotinous pine) and P. canariensis (right, resprouting) two years after fire (Alicante, eastern Spain).
· Top-, bottom-right: epicormic resprouts 3 months after fire (Tenerife, Canary Is.).
· Mid-right: a fire plume from a wildfire in La Palma (Canary Is.).

References
[1] Pinus brutia, jgpausas.blogs.uv.es, 19 Apr 2017

[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: 406-411. [doi | sciencedirect | trends | pdf | For managers]

[3] Pausas, J.G. 2015. Evolutionary fire ecology: lessons learned from pines. Trends Plant Sci. 20: 318-324. [doi | sciencedirect | cell | pdf]

 

Manual para la medición de caracteres funcionales de plantas

January 4th, 2017 No comments

En 2003 se publicó el  primer manual para la medición estandarizada de caracteres funcionales de plantas [1], y en 2013 se realizó una segunda versión más completa y actualizada [2]. Estas dos versiones se publicaron en inglés en la revista Australian Journal of Botany. Ahora, la misma revista, publica la traducción de la segunda versión (2013) en español y la pone disponible a todo el mundo.

Versión en inglés: Handbook  y  supplementay material  [2]

Versión en español: Manual  y  material suplementario  [3]

Original en:  Australian Journal of Botany

 

Emas2009Fotografía: Midiendo caracteres funcionales en plantas de la sabana brasileña (cerrdao) [4]

 

Referencias

[1] Cornelissen, J.H.C., Lavorel, S., Garnier, E., Díaz, S., Buchmann, N., Gurvich, D.E., Reich, P.B., Ter Steege, H., Morgan, H.D., van der Heijden, M.G.A., Pausas, J.G. & Poorter, H. 2003. Handbook of protocols for standardised and easy measurement of plant functional traits worldwide. Aust. J. Bot. 51: 335-380. [doipdf | CSIRO pub]

[2] Pérez-Harguindeguy N, Díaz S, Garnier E, Lavorel S, Poorter H, Jaureguiberry P, Bret-Harte MS, Cornwell WK, Craine JM, Gurvich DE, Urcelay C, Veneklaas EJ, Reich PB, Poorter L, Wright IJ, Ray P, Enrico L, Pausas JG, de Vos AC, Buchmann N, Funes G, Quetier F, Hodgson JG, Thompson K, Morgan HD, ter Steege H, van der Heijden MGA, Sack L, Blonder B, Poschlod P, Vaieretti V, Conti G, Staver AC, Aquino S, Cornelissen JHC. 2013. New handbook for standardised measurement of plant functional traits worldwide. Australian Journal of Botany 61(3): 167-234. [doi | pdf | Suppl. Mat.]

[3] Pérez-Harguindeguy N, Díaz S, Garnier E, Lavorel S, Poorter H, Jaureguiberry P, Bret-Harte MS, Cornwell WK, Craine JM, Gurvich DE, Urcelay C, Veneklaas EJ, Reich PB, Poorter L, Wright IJ, Ray P, Enrico L, Pausas JG, de Vos AC, Buchmann N, Funes G, Quetier F, Hodgson JG, Thompson K, Morgan HD, ter Steege H, van der Heijden MGA, Sack L, Blonder B, Poschlod P, Vaieretti V, Conti G, Staver AC, Aquino S, Cornelissen JHC. 2013. Nuevo manual para la medición estandarizada de caracteres funcionales de plantas. Australian Journal of Botany 61(3): 167-234. [doi | pdf | Mat. Supl.]

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

Heritability of serotiny (2): a molecular approach

December 2nd, 2015 No comments

Not long ago we demonstrated that serotiny (i.e., the capacity to accumulate a seed bank in the canopy until the seeds are released by fire) is an heritable trait in pines [1]. This analysis was based on a classical provenance – progeny common garden experiment. However, trait variability under controlled environmental conditions may not fully reflect the variability observed in the field, and thus this estimate of heritability may not reflect how traits respond to natural selection. This is because there is higher environmental variability in the field and also because garden experiments typically include individuals that would not survive in the field (i.e., artificially increases progeny survival) [2]. With the aim of obtaining a more realistic estimate of heritability of serotiny, we have recently estimate it directly in the field for two pine species (P. halepensis, P. pinaster) [3]. Because in the field it is not possible to construct a pedigree, we used the relatedness among individuals estimated from molecular markers (SNPs) for the same individuals from which we had estimated serotiny previously [4]. The variance in serotiny was modelled incorporating the environmental variability (climate and fire regime) using a Bayesian ‘animal model’. As expected, field heritability was smaller (around 0.10 for both species) than previous estimates under common garden conditions (0.20). The difference is not surprising because wild P. halepensis and P. pinaster populations extend over heterogeneous landscapes with large environmental variations. Our results highlight the importance of measuring quantitative genetic parameters in natural populations, where environmental heterogeneity is a critical aspect. The heritability of serotiny, although not high, combined with high phenotypic variance within populations, confirms the potential of this fire-related trait for evolutionary change in the wild [2].

Pinus patula
Fig: Serotinous cones of P, halepensis and P. pinaster can be observed in previous posts (P, halepensis, P. pinaster). The photo here shows serotinous cones of Pinus patula from central Mexico (in a foggy rainy day).

References

[1] Hernández-Serrano, A., Verdú, M., Santos-Del-Blanco, L., Climent, J., González-Martínez, S.C. & Pausas, J.G. 2014. Heritability and quantitative genetic divergence of serotiny, a fire-persistence plant trait. Annals of Botany 114: 571-577.  [doi | pdf | suppl. | blog]

[2] Pausas, J.G. 2015. Evolutionary fire ecology: lessons learned from pines. Trends in Plant Science 20: 318-324. [doi | sciencedirect | cell | pdf]

[3] Castellanos, M.C., González-Martínez, S. & Pausas, J.G. 2015. Field heritability of a plant adaptation to fire in heterogeneous landscapes. Molecular Ecology 24: 5633–5642 [doi | pdf | suppl.]

[4] Hernández-Serrano A., Verdú M., González-Martínez S.C., Pausas J.G. 2013. Fire structures pine serotiny at different scales. American Journal of Botany 100: 2349-2356. [doi | amjbot | pdf | supp. | blog]

 

Lignotubers

November 17th, 2015 1 comment

Lignotubers are swollen woody structures located at the root-shoot transition zone of some plants; they contain numerous dormant buds and starch reserves [1]. They are ontogenetically programmed, that is, they are not the product of repeated disturbances; and thus they can be observed at very early stages of the plant development (other types of basal burls may be a response to multiple disturbances). Lignotubers enables the plant to resprout prolifically after severe disturbances that remove the aboveground biomass, thus they are considered adaptive in fire-prone ecosystems [2]. Lignotubers are not well-known in many floras because they are often below-ground (i.e., detected only after excavation) and because they are often confused by other non-ontogenetically determined basal burls; thus some reports of lignotubers in the literature are mistakes. In a recent review [1] we provide examples of species with a clear evidence of lignotubers in the Mediterranean basin, together with detailed morphological and anatomical description of lignotubers in saplings. The species with lignotuebers in the Mediterranean basin include many Erica species (e.g. E. arborea, E. scoparia, E. australis, E. lusitanica, E. multiflora), the two Arbutus species (A. unedo, A. andrachne), Rhododendron ponticum, Viburnum tinus, Phillyera angustifolia, Quercus suber (not obvious macroscopically!), Tetraclinis articulata and Juniperus oxycedrus (but not in all populations!). Please let me know (email address here) if you know of other Mediterranean basin species with lignotubers! Thanks

lignotubers
Figures: Examples of lignotubers for Mediterranean basin species. A Juniperus oxycedrus (resprouting after fire). B Viburnum tinus. C Arbutus unedo. D Quercus suber (not a clear basal swelling). E Olea europaea. F Phillyrea angustifolia (adult), G Phillyrea angustifolia (saplings). In many species (e.g., V. tinus, A. unedo and P. angustifolia) the lignotuber is only evident after excavating the root-shoot transition zone.

References

[1] Paula S., Naulin P.I., Arce C., Galaz C. & Pausas J.G. 2016. Lignotubers in Mediterranean basin plants. Plant Ecology  [doi | pdf | suppl.]

[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: 406-411.  [doi | sciencedirect | pdf | For managers]

 

Resprouting at the global scale

November 2nd, 2015 No comments

Plant resprouting (i.e., the ability to form new shoots after destruction of living tissues from disturbance) is often considered a simple qualitative trait and used in many ecological studies. However, resprouting is a trait that increases fitness under many different disturbance types, occurs in a wide range of environments, is widespread in many lineages, and is morphologically very diverse. In a recent paper we review some of the complexities and misunderstandings of resprouting and highlight that cautions is needed when using resprouting ability to predict vegetation responses across disturbance types and biomes [1]. There are marked differences in resprouting depending on the disturbance type, and fire is often the most severe disturbance because it includes both defoliation and lethal temperatures. In the mediterranean biome, there are differences in functional strategies to cope with water deficit between reprouters (dehydration avoiders) and non-resprouters (dehydration tolerators) [1,2]; however, there is little research to unambiguously extrapolate these results to other biomes, and some of the extrapolations seems to be incorrect. In addition, resprouting in the mediterranean biome tends to be binary, that is, species are either resprouters or non-resprouters [3], and intermediate cases are evolutionary unstable [4]; however this is not necessary true in other biomes (e.g., in the tropics). Furthermore, predictions of vegetation responses to changes in disturbance regimes require consideration of not only resprouting but also other relevant traits (e.g., seeding, bark thickness) and the different correlations among traits observed in different biomes [5]; models lacking these details would behave poorly at the global scale. Overall, the lessons learned from a given disturbance regime and biome, like crown-fire mediterranean ecosystems, can guide research in other ecosystems but should not be extrapolated at the global scale.

 

Cistus-Quercus

Fig: Fire allows the coexistence of species with very different strategies: Cistus albidus seedling (left) and Quercus coccifera resprout (right) 10 months after a high intensity fire in eastern Spain (Cortes de Pallás fire, 2012, Valencia). Cistus albidus  is a drought semi-deciduous nonresprouter (obligate postfire seeder ) with a physiological drought-tolerant behavior; Quercus coccifera  is an sclerophyllous (evergreen) obligate resprouter with drought-avoiding traits [2].

References
[1] Pausas, J.G., Pratt, R.B., Keeley, J.E., Jacobsen, A.L., Ramirez, A.R., Vilagrosa, A., Paula, S., Kaneakua-Pia, I.N. & Davis, S.D. 2016. Towards understanding resprouting at the global scale. New Phytologist 209:945-954. [doi | pdf] — New paper!

[2] Vilagrosa A., Hernández E.I., Luis V.C., Cochard H., Pausas, J.G. 2014. Physiological differences explain the co-existence of different regeneration strategies in Mediterranean ecosystems. New Phytologist 201: 1277-1288. [doi | pdf ]

[3] Pausas, J.G., Bradstock, R.A., Keith, D.A., Keeley, J.E. & GCTE Fire Network. 2004. Plant functional traits in relation to fire in crown-fire ecosystems. Ecology 85: 1085-1100. [pdf | esa | doi]

[4] Pausas J.G. & Keeley J.E., 2014. Evolutionary ecology of resprouting and seeding in fire-prone ecosystems. New Phytologist 204: 55-65. [doi | wiley | pdf]

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

 

Fire adaptations in Mediterranean Basin plants

September 7th, 2015 No comments

Few days ago a botanist colleague ask me whether there were some fire adaptations in the plants of the Mediterranean Basin, similar to those reported in other mediterraenan-climate regions. So I realised that researchers working on other topics may not be aware of the recent advances in this area. Here is my brief answer, i.e., some examples of species growing in Spain that show fire adaptations; this is by no means an exhaustive list, but a few examples of common species for illustrative purpose. You can find a description of these adaptations and further examples elsewhere [1, 2, 3, 4]. It is also important to note that plants are not adapted to fire per se, but to specific fire regimes, and thus some adaptations my provide persistence to some fire regimes but not to all [1]. That is, species that exhibit traits that are adaptive under a particular fire regime can be threatened when that regime changes.

  • Serotiny (canopy seed storage): Pinus halepensis, Pinus pinaster, with variability in serotiny driven by different fire regimes [5, 6]
  • Fire-stimulated germination: There are examples of heat-stimulated germination, like many Cistaceae (e.g., Cistus, Fumana [7, 8]) and many Fabaceae (e.g., Ulex parviflorus, Anthyllis cytisoides [7, 8]), as well as examples of smoke-stimulated germination like many Lamiaceae (e.g., Rosmarinus officinalis, Lavandula latifolia [7]) or Coris monspeliensis (Primulaceae [7]). There are also examples of species with smoke-stimulated seedling growth (Lavandula latifolia [7])
  • Resprouting from lignotubers: Arbutus unedo, Phillyrea angustifolia, Juniperus oxycedrus, many Erica species (e.g., E. multiflora, E. arborea, E. scoparia, E. australis) [4, 17]
  • Epicormic resprouting: Quercus suber [9, 10], Pinus canariensis [4]
  • Fire-stimulated flowering: Some monocots like species of Asphodelus, Iris, Narcissus [11, 12]
  • Enhanced flammability: Ulex parviflorus shows variability of flammability driven by different fire regimes [13] and under genetic control [14]. Many Lamiaceae species have volatile organic compounds that enhance flammability (e.g., Rosmarinus officinalis [16]).
  • Thick bark and self-pruning (in understory fires): Pinus nigra [3,15]

 

fireadaptations2

References

[1] Keeley et al. 2011. Fire as an evolutionary pressure shaping plant traits. Trends Plant Sci 16:406-411. [doi | pdf]

[2] Keeley et al. 2012. Fire in Mediterranean Ecosystems. Cambridge University Press. [book]

[3] Pausas JG. 2012. Incendios forestales. Catarata-CSIC. [book]

[4] Paula et al. 2009. Fire-related traits for plant species of the Mediterranean Basin. Ecology 90:1420-1420. [doi | pdf | BROT database]

[5] Hernández-Serrano et al. 2013. Fire structures pine serotiny at different scales. Am J Bot 100:2349-2356. [doi | pdf]

[6] Hernández-Serrano et al. 2014. Heritability and quantitative genetic divergence of serotiny, a fire persistence plant trait. Ann Bot 114:571-577. [doi | pdf]

[7] Moreira et al. 2010. Disentangling the role of heat and smoke as germination cues in Mediterranean Basin flora. Ann Bot 105:627-635. [doi | pdf]

[8] Moreira B and Pausas JG. 2012. Tanned or Burned: the role of fire in shaping physical seed dormancy. PLoS ONE 7:e51523. [doi | plos | pdf]

[9] Pausas JG. 1997. Resprouting of Quercus suber in NE Spain after fire. J Veget Sci 8:703-706. [doi | pdf]

[10] Catry et al. 2012. Cork oak vulnerability to fire: the role of bark harvesting, tree characteristics and abiotic factors. PLoS ONE 7:e39810. [doi | pdf ]

[11] Postfire flowering: Narcissusjgpausas.blogs.uv.es 2 May 2015

[12] Postfire blooming of Asphodelous, jgpausas.blogs.uv.es 5 Apr 2014

[13] Pausas et al. 2012. Fires enhance flammability in Ulex parviflorus. New Phytol 193:18-23. [doi | pdf]

[14] Moreira et al. 2014. Genetic component of flammability variation in a Mediterranean shrub. Mol Ecol 23:1213-1223. [doi | pdf]

[15] He et al. 2012. Fire-adapted traits of Pinus arose in the fiery Cretaceous. New Phytol 194:751-759. [doi | pdf | picture]

[16] Flammable organic compounds: Rosmarinus officinalis, jgpausas.blogs.uv.es 2-Oct-2015

[17] Paula et al. 2016. Lignotubers in Mediterranean basin plants. Plant Ecology [doi | pdf | suppl. | blog]

 

Evolutionary fire ecology in pines

April 1st, 2015 No comments

Fire is an ancient and recurrent disturbance factor in our planet and has been present since the origin of terrestrial plants [1]. However, demonstrating whether fire has acted as an evolutionary force is not an easy task [2]. In this context, the emerging discipline of evolutionary fire ecology aims to understand the role of wildfires in shaping biodiversity. In a recent review paper I summarize what we have learned on evolutionary fire ecology by studying the iconic genus Pinus [3]. I suggest that the study of pines has greatly increased our understanding of the role of fire as an evolutionary pressure on plants.

Macro-evolutionary studies of the genus Pinus provide the oldest current evidence of fire as an evolutionary pressure on plants and date back to ca. 125 Million years ago (Ma). Micro-evolutionary studies show that fire traits are variable within and among populations, and especially among populations subject to different fire regimes. In addition, there is increasing evidence of an inherited genetic basis to variability in fire traits. Added together, pines provide compelling evidence that fire can exert an evolutionary pressure on plants and thus shape our biodiversity. In addition, evolutionary fire ecology is providing insights to improve the management of our pine forests under changing conditions. The lessons learned from pines may guide research on the evolutionary ecology in other taxa.

pinus-serotiny
Figure: Example of trait divergence among populations living under different fire regime. Serotiny (as % of closed cones) in populations living under frequent crown fires (red boxes) and in populations where crown-fires are rare (green boxes) for two pine species, Pinus halepensis (Allepo pine, left) and P. pinaster (maritime pine, right).

References
[1] Pausas, J.G. and Keeley, J.E. 2009. A burning story: The role of fire in the history of life. Bioscience 59: 593-601. [doi | jstor | BioOne | pdf]

[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 in Plant Science 16: 406-411. [doi | sciencedirect | trends | pdf]

[3] Pausas, J.G. (2015) Evolutionary fire ecology: lessons learned from pines. Trends in Plant Science 20(5): 318-324. [doi | sciencedirect | pdf]

 

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.

thick-bark2

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].

References

[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]

Trait databases: BROT to EOL

October 26th, 2014 No comments

Some years ago we complied and published a database on plant traits related to fire for the Mediterranean basin, the BROT database [1, 2]. Now the Encyclopedia of Life (EOL, eol.org), which is an initiative to gather scientific knowledge about all animal and plant life on Earth, has incorporated the BROT database [link]! We are very happy that EOL consider BROT as a reliable source of information; this implies that our compilation effort is now much more widely accessible, with a friendly interface, and integrated with other sources of information. For instance, if you search a Mediterranean plant species in the EOL search engine (e.g., Cistus albidus), you get, a part from pictures, a description, and other details, a window with the trait information extracted from BROT (see the overview result here; you can also go to the full trait data). It is aslo possible to search by trait in all EOL databases (eol.org/traitbank). Note however that we were not responsible for translating the BROT database to the EOL format, so any error or misinterpretation during this process is not our fault! In fact we have never been asked or notified that EOL was going to incorporate BROT, I found it just by chance …

eol

References:

[1]  Paula S, Arianoutsou M, Kazanis D, Tavsanoglu Ç, Lloret F, Buhk C, Ojeda F, Luna B, Moreno JM, Rodrigo A, Espelta JM, Palacio S, Fernández-Santos B, Fernandes PM, and Pausas JG. 2009. Fire-related traits for plant species of the Mediterranean Basin. Ecology 90: 1420. [doi] [ESA journals] [Ecological Archives E090-094] [pdf]

[2] Paula S. & Pausas J.G. 2013. BROT: a plant trait database for Mediterranean Basin species. Version 2013.06. URL: http://www.uv.es/jgpausas/brot.htm

 

Heritability of serotiny

September 29th, 2014 No comments

Evolution by mean of natural selection requires three conditions: there is variation in the trait, this variation is linked to differences in fitness, and the variation is heritable (Darwin!). In many traits we do not have reliable information for the three processes. For a serotinous species, there is evidence that the level of serotiny is variable, and specially it varies in relation to the fire regime of the population. This is because serotiny increases fitness in crown-fire ecosystems and it is not advantageous in ecosystems that do not suffer frequent fires or in ecosystems with understory fires. We recently studied how serotiny of two pine species (Pinus halepensis and Pinus pinaster) varies within population and between populations with different fire regimes and also provided a meta-analysis of the relation between serotiny and fire from other published studies [1]. We also performed a genetic association study for serotiny using SNPs and showed that 17 loci explained ca. 29% of the serotiny variation found in the field in Pinus pinaster [2], suggesting that serotiny variation have a genetic basis. In our most recent paper we provide the first estimate of heritability for a fire trait; specifically we computed the norrow-sense heritability (h2) of serotiny in Pinus halepensis using the common garden approach [3]. We also evaluated whether fire has left a selection signature on the level of serotiny among populations by comparing the genetic divergence of serotiny with the expected divergence of neutral molecular markers (QST – FST comparison). Serotiny showed a significant heritability (h2 = 0.20). The quantitative genetic differentiation among provenances for serotiny (QST= 0.44) was significantly higher than expected under a neutral process (FST = 0.12), suggesting adaptive differentiation. Overall we showed that serotiny is a heritable trait and that it has been shaped by natural selection driven by fire.

ph-serotiny
Figure: Serotinous cones of Pinus halepensis (Foto: J.G. Pausas)

References:

[1] Hernández-Serrano A., Verdú M., González-Martínez S.C., Pausas J.G. 2013. Fire structures pine serotiny at different scales. American Journal of Botany 100 (12): 2349-2356. [doi | amjbot | pdf | supp. | blog]

[2] Budde, K. B., Heuertz, M., Hernández-Serrano, A., Pausas, J.G., Vendramin, G.G., Verdú, M. & González-Martínez, S.C. 2014. In situ genetic association for serotiny, a fire-related trait, in Mediterranean maritime pine (Pinus pinaster Aiton). New Phytologist 201: 230-241.  [doi | pdf | supp1 | supp2]

[3] Hernández-Serrano, A., Verdú, M., Santos-Del-Blanco, L., Climent, J., González-Martínez, S.C. & Pausas, J.G. 2014. Heritability and quantitative genetic divergence of serotiny, a fire-persistence plant trait. Annals of Botany 114: 571-577. [doi | pdf | suppl.]

 

Evolutionary ecology of resprouting and seeding

July 15th, 2014 No comments

There are two broad mechanisms by which plant populations persist under recurrent fires: resprouting from surviving tissues, and seedling recruitment [1]. Species that live in fire-prone ecosystems can have one of these mechanisms or both [1]. In a recent review paper [2], we propose a model suggesting that changes in evolutionary pressures that modify adult (P) and juvenile (C) survival in postfire conditions (Fig. 1 below) determine the long-term success of each of the two regeneration mechanisms, and thus the postfire regeneration strategy: obligate resprouters, facultative species and obligate seeders (Fig. 2). Specifically we propose the following three hypotheses: 1) resprouting appeared early in plant evolution as a response to disturbance, and fire was an important driver in many lineages; 2) postfire seeding evolved under conditions where fires were predictable within the life span of the dominant plants and created conditions unfavorable for resprouting; and 3) the intensification of conditions favoring juvenile survival (C) and adult mortality (P) drove the loss of resprouting ability with the consequence of obligate-seeding species becoming entirely dependent on fire to complete their life cycle, with one generation per fire interval (monopyric life cyle). This approach provides a framework for understanding temporal and spatial variation in resprouting and seeding under crown-fire regimes. It accounts for patterns of coexistence and environmental changes that contribute to the evolution of seeding from resprouting ancestors. In this review, we also provide definitions and details of the main concepts used in evolutionary fire ecology: postfire regeneration traits, postfire strategies, life cycle in relation to fire, fire regimes (Box 1), costs of resprouting (Box 2), postfire seeding mechanisms (Box 3), and the possible evolutionary transitions (Box 4).

 

Fig2_sm
Fig. 1 : Main factors affecting adult and offspring seedling survival (P and C, respectively), and thus the P/C ratio, in fire-prone ecosystems (from Pausas & Keeley 2014 [2]).

 

Fig3_sm

Fig. 2: The changes in the probability of resprouting along an adult-to-offspring survival (P/C) gradient are not linear but show two turning points related to the acquisition of key innovations: the capacity to store a fire-resistant seed bank (postfire seeding), and the loss of resprouting capacity. Changes in P/C ratio may be produced by different drivers (Fig. 1) which drove the rise of innovations during evolution, e.g., during the increasing aridity from the Tertiary to the Quaternary (from Pausas & Keeley 2014 [2]).

 

Refecences

[1] Pausas, J.G., Bradstock, R.A., Keith, D.A., Keeley, J.E. 2004. Plant functional traits in relation to fire in crown-fire ecosystems. Ecology 85: 1085-1100. [doi | pdf | esa | jstor]

[2] Pausas J.G. & Keeley J.E. 2014. Evolutionary ecology of resprouting and seeding in fire-prone ecosystems. New Phytologist 204: 55-65 [doi | wiley | pdf]

 

Fire drives trait divergence: smoke-induced germination

April 3rd, 2014 No comments

There is an increasing evidence that recurrent fires are driving within species phenotypic variability, and that different fire regimes can generate trait divergence among populations [1]. For instance, populations of the annual species Helenium aromaticum (Asteraceae) growing under different fire histories in Chile have different seed traits in such a way that the anthropogenic increase in fire frequency selected for an increasing in seed pubescence [2]. In the Mediterranean Basin there is also evidence of phenotypic divergence among populations under different fire regimes: Ulex parviflorus (Fabaceae) plants living under high fire frequency are more flammable than those growing in sites that have not suffered fires [3-5]; Pinus halepensis and P. pinaster living under high crown-fire frequency have higher serotiny that those living in areas that rarely burn in crown fires [6]

A recent paper add further examples of this fire-driven trait divergence: Vandvik et al. show that smoke-induced germination is observed in populations of Calluna vulgaris (Ericaceae) from traditionally burnt coastal heathlands of Norway but it is lacking in populations from other habitats with infrequent fires [7]. The results are also consistent with the suggestion that smoke-induced germination is a fire adaptation [8-9].

Calluna-smoke-germination

Figure: Probability of germination of Calluna vulgaris in relation to time (days) since sowing for smoke-treated (pink) and control (grey) seeds, in coastal and inland heathlands of Norway. From Vandvik et al. 2014 [7].

References:

[1] Pausas, J. G. and D. W. Schwilk. 2012. Fire and plant evolution. New Phytologist 193 (2). [doi | pdf | blog]

[2] Gómez-González S, Torres-Díaz C, Bustos-Schindler C, Gianoli E, 2011. Anthropogenic fire drives the evolution of seed traits. PNAS 108: 18743-18747. [doi blog]

[3] Pausas J.G., Alessio G., Moreira B. & Corcobado G. 2012. Fires enhance flammability in Ulex parviflorusNew Phytologist 193: 18-23. [doi | pdf | blog]

[4] Pausas J.G. & Moreira B. 2012. Flammability as a biological concept. New Phytologist 194: 610-613. [doi | wiley | pdf]

[5] Moreira B., Castellanos M.C., Pausas J.G. 2014. Genetic component of flammability variation in a Mediterranean shrub. Molecular Ecology 23: 1213-1223. [doi | pdf | suppl. | data:dryad | blog]

[6] Hernández-Serrano A., Verdú M., González-Martínez S.C., Pausas J.G. 2013. Fire structures pine serotiny at different scales. American Journal of Botany 100 (12): 2349-2356. [doi | amjbot | pdf | supp. | blog]

[7] Vandvik, V., J. P. Töpper, Z. Cook, M. I. Daws, E. Heegaard, I. E. Måren, and L. G. Velle. 2014. Management-driven evolution in a domesticated ecosystem. Biology Letters 10 (2): 20131082. [doi]

[8] Pausas J.G. & Keeley J.E. 2009. A burning story: The role of fire in the history of life. BioScience 59: 593-601 [doi | jstor | BioOne | pdf | scribd | ppt slides | post]

[9] 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 in Plant Science 16(8): 406-411. [doi | trends | pdf]

 

Ulex born to burn (II): genetic basis of plant flammability

January 25th, 2014 No comments

In an previous study we found that Ulex parviflorus (Fabaceae) populations that inhabit in recurrently burn areas (HiFi populations) were more flammable than populations of this species growing in old-fields where the recruitment was independent of fire (NoFi populations) [1,2, 3]. That is, HiFi plants ignited quicker, burn slower, released more heat and had higher bulk density than NoFi plants. Thus, it appeared that repeated fires selected for individuals with higher flammability, and thus driving trait divergence among populations living in different fire regimes. These results were based on the study of plant flammability (phenotypic variability) without knowing whether plant flammability was genetically controlled. In a recent study using the same individuals [4], we show that phenotypic variability in flammability was correlated to genetic variability (estimated using AFLP loci) [figure below]. This result provide the first field evidence supporting that traits enhancing plant flammability have a genetic component and thus can be responding to natural selection driven by fire [5]. These results highlight the importance of flammability as an adaptive trait in fire-prone ecosystems.

Ulex-flam-AFLP

Figure: Relationship between flammability and genotypic variability at individual level in Ulex parviflorus (red symbols: individuals in HiFi populations; green symbols: individuals in NoFi populations). Variations in flammability are described using the first axis of a Principal Component Analysis (PCA1) performed from different flammability traits, and genetic variability is described using the first axis of a Principal Coordinate Analysis (PCo1) from the set of AFPL loci that were significantly related to flammability. See details in [4].

References
[1] Ulex born to burn, jgpausas.blogs.uv.es, 9/Nov/2011

[2] Pausas J.G., Alessio G., Moreira B., Corcobado G. 2012. Fires enhance flammability in Ulex parviflorusNew Phytologist 193:18-23 [doi | wiley | pdf]

[3] Pausas J.G. & Moreira B. 2012. Flammability as a biological concept. New Phytologist 194: 610-613.  [doi | wiley | pdf]

[4] Moreira B., Castellanos M.C., Pausas J.G. 2014. Genetic component of flammability variation in a Mediterranean shrub. Molecular Ecology 23: 1213-1223 [doi | pdf | data:dryad]

[5] 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 in Plant Science 16: 406-411. [doi | trends | pdf]

 

Serotiny

November 16th, 2013 No comments

Serotiny is the delayed seed release for more than a year by retaining the seeds in a woody structure [1]. This implies an accumulation of a canopy seed bank. Serotiny confer fitness benefits in environments with frequent crown-fires, as the heat opens the cones and seeds are dispersed in the post-fire bed which is rich in resource and the competition and predation are low. It is typical of many Proteaceae and some conifers, like some pine species [1, 2; figure below].

Two recent papers analyse the serotiny of two mediterranean pines Pinus halepensis and Pinus pinaster [3, 4]. P. halepensis show higher proportion of serotinous cones than P. pinaster, but the latter retain the cones for longer [3]. The two species show high variability of serotiny within and between populations, but they show a clear pattern of higher serotiny in populations subject to high frequency of crown-fires than those living in areas where crown-fires are rare or absent. This is true either considering serotiny as the proportion of serotinous cones or as the age of the cones stored. Compared with other pines worldwide, the strength of the fire-serotiny relationship in P. pinaster is intermediate, and in P. halepensis is among the highest known [3]. For P. halepensis (the species with higher % serotiny), populations in high fire recurrence regimes have higher fine-scale spatial aggregation of serotiny than those inhabiting low fire recurrence systems. This phenotypic spatial structure generated by fire could be a consequence of the spatial genetic structure of the population. The second study used genomic tools to search for a genetic association for serotiny [4]. The analysis of 384 SNPs of 199 individuals of P. pinaster (in 3 populations included in the previous study [3])  shows that 17 loci were associated with serotiny and explain all together ca. 29% of the serotiny variation found in the field. All these results adds further evidence to the emerging view that fire shapes intraspecific variability of traits and generates phenotypic divergence between populations [5, 6, 7].

Figure: Serotinous cones of Pinus pinaster (Foto: K.B. Budde)

References:

[1] 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]

[2] He T, Pausas JG, Belcher CM, Schwilk DW, Lamont BB. 2012. Fire-adapted traits of Pinus arose in the fiery Cretaceous. New Phytologist 194: 751-759. [doi | wiley | pdf (suppl.)]

[3] Hernández-Serrano A., Verdú M., González-Martínez S.C., Pausas J.G. 2013. Fire structures pine serotiny at different scales. American Journal of Botany 100: 2349-2356 [doi | amjbot | pdf | supp.]

[4] Budde, K. B., Heuertz, M., Hernández-Serrano, A., Pausas, J.G., Vendramin, G.G., Verdú, M. & González-Martínez, S.C. 2014. In situ genetic association for serotiny, a fire-related trait, in Mediterranean maritime pine (Pinus pinaster Aiton). New Phytologist  201: 230-241 [doi | pdf]

[5] 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 in Plant Science 16(8): 406-411. [doi] [trends] [pdf]

[6] Pausas, J. G., Schwilk, D. W. 2012. Fire and plant evolution. New Phytologist, 193:301-303. [doi | wiley | pdf]

[7] Pausas J.G., Alessio G., Moreira B. & Corcobado G. 2012. Fires enhance flammability in Ulex parviflorusNew Phytologist 193: 18-23. [doi | wiley | pdf]

 

Physiological differences between resprouters and seeders

November 9th, 2013 No comments

The ability to resprout and to recruit after fire are two extremely important traits for the persistence in fire-prone ecosystems [1,2], and they define three life histories: obligate resprouters, obligate seeders (non-resprouters), and facultative seeders. After a fire, obligate seeders die and recruit profusely from the seeds stored in the seed bank [3-5]. In contrast, resprouters survive after fire and their above-ground tissues regenerate from protected (often below-ground) buds by using stored carbohydrates [6]. Facultative seeders not only recruit profusely after fire, but are also able to resprout. In fact, seeders and resprouters have different regeneration niches: seedling regeneration of obligate resprouters is not linked to fire, and they recruit during the inter-fire period under sheltered conditions (i.e., under vegetation cover), while seedling regeneration of seeders occurs in open postfire environments. Given the marked difference in water availability between microsites under vegetation and microsites open to the sun under Mediterranean conditions, seedlings of resprouters and seeders are subjected to different water-stress conditions, and thus they are expected to have different physiological attributes. Despite these differences, resprouters and seeders co-exist, are often well-mixed on local and landscape scales [7,8], and represent the two main types of post-fire regeneration strategies in Mediterranean ecosystems [2].

A recent study demonstrates marked differences in physiological attributes between seedlings of seeders and resprouters [9]: Seeders show a range of physiological traits that better deal with water-limited and highly variable conditions (e.g., higher resistance to xylem cavitation, earlier stomatal closure with drought, higher leaf dehydration tolerance), but they are also capable of taking full advantage of periods with high water availability (greater efficiency in conducting water through the xylem to to sustain high gas exchange rates when water is available). Conversely, resprouter species are adapted to more stable water availability conditions, favoured by their deeper root system, but they also display traits that help them resist water shortages during long summers.

Previous studies already showed marked differences between seeders and resprouters in a range of attributes: resprouters tend to exhibit a deeper root-system, while seedling root structure of seeders are more efficient in exploring the upper soil layer [10]. Leaves of seeders show higher water use efficiency (WUE) and higher leaf mass per area (LMA; i.e., higher sclerophylly, lower SLA) [11]. Seeds of seeder species are more tolerant to heat shocks and have greater heat-stimulated germination [3]. All these differences support the idea that they are distinct syndromes with different functioning characteristics at the whole plant level and suggest that they undertook different evolutionary pathways [12].

Figure: Coexistence of resprouters (R+) and seeders (R-) in postfire conditions near Valencia, Spain. (Foto: A. Vilagrosa).

 

References:

[1] Pausas, J.G., Bradstock, R.A., Keith, D.A., Keeley, J.E. & GCTE Fire Network. 2004. Plant functional traits in relation to fire in crown-fire ecosystems. Ecology 85: 1085-1100. [jstor |[pdf | Ecological Archives E085-029]

[2] 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]

[3] Paula S. & Pausas J.G. 2008. Burning seeds: Germinative response to heat treatments in relation to resprouting ability. Journal of Ecology 96 (3): 543 – 552. [doi | pdf]

[4] Moreira B., Tormo J., Estrelles E., Pausas J.G. 2010. Disentangling the role of heat and smoke as germination cues in Mediterranean Basin flora. Annals of Botany 105: 627-635. [doi | pdf | blog]

[5] Moreira B. & Pausas J.G. 2012. Tanned or burned: The role of fire in shaping physical seed dormancy. PLoS ONE 7(12): e51523. [doi | plos | pdf | blog]

[6] Moreira B., Tormo J, Pausas J.G. 2012. To resprout or not to resprout: factors driving intraspecific variability in resprouting. Oikos 121: 1577-1584. [doi | pdf]

[7] Verdú M, & Pausas JG 2007. Fire drives phylogenetic clustering in Mediterranean Basin woody plant communities Journal of Ecology 95 (6), 1316-323 [doi | pdf]

[8] Ojeda, F., Pausas, J.G., Verdú, M. 2010. Soil shapes community structure through fire. Oecologia 163:729-735. [doi | pdf | blog]

[9] Vilagrosa A., Hernández E.I., Luis V.C., Cochard H., Pausas, J.G. 2014. Physiological differences explain the co-existence of different regeneration strategies in Mediterranean ecosystems. New Phytologist 201 : xx-xx [doi | pdf | suppl.] – NEW

[10] Paula S. & Pausas J.G. 2011. Root traits explain different foraging strategies between resprouting life histories. Oecologia 165:321-331. [doi | pdf | blog]

[11] Paula S. & Pausas J.G. 2006. Leaf traits and resprouting ability in the Mediterranean basin. Functional Ecology 20: 941-947. [doi | pdf | blog]

[12] Verdú M. & Pausas J.G. 2013. Syndrome-driven diversification in a Mediterranean ecosystem. Evolution 67: 1756-1766. [doi | pdf | blog]

 

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].

References:
[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.

 

Fire-stimulated flowering

May 25th, 2013 No comments

Some plant species flower profusely and quickly after fire (fire-stimulated flowering). Compared with resprouting or postfire seeding, this trait is relatively unknown outside of South Africa and Australia [1, 2]. It is considered one of the adaptations of some resprouting species to live in recurrently burn environments. There are some of these species that rarely flower without a fire (obligate postfire flowering) while others can flower in the absence of fire but they produce more flowers after it (facultative postfire flowering). One example I had the chance to observe recently in Central America is Bulbostylis paradoxa (Cyperaceae; Figure below); it is a very flammable plant that grow in savannas and dry forest of Central/South America and the Caribbean. Local foresters told me that they have never seen this species flowering in absence of fire, and that they start flowering next day after the fire.


Figure: Bulbostylis paradoxa (Cyperaceae) one month after a fire in Santa Rosa National Park, Costa Rica (fotos: J.G. Pausas, May 2013).

References:
[1] Bytebier B., Antonelli A., Bellstedt D.U., Linder H. P. 2011. Estimating the age of fire in the Cape flora of South Africa from an orchid phylogeny. Proc. R. Soc. B, 278: 188-195.

[2] Lamont B.B., Downes K.S. 2011. Fire-stimulated flowering among resprouters and geophytes in Australia and South Africa. Plant Ecol. 212: 2111-2125.

 

Fire shapes savanna-forest mosaics in the Brazilian cerrado

May 14th, 2013 No comments

Cerrado is the name of a tropical fire-prone mosaic of savanna and forest in Brazil. In a recent paper [1], we showed that in cerrado landscapes, despite the existence of a great variety of community structure (from open savannas to closed forests; Figure below), there are two well-defined stable states of community function, each associated with contrasting levels of community closure (open and closed environments) and maintained by different fire regimes. Soil properties, phylogenetic and non-phylogenetic beta-diversities, and most of the plant functional traits presented a threshold pattern along the community closure gradient with coinciding breakpoints, providing strong evidence of a functional threshold along the forest-savanna gradient. Open environments consisted of communities growing on poor soil and dominated by short species with early investments in thick barks, low wood density and with thick and tough leaves (high toughness and low specific area). In contrast, closed communities grow in more fertile soils and include plants having the opposite functional attributes. Moreover, we found contrasting fire regimes on the two sides of the threshold, with open formations showing shorter fire intervals than closed formations and a switch from communities dominated by fire-resistant plants to communities dominated by shade tolerant species that compensate for their lack of fire resistance by efficiently closing the canopy (i.e., reducing flammability). Overall, these results are consistent with the theoretical model of fire-plant feedbacks as main drivers of the coexistence of two stable states, savanna and forest. In this context, we provide the first field-based evidence for a community-level threshold separating two vegetation states with distinct functional and phylogenetic characteristics and associated with different fire regimes.

Top: A woodland cerrado (cerrado sensu stricto) six months after a fire, with several top-killed trees and a developed layer of resprouting vegetation; and one of the sampled closed forests.
Middle: A dense woodland cerrado (cerrado denso); one example of a typical thick-barked species found in open communities (Anadenanthera peregrina (Benth.) Reis, Fabaceae); a transitional zone between dense savannas and forests.
Bottom: A typical open savanna at the early rainy season, with tall flammable grasses and small trees and shrubs.
Photo credits: V. Dantas, G. Sartori, V. Cadry, J.G. Pausas, F. Noronha, A. Favari. See [1].

References

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

 

Mediterranean diversification and plant syndromes

January 21st, 2013 No comments

Woody plants of the Mediterranean Basin can be classified in two contrasted morpho-functional syndromes [1]: a) plants with sclerophyllous, evergreen leaves and small, unisexual greenish or brownish flowers with a reduced perianth, and large seeds dispersed by vertebrates; and b) plants with alternative character states (non-sclerophyllous deciduous, semi-deciduous or summer deciduous species with large and conspicuous flowers pollinated by insects, and small seeds). The sclerophyllous syndrorme (a) occurs in clades whose characteristics pre-date the appearance of the mediterranean climate while the non-sclerophyllous syndrome (b) arose in clades that have evolved after the emergence of this distinctive climate (Tertiary – Quaternary transition).

A recent phylogenetic study [2] show that during the time with prevalent mediterranean climate, lineages with the non-sclefophyllous syndrome showed a higher speciation rate than the sclerophyllous lineages, suggesting that a syndrome-driven local diversification has occurred in shrublands under mediterranean conditions. The processes behind this result might be divers, but fire might had an important role. The rise of mediterranean climate increased fire activity [3] and traits defining these two syndromes are related to post-fire regeneration traits and to the age to maturity [4,5]. The non-sclerophyllous syndrome is associated with species considered post-fire seeders (i.e., killed by fire in which populations regenerate from a persistent seed bank; fire-stimulated germination [6,7]) and to species with early maturation. In fire-prone ecosystems, these characteristics reduce the generation time and the overlap between generations and thus they provide more opportunities for diversification.

Overall, the results provide an example of how the integration of the environmental filter in a dated phylogeny may recreate the local history of lineages and help to explain assembly processes in mediterranean ecosystems.

Figure: Frequency distribution of differences in local speciation rate (λ) between non-sclerophyllous (n) and sclerophyllous (s) syndromes in the Valencia woody flora for 3 different post cut temporal slices (cutoff of 10, 6, and 3.6 My) related to the increasing aridity associated with the rise of mediterranean climate. For all alternative phylogenies (i.e., accounting for the undertainity in node age), speciation rate of the non-sclerophyllous syndrome is greater than for the sclerophyllous one. See Verdú & Pausas (2013) for details [2].

References
[1] Herrera, CM. 1992. Historical effects and sorting processes as explanations for contemporary ecological patterns: character syndromes in Mediterranean woody plants. Am. Nat. 140:421-446.

[2] Verdú M. & Pausas J.G. 2013. Syndrome-driven diversification in a Mediterranean ecosystem. Evolution. [doi | pdf]

[3] Keeley JE., Bond WJ., Bradstock RA., Pausas JG. & Rundel PW. 2012. Fire in Mediterranean Ecosystems: Ecology, Evolution and Management. Cambridge University Press [the book]

[4] Pausas, J.G., Bradstock, R.A., Keith, D.A., Keeley, J.E. & GCTE Fire Network. 2004. Plant functional traits in relation to fire in crown-fire ecosystems. Ecology 85: 1085-1100. [pdf | jstor]

[5] Pausas J.G. & Verdú M. 2005. Plant persistence traits in fire-prone ecosystems of the Mediterranean Basin: A phylogenetic approach. Oikos 109: 196-202. [doi| pdf]

[6] Moreira B., Tormo J., Estrelles E., Pausas J.G. 2010. Disentangling the role of heat and smoke as germination cues in Mediterranean Basin flora. Annals of Botany 105: 627-635. [doi| pdf]

[7] Moreira B. & Pausas J.G. 2012. Tanned or burned: The role of fire in shaping physical seed dormancy. PLoS ONE 7(12): e51523.  [doi | plos | pdf]

 

Seed dormancy as a fire adaptation in Mediterranean ecosystems

December 6th, 2012 1 comment

Plant species with physical seed dormancy are common in mediterranean fire-prone ecosystems. Because fire breaks seed dormancy and enhances the recruitment of many species, this trait might be considered adaptive in fire-prone environments [1]. However, to what extent the temperature thresholds that break physical seed dormancy have been shaped by fire (i.e., for post-fire recruitment) or by summer temperatures in the bare soil (i.e., for recruitment in fire-independent gaps) remains unknown [1]. In a recent paper published in PLoS ONE [2], we tested these two alternatives in six woody species (21 populations) occurring in fire-prone areas across the Mediterranean Basin (Spain and Turkey). Seeds from different populations of each species were subject to heat treatments simulating fire (i.e., a single high temperature peak of 100ºC, 120ºC or 150ºC for 5 minutes) and heat treatments simulating summer (i.e., temperature fluctuations; 30 daily cycles of 3 hours at 31ºC, 4 hours at 43ºC, 3 hours at 33ºC and 14 hours at 18ºC).

The results showed that fire treatments broke dormancy and stimulated germination in all populations of all species. In contrast, summer treatments had no effect over the seed dormancy for most species and only enhanced the germination in Ulex parviflorus, although less than the fire treatments. That is, the results suggest that in Mediterranean species with physical dormancy, the temperature thresholds necessary to trigger seed germination are better explained as a response to fire than as a response to summer temperatures (see Figure below). The high level of dormancy release by the heat produced by fire might enforce most recruitment to be capitalized into a single post-fire pulse when the most favorable conditions occur. This supports the important role of fire in shaping seed traits [3]. Given that seed dormancy is heritable, demonstrating that it provides higher chances of recruitment (i.e., higher potential fitness benefits) in response to fire than in response to summer temperatures suggests the temperature threshold for breaking dormancy might be an adaptation to fire [1, 4].

Figure: Germination (%) in fire conditions (y axis) versus germination (%) in summer conditions (x axis) for 6 species (21 populations across the Mediterranean basin). Intraspecific variability (i.e., among populations) is indicated by small symbols (mean population value) emerging from the large symbol (mean species value). The 1:1 line is also shown (dotted line). Species considered are: Cistus albidus, Cistus creticus, Cistus parviflorus, Cistus salviifolius, Fumana thymifolia, and Ulex parviflorus.

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

[2] Moreira, B. and J. G. Pausas. in press. Tanned or burned: The role of fire in shaping physical seed dormancy. PLoS ONE 7(6): e39810. [doi | pdf]

[3] Moreira B., Tavsanoglu Ç., Pausas J.G. 2012. Local versus regional intraspecific variability in regeneration traits. Oecologia 168: 671-677. [doi | pdf]

[4] Pausas J.G. & Schwilk D.W. 2012. Fire and plant evolution. New Phytol., 193, 301-303. [doi | wiley | pdf]

Fire generates intraspecific trait variability in neotropical savannas

August 28th, 2012 No comments

“Cerrado” are neotropical savannas from Brazil. As in most savannas, fire is very frequent in cerrado, and fires has been occurring in these ecosystems during the last few millions years. Consequently, cerrado communities are strongly filtered by fire and are composed by species capable of succeed under frequent fires (e.g., resprouters, with very thick bark, etc). A recent study [1] comparing zones with different fire regimes (annual fires, biennial fires, and protected from fires) within the cerrado (in Emas National Park) suggests that most plant trait variability is found within species (intraspecific) and little trait variability is due to changes in species composition (interspecific) between fire regimes. Thus, at community scale, fire act more as an filter, preventing some of the species from outside cerrado to colonize the cerrado (e.g., from nearby non-flammable forests), than as an internal factor structuring species composition in the already filtered cerrado communities with different fire regimes. However, fire acts as an important factor generating intraspecific variability. These results support the hypothesis of the prominent importance of intraspecific variability in strongly fire-filtered communities [2,3].

Figure: The rhea (emas in Portuguese; Rhea americana) are a flightless birds that give the name to the Emas National Park (Parque Nacional das Emas), a World Natural Heritage site located in the Brazilian Central Plateau (Photo: JG Pausas, 2009, during the field sampling [1]).

References

[1] Dantas V.L., Pausas J.G., Batalha M.A., Loiola P.P. & Cianciaruso M.V. 2013. The role of fire in structuring trait variability in Neotropical savannas. Oecologia, 171: 487-494. [doi | pdf]

[2] Moreira B., Tavsanoglu Ç. & Pausas J.G. 2012. Local versus regional intraspecific variability in regeneration traits. Oecologia, 168, 671-677. [doi | pdf | post]

[3] Pausas J.G., Alessio G., Moreira B. & Corcobado G. 2012. Fires enhance flammability in Ulex parviflorusNew Phytologist 193: 18-23. [doi | wiley | pdf]

 

Fire and the evolution of pine life histories

August 15th, 2012 No comments

Many pines species are fire adapted. In 1998, JE Keeley & PH Zedler provided a seminal paper showing the various fire adaptations of pines, and the relation between the different adaptations and the different fire regimes [1]. Recent phylogenetic [2,3] and conceptual [4,5] advances in fire ecology have allowed to better understand the evolutionary role of fire in plants, and specifically in pines [2-6]. In a recent paper, JE Keeley provides a new review on the ecology and evolution of pine life histories [7]. Pinus originated ~150 Ma in the mid-Mesozoic Era and radiated across the northern continent of Laurasia during the Cretaceous period, when fire activity was high [3]. Pines have followed two evolutionary strategies interpreted as responses to competition by the newly emerging angiosperms: 1) The Strobus lineage mostly has radiated into stressful sites of low nutrient soils and extremes in cold or heat; ans 2) The Pinus (subgenus) lineage has radiated into fire-prone landscapes with diverse fire regimes. Based on the life history traits associated to fire, JE Keeley define four pine syndromes [7]: fire-avoiders (no fire-adapted; with thin bark), fire-toleraters (adapted to surface fires; with thick bark and self-pruning of dead branches; tall pines), fire-embracers (adapted to crown fires; with retention of dead branches and serotinous cones), and fire-refugia (with marked metapopulation dynamics) strategies.

Figure: Basal fire scar (a) and cross-section of pine with previous fires delineated (b) demonstrating fire survival after recurrent fires. Photos by JE Keeley from [7].

References
[1] Keeley J.E. & Zedler P.H. 1998. Evolution of life histories in Pinus. In: Ecology and biogeography of Pinus (ed. Richardson DM). Cambridge University Press Cambridge (UK), pp. 219-250.

[2] Schwilk D.W. & Ackerly D.D. 2001. Flammability and serotiny as strategies: correlated evolution in pines. Oikos, 94, 326-336. [doi]

[3] He T., Pausas J.G., Belcher C.M., Schwilk D.W. & Lamont B.B. 2012. Fire-adapted traits of Pinus arose in the fiery Cretaceous. New Phytol., 194, 751-759. [doi | wiley | pdf ]

[4] Pausas, J. G. and J. E. Keeley. 2009. A burning story: The role of fire in the history of life. Bioscience 59: 593-601. [doi | jstor | pdf]

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

[6] Pausas J.G. & Schwilk D.W. 2012. Fire and plant evolution. New Phytol., 193, 301-303.  [doi | wiley | pdf]

[7] Keeley J.E. 2012. Ecology and evolution of pine life histories. Ann. For. Sci., 69, 445–453. [doi]

 

New Book: Fire in Mediterranean Ecosystems

March 13th, 2012 No comments

Finally the new fire ecology book by Keeley et al. (2012) has been published:



For more information, table of contents, etc, see here.

Cambridge UP (ukusaau), Amazon (ukusajp), eBooks

The age of fire adaptations

February 20th, 2012 No comments

How old are wildfires? Probably as old as terrestrial ecosystems [1]. The origin of fire is tied to the origin of land plants, which are responsible for two of the three elements essential to the existence of fire: oxygen and fuel. The third element, a heat source, has probably been available throughout the history of the planet (mainly through lightning). There is charcoal evidencs of fires already in the Silurian (440 Ma). However, the existence of fire does not necessarily mean that fire was playing an evolutionary role at that time. So when did fire start to play an evolutionary role generating fire adaptations [2, 3]? By mapping fire adaptation onto a dated phylogeny of Pinaceae, we recently demonstrated [4] that at least, and for this family, fire was an agent of natural selection since about 90-125 Ma! This is far back from what was known until now [4]. At this time, fire-protective thick barks were originated in Pinus species as response to surface fires. With increasing fire intensity, thicker barks and serotiny appeared by 70-90 Ma. These innovations appear at the same time as the Earth’s paleoatmosphere experienced elevated oxygen levels that led to high burn probabilities (mid-Cretaceous). That is, the fiery environments of the Cretaceous strongly influenced trait evolution in Pinus. Whether fire had an evolutionary role prior to this is a challance for future research.

Fotos: In many pines, the thick bark and the discontinuity between the canopy and the understory (self-pruning) allows survival after surface fires (left: Pinus nigra, eastern Spain). Serotinous cones allow a quick seed regeneation after crown fire (right: P. halepensis, eastern Spain). Photos: JG Pausas

References
[1] Pausas, J. G. and J. E. Keeley. 2009. A burning story: The role of fire in the history of life. Bioscience 59: 593-601. [doijstor | BioOne | pdf]

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

[3] Pausas, J. G. and D. W. Schwilk. 2012. Fire and plant evolution. New Phytologist 193:301-303. [doiwileypdf]

[4] He T., Pausas J.G., Belcher C.M., Schwilk D.W., Lamont B.B. (2012). Fire-adapted traits of Pinus arose in the fiery Cretaceous. New Phytologist 194: 751-759 [doi | wileypdf]

Ulex born to burn

November 9th, 2011 No comments

Recurrent fires are a strong evolutionary pressure shaping plants [1,2]. It has been hypothesized that in fire prone-ecosystems, natural selection has favoured the development of traits that enhance flammability [3]. Consistent with this idea, in a recent study [4] we found that Ulex parviflorus (Fabaceae) populations that inhabit in recurrently burn areas (HiFi populations) are more flammable than populations of this species growing in old-fields where the recruitment was independent of fire (NoFi populations). That is, HiFi plants ignite quicker, burn slower, release more heat and have higher bulk density than NoFi plants. Thus, it appears that repeated fires select for individuals with higher flammability, and thus driving trait divergence among populations living in different fire regimes. These results provide some field support for the ‘kill thy neighbour’ hypothesis [3], but they also highlighted the need for heritability studies to unambiguously demonstrate natural selection driven by fire. This study together with other studies recently commented in this blog [5, 6] are placing flammability as a fundamental trait in plant evolution.

Figure: Flammability experiments using an epiradiator [4].

References

[1] Keeley, J. E., J. G. Pausas, P. W. Rundel, W. J. Bond, and R. A. Bradstock. 2011. Fire as an evolutionary pressure shaping plant traits. Trends in Plant Science 16:406-411. [doi | pdf]

[2] Pausas J.G. & Keeley J.E. 2009. A burning story: The role of fire in the history of life. BioScience [doi | jstore | pdf]

[3] Bond, W. J. and J. J. Midgley. 1995. Kill thy neighbour: an individualistic argument for the evolution of flammability. Oikos 73:79-85.

[4] Pausas J.G., Alessio G., Moreira B., Corcobado G. 2012. Fires enhance flammability in Ulex parviflorus. New Phytologist 193:18-23 [doi | pdf]

[4′] Pausas J.G. & Moreira B. 2012. Flammability as a biological concept. New Phytologist 194: 610-613. [doipdf]

[5] Pausas JG. 2011. Australia born-to-burn: a phylogenetic approach. jgpausas.blogs.uv.es, 18/March/2011 [link]

[6] Pausas JG. 2011. Fire and evolution: Cretaceous fires and the spread of angiosperms. jgpausas.blogs.uv.es, 9/Sept/2011 [link]

Differences between resprouters and non-resprouters

October 1st, 2011 No comments

Resprouting is a very important process in plants living in disturbance-prone ecosystems, and the December issue of the journal Plant Ecology is going to be dedicated to this topic (Ecology of plant resprouting in fire-prone ecosystems). During the recent years, and starting from the PERSIST project, we have been comparing functional traits between resprouters and non-resprouters in Mediterranean fire-prone ecosystems, and the last comparison (physiological traits [5]), is included in this special issue. Resprouters and non-resprouters are two plant syndromes in Mediterranean ecosystems that also differ in their evolutionary history [1]. Resprouters tend to exhibit a deeper root-system than non-resprouters that inverse less resources on roots. So one could think that resprouters are better adapted to drought. However, both resprouters and non resprouters coexist, and non-resprouters counteract their lower root allocation by different traits that confer higher drought resistance [2]. Non-resprouters have higher drought resistance at leave level because they have higher water use efficiency (WUE) and higher leaf mass per area (LMA; i.e., higher sclerophylly, lower SLA) [3]. The seedling root structure of non-resprouters also allows them to more efficiently explore the upper soil layer [4]. A recent paper also shows that, when water is non-limiting, non-resprouters showed a better performance of leaf gas exchange traits (higher assimilation, stomatal conductance and transpiration) than resprouters [5]; that is non-resprouters have higher efficiency in resource capture, and thus a better capacity to take advantage of water when it is freely available. In addition, resprouters and non-resprouters also differ in their post-fire germination, as non-resprouters tend to have a greater capacity to both (i) persist after fire by means of recruiting (greater heat-tolerance) and (ii) increase their population after fire (greater heat-stimulated germination), than resprouters [4]. All these results suggest that resprouters and non-resprouters are two contrasted syndromes or functional types in the Mediterranean Basin [6].

Figure: Arbutus unedo resprouting after a fire.

References:

[1] Pausas J.G. & Verdú M. 2005. Plant persistence traits in fire-prone ecosystems of the Mediterranean Basin: A phylogenetic approach. Oikos 109: 196-202. [pdf |doi]

[2] Pausas J.G. 2010. Fire, drought, resprouting: leaf and root traits. URL: jgpausas.blogs.uv.es, 22/Oct/2010.

[3] Paula S. & Pausas J.G. 2006. Leaf traits and resprouting ability in the Mediterranean basin. Functional Ecology 20: 941-947. [pdf | [doi]

[4] Paula S. & Pausas J.G. 2011. Root traits explain different foraging strategies between resprouting life histories. Oecologia 165:321-331. [doipdfblog]

[5] Hernández E.I., Pausas J.G. & Vilagrosa A. 2011. Leaf physiological traits in relation to resprouter ability in the Mediterranean Basin. Plant Ecology 212:1959-1966 [doi| pdf]

[4] Paula S. & Pausas J.G. 2008. Burning seeds: Germinative response to heat treatments in relation to resprouting ability. Journal of Ecology 96 (3): 543 – 552. [pdf | doi]

[6] Pausas, J.G., Bradstock, R.A., Keith, D.A., Keeley, J.E. & GCTE Fire Network. 2004. Plant functional traits in relation to fire in crown-fire ecosystems. Ecology 85: 1085-1100. [pdfjstor] [Ecological Archives E085-029]

Intraspecific plant variability and the spatial scale

September 24th, 2011 No comments

Variability is a fundamental characteristic of life and the raw material for natural selection, driving speciation and diversification processes. Traditional biogeographical theory would predict that plants in populations that are close each other (e.g., few km) should be more similar among them, than plants in distant populations (e.g., 100s or 1000s km). This is because biogeographical processes such as migration, glacial/interglacial climatic fluctuations and isolation should cause distant plant populations to diverge, and thus enhance intraspecific variability at large scales, while gene flow through close populations should reduce divergences. In contrast, in a recent paper we suggest that in fire prone-ecosystems, where fire may generate local heterogeneity, local variability in traits related to regeneration are quite large, overriding the variability at the larger scale [1]. Studying post-fire regeneration traits in Cistus salviifolius and Lavandula stoechas, in eastern Iberia (IB, Spain) and in south-western Anatolia (AN, Turkey), we found that the trait variability within each region is larger than between regions (separated by about 2600 km, with the sea in the middle). The traits studied were seed size, seed dormancy and germination stimulation by head and by smoke. The two studied species exhibited germination stimulated by the fire-related cues; and independently of the region, the different populations of each species had a similar pattern of response. That is, Cistus salviifolius was stimulated by heat and Lavandula stoechas was mainly stimulated by smoke, although heat also exhibited a positive effect on the latter species (see also [2] for more details on heat- and smoke- stimulated germination). All these results supports the prominent role of fire as an ecological and evolutionary process across the Mediterranean Basin, producing trait variability and shaping biodiversity [3, 4].

References

[1] Moreira B., Tavsanoglu Ç., Pausas J.G. 2012. Local vs regional intraspecific variability in regeneration traits. Oecologia 168: 671-677 [doi | pdf]

[2] Moreira B., Tormo J., Estrelles E., Pausas J.G. 2010. Disentangling the role of heat and smoke as germination cues in Mediterranean Basin flora. Annals of Botany 105: 627-635.[pdf| doiblog]

[3] Pausas J.G. & Keeley J.E. 2009. A burning story: The role of fire in the history of life. BioScience 59: 593-601 [doi | pdfpost]

[4] 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 in Plant Science 16(8): 406-411. [doi | pdf | For managers]

Plant traits databases

July 23rd, 2011 No comments

Some time ago we complied an extensive database on plant traits for species of the Meditrranean Basin (the BROT database [1]). Simulatneously, there were other researches compiling similar databases for other regions. Now we have put all these databases together and obtained the world largest database on plant traits (the TRY database [2]). TRY combines 93 databases from 198 scientists, to produce a global database that contains 3 million trait records for about 69000 plant species. This compilation makes global trait data available to the scientific community and it is really stimulating research; currently there are about 50 research projects using the data.

References
[1] Paula S, Arianoutsou M, Kazanis D, Tavsanoglu Ç, Lloret F, Buhk C, Ojeda F, Luna B, Moreno JM, Rodrigo A, Espelta JM, Palacio S, Fernández-Santos B, Fernandes PM, and Pausas JG. 2009. Fire-related traits for plant species of the Mediterranean Basin. Ecology 90: 1420. [doi | BROT web]

[2] Kattge et al. (2011) TRY – a global database of plant traits. Global Change Biology  17: 2905–2935. [doi | TRY web]

Fire and plants: adaptations and exaptations

May 18th, 2011 No comments

There are still people believing that wildfires are a catastrophic disturbance to ecosystems, and that are the product of humans. However there is an increasing evidence from paleoecological records and from phylogenetic analyzes suggesting that fire is a very old process in the history of life, dating back to the origin of land plants [1, 2, 3]. As a consequence many plants have evolved in the presence of recurrent wildfires and acquired adaptive traits to persist and reproduce in those conditions. Examples of these traits are the resprouting ability, germination by head or smoke, and serotiny; all of these confer fitness advantage in fire-prone ecosystems. However, plants are not adapted to fire per se but to fire regimes. Species that exhibit traits adaptive under a particular fire regime can be threatened when that regime changes, like the recent human-induced fire regime changes (e.g., increasing or decreasing fire frequency or severity in relation to the historic fire regime).

In a recent paper, Keeley et al. [4] proposed five scenarios of change in a trait state (Figure 1). An adaptive trait might not change through time regardless of the selective environment (scenarios 1 and 2 in Figure 1). Such traits cannot be described as adaptations to the current selective (fire-prone) environment as there is no evidence that natural selection shaped this trait. Other adaptive traits that were shaped by natural selection under a previous evolutionary pressure, but not under the current (fire-prone) environment (scenario 3 in Figure 1) would be adaptations to previous evolutionary pressures and exaptations to the current (fire) environment [4, 5]. Fire adaptations are those adaptive traits in which natural selection is acting under the current fire-prone environment to shape the trait, and it is independent of how long this pressure has been present (scenarios 4 and 5 in Figure 1). For instance, there are clear examples of lineages that resprout after fire, but their origin and evolution is hardly liked to fire. However the most plausible scenario of lineages that resprouting from lignotubers is the number 4 in Fig. 1 (old origin of resprouting reshaped by current recurrent fires). Similarly serotiny and thick barks are traits that has been reshaped by natural selection under the framework of recurrent fires and thus they also fit under the concept of adaptation to fire (scenario 4 or 5 in Fig. 1).

References
[1] Pausas J.G. & Keeley J.E. 2009. A Burning Story: The role of fire in the history of life. BioScience 59: 593-601. [doi pdfpostslides]

[2] Pausas J.G. 2011. Australia born to burn – phylogenetic evidences. URL: jgpausas.blogs.uv.es, 18/03/2011.

[3] Pausas J.G. 2010. Fire and evolution: Cretaceous fires and the spread of angiosperms. URL: jgpausas.blogs.uv.es, 9/Sep/2010.

[4] 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 in Plant Science 16: 406-411 [doi | trends | pdf]

[5] Endler J.A. (1986) Natural selection in the wild. Princeton University Press.

Figure. 1. Five possible evolutionary scenarios of change in a trait state along the evolutionary time (simplified). For each scenario, different line types are periods under different dominant evolutionary pressures (e.g., the continuous line represents a period in which fire acted as an evolutionary pressure; dashed line the period with a different previous selective environment). The scenarios are:
1) and 2) No change along the time axis and no sign of natural selection (no adaptation to fire);
3) trait shaped during the first evolutionary pressure, but no change (with persistence of the state of the trait) during the second evolutionary pressure; natural selection acted during the first period only (no adaptation to fire, but exaptation);
4) and 5) Trait shaped during the whole period; natural selection acts during the whole period even if the dominant evolutionary pressure changed (adaptation to fire).
From Keeley et al. (2011, [4]).

Australia born to burn – phylogenetic evidence

March 18th, 2011 1 comment

Traditionally wildfires were considered a disturbance linked to the recent history of the Quaternary, and specially linked to the humans. However, evidence are accumulating about the ancient role of wildfires in terrestrial ecosystems [1]. In Australia, the flammable continent, the current believe is that fires started to be important during the onset of seasonal aridity in the Miocene (25 Ma). However, two recent and independent papers demonstrate, using phylogenetic techniques, that fire-dependent traits appeared about 60 Ma ago (early Paleocene), implying that fire was already an effective agent of selection by then. Crisp et al [2] studied the Myrtaceae family and showed that post-fire epicormic resprouting (typical of many eucalypts) is an ancient trait linked to the flammable sclerophyll biomes originated about 60-62 Ma. He et al. [3] studied the Banksia genus (Proteaceae) and showed that serotiny (fire dependent dispersal; figure below) and dead floret retention around the cones (enhanced flammability around serotinous cones) co-originate with the first appearance of Banksia 60.8 Ma ago. The coincidence of the two independent papers, using two different taxa (Myrtaceae and Banksia) is amazing, and clearly suggests that fire was a selective force in Australia during the Paleocene.  These papers are part of the accumulating research on the prominent and ancient role of fire in shaping plant species and biodiversity [1, 4 ].

References
[1] Pausas J.G. & Keeley J.E. 2009. A Burning Story: The role of fire in the history of life. BioScience 59: 593-601. [doipdfpost slides]

[2] Crisp MD, Burrows GE, Cook LG, Thornhill AH, Bowman DMJS. 2011. Flammable biomes dominated by eucalypts originated at the Cretaceous-Palaeogene boundary. Nature Communications 2: 193. [doi]

[3] He T, Lamont BB, Downes KS. 2011. Banksia born to burn. New Phytol. [doi]

[4] Bond, W. J. and Scott, A. C. 2010. Fire and the spread of flowering plants in the Cretaceous. New Phytol. 188: 1137–1150 [post]

Figure: Banksia cone opened by the fire to release seeds (serotiny).

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).

References:

[1] Pausas J.G. 2009. Convergent evolution. jgpausas.blogs.uv.es, 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.

Fire, drought, resprouting: leaf and root traits

October 22nd, 2010 No comments

Drought and fire are prevalent disturbances in Mediterranean ecosystems. Plant species able to regrow after severe disturbances (i.e. resprouter life history) have higher allocation to roots and higher water potential during the dry season than coexisting non-resprouting species. However, non-resprouters have higher survival rate after summer drought. We expect that, to counteract their shallow-rooting systems and to maximize seedling survival, non-resprouters have traits that confer higher water-use efficiency and higher efficiency in soil resource acquisition than resprouters.

Some time ago we tested this prediction in relation to leaf traits [1] and found that non-resprouters have higher leaf mass per area (LMA; i.e. lower specific leaf area, SLA), leaf dry matter content (LDMC), area-based leaf nitrogen content (LNCa) and integrated water-use efficiency (δ13C) than resprouters, suggesting that they have higher potential for structural resistance to drought and higher water-use efficiency than resprouters.

In a recent paper we have now tested the prediction for root traits in seedlings [2]. We found that non-resprouters have higher specific root length (SRL) and longer, thinner and more branched lateral roots, especially in the upper soil layers. The external links (i.e. the most absorptive root region) are also more abundant, longer, thinner and with higher SVR for non-resprouters. Thus seedling root structure of non-resprouters species allows them to explore more efficiently the upper soil layer, whereas seedling roots of resprouters will permit both carbon storage and deep soil penetration.

Whereas resprouters tend to maximize the surface and the efficiency of the organs for carbon uptake to ensure carbohydrate storage for resprouting (eg, higher SLA), non-resprouters maximize the root surface (eg, higher SRL), since their survival and growth may be limited by soil resources.

[1] Paula S. & Pausas J.G. 2006. Leaf traits and resprouting ability in the Mediterranean basin. Functional Ecology 20: 941-947. [doi | pdf]

[2] Paula S. & Pausas J.G. 2011. Root traits explain different foraging strategies between resprouting life histories. Oecologia 165:321-331 [doi | pdf]

Specific Root Length

Relationship between average root diameter and specific root length (SRL; log-scale) for resprouting (R+, closed symbols) and non-resprouting (R-, open symbols) species. Intraspecific variability is indicated by segments emerging from each symbol. From Paula & Pausas (Oecologia [2]).

Methods for community ecology

September 2nd, 2010 No comments

Why so many desert plant communities are dominated by spiny species, most of them cacti? The first observation refers to the community’s phenotypic structure and the second to its phylogenetic structure. What do these observations tell us about the mechanisms assembling these communities?

The way communities are assembled is an old ecological question currently experiencing renewed interest thanks to the recent advances in molecular biology and phylogenetics. The generality of these new methods has allowed us to understand the structure of communities of organisms from different kingdoms and at different scales. Concomitant with this growing interest, new methods, metrics, terms, and software have appeared that independently solve similar questions, but with different approaches. In this new paper we provide a unifying framework on methods for community structure based on the relationships between four key concepts: phylogeny, phenotype, environment, and co-occurrence. The different approaches are based on different community representations of traits, the phylogenetic relationships of species in the community, or species occurrence along the environmental gradients. We finally provide insights on future directions of this emerging discipline.

Pausas, J.G., Verdú, M. 2010. The jungle of methods for evaluating phenotypic and phylogenetic structure of communities. BioScience, 60: 614-625. [doi | pdf | slides]

Pausas-Verdu-BioScience

Figure 1. Methods for analyzing community structure can be represented in a simple framework in which the relationships (arrows) between the four key concepts (phylogeny, phenotype, environment, and co-occurrence) are integrated. The numbers in brackets refer to:

(1) Co-occurrence pattern versus the random expectation

(2) Phenotype-based approach:

(2.1) Relationship between the species’ phenotypes and their co-occurrence (phenotypic community structure).

(2.2) Relationship between species response to the environment and the species phenotypes, controlling by the species’ phylogenetic relatedness (phenotypic community structure).

(3) Phylogeny-based approach:

(3.1) Relationship between the species’ phylogenetic relationships and their co-occurrence (phylogenetic community structure).

(3.2) Relationship between the species’ phenotypes and their phylogenetic relationships (trait evolution).

(4) Environment-based approach: relationship between species response to the environment and the co-occurrence, considering the phylogenetic relatedness (phylogenetic community structure).

See previous post [link] on the effect of fire in phenotypic and phylogenetic structure of communities.

Soil shapes community structure through fire

January 21st, 2010 No comments

Recurrent wildfires constitute a major selecting force in shaping the structure of plant communities. At the regional scale, fire favours phenotypic and phylogenetic clustering in Mediterranean woody plant communities. Nevertheless, the incidence of fire within a fire-prone region may present strong variations at the local, landscape scale. This study tests the prediction that woody communities on acid, nutrient-poor soils should exhibit more pronounced phenotypic and phylogenetic clustering patterns than woody communities on fertile soils, as a consequence of their higher flammability and, hence, presumably higher propensity to recurrent fire. Results confirm the predictions and show that habitat filtering driven by fire may be detected even in local communities from an already fire-filtered regional flora. They also provide a new perspective from which to consider a preponderant role of fire as a key evolutionary force in acid, infertile Mediterranean heathlands.

Ojeda, F., Pausas, J.G., Verdú, M. 2010. Soil shapes community structure through fire. Oecologia 163:729-735. [doi]  [pdf]

IMG_0270_Ojeda en su brezal_sm The first author in the flammable, low fertility community.

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