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Posts Tagged ‘Mediterranean’

Pinus brutia

April 19th, 2017 No comments

Pinus halepensis is a strongly serotinous pine [1,2] occurring mainly in the western part of the Mediterranean Basin (especially in Spain; see map below). The most phylogenetically closely related species to P. halepensis is Pinus brutia that occurs in the eastern Mediterranean Basin (mainly in Turkey). P. brutia is called ‘red pine’ in Turkish (Kızıl çam) because sometimes the upper part of the trunk is reddish (as in P. sylvestris); the leaves are pale green as in P. halepensis. In relation to their fire response strategy [3], the main differences between the two species are that P. brutia is taller, the bark is thicker and the serotiny level is lower. Our observations suggest that P. brutia have relatively few serotinous cones, and most of then are less than 4 years old; P. halepensis have a higher proportion of serotinous cones, with many of them over 5 year old (and some more than 20 year old) [1].

Distribution maps of P. halepensis and P. brutia (from Wikipedia)

 
Pinus brutia forest, serotinous cones (2 serotinous and one non-serotinous (open)), and an example of a bark of more than 5 cm thick (the gaude was too short!). Photos from SW Turkey (by JG Pausas). For an example of serotinous cones in P. halepensis, see here.

 

References
[1] Hernandez-Serrano A., Verdú M., González-Martínez S.C., Pausas J.G. 2013. Fire structures pine serotiny at different scales. Am. J. Bot. 100: 2349-2356. [doi | pdf | supp.]

[2] Castellanos, M.C., González-Martínez, S. & Pausas, J.G. 2015. Field heritability of a plant adaptation to fire in heterogeneous landscapes. Mol. Ecol. 24, 5633-5642. [doi | pdf | suppl. | blog]

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

Eastern Mediterranean tour, 2017

April 18th, 2017 No comments

It is a pleasure to visit the different ecosystems of Lebanon and Turkey, and to meet old students and colleagues.

Cedar (Cedrus libani) forest, Al Shouf Biosphere Reserve, Lebanon

Anatolian steppe (bushes are Astragalus), central Turkey

Pinus brutia forest, SW Turkey

MEDECOS XIV

February 6th, 2017 1 comment

The XIV International Conference on Mediterranean Ecosystems (MEDECOS), has been successfully held on Sevilla, Spain, 31st Jan – 4th Feb, 2017, together with the conference of the Spanish Society of Terrestrial Ecology (AEET). For details, see the web page of the meeting and the post conference comments in the J. Ecol. blog. My contributions to this MEDECOS include two talks on fire and biotic interactions, two on resprouting, and one on fire hazard:

  • Pausas J.G – Fire and biotic interactions: the benefits of the disruption
  • García Y., Castellanos M.C., Pausas J.G. – Fires do not jeopardize reproduction of Chamaerops humilis despite disrupting its pollination
  • Tavsanoglu C. & Pausas J.G. – Resprouting ability encapsulates the most functional variability in the Mediterranean Basin flora
  • Paula, S. & Pausas J.G – Worldwide geographic and phylogenetic distribution of lignotubers
  • Cáceres M. de, Casals P., Álvarez D., Pausas J.G., Vayreda J., Beltrán M. – The role of understory fuel characteristics in the fire hazard of Mediterranean forests

The last day I attended to the field trip to Los Alcornocales Natural Park (a mosaic of cork oak forests and heathlands), where I enjoyed a long conversation on alternative vegetation states in non-tropical ecosystems (e.g., PDF) with William Bond (photo below).

Thank you very much to the organizers of MEDECOS, especially to Juan Arroyo (Universidad de Sevilla) and Montse Vilà (Doñana-CSIC) for this nice and smooth conference.

Bond-Pausas_sm

William Bond (left) and myself (right) in a Cork oak forest (photo: F. Ojeda)

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]

 

Flammable organic compounds: Rosmarinus officinalis

October 2nd, 2015 No comments

Given an ignition source and the right environmental conditions, all plants can potentially burn. However, some plants have characteristics that make them burn more easily. The capacity to store volatile organic compounds (VOCs) such as aromatic terpenes, can be considered one of these flammability-enhancing traits (flammable organic compounds, FOCs), as has now been demonstrated for Rosmarinus officinalis [1]: The more terpenes in the leaves, the more quickly they ignite (i.e., less time to ignition) (Figure below). Other species enhance flammability by having a very fine fuel, retaining dead fuel or having a flammable canopy structure [2-5]. There is growing evidence that flammability-enhancing traits are adaptive in Mediterranean fire-prone ecosystems [2-4]. To what extent the evolutionary pressure exerted by fire could have contributed to the abundance of aromatic plants in many fire-prone ecosystems (mints, rosemary, thyme, eucalypts, etc…) remains unknown. But certainly Mediterranean ecosystems are probably the most aromatic and among the most flammable ecosystems in the world.

Rosmarinus-flammability
Figure: relation between time to ignition (given a heat source, corrected by the differences in moisture) and the contents of terpenes (here the sum of camphene, para-cymene, borneol, limonene) in leaves of a wild population of rosmary (Rosmarinus officinalis), in Eastern Spain (from [1]). The top right corner shows the epiraditor, the device for  testing for time-to-ignition (see [2]).

References
[1] Pausas J.G., Alessio G.A., Moreira B., Segarra-Moragues J.G. (in press). Secondary compounds enhance flammability in a Mediterranean plant. Oecologia. [doi | pdf]

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

[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] [Ulex born to burn (II)]

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

 

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]

 

How plants survive the harsh environment of Australia

June 1st, 2015 No comments

New book: Groom, P. K., and Lamont, B. B. (2015). Plant Life of Southwestern Australia. Adaptations for Survival. De Gruyter Open

Early explorers described Western Australia as ‘the most barren spot on the face of the earth’. In this book we learn that south-western Australia is one of the world’s biodiversity hotspots – not despite but because of its harsh environment. Nutrient-poor soils, frequent droughts, and recurrent fires, together with adverse fauna interactions (e.g., strong-billed cockatoos, voracious kangaroos, and the lack of efficient pollinating bees and hummingbirds) have made this region the perfect evolutionary scenario for developing a plethora of plant adaptations and assembling an hyperdiverse flora. The authors nicely describe this scenario and offer an impressive wealth of knowledge on the natural history of the region in an attractive book with abundant tables and quality full-colour pictures. One of the strengths of the book is that it brings together both biotic and abiotic factors to explain biodiversity, something uncommon in most specialised books.

Overall this is a must-read book for Australian naturalists but will also be a key reference for international ecologists interested in how plants thrive and evolve in dry, nutrient-poor, fire-prone environments. The lessons learned from this region help us understand evolutionary pathways in other dry regions worldwide.

Groom-Lamont

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

 

Cultural trees: Cupressus sempervirens

January 4th, 2015 3 comments

 

Cupressus_sempervirens-Roma-01.2015_sm
Cupressus sempervirens (Mediterranean cypress; ciprés) and Colosseum, Rome,
1 January 2015 (photo: J.G. Pausas)

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]

 

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]

 

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]

 

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