Local survival after fire in Mediterranean shrublands: combining capture-recapture data over several bird species

Post-disturbance survival is a key factor in the onset of secondary successions. Here we analyse capture-recapture data from two before/after disturbance studies to estimate the effect of fire on local bird survival. Analyses of six bird species at two Mediterranean shrubland sites were combined using a meta-analysis approach. Two warblers, Sylvia undata and S. melanocephala, were studied at one site altered by prescribed burning, and five passerines (Luscinia megarhynchos, Turdus merula, Parus major, P. caeruleus and S. melanocephala) at one site disturbed by wildfire. Based on the combined analysis, annual survival probability significantly decreased from 0.49 to 0.18 (i.e. a 64% decline) after the fire. Our results further suggest a trend for a higher decrease in annual survival associated with wildfire (–72%, from 0.51 to 0.14) than with prescribed burning (–35%, from 0.41 to 0.27), although this should be properly tested with a specific experimental design. In S. undata, a decline in survival in the long-term cannot account for the drop in density observed the first spring after fire. We suggest that a decrease in recruitment rate and an increase in the proportion of non-breeders immediately after the fire may contribute more strongly to the decline in the breeding population. Our results tend to support the idea that bird populations may respond to moderate disturbances with noticeable time lags, because of individual site tenacity.     

Early root growth plasticity in seedlings of three Mediterranean woody species

Using a 141 F₂ population generated from maize inbred B64 × teosinte Zea nicaraguensis cross, quantitative trait loci (QTLs) controlling aerenchyma formation in roots under non-flooding drained soil conditions were identified. Seedlings of Z. nicaraguensis formed clear aerenchyma in the cortex of adventitious roots in non-flooding conditions, whereas the maize inbred line B64 did not. In the F₂ population, the capacity to develop aerenchyma exhibited wide and continuous variation, suggesting the trait was controlled by multiple genes. A linkage map was developed using 85 SSR markers, covering 1,224 cM across all ten chromosomes. Composite interval mapping analysis revealed that four QTLs for aerenchyma formation under non-flooding conditions were located to two regions of chromosome 1 (identified as Qaer1.02-3 and Qaer1.07), chromosome 5 (Qaer5.09) and chromosome 8 (Qaer8.06-7), and these explained 46.5% of the total phenotypic variance. The multiple interval mapping approach identified additional QTLs on chromosomes 1 (Qaer1.01) and 5 (Qaer5.01). Using these results, it may be possible to use SSR markers linked to aerenchyma formation in a marker assisted selection approach to introduce aerenchyma formation in drained soil conditions into maize for the eventual development of flooding tolerant maize hybrids.     

Functional traits predict drought performance and distribution of Mediterranean woody species

Water availability is one of the key environmental factors that affect plant establishment and distribution. In many regions water availability will decline with climate change, exposing small seedlings to a greater likelihood of drought. In this study, 17 leaves, stem, root, and whole-plant traits of ten woody Mediterranean species were measured under favourable growing conditions and seedling drought survival was evaluated during a simulated dry-down episode. The aims of this study were: i) to assess drought survival of different species, ii) to analyse which functional traits predict drought survival time, and iii) to explain species distribution in the field, based on species drought survival and drought strategies. Drought survival time varied ten-fold across species, from 19 to 192 days. Across species, drought survival was positively related to the rooting depth per leaf area, i.e., the ability to acquire water from deeper soil layers while reducing transpiring leaf area. Drought survival time was negatively related to species ability to grow quickly, as indicated by high relative growth and net assimilation rates. Drought survival also explained species distribution in the field. It was found that species were sorted along a continuum, ranging between two contrasting species functional extremes based on functional traits and drought performance. One extreme consisted of acquisitive fast-growing deciduous species, with thin, soft metabolically active leaves, with high resource use and vulnerability to drought. The opposite extreme consisted of conservative slow-growing evergreen species with sclerophyllous leaves, deep roots, a low transpiring area, and low water use, resulting in high drought survival and drought tolerance. The results show that these drought strategies shape species distribution in this Mediterranean area.     

Resprouting of Erica multiflora after experimental fire treatments

Abstract. The effect of fire intensity - both temperature and duration - on the resprouting pattern of the evergreen Mediterranean shrub Erica multiflora in relation to plant size, was experimentally investigated by subjecting plants to the flame of a propane torch, and observing mortality and resprouting after 5 and 20 months. Pre-treatment plant size was not important in determining post-fire plant survival, but it did influence the resprouting vigour, increasingly so with time. High temperatures induced higher mortality rates within populations, but duration of fire did not effect mortality. Biomass of resprouts was lower following more intense fire treatments, but this effect progressively disappeared over time, except in plants subjected to the most intense fire treatment. This is probably because of the increasing importance of the below-ground organs for the regeneration of the above-ground biomass. Some of the plants which were clipped but not exposed to fire also died 20 months after treatment. The effect of clipping onmortality andresprout-ing response, estimated as biomass of resprouts, was not significantly different from the effect induced by either low or medium temperature treatments, but was significantly different when compared with the effect of high-temperature treatments. Field observations show that the establishment of seedlings of E. multiflora is rare both after fire and between fires. Thus, our data support the idea that both a single fire or clipping can diminish the population size of a resprouting species. We conclude that fire may have a stochastic effect on E. multiflora populations, due to the variation in fire intensity existing within a single burning stand.     

Fire regenerative syndromes of forest woody species across fire and climatic gradients

Community resilience after fire is determined by species’ ability to regenerate through two main mechanisms growth of new sprouts (resprouter species) and germination from surviving seed banks or from seeds arriving from neighbouring populations (seeder species). Both the mechanisms are present in Mediterranean communities. The occurrence of both the types in a community depends on fire history and the bio-geographical history determining the available species pool. Regenerative traits also covary with other functional attributes associated with resource acquisition and stress tolerance. As post-fire regenerative responses can be related to various ecological factors other than fire, we tested the hypothesis of a different proportional representation of post-fire regenerative syndromes in forest woody species along a climatic gradient in Catalonia (NE Spain) ranging from Mediterranean to temperate-boreal climates. Specifically, we expected seeder species to become less common with colder and moister conditions while resprouters would not be so influenced by the climatic gradient. We also tested the hypothesis of change in the relative abundance of regenerative syndromes in relation to recent fire history. We analysed a large database obtained from extensive forestry surveys and remote sensing fire records. After correction for spatial autocorrelation, we found an increase in the proportion of seeder species under more Mediterranean conditions and a decrease in fire-sensitive species (with no efficient mechanisms of post-fire recovery) in moister conditions. Resprouter species were similarly present across the whole gradient. A similar pattern was observed after excluding recently burnt plots. Therefore, post-fire regenerative syndromes segregate along the climatic gradient. Recent fires reduced the occurrence of fire-sensitive species and increased the proportion of seeder species. No significant effect was observed on resprouter species. Fire has a sorting effect, shaping the occurrence of species with different regenerative traits. Overall, fire seems to explain better the variability of the proportion of fire-sensitive species and climate the variability of seeder species. In addition, other factors (forestry practices and the covariation between regenerative and functional attributes) are likely to contribute to the regional pattern of regenerative syndromes.     

Land‐use and fire history effects on post‐fire vegetation dynamics in eastern Spain

Question: Is post‐fire, medium‐term vegetation dynamics determined by land‐use or fire history prior to fire? Location: South‐facing slope in the Gallinera valley, Alicante province, eastern Spain. Methods: After mapping the land‐use and fire history of the study site using photo‐interpretation, we sampled vegetation structure on a set of plots representing the most frequent land‐use and fire history combinations on an area burned six years before sampling. We studied the effects of land‐use history, comparing the one‐fire land‐use trajectories. We analysed the effects of fire history; comparing one‐ and two‐fire plots for both previously cropped and uncropped areas. Results: Most variables were not significantly different between the earliest abandoned plots (abandoned at least 38 years before the fire) and the uncropped plots. On the most recently abandoned plots (abandoned between one and four years before the fire), the therophyte richness and the ratio of seeder: resprouter richness were significantly greatest. Different fire recurrences did not determine different post‐fire vegetation on either the uncropped or the early abandoned plots (all dominated by fire‐recruited seeder shrubs). The most recently abandoned plots had a lower resilience to fire. Conclusions: Land‐use history and recent pre‐fire land use, in particular, determined the post‐fire vegetation in the medium term. The vegetation composition converged during secondary succession among land‐use histories. Increasing fire recurrence had a small effect on mature plant communities, due to the combination of life‐history traits determining the response to fire of the dominant species.     

Seasonality and life cycles of woody plant-feeding noctuid moths (Lepidoptera: Noctuidae) in Mediterranean habitats

Abstract.

• 1 The annual cycle of noctuid moths feeding as larvae on the foliage of eighteen woody plant species (belonging to ten different families) was studied at two southeastern Spanish Mediterranean forest habitats. Two questions were addressed in this study. (1) Does the rigorous (hot and dry) summer season characteristic of the Mediterranean-type climate impose a constraint on noctuid life histories? (2) Are there detectable differences in abundance or phenological patterns between the noctuid assemblages feeding on evergreen and winter-deciduous trees and shrubs?
• 2 Regardless of their leaf persistence habit, the majority of woody species studied had short shoot growth and leaf production periods in spring. Abundance of noctuid larvae peaked in May-June, shortly after host plants started to produce new leaves. Virtually no noctuid larvae were found on the foliage of the woody species studied at other times of year. Abundance and seasonal pattern of occurrence of larvae were similar on evergreen and deciduous food plants.
• 3 In contrast with the unimodal seasonal pattern exhibited by larvae, the abundance of adult moths (assessed by light trapping) showed two distinct peaks in early summer and early autumn, and a marked minimum in mid-summer.
• 4 Most woody plant-feeding noctuids recorded in this study (86% of species, about 95% of individuals) were univoltine. Univoltine species fell into one of two distinct life history categories, corresponding to pre- and post-summer flight periods. These two contrasting phenologies were associated with a bimodality in the duration of the pupal stage (‘short’ versus ‘long’), and were closely related to taxonomical affiliation at the subfamily level.
• 5 Within the ‘short’ pupal duration group, some species mate and oviposit shortly after emergence and pass the dry season in the egg stage, while others remain as potentially active, non-reproductive adults over most of the summer and mate by the end of that season. Species in the ‘long’ pupal duration group pass the summer in pupal or prepupal stage.
• 6 In the Mediterranean habitats studied, the adversity of the summer dry season seems to have led to a woody plant-feeding noctuid species assemblage almost entirely made up of univoltine species that concentrate their larval phases when suitable food (young leaves) is most abundant, and ‘avoid’ mid-summer as a flight time by emerging either shortly before or shortly after the adverse summer drought period. Year-round foliage availability afforded by dominant evergreen plants does not seem to have influenced the seasonal organization of noctuid cycles in any substantial way.

     

Seasonal and inter-annual variations of gas exchange in thirteen woody species along a climatic gradient in the Mediterranean island of Mallorca

We studied the influence of summer drought and winter temperatures on seasonal and spatial variations of light-saturated net photosynthesis and stomatal conductance in Mediterranean woody species. We measured variations in leaf gas exchange over 3 years in 13 Mediterranean trees and shrubs, located at four different sites along a climate gradient of temperature and precipitation in the island of Mallorca (West Mediterranean basin). Net photosynthesis and stomatal conductance were at a maximum during spring, autumn or winter and at a minimum during summer in most sites, species and years. Nevertheless, important spatial, temporal and species-specific variations were observed. Summer drought limitation to gas exchange was greatest in the dry part of the transect, where many species showed their maximum gas exchange rate during winter. In contrast, winter temperatures limited gas exchange of many species at the wet and cool end of the transect, while summer depression of gas exchange was shorter and less pronounced. These results suggest that the effect on carbon fixation and productivity by the predicted future increase of aridity in the Mediterranean basin will depend on whether gas exchange is mostly limited by summer drought or by low winter temperatures.     

The challenge of plant regeneration after fire in the Mediterranean Basin: scientific gaps in our knowledge on plant strategies and evolution of traits

Though observations on re-colonisation of post-fire sites in the Mediterranean Basin are plentiful, there still is an ongoing debate on the interrelation of fire regimes and species traits related to fire adaptation. Most of the studies found are restricted to particular species or claim to present community attributes. Therefore they often lack information for the evaluation of evolutionary evidence and historical contingency of the local fire regime and other abiotic conditions, which may act as selective pressure for plant regeneration strategies. Indeed, knowledge about the success of regeneration mechanisms and their interrelation with ecological factors is essential for the interpretation of the high spatio-temporal variability found in post-fire species performance. Such knowledge would be necessary to assess the potential of different regeneration mechanisms to cope with ongoing land-use and climate change—a crucial scientific challenge. A summary is given of the knowledge about the limits and potential of plant regeneration mechanisms after fire in the Mediterranean Basin, along with corresponding studies conducted in other parts of the world with similar climatic conditions in order to present the fullest possible picture. Moreover, the positive or negative impacts of particular parameters of a fire regime on different regeneration strategies (post-fire seeders, resprouters, and facultative resprouters) are explained and discussed in the light of published literature. To conclude, reference is made to scientific gaps that need to be filled in order to analyse species resistance and community resilience absorbing possible climate or land use changes.     

Comparative water relations of four Mediterranean oak species

The water relations and responses of two evergreen (Quercus ilex L. and Q. suber L.) and two deciduous (S. afares Pomel. and Q. faginea Will.) Quercus species were studied under experimental conditions. Two-year old seedlings grown in 30 l. pots were subjected to a drying period during which stomatal conductance, pre-dawn potential and minimum foliar potential were measured.The results shows that, for all species, the daily course of stomatal conductance agrees with the patterns proposed by Hinckley et al. (1978 & 1983). Concurrent with the species responses to short-term variation in water availability, it was found that pre-dawn leaf water potential controlled the maximum daily leaf conductance. There was a strong correlation between pre-dawn leaf potential and maximum daily conductance as described by the reciprocal function g_{srmax for}=(-0.47+2.61._p)^-1 the evergreen oaks and g_{srmax for}=(-1.94+7.39._p)^-1 for the deciduous species. These differences between the two groups may partialy explain their geograhic distributions, and suggest general questions concerning the mechanisms which optimize water-use efficiency in Mediterranean oak species.     

Method to classify xylem cell types using cross sectionsof 1-year-old branches in Mediterranean woody species

Measurements of anatomical parameters of wood are of great interest both for eco-physiological purposes and for technological applications. The aim of this paper is to describe a new method for classifying and measuring cell lumen of xylem, analysing cross sections under the light microscope. The proposed method is based on the application of digital image analysis on images of the cross sections of xylem in combination with graphical and statistical methods. The methodology was tested on 1-year-old branches of several woody species, both trees and shrubs, occurring in a Mediterranean natural ecosystem in southern Italy. The development of the procedure was based on statistical comparison between data collected according to four procedures: (a) manual identification and measurement of lumen diameter of conduits on longitudinal sections; (b) manual identification and measurement of lumen diameter of conduits on cross sections; (c) manual identification and measurement of lumen area of conduits on cross sections; and (d) automatic measurement of lumen area of conduits on cross sections. The influence of image resolution and that of the position of the selected area on cell classification and measurements were ascertained. The proposed method was proved to be specific to woody structures and allowed the construction of a model-graph that is species-specific. Interpretation of the model-graphs allows classification and hence measurement of identified cells.     

Life‐history traits predict perennial species response to fire in a desert ecosystem

The Mojave Desert of North America has become fire‐prone in recent decades due to invasive annual grasses that fuel wildfires following years of high rainfall. Perennial species are poorly adapted to fire in this system, and post‐fire shifts in species composition have been substantial but variable across community types. To generalize across a range of conditions, we investigated whether simple life‐history traits could predict how species responded to fire. Further, we classified species into plant functional types (PFTs) based on combinations of life‐history traits and evaluated whether these groups exhibited a consistent fire‐response. Six life‐history traits varied significantly between burned and unburned areas in short (up to 4 years) or long‐term (up to 52 years) post‐fire datasets, including growth form, lifespan, seed size, seed dispersal, height, and leaf longevity. Forbs and grasses consistently increased in abundance after fire, while cacti were reduced and woody species exhibited a variable response. Woody species were classified into three PFTs based on combinations of life‐history traits. Species in Group 1 increased in abundance after fire and were characterized by short lifespans, small, wind‐dispersed seeds, low height, and deciduous leaves. Species in Group 2 were reduced by fire and distinguished from Group 1 by longer lifespans and evergreen leaves. Group 3 species, which also decreased after fire, were characterized by long lifespans, large non‐wind dispersed seeds, and taller heights. Our results show that PFTs based on life‐history traits can reliably predict the responses of most species to fire in the Mojave Desert. Dominant, long‐lived species of this region possess a combination of traits limiting their ability to recover, presenting a clear example of how a novel disturbance regime may shift selective environmental pressures to favor alternative life‐history strategies.     

Seed germination of high mountain Mediterranean species: altitudinal,interpopulation and interannual variability

The germination response of 20 species from high altitude Mediterranean climates, most of them rare endemics, was studied. Our main goal was to model the germination response of a complete set of Iberian high mountain species. The effect of temperature and other parameters, such as spatial and temporal short gradients, on germination were also evaluated. Some seed features (mass and size) were also related to the germination response. Finally, we tested the effect of cold-wet stratification pretreatment when germination was low under natural conditions. Seeds were collected at four locations from 1,900 to 2,400 m a.s.l. in the Sierra de Guadarrama (Spanish Central Range) over two consecutive growing seasons (2001–2002) and submitted to different temperatures and a constant photoperiod of 16 h light/8 h darkness. Most plants readily germinate without treatment, reaching an optimum at relatively high temperatures in contrast to lowland Mediterranean species. Seeds seem to be physiologically prepared for rapid germination even though these plants usually face very intense summer droughts after ripening and dispersal. Germination was also highly variable among altitudes, populations and years, but results were inconsistent among species. Such flexibility could be interpreted as an efficient survival strategy for species growing under unpredictable environments, such as the Mediterranean climate. Finally cold-wet stratification increased germination capacity in five of nine dormant species, as widely reported for many arctic, boreal and alpine species. In conclusion, high mountain Mediterranean species do not differ from alpine species except that a relatively high number of species are ready to germinate without any treatment.     

Stomatal encryption by epicuticular waxes as a plastic trait modifying gas exchange in a Mediterranean evergreen species (Quercus coccifera L.)

The adaptive benefit of stomatal crypts remains a matter of controversy. This work studies the effect on gas exchange of cuticular rims that overarch the stomatal pore in the Mediterranean species Quercus coccifera L. growing under Mediterranean (lower relative humidities and high summer temperatures) or oceanic conditions (higher daily relative humidities and mild temperatures). After microscopic assessment of the leaf surfaces and stomatal architecture, the impact of the cuticular ‘cup’ on gas exchange was evaluated by employing three‐dimensional finite element models. Here, we provide evidence for a high plasticity of the Q. coccifera cuticular cup, with much larger vents under oceanic conditions compared to small vents under Mediterranean conditions. This structure adds a substantial fixed resistance thereby strongly decreasing gas exchange under Mediterranean conditions. The cuticular cup, which also increases leaf internal humidity, might buffer the rapid changes in vapour pressure deficit (VPD) often observed under Mediterranean conditions. Since water loss of guard and adjacent epidermal cells regulates stomatal aperture, we suggest that this structure allows an efficient regulation of stomatal conductance and optimum use of resources under high VPD. This study provides evidence that plasticity of stomatal architecture can be an important structural component of hydraulic adaptation to different climate conditions.     

Germination behaviour of 14 Mediterranean species in relation to fire factors: smoke and heat

Fire is an ecological factor that has been present in the ecosystems of the Mediterranean region for thousands of years. Our study was undertaken to acquire knowledge of the effect of fire on the germination of Mediterranean species. We used high temperatures (up to 60°C) and smoke to determine the effect of these factors on the germination of species from the Mediterranean region. The species selected are characteristic of the central Mediterranean basin and are representative of both woody and herbaceous species: Rhamnus alaternus L., Cistus albidus L., Cistus monspeliensis L., Fumana ericoides (Cav.) Gand., Rosmarinus officinalis L., Melica ciliata L., Avena sterilis L., Bituminaria bituminosa (L.) C.H. Stirt., Anthyllis vulneraria L., Coronilla glauca L., Argyrolobium zanonii (Turra) P.W. Balland, Emerus major Mill., Genista scorpius (L.) D.C. and Spartium junceum L. The seeds were collected in Mediterranean shrubland (8) and woodland (6) ecosystems, around Montpellier, France (24°45′N and 3°50′E). Ten treatments were tested: a control, three smoke treatments and six heat treatments. The average germination level (germination percentage) and the average T₅₀ rates (time taken to reach 50% of germination) were calculated. The smoke and heat act in a different way on each of the species. The smoke enhanced the germination of two species, whilst moderate heat increased germination in all of the species excepting R. officinalis, F. ericoides, A. sterilis, A. vulneraria, and G. scorpius. Germination was fastest in M. ciliata and S. junceum and slowest in A. sterilis, E. major and C. albidus. The cues did not significantly affect the rate of germination. Fire modified the germination response of 12 of the 14 species studied.     

Comparative phenology and growth in different species of the Mediterranean maquis of central Italy

A phenomorphological survey was carried out in central Italy to study the effects of increasing water stress on some characteristic species of the Mediterranean maquis. Nutrient content and leaf water potential were examined. The results show that three different groups exist which diverge in the modulation of growth activity. 1) Evergreen sclerophyllous species (e.g. Pistacia lentiscus, Phillyrea media, Arbutus unedo, Ruscus aculeatus), which were supposed to be drought-tolerant, in fact limited their growth activity to a brief period before aridity increased. A similar growth pattern was exhibited by those species (e.g. Quercus ilex, Erica arborea, Smilax aspera) that stopped producing new leaves and branches during the driest season and that recovered after the first rain; i.e., their growth period lasted longer. 2) Drought-deciduous species (e.g. Calicotome villosa) that adopted the drought-avoidance strategy had two vegetative periods interrupted by a phase during which they completely shed their leaves. 3) Semideciduous species (Cistus monspeliensis) with mesophitic leaves adopted an intermediate response. These grew even in the dry and cold season.     

Seasonal patterns of terpene content and emission from seven Mediterranean woody species in field conditions

The seasonal pattern of terpene content and emission by seven Mediterranean woody species was studied under field conditions. Emission rates were normalized at 30°C and 1000 μmol·m·s PFD (photosynthetic photon flux density). Bupleurum fruticosum, Pinus halepensis, and Cistus albidus stored large amounts of terpenes (0.01-1.77% [dry matter]) with maximum values in autumn and minimum values in spring. They emitted large amounts of terpenes (2-40 μg·g DM·h), but with no clear seasonal trend except for Cistus albidus, which had maximum values in spring and minimum values in autumn. The nonstoring species Arbutus unedo, Erica arborea, Quercus coccifera and Quercus ilex also emitted large amounts of terpenes (0-40 μg·g DM·h) and also tended to present maximum emission rates in spring, although this trend was significant only for A. unedo. At the seasonal scale, emission rates did not follow changes in photosynthetic rates; instead, they mostly followed changes in temperature. From autumn to spring, the least volatile monoterpenes such as limonene were emitted at highest rates, whereas the most volatile monoterpenes such as α-pinene and β-pinene were the most emitted in summer. The monoterpene emission rates represented a greater percentage of the photosynthetic carbon fixation in summer (from 0.51% in Arbutus unedo to 5.64% in Quercus coccifera) than in the rest of the seasons. All these seasonality trends must be considered when inventorying and modeling annual emission rates in Mediterranean ecosystems.     

The influence of fire on the phenological behaviour of Mediterranean plant species in Bas-Languedoc (southern France)

Fire is an ancient ecological factor influencing the Mediterranean vegetation of southern France. The study was carried out on three areas to determine the phenological behaviour of plants with regard to fire. First we studied the flowering responses of perennials in relation to the time since fire: in a Quercus coccifera garrigue most species flower during the year following burning. In comparing species by species between burned and unburned areas most species did not show major differences in the phenological stages. However, fire did increase the number of inflorescences of grasses. A phenological synthesis showed that differences at the community level existed for the flowering stages between the burned areas and the unburned control sites during the first and second years following fire. The growth of some woody species was also studied; the elongation and growth of the plants were biggest during the first or second year after fire. The lack of differences in phenological response between burned and unburned plants may be an adaptive trait to fire.