Jesús Julio Camarero

Response of vegetation to drought time-scales across global land biomes

We evaluated the response of the Earth land biomes to drought by correlating a drought index with three global indicators of vegetation activity and growth: vegetation indices from satellite imagery, tree-ring growth series, and Aboveground Net Primary Production (ANPP) records. Arid and humid biomes are both affected by drought, and we suggest that the persistence of the water deficit (i.e., the drought time-scale) could be playing a key role in determining the sensitivity of land biomes to drought. We found that arid biomes respond to drought at short time-scales; that is, there is a rapid vegetation reaction as soon as water deficits below normal conditions occur. This may be due to the fact that plant species of arid regions have mechanisms allowing them to rapidly adapt to changing water availability. Humid biomes also respond to drought at short time-scales, but in this case the physiological mechanisms likely differ from those operating in arid biomes, as plants usually have a poor adaptability to water shortage. On the contrary, semiarid and subhumid biomes respond to drought at long time-scales, probably because plants are able to withstand water deficits, but they lack the rapid response of arid biomes to drought. These results are consistent among three vegetation parameters analyzed and across different land biomes, showing that the response of vegetation to drought depends on characteristic drought time-scales for each biome. Understanding the dominant time-scales at which drought most influences vegetation might help assessing the resistance and resilience of vegetation and improving our knowledge of vegetation vulnerability to climate change.

Performance of Drought Indices for Ecological, Agricultural, and Hydrological Applications

AbstractIn this study, the authors provide a global assessment of the performance of different drought indices for monitoring drought impacts on several hydrological, agricultural, and ecological response variables. For this purpose, they compare the performance of several drought indices [the standardized precipitation index (SPI); four versions of the Palmer drought severity index (PDSI); and the standardized precipitation evapotranspiration index (SPEI)] to predict changes in streamflow, soil moisture, forest growth, and crop yield. The authors found a superior capability of the SPEI and the SPI drought indices, which are calculated on different time scales than the Palmer indices to capture the drought impacts on the aforementioned hydrological, agricultural, and ecological variables. They detected small differences in the comparative performance of the SPI and the SPEI indices, but the SPEI was the drought index that best captured the responses of the assessed variables to drought in summer, the seas...

Plastic bimodal xylogenesis in conifers from continental Mediterranean climates

*Seasonal radial-increment and xylogenesis data can help to elucidate how climate modulates wood formation in conifers. Few xylogenesis studies have assessed how plastic xylogenesis is in sympatric conifer species from continental Mediterranean areas, where low winter temperatures and summer drought constrain growth. *Here, we analysed intra-annual patterns of secondary growth in sympatric conifer species (Juniperus thurifera, Pinus halepensis and Pinus sylvestris). Two field sites (xeric and mesic) were evaluated using dendrometers, microcores and climatic data. *A bimodal pattern of xylogenesis characterized by spring and autumn precipitation and subsequent cambial reactivation was detected in J. thurifera at both study sites and in P. halepensis at the xeric site, but was absent in P. sylvestris where growth was largely controlled by day length. In the xeric site J. thurifera exhibited an increased response to water availability in autumn relative to P. halepensis and summer cambial suppression was more marked in J. thurifera than in P. halepensis. *Juniperus thurifera exhibited increased plasticity in its xylogenesis pattern compared with sympatric pines, enabling this species to occupy sites with more variable climatic conditions. The plastic xylogenesis patterns of junipers in drought-stressed areas may also provide them with a competitive advantage against co-occurring pines.

Effects of a severe drought on Quercus ilex radial growth and xylem anatomy

We assessed the response of Quercus ilex subsp. ballota to the severe summer drought recorded in 1994 in NE Spain through the study of changes in radial growth and wood anatomy. We selected a coppice stand in the Iberian Peninsula, which is characterized by a Mediterranean climate under continental influence. We measured internode length, tree-ring width, mean and maximum vessel diameter, and vessel density for 1981–1997. The annual predicted hydraulic conductance (Kh) was calculated following Hagen-Poisseuilles law. We compared the tree-ring width, vessel diameter and Kh of Q. ilex subsp. ballota and co-existing ring-porous oaks (Q. faginea, Q. pyrenaica) for a dry summer (1994) and a wet summer (1997). To evaluate the drought-resistance of xylem for Q. ilex subsp. ballota (dominant under continental conditions) and Q. ilex subsp. ilex (dominant in mild areas) we determined vulnerability curves. Dimensionless indices of internode length, tree-ring width, and vessel density were compared with climatic data (monthly total precipitation and mean temperature) using correlation analyses. Internode length, tree-ring width, Kh, and mean and maximum vessel diameter declined in 1994. According to vulnerability curves, Q. ilex subsp. ballota showed a greater drought resistance than Q. ilex subsp. ilex. During the year of growth, we found a positive influence of January and June–August precipitation on the internode length, tree-ring width, and vessel density. The response of Q. ilex subsp. ballota radial-growth to summer drought was comparable to that of Q. faginea latewood. Overall, growth and wood anatomy of Q. ilex subsp. ballota showed a plastic response to drought.

Functional groups in Quercus species derived from the analysis of pressure-volume curves

Abstract. Oaks that occupy Mediterranean phytoclimates share common leaf features (evergreenness, high leaf dry mass per unit area, LMA). Due to this phytoclimatic, morphological, and phenological convergence it has been suggested that they might constitute a coherent functional group. To confirm this hypothesis, some physiological parameters were determined after calculating pressure–volume curves (P–V curves) using the free-transpiration method. Seventeen Quercus species from contrasting phytoclimates were studied: six Mediterranean evergreen species (Q. agrifolia, Q. chrysolepis, Q. coccifera, Q. ilex ssp. ballota, Q. ilex ssp. ilex, and Q. suber); seven nemoral deciduous species (Q. alba, Q. laurifolia, Q. nigra, Q. petraea, Q. robur, Q. rubra and Q.velutina), and four nemoro-Mediterranean deciduous species (Q. cerris, Q. faginea, Q. frainetto, and Q. pyrenaica). Two-year-old seedlings growing under homogeneous environmental conditions (no water restrictions, uniform light, and nutrient supply) were used. The statistical analyses (correlation, mean-value comparisons, principal component analysis) of leaf features and parameters derived from the P–V curves confirmed the functional homogeneity of the three phytoclimatic groups, which were characterized by their contrasting ecophysiological response to water stress. The Mediterranean oak species developed mechanisms to avoid an excessive loss of cell water (e.g. high cell-wall rigidity). Conversely, the nemoral oaks showed the opposite. The nemoro-Mediterranean oaks perform better than nemoral oaks under water-stress conditions, but they cannot perform as well as the Mediterranean oaks on relatively dry soils.

EFFECTS OF A SEVERE DROUGHT ON GROWTH AND WOOD ANATOMICAL PROPERTIES OF QUERCUS FAGINEA

We studied the growth response to drought of a Quercus faginea Lam. stand in a xeric site in NE Spain, that experienced an intense defoliation in 1993–94. This event coincided with very low precipitation from November to February, the period when total monthly precipitation exceeds evapotranspiration. We evaluated the effects of November–February precipitation (recharge precipitation, RP) on internode length, radial growth, and wood anatomy. Quercus faginea showed reduced longitudinal and radial growth during the years with low RP, and most sampled trees did not produce latewood in 1993–94 but showed wide earlywood vessels. We observed the reverse for years with a high RP. Radial growth was enhanced by increased precipitation during January and May of the growth year. If severe droughts become more frequent, due to a greater climatic variability, extensive dieback of marginal Q. faginea populations may be expected.

Selective drought-induced decline of pine species in southeastern Spain

The negative impacts of severe drought on the growth and vigor of tree species and their relationship with forest decline have not been properly evaluated taking into account the differential responses to such stress of trees, sites and species. We evaluated these responses by quantifying the changes in radial growth of plantations of four pine species (Pinus sylvestris, Pinus nigra, Pinus pinaster, Pinus halepensis) which showed distinct decline and defoliation levels in southeastern Spain. We used dendrochronological methods, defoliation records, linear mixed models of basal area increment and dynamic factor analysis to quantify the responses of trees at the species and individual scales to site conditions and drought stress. In the region a temperature rise and a decrease in spring precipitation have led to drier conditions during the late twentieth century characterized by severe droughts in the 1990s and 2000s. As expected, the defoliation levels and the reductions in basal area increment were higher in those species more vulnerable to drought-induced xylem embolism (P. sylvestris) than in those more resistant (P. halepensis). Species adapted to xeric conditions but with high growth rates, such as P. pinaster, were also vulnerable to drought-induced decline. The reduction in basal area increment and the defoliation events occurred after consecutive severe droughts. A decrease in spring precipitation, which is the main driver of radial growth, is the most plausible cause of recent forest decline. The sharp growth reduction and widespread defoliation of the most affected pine plantations of Scots pine make their future persistence in drought-prone sites unlikely under the forecasted warmer and drier conditions.

Woody biomass production lags stem-girth increase by over one month in coniferous forests

Wood is the main terrestrial biotic reservoir for long-term carbon sequestration(1), and its formation in trees consumes around 15% of anthropogenic carbon dioxide emissions each year(2). However, the seasonal dynamics of woody biomass production cannot be quantified from eddy covariance or satellite observations. As such, our understanding of this key carbon cycle component, and its sensitivity to climate, remains limited. Here, we present high-resolution cellular based measurements of wood formation dynamics in three coniferous forest sites in northeastern France, performed over a period of 3 years. We show that stem woody biomass production lags behind stem-girth increase by over 1 month. We also analyse more general phenological observations of xylem tissue formation in Northern Hemisphere forests and find similar time lags in boreal, temperate, subalpine and Mediterranean forests. These time lags question the extension of the equivalence between stem size increase and woody biomass production to intra-annual time scales(3, 4, 5, 6). They also suggest that these two growth processes exhibit differential sensitivities to local environmental conditions. Indeed, in the well-watered French sites the seasonal dynamics of stem-girth increase matched the photoperiod cycle, whereas those of woody biomass production closely followed the seasonal course of temperature. We suggest that forecasted changes in the annual cycle of climatic factors(7) may shift the phase timing of stem size increase and woody biomass production in the future.

Relationship between hydraulic resistance and leaf morphology in broadleaf Quercus species: a new interpretation of leaf lobation

Abstract. We investigated the relationship between leaf shape and leaf hydraulic resistance in a set of broadleaf Quercus tree species (Q. cerris, Q. frainetto, Q. petraea, Q. pyrenaica, Q. robur, Q. rubra, Q. velutina). Seedlings of all the studied species were grown under uniform environmental conditions. A new high-pressure flowmeter was designed to measure leaf-blade hydraulic resistance. Leaf shape was characterised by the complexity of leaf outline which was regarded as an estimate of leaf lobation. This was done using the box-counting fractal dimension of the leaf silhouette. Leaf hydraulic resistance was negatively related to leaf lobation. It is suggested that the lower hydraulic resistance in deeply lobed leaves may constitute a mechanism for improving water balance under dry atmospheric conditions.

Plant species distribution across two contrasting treeline ecotones in the Spanish Pyrenees

We describe the structure of two contrasting (elevation, topography,climate, vegetation, soil) alpine forest–pasture ecotones located in theCentral Pyrenees (sites Ordesa, O, and Tessó, T). We define ecotonestructure as the spatial distribution of trees of different size classes andgrowth-forms and the relationship between these aspects and the spatialdistribution of understory vegetation and substrate. The studied ecotones aredominated by Pinus uncinata Ram. and have been littleaffected by anthropogenic disturbances (logging, grazing) during this century.One rectangular plot (30 × 140 m) was located within eachsite with its longest side parallel to the slope and encompassing treeline andtimberline. The distribution of size and growth-form classes at site O followeda clear sequence of increasing size downslope from shrubby multistemmedkrummholz individuals to bigger arborescent trees. At site O, regeneration wasconcentrated near the krummholz area and over rocky substrates. This suggeststhat krummholz may modify microenvironment conditions and increase seedlingsurvival. At site T, regeneration was abundant above the treeline where thecover of the dominant understory shrub (Rhododendronferrugineum) decreased. In both ecotones the diversity of plants washigher above the treeline than in the forest and decreased going downslopecoinciding with the increase of P. uncinata cover. Thereduction of plant diversity appeared above the current timberline. At site O,the decrease was steep and spatially heterogeneous what may be due in part tothe edaphic heterogeneity. At site T the change was abrupt though smaller. Therelationships between the plant community and tree regeneration should be takeninto account in future ecological studies of treeline pattern.