María Valbuena-Carabaña

Fine‐scale spatial genetic structure in mixed oak stands with different levels of hybridization

Oaks are model species for the study of natural introgressive hybridization. High interfertility among oak taxa might result in collective evolution, through transpecific spread of advantageous alleles, challenging the standard concept of species. Nine highly polymorphic microsatellite (nuSSR) loci were analysed in three mixed oak populations of Quercus pyrenaica and Quercus petraea (Montejo, Somosierra and Robregordo) with different density and hybridization rates. Both leaf morphology and molecular markers were used to assess individual admixture rates. Insights about the relative effect of density and hybridization rates on fine‐scale spatial genetic structure (SGS) were obtained from autocorrelograms and Sp statistics. Differences in SGS among populations were higher than between species. These differences cannot be attributed solely to census densities but also relate to other factors, such as the spatial configuration of the population. Hybridization was an important factor shaping within‐population spatial genetic structure, and an interspecific component of SGS was found in Somosierra. Indirect estimates of historical gene flow in Montejo were compared with actual values of gene dispersal assessed by parentage analysis in a former study. Similar values were found for current and historical gene flow in both species, which might reflect demographical stability.

Functional performance of oak seedlings naturally regenerated across microhabitats of distinct overstorey canopy closure

The extent to which seedling recruitment is limited by summer drought in Mediterranean-type ecosystems depends on the light microsite, yet the relationship between light availability and water status, functional performance, and survival of seedlings in these systems is still unclear. Over a 3-year period, we studied the pattern of survival and functional performance of seedlings of Quercus petraea (Matt.) Liebl. and Quercus pyrenaica Willd. in a montane forest in central Spain, which is the southern edge of the natural range of Q. petraea. After a mast year of the two species, 72 plots were established in six microhabitats spanning a range of overstorey canopy closure: closed, partial and open canopies dominated by either Q. petraea or Q. pyrenaica adult trees. Seedlings of each species naturally emerged beneath the conspecific-dominated canopies. The second and third years of study were extremely dry. Three years after emergence, the greatest seedling survival occurred beneath the partial canopy of Q. pyrenaica trees (8%) and the lowest (0%) beneath the closed canopies of Q. pyrenaica and Q. petraea. Survival for Q. pyrenaica increased linearly with understorey light across the range of 10–35% Global Site Factor. Plant water deficit (estimated by leaf water potential) was high across microhabitats, and increased with light availability for Q. pyrenaica. Potential for photosynthesis (estimated by the electron transport rate of photosystem II) decreased with canopy closure; and potential for light harvesting (e.g. specific leaf area (SLA) and chlorophyll concentration) increased with closure. Extreme water deficit could be the main contributor to seedling death in the more open microhabitats, whereas light level was insufficient to maintain carbon balance under the water-stressful conditions existing beneath the closed tree canopies. Seedling establishment appears to be a limiting factor for the recruitment of both oaks within this forest in a wide range of microhabitats, especially for the more drought-sensitive Q. petraea. Moderate reductions of tree canopy cover can improve seedling establishment, but extreme summer droughts can prevent the success of any silvicultural practice made.

Stem CO2 efflux in six co-occurring tree species: underlying factors and ecological implications

Stem respiration plays a role in species coexistence and forest dynamics. Here we examined the intra- and inter-specific variability of stem CO2 efflux (E) in dominant and suppressed trees of six deciduous species in a mixed forest stand: Fagus sylvatica L., Quercus petraea [Matt.] Liebl, Quercus pyrenaica Willd., Prunus avium L., Sorbus aucuparia L. and Crataegus monogyna Jacq. We conducted measurements in late autumn. Within species, dominants had higher E per unit stem surface area (Es ) mainly because sapwood depth was higher than in suppressed trees. Across species, however, differences in Es corresponded with differences in the proportion of living parenchyma in sapwood and concentration of non-structural carbohydrates (NSC). Across species, Es was strongly and NSC marginally positively related with an index of drought tolerance, suggesting that slow growth of drought-tolerant trees is related to higher NSC concentration and Es . We conclude that, during the leafless period, E is indicative of maintenance respiration and is related with some ecological characteristics of the species, such as drought resistance; that sapwood depth is the main factor explaining variability in Es within species; and that the proportion of NSC in the sapwood is the main factor behind variability in Es among species.

Variation components in leaf morphology of recruits of two hybridising oaks (Q. petraea (Matt.) Liebl. and Q. pyrenaica Willd.) at small spatial scale

Leaf morphological variation was examined in recruits of two hybridising oaks in a small sympatric area from Central Spain. Nuclear microsatellites were used to identify hybrids and assess the parental lineage. By Bayesian clustering analysis, 5% of hybrids were found. Principal component analysis was used to reduce 15 morphometric variables to four components associated with leaf size, lobation/pubescence and overall shape of the leaf. The percentage of variance due to genetic factors was evaluated through nested analysis of variance. As much as 70% of variance component was due to the factor “species” for lobation/pubescence, suggesting high adaptive value for these traits, possibly related to ecological constraints of the species. The genetic component of variance for leaf size and overall shape of the leaf was below 33%. Age and height of the recruits did not correlate with sun-leaf morphology. Competition indexes and diameter of the recruits showed slight, although significant, correlations with leaf size and lobation/pubescence components, pointing to some trade-offs between competition for light and leaf morphology of Q. petraea and Q. pyrenaica recruits.

Seasonal and diel variation in xylem CO2 concentration and sap pH in sub-Mediterranean oak stems

Since a substantial portion of respired CO2 remains within the stem, diel and seasonal trends in stem CO2 concentration ([CO2]) are of major interest in plant respiration and carbon budget research. However, continuous long-term stem [CO2] studies are scarce, and generally absent in Mediterranean climates. In this study, stem [CO2] was monitored every 15min together with stem and air temperature, sap flow, and soil water storage during a growing season in 16 stems of Quercus pyrenaica to elucidate the main drivers of stem [CO2] at different temporal scales. Fluctuations in sap pH were also assessed during two growing seasons to evaluate potential errors in estimates of the concentration of CO2 dissolved in xylem sap ([CO2*]) calculated using Henrys law. Stem temperature was the best predictor of stem [CO2] and explained more than 90% and 50% of the variability in stem [CO2] at diel and seasonal scales, respectively. Under dry conditions, soil water storage was the main driver of stem [CO2]. Likewise, the first rains after summer drought caused intense stem [CO2] pulses, suggesting enhanced stem and root respiration and increased resistance to radial CO2 diffusion. Sap flow played a secondary role in controlling stem [CO2] variations. We observed night-time sap pH acidification and progressive seasonal alkalinization. Thus, if the annual mean value of sap pH (measured at midday) was assumed to be constant, night-time sap [CO2*] was substantially overestimated (40%), and spring and autumn sap [CO2*] were misestimated by 25%. This work highlights that diel and seasonal variations in temperature, tree water availability, and sap pH substantially affect xylem [CO2] and sap [CO2*].

Population clustering and clonal structure evidence the relict state of Ulmus minor Mill. in the Balearic Islands

Field elm (Ulmus minor) is a riparian tree that grows in rare, small populations scattered along temporary watercourses in the Balearic Islands, nowadays mostly covered with Mediterranean vegetation. Agriculture and farming on the fertile land along the periodically flooded plains have reduced the elm populations to sparse tree lines along the creek beds. The presence of field elm in this very anthropic landscape has led some authors to consider it as an introduced species in the Balearics. However, pollen data suggest these elms may be the remains of larger populations experiencing continuous population shrinkage during the Holocene, and hence be native to the isles. In this paper, we apply genetic markers to assess whether field elm is or is not indigenous to the Balearic Islands. We compare the genetic variation in nine nuclear microsatellites of six Balearic populations (three in each of the largest islands, Majorca and Minorca) with that of three natural Iberian populations located in two regions, one geologically (Baetic mountains, SE Iberia) and another historically (Catalonia, NE Iberia) related to the islands. Principal coordinates analysis and Bayesian clustering methods reveal a strong genetic differentiation of the Balearic populations from the Iberian ones, and even among islands, which support their native origin. Genotypic variation in the islands is very low and clonal reproduction is very high compared with the mainland, as it is frequently observed in populations of clonal species where sexual reproduction is limited. We discuss the practical implications of these findings for the conservation of elm genetic resources of these findings.

On the general failure of coppice conversion into high forest in Quercus pyrenaica stands: a genetic and physiological approach

Genetic, dendrochronological and physiological studies are reviewed to investigate causes of tree stagnation of abandoned Quercus pyrenaica coppices. Tree decline – displayed by slow stem growth, branch dieback and scarce acorn production – has led to advice on the conversion of abandoned oak coppices into high forests by thinning. However, this practice has often failed. The shortage of genetic diversity and disequilibrium between above- and below-ground organs in large, over-aged stools (clones) are adduced consequences of centenary coppicing driving stand stagnation in Q. pyrenaica. Our results evidence that historical coppicing has not enhanced genetic diversity losses; on the contrary, it has allowed for the recruitment of new genotypes and the existence of uneven-aged stands of heterogenic clonal structure. Tree excavations have revealed widely spread root systems. High carbon expenditures in root respiration in large and centennial clones may constrain their aboveground development, and thinning practices aiming at converting Q. pyrenaica coppices into high forests might not succeed or even enhance a physiological root-to-shoot imbalance.

Centenary coppicing maintains high levels of genetic diversity in a root resprouting oak (Quercus pyrenaica Willd.)

Studies disentangling the anthropogenic influences of traditional forest uses are crucial to assess the current conservation value of cultural landscapes. By promoting asexual regeneration, centenary coppicing in the predominantly root resprouter Quercus pyrenaica is assumed to have reduced genetic diversity levels contributing to the decline of abandoned coppices and the common lack of acorn production. This work aims to test the widespread assumption that historical coppicing in Q. pyrenaica has caused depleted levels of genetic diversity. Seven microsatellite markers were used to assess clonal structure and population genetic diversity levels in six abandoned coppices of Q. pyrenaica, which were compared to three open woodlands in national parks in Spain. Asexual regeneration was higher in coppices, leading to more frequent and larger clonal assemblies. Clonal diversity parameters (genotypic richness and proportion of unique genotypes) were significantly lower in coppices, although density of genotypes per surface area and levels of population genetic diversity were comparable to those observed in open parklands. Heterogenic clonal structures were found both within and among stands, hindering the inference of concrete anthropogenic disturbances. Despite promoting asexual reproduction, coppicing maintains high levels of genotypic and genetic diversity and allows the incorporation of new genotypes by seed recruitment. The natural resprouting capacity of Q. pyrenaica preserved the species in face of long-lasting anthropogenic disturbances, fostering ecosystem resilience and harbouring high conservation values.