Fernando Pulido

Regeneration of a Mediterranean oak: A whole-cycle approach

ABSTRACT Using the holm oak (Quercus ilex) as a model system, we quantified the losses in the potential number of recruits from the flower to the sapling stage caused by abiotic factors, pre- and post-dispersal seed predators, and invertebrate and ungulate herbivores in one well-preserved forest plot (F) and in one dehesa (savanna-like) plot (D). Tree fecundity was an order of magnitude higher in D than in F. Predispersal (flower plus fruit) losses due to abiotic factors were much larger (71% in F and 90% in D) than those originated by biotic factors (29% in F and 10% in D). Post-dispersal predation of acorns under trees led to crop depletion in F, while predators did not deplete the acorn crop despite a much higher predation level in D. Surface acorns were invariably eaten, while most buried acorns survived to germination in both plots. Ninety-five percent of seedlings emerged under oaks in D and only 20% in F, the density of newly emerged seedlings being ten times greater in F. Emergence from experimentally sown acorns was very high on average (73%), while seedling survival to the 2-y sapling stage was primarily affected by water stress in the first summer, especially in the dehesa stand. A 75-fold difference in recruitment rates between populations was found (0.00150 in F and 0.00002 in D). This whole-cycle disparity was the result of differences in the conversion rate from viable fruits to newly emerged seedlings (two orders of magnitude larger in F). The inability to direct acorns to safe (shaded) sites by means of efficient dispersers appears to limit recruitment in open dehesa stands. This finding could explain the general lack of natural regeneration of dehesas as compared to the forests from which they developed.

Effects of land-use history on size structure of holm oak stands in Spanish dehesas: implications for conservation and restoration

Dehesas , rangelands occupied by scattered oak trees and characterized by silvopastoral uses, cover about 3.1 million ha in south-western Spain. There is considerable debate about the long-term persistence of holm oak ( Quercus ilex ) populations in dehesas, since most stands are overaged and seedlings and saplings are sparse. The forest cycle has been disrupted in most dehesas. Regeneration has been inhibited since stands were opened for agriculture and grazing. Oak diameters from three land-use groups (young dehesa [YD], middle-aged dehesa [MD], and old dehesa [OD]) in Caceres Province, Spain, were compared. These groups differed in the age of the land-use system, i.e. time since the original Mediterranean forest was cleared. The dehesa systems were established about 80 (YD), 150 (MD) and 500 (OD) years ago. An analysis of 66 holm oak cross sections revealed a close correlation ( r 2 = 91.2%) between tree rings and diameters, so that diameter seems to be a reliable indicator of tree age. Nested analysis of variance showed significant variation in diameters between the land-use groups. There is generally a positive relationship between tree age and the age of agrosilvopastoral use of the dehesas. Sparse holm oaks in the dehesas are primarily remnants from the first forest cycle. Local differences in growth conditions (for example soil quality and tree density) contribute further significant diameter variation on a between-plot level. Diameter structure of abandoned dehesas showed two peaks and a high proportion of trees in the smallest size class. This indicates that the forest degradation process is reversible. An effective regeneration policy should promote a rotating 20- to 30-year set-aside of dehesa parcels.

Gains to species diversity in organically farmed fields are not propagated at the farm level

Organic farming is promoted to reduce environmental impacts of agriculture, but surprisingly little is known about its effects at the farm level, the primary unit of decision making. Here we report the effects of organic farming on species diversity at the field, farm and regional levels by sampling plants, earthworms, spiders and bees in 1470 fields of 205 randomly selected organic and nonorganic farms in twelve European and African regions. Species richness is, on average, 10.5% higher in organic than nonorganic production fields, with highest gains in intensive arable fields (around +45%). Gains to species richness are partly caused by higher organism abundance and are common in plants and bees but intermittent in earthworms and spiders. Average gains are marginal +4.6% at the farm and +3.1% at the regional level, even in intensive arable regions. Additional, targeted measures are therefore needed to fulfil the commitment of organic farming to benefit farmland biodiversity.

Herbivore effects on developmental instability and fecundity of holm oaks

Plants are able to compensate for loss of tissue due to herbivores at a variety of spatial and temporal scales, masking detrimental effects of herbivory on plant fitness at these scales. The stressing effect of herbivory could also produce instability in the development of plant modules, and measures of such instability may reflect the fitness consequences of herbivory if instability is related to components of plant fitness. We analyse the relationships between herbivory, developmental instability and production of female flowers and fruits of holm oak Quercus ilex trees by means of herbivore removal experiments. Removal of leaf herbivores reduced herbivory rates at the tree level, but had no effect on mean production of female flowers or mature fruits, whereas herbivory tended to enhance flower production and had no effect on fruit abortion at the shoot level. Differences in herbivory levels between shoots of the same branch did not affect the size and fluctuating asymmetry of intact leaves. These results indicate compensation for herbivory at the tree level and over-compensation at the shoot level in terms of allocation of resources to female flower production. Removal of insect herbivores produced an increase in the mean developmental instability of leaves at the tree level in the year following the insecticide treatment, and there was a direct relationship between herbivory rates in the current year and leaf fluctuating asymmetry the following year irrespective of herbivore removal treatment. Finally, the production of pistillate flowers and fruits by trees was inversely related to the mean fluctuating asymmetry of leaves growing the same year. Leaf fluctuating asymmetry was thus an estimator of the stressing effects of herbivory on adult trees, an effect that was delayed to the following year. As leaf fluctuating asymmetry was also related to tree fecundity, asymmetry levels provided a sensitive measure of plant performance under conditions of compensatory responses to herbivory.

Biología reproductiva y conservación: el caso de la regeneración de bosques templados y subtropicales de robles (Quercus spp.)

This study presents a review of 21 experimental studies on oak ( Quercus) regeneration analysing at least three phases of the reproductive cycle, namely fertilization, viable fruit production, acorn dispersal, postdispersal predation, seedling emergence, sapling establishment, and adult recruitment. Most study cases came from North American and European temperate forests representing a continuum from xeric sites with evergreen oaks (usually managed by humans) to mesic, well-preserved sites with deciduous oaks. Studies were classified according to three types of regeneration limitation, that is, viable seed production, herbivory, and safe sites availability. Results show that regeneration is usually limited by a combination of factors, tipically acorn consumption and unfavourable abiotic conditions for establishment. Tree fecundity, as well as seed dispersal by animals, are two important, usually neglected sources of recruitment limitation. On the other hand, anthropogenic disturbances negatively affects forest regeneration by changing the abundance or behavior of mutualistic or phytophagous organisms, and by modifying the abiotic scenario for seedling establishment. Finally, the application of ecological information for solving regeneration failures of oak forests is discussed, especially in relation to the alternatives of natural regeneration versus planting.

REPRODUCTIVE BEHAVIOR IN FEMALE IBERIAN RED DEER: EFFECTS OF AGGREGATION AND DISPERSION OF FOOD

Abstract Resource dispersion influences mammalian mating systems by affecting space use by females and in turn the strategies employed by males. Other elements related to mating interactions, however, also may affect female decisions, and it usually becomes difficult to discriminate between them. Here we present the results of experimental food supplementation during rut in the red deer (Cervus elaphus hispanicus). We provided food at 2 levels of spatial dispersion in order to test whether changes in the distribution of food within a short period of time affect spatial behavior of females or other features of the mating system. Food clumping produced an increase in local density of females using an area and in the size of female groups. It also produced an increase in number of adult males, male–male interactions, and male harassment toward females. Despite the overall rate of harassment being higher with food clumping, females received a lower per capita rate of harassment because they gathered into much larger harems. Variance in harem size increased with higher levels of food clumping. Our data show that females were highly sensitive to food distribution when joining harems, which stresses the relative importance of direct benefits (food and sexual harassment avoidance) in molding the conditions under which selection operates during rut in Iberian red deer.

Fruit abortion, developmental selection and developmental stability in Quercus ilex

Fruit abortion has been hypothesized to be a parental means of selective removal of propagules with low viability. In particular, aborted zygotes have been suggested to have developmentally deviant phenotypes, and surviving offspring may therefore give rise to adults with a developmentally stable phenotype. We tested predictions from this hypothesis using acorns of holm oaks Quercus ilex as a model system. Fecundity of oak trees was negatively related to mean fluctuating asymmetry of leaves, and abortion rates were positively related to leaf fluctuating asymmetry in at least one population. Aborted acorns were asymmetric in 83–99% of cases in three samples, while mature acorns were only asymmetric in 57–78% of cases. Acorn asymmetry was unrelated to germination probability and germination date, and had no significant effect on number of leaves, leaf mass, stem mass, seedling height or leaf area of seedlings. However, acorn asymmetry affected the trade-off between number and size of leaves in seedlings. Seedlings from asymmetric acorns showed a positive relationship between acorn size and number of leaves, but no relationships between acorn size and leaf area, while symmetric acorns showed the opposite. A positive relationship between acorn size and number of leaves in spring was found for naturally emerged seedlings that died during their first summer, whereas the number of leaves produced by surviving seedlings did not depend on acorn size. These findings are consistent with the hypothesis that stressed trees selectively abort propagules of low viability, and that developmental selection acts on a measure of developmental instability of fruits.

Oak Regeneration: Ecological Dynamics and Restoration Techniques

The acreage of oak woodlands has decreased in California and Spain, especially in the twentieth century. Currently, most surviving stands in Spain suffer from oak regeneration failure and it has been noted as a problem in many stands in California. A lack of dispersers transferring acorns to safe (shaded) sites is the main recruitment limitation in dehesas, where shrub encroachment generally results in higher oak recruitment rates. In California, recruitment failure is due to a combination of factors. The effects of introduced Mediterranean annuals, heavy livestock grazing, fire suppression, and predation by native and non-native wildlife on acorns and seedlings are all implicated, depending on locale and time period. Afforestation has been the main instrument for addressing the regeneration problem, especially in Spain. Natural regeneration at local scales is favored by shrubs, but also supported by protecting seedlings and by modifying the environment so young oaks can grow to a safe height. Complete livestock exclusion is of limited value in California as it hampers seedling establishment due to increasing rodent density. In dehesas, however, natural regeneration can only occur in seasonally grazed or wholly ungrazed sites, though livestock-dependent landowners are generally reluctant to carry out these measures.

Acorn Production Patterns

Acorns—the fruits of oaks—are a key resource for wildlife in temperate forests throughout the Northern Hemisphere. Acorns are also economically important for extensive livestock rearing, and as a staple food have supported indigenous human populations. Consequently, differences in how individual trees and populations of oaks invest in acorn production, both in terms of the size of the acorn crop and of the size of individual acorns, are of interest both ecologically and economically. Acorn production by oaks in both California and Spain tends to be highly variable and spatially synchronous. We summarize studies conducted in the two regions that investigate the factors influencing acorn production. One hypothesis explored is that, as a consequence of management, acorn production tends to be affected by different environmental factors in the two regions; another hypothesis is that acorn production in oaks in Spanish dehesas produce larger and more predictable acorn crops than trees in less managed Spanish forests or in California woodlands. Other factors potentially influencing acorn production are summarized, including biotic factors, trade-offs with growth, trade-offs with acorn size, and pollen limitation. We conclude with a discussion of spatial synchrony and acorn production at the community level. There remain many questions concerning the mating systems of oaks, trade-offs between different oak life-history characters, and the patterns and drivers of spatial synchrony. Environmental conditions in the two regions are similar, but understanding how their subtle differences influence acorn production is likely to yield important insights about the proximate and ultimate factors affecting acorn production and masting behavior.

Are silvopastoral systems compatible with forest regeneration? An integrative approach in southern Patagonia

We used an integrated approach to analyze the influence of silvopastoral management on the biotic and abiotic factors influencing on the natural recruitment of native forests in Tierra del Fuego (Argentina). The probabilities of transition from flowers to seeds to seedlings were estimated in forests with silvopastoral management (i.e., stands that had been both grazed and thinned), secondary growth stands, and unmanaged old-growth areas. Pre-dispersal losses were caused by microclimate and insect predation acting on flowers and immature fruits. Post-dispersal losses resulted from the impact of reduced litterfall and microclimate. But the most critical stage of the overall cycle was seedling establishment. Silvopastoral practices and stand age modified the main drivers of regeneration. While flowering was unaffected by management, fruiting and seed production were more successful in unmanaged forests. Seedling establishment and survival were favored by canopy cover reduction in silvopastoral stands. The increase of solar radiation and soil moisture in managed forests positively influenced the seedling establishment and survival, while in second-growth forests it was limited by suitable micro-site availability. Thinning practices aimed at increasing the understory heterogeneity in mature forests and restoring canopy complexity of second-growth stands could be suitable for sustainable management of these temperate forests.