Effects of <Emphasis Type="Italic">Quercus suber</Emphasis> Decline on Woody Plant Regeneration: Potential Implications for Successional Dynamics in Mediterranean Forests

In the last two decades, widespread tree decline and mortality have been documented in forests worldwide. These mortality events usually show certain level of host-specificity, translating into rapid changes in the relative abundance of the adult community. Despite these short-term changes, it is poorly understood whether the decline and mortality of certain tree species are likely to result in long-term vegetation shifts. Trajectories of forest recovery and the probability of occurrence of permanent vegetation shifts are to a large extent determined by post-mortality regeneration dynamics. Using a spatially explicit neighborhood approach, we evaluated the spatial patterns of natural regeneration of the woody plant community in mixed Mediterranean forests affected by the decline of their dominant tree species, Quercus suber. We predicted the abundance, survival, and richness of the seedling and sapling bank as a function of the distribution and health status of the tree and shrub community. Results indicated that Q. suber decline had detectable effects on seedlings and saplings of coexistent woody species from very different functional groups (trees, shrubs, and lianas). The sign and magnitude of these effects varied substantially among coexistent species, which could imply shifts in the species ranking of seedling and sapling abundance, affecting successional trajectories and potentially leading to vegetation shifts. Because most of these changes pointed towards a loss of dominance of Q. suber, management strategies are urgently needed in order to attenuate adult mortality or promote its regeneration, counteracting the negative effects of global change drivers (exotic pathogens, climate change) on these valuable forests.     

More than just drought: complexity of recruitment patterns in Mediterranean forests

Understanding community dynamics during early life stages of trees is critical for the prediction of future species composition. In Mediterranean forests drought is a major constraint for regeneration, but likely not the only factor determining the observed spatial patterns. We carried out a sowing experiment aimed at identifying main filters during seed-seedling transition. Specifically, we studied seed fate (predation, fungi infection, emergence) and subsequent seedling performance (mortality during the first summer and overall recruitment after 2 years) of four co-occurring Mediterranean tree species (Quercus ilex, Quercus faginea, Juniperus thurifera, Pinus nigra). We related these processes to the dominant species composition, microhabitat heterogeneity, herb cover and seed mass. The identity of the dominant species in the forest canopy was more important for recruitment than the forest canopy being dominated by conspecific vs. heterospecific species. The patterns we found suggest that biotic interactions such as facilitation (lower mortality under the canopies) and herb competition (during emergence of J. thurifera) are relevant during recruitment. Moreover, our results pointed to ontogenetic conflicts regarding the seed mass of Q. faginea and to density-dependent seed mortality for Q. ilex, rarely described in Mediterranean ecosystems. We propose that our study species experience population growth in forests dominated by heterospecifics where the recruitment success depends on habitat heterogeneity and on moderated biotic and abiotic stresses created by each species. Our results reveal patterns and mechanisms involved in recruitment constraints that add complexity to the well-known drought-related processes in Mediterranean ecosystems.     

Competitive suppression of Quercus douglasii (Fagaceae) seedling emergence and growth

Reduced recruitment of blue oak (Quercus douglasii) seedlings in California grasslands and woodlands may result from shifts in seasonal soil water availability coincident with replacement of the native perennial herbaceous community by Mediterranean annuals. We used a combination of container and field experiments to examine the interrelationships between soil water potential, herbaceous neighborhood composition, and blue oak seedling shoot emergence and growth. Neighborhoods of exotic annuals depleted soil moisture more rapidly than neighborhoods of a perennial grass or "no-neighbor" controls. Although effects of neighborhood composition on oak seedling root elongation were not statistically significant, seedling shoot emergence was significantly inhibited in the annual neighborhoods where soil water was rapidly depleted. Seedling water status directly reflected soil water potential, which also determined the extent and duration of oak seedling growth during the first year. End-of-season seedling height significantly influenced survival and growth in subsequent years. While growth and survival of blue oak seedlings may be initially constrained by competition with herbaceous species, subsequent competition with adult blue oak trees may further contribute to reduced sapling recruitment.     

Facilitation and interference at the intraspecific level: Recruitment of Kageneckia angustifolia D. Don (Rosaceae) in the montane sclerophyllous woodland of central Chile

Most of the studies that have evaluated the interplay between interference and facilitation have been done at the interspecific level, whereas studies at the intraspecific level are scarce. The montane sclerophyllous forests of central Chile are dominated by the tree Kageneckia angustifolia, a semi-deciduous species that lose part of its foliage during summer. It has been reported that during winter snow accumulates in lower amounts beneath the canopy of K. angustifolia favoring the recruitment of new individuals compared to open areas (i.e., facilitation effect). However, it has also been reported that the leaf litter accumulated beneath parental trees contains allelopathic compounds that decrease seed germination, suggesting that recruitment beneath parental plants can be disfavored (i.e., interference effect). Hence, this system seems appropriate to assess the net-outcome between facilitative and negative effects during the emergence and survival of seedlings during the first year. In this study, we asked (i) what is the net-outcome between facilitative and interfering effects for K. angustifolia? (ii) does this net-outcome varies with the distance to parental trees? (iii) are positive and negative effects consistent through the seedling emergence and first year seedling survival phases? (iv) what are the main mechanisms behind the observed net-outcome? and (v) which is the optimal microhabitat for successful recruitment of this species? In an experimental plot of 10,000 m², we selected ten K. angustifolia trees and evaluated the effect of leaf litter on the emergence and survival of seedlings produced by experimentally sown seed seeds in three different microhabitats: beneath adult trees, edge of canopy and in open areas. In addition, we sampled three K. angustifolia stands to evaluate the microhabitat where the natural recruitment of this species actually occurring. Results showed that (1) seedling emergence was greater beneath canopy, intermediate in canopy edge and low in open areas, (2) whilst leaf litter significantly reduced seed germination, the magnitude of this negative effect was lower than the positive effect of beneath canopy microhabitat, (3) seedling survival was affected by microhabitats but not by the presence of leaf litter, (4) that the main mechanisms behind the observed patterns are the lower and delayed emergence of seedlings in open areas due to the longer duration of snow cover, decreasing the time to growth before the onset of summer drought, and (5) the greatest natural recruitment of K. angustifolia seedlings occurs beneath parental plants. Therefore, our findings suggest that the net-outcome between facilitative and interfering effect during the first year is mostly facilitative, indicating that adult trees of K. angustifolia are exerting a conspecific nurse effect on the recruitment of new individuals, a form of parental care in plants.     

Density‐dependent Survival in Seedlings Differs among Woody Life‐forms in Tropical Wet Forests of a Caribbean Island

Negative density dependence contributes to seedling dynamics in forested ecosystems, but the relative importance of this factor for different woody plant life‐forms is not well‐understood. We used 1 yr of seedling survivorship data for woody seedlings in 17 different plots of lower to mid‐montane rain forests on the island of Dominica to examine how seedling height, abiotic factors, and biotic factors such as negative density dependence are related to seedling survival of five different life‐forms (canopy, midstory, and understory trees; shrubs; and lianas). Across 64 species, taller seedlings in seedling plots with higher canopy openness, greater seedling density, lower relative abundance of conspecific seedlings, and lower relative abundance of conspecific adults generally had a greater probability of surviving. Height was the strongest predictor of seedling survival for all life‐forms except lianas. Greater seedling density was positively related to survival for canopy and midstory trees but negatively related to survival for the other life‐forms. For trees, the relative abundance of conspecific seedling and adult neighbors had weak and strong negative effects on survival respectively. Neither shrub nor liana seedling survival was affected by the relative abundance of conspecific neighbors. Thus, negative density dependence is confirmed as an important structuring mechanism for tree seedling communities but does not seem to be important for lianas and shrubs in Dominican rain forests. These results represent the first direct assessment of controls on seedling survival of all woody life‐forms – an important step in understanding the dynamics and structure of the entire woody plant community.     

Regeneration of a marginal Quercus suber forest in the eastern Iberian Peninsula

Question: Small and marginal forest populations are a focus of attention because of their high biodiversity value as well as the risk of population decline and loss. In this context, we ask to what extent a small, marginal Quercus suber (Cork oak) population located in the eastern Iberian Peninsula (Valencia, Spain) has the capacity for self‐regeneration and what are the factors that determine its recruitment variability. Location: Quercus suber forest in Pinet (Valencia, Spain). Methods: We performed a spatially explicit sampling both of the recruitment and of the potential parameters that could account for the recruitment variability. Using regression techniques we model the recruitment occurrence and abundance, and then we test to what extent the model obtained is still constrained by the spatial dependence. Results: Quercus suber recruitment density ranges from 0 to 18.66 individuals/25m² (mean = 1.46, SD = 2.8), with a very skewed distribution. Recruitment is similar under Q. suber forests and under Pinus forests, but it is almost absent under shrublands. Thus the parameters that explain most of the recruitment variability in local vegetation types are: the presence and cover of shrubs (negative relationship with recruitment), the basal area of Q. suber and Pinus and the amount of bare soil (all positively related to recruitment). These parameters are strongly related to the ecological processes driving recruitment (i.e. dispersal and predation) and they remove most of the spatial dependence of recruitment. Most recruiters, however, are small, forming a seedling bank rather than growing to successfully colonize new habitats. Conclusion: The results suggest that although recruitment densities are not very high, they do not limit potential regeneration in the Pinet Q. suber forest. However, successful regeneration is not observed. If we aim to increase the Pinet Q. suber population size, land management measures need to provide appropriate conditions for both seedling establishment in shrublands (e.g. shrub clearing) and seedling growth in woodlands (e.g. Pinus logging).     

Herbivores limit the population size of big‐leaf mahogany trees in an Amazonian forest

The Janzen–Connell hypothesis proposes that specialized herbivores maintain high numbers of tree species in tropical forests by restricting adult recruitment so that host populations remain at low densities. We tested this prediction for the large timber tree species, Swietenia macrophylla, whose seeds and seedlings are preyed upon by small mammals and a host‐specific moth caterpillar Steniscadia poliophaea, respectively. At a primary forest site, experimental seed additions to gaps – canopy‐disturbed areas that enhance seedling growth into saplings – over three years revealed lower survival and seedling recruitment closer to conspecific trees and in higher basal area neighborhoods, as well as reduced subsequent seedling survival and height growth. When we included these Janzen–Connell effects in a spatially explicit individual‐based population model, the caterpillar's impact was critical to limiting Swietenia's adult tree density, with a > 10‐fold reduction estimated at 300 years. Our research demonstrates the crucial but oft‐ignored linkage between Janzen–Connell effects on offspring and population‐level consequences for a long‐lived, potentially dominant tree species.     

Facilitation of Quercus ilex recruitment by shrubs in Mediterranean open woodlands

Question: Insufficient tree regeneration threatens the long‐term persistence of biodiverse Mediterranean open oak woodlands. Could shrubs, scarce due to decades of management (clearing and ploughing), facilitate holm oak recruitment at both acorn and seedling stages? Location: Open oak woodlands in Central Spain. Methods: Plots with four acorns were planted: (1) under the canopy of the spiny shrub Genista hirsuta, (2) in a small cage, protecting against ungulates, (3) in a shaded cage, protecting against ungulates and sun, and (4) in open grassland. Sets of these four treatments were spatially grouped according to a randomised block design, with 16 blocks near (< 10 m) and 16 away from (> 20 m) parent trees to test for distance‐related survival. Plots were regularly checked for seed removal. After emergence one seedling per plot (97 in total) was selected and its survival monitored. Results: Three months after sowing, 199 of 512 acorns were removed, predominantly by rodents. Acorn removal occurred at each treatment but was highest under shrubs. Eight months after sowing, seedling survival was highest under shrubs (50%), followed by shaded cages (16%), open grassland (4%) and cages (0%). Main mortality cause was drought (90%), killing most seedlings between June and July. No seedlings died from ungulate browsing. Conclusion: Shrubs demonstrated clear net facilitative effects for Quercus ilex recruitment, despite higher seed removal. Shading appears the crucial factor facilitating seedling survival. We therefore propose that lack of shrubs contributes largely to tree recruitment failure in Mediterranean open woodlands; management should aim at conserving shrubs.     

Feedbacks between canopy composition and seedling regeneration in mixed conifer broad‐leaved forests

To address the role of canopy‐seedling feedbacks in the structure and dynamics of mixed conifer broad‐leaved forests in the eastern US, we monitored seedling regeneration patterns and environmental conditions in the understorey of stands dominated by either hemlock (Tsuga canadensis) or red oak (Quercus rubra) for three years. Hemlock seedlings were favoured over other species’ seedlings in hemlock stands (a true positive feedback), due to a combination of high seed inputs, high seedling emergence and relatively high seedling survival during the growing season, which allowed hemlock to remain dominant under its own canopy. Red oak stands favoured a suite of mid‐successional broad‐leaved species over hemlock. A more even age structure of broad‐leaved species in red oak stands revealed that high seedling survival in such stands were driving this feedback. Canopy‐mediated variations in both understorey light availability (1.5% for hemlock vs 3.5% for red oak) and soil pH (3.9 for hemlock vs 4.4 for red oak) were found to be the primary correlates of stand‐level differences in seedling regeneration dynamics. In mixed temperate forests in the eastern US, canopy‐seedling feedbacks could act to slow successional trajectories and contribute to the maintenance of a stable landscape structure over many generations.     

Large-Scale Patterns of Quercus ilex, Quercus suber, and Quercus pyrenaica Regeneration in Central-Western Spain

In Central-Western Spain, forests and woodlands composed of Quercus sp. support outstanding levels of biodiversity, but there is increasing concern about their long-term persistence due to a lack of regeneration. We hypothesize that this regenerative lack is operating on a large geographic scale; that there are differences in the abundance of regeneration between three oak species; that oak regeneration is governed mainly by forest management and structure; and that shrubs act as important physical protectors of seedlings and saplings. We analyzed whether densities of oak seedlings and saplings in several size classes were related to stand-structure, understory, and physiographic variables potentially affecting regeneration. Data collected at a regional level (1 km × 1 km grid) by the Spanish Forest Inventory were evaluated from 2,816 plots. Results revealed that regeneration failure was common for all size categories, from small seedlings to large saplings, and for the three oak species studied, especially the evergreens. Of the Quercus ilex, Q. suber, and Q. pyrenaica plots studied, 49%, 62%, and 20% were lacking any small seedlings, and 82%, 96%, and 56% did not have any large saplings, respectively. Regeneration was positively correlated with tree cover and density, especially of small and medium-sized trees, and negatively correlated with the presence of large trees, indicating that regeneration failure is mostly associated with more open, uniform, and/or aged woodlands. Regeneration densities of Q. ilex and Q. suber were positively correlated with all understory variables, suggesting that the presence of pioneer shrubs represent a major safe site for early tree recruitment, independent from specific shrub species.