Do consumer‐mediated negative effects on plant establishment outweigh the positive effects of a nurse plant?

Many studies demonstrated the importance of facilitative effect by nurse plant on seedling establishment. Few studies evaluated the negative effects of consumers on plant establishment under nurse plants by dealing with them during multiple demographic processes. We investigated the balance between the facilitative effect and negative effects of consumers during multiple demographic processes in Malawi in southeastern Africa. We chose Ficus natalensis as a nurse plant and compared it with three other microsites in tropical woodlands: Brachystegia floribunda (a dominant woodland species), Uapaca kirkiana (a woodland species), and a treeless site. We quantified the seed rain, postdispersal seed predation, germination, and seedling survival of Syzygium guineense ssp. afromontanum (a common forest species). Within each microsite, we quantified the overall probability of recruitment. We also measured seedling abundance of S. guineense ssp. afromontanum. We found that Ficus natalensis exerted both positive and negative impacts on the establishment of S. guineense ssp. afromontanum. Ficus natalensis facilitated seed deposition, seed germination, and seedling survival. On the other hand, seed removal at postdispersal stage was highest under F. natalensis. Interestingly, B. floribunda also had positive effects on germination and seedling survival, but not on seed deposition. When we excluded the seed arrival stage from our estimation of the recruitment probability, the highest value was found under B. floribunda, not under F. natalensis. When we included the seed arrival stage, however, the order of recruitment probability between F. natalensis and B. floribunda was reversed. The probability was one order of magnitude higher under F. natalensis than under B. floribunda. Our estimation of the probability which included the seed arrival stage was consistent with natural patterns of S. guineense ssp. afromontanum establishment. Despite the presence of opposite effects, the net effects of F. natalensis on S. guineense ssp. afromontanum recruitment in tropical woodlands can be positive.     

Stress‐Driven Changes in the Strength of Facilitation on Tree Seedling Establishment in West African Woodlands

The strength of competitive and facilitative interactions in plant communities is expected to change along resource gradients. Contrasting theoretical models predict that with increasing abiotic stress, facilitative effects are higher, lower, or similar than those found under more productive conditions. While these predictions have been tested in stressful environments such as arid and alpine ecosystems, they have hardly been tested for more productive African woodlands. We experimentally assessed the strength of tree seedling facilitation by nurse trees in mesic and dry woodlands in Benin, West Africa. We planted seedlings of the drought‐sensitive Afzelia africana and the drought‐tolerant Khaya senegalensis under three microsite conditions (closed woodland, woodland gap, and open fields). Seedling survival was greater within woodlands compared with open fields in both the mesic and dry woodlands. The relative benefits in seedling survival were larger at the dry site, especially for the drought‐sensitive species. Nevertheless, plant interactions became neutral or negative during the dry season in the drier woodland, indicating that the net positive effects may be lost under very stressful abiotic conditions. We conclude that facilitation also occurs in the relatively more productive conditions of African woodlands. Our results underscore the role of environmental variation in space and time, and the stress tolerance of species, in explaining competitive and facilitative interactions within plant communities. Abstract in French is available at http://www.blackwell‐synergy.com/loi/btp .     

Rain forest invasion of eucalypt-dominated woodland savanna, Iron Range, north-eastern Australia: I. Successional processes

Aim To explore successional processes associated with rain forest expansion in Eucalyptus‐dominated woodland savanna vegetation in north‐eastern Australia. Location Iron Range National Park and environs, northeast Queensland, Australia. This remote region supports probably the largest extent of lowland (< 300 m) rainforest remnant in Australia. Rainfall (c. 1700 mm p.a.) occurs mostly between November and June, with some rain typically occurring even in the driest months July–October. Methods (1) Sampling of rain forest seedling distributions, and other vegetation structural attributes, in fifteen 10 × 10 m quadrats distributed equi‐distantly between mature rain forest margins (range: 70–840 m), at each of 10 sites which were open‐canopied vegetation in 1943. (2) Assessment of relationships between rain forest seedling densities and structural characteristics, including distance‐to‐rain forest‐margin, canopy height, stem density. (3) Assessment of lifeform and dispersal spectra for defined vegetation structural types. Results Rates of rain forest invasion were found to be substrate‐mediated. Transects established on hematite schist, diorite, riverine alluvium, and granite developed closed canopies (termed phase III sites) by 1991. The remainder (four transects on poorly drained colluvial/alluvial sediments; one on dune sands) continued to occur either as grassy woodland (phase I), or with developing rain forest understoreys (phase II). Rain forest seedlings were observed at maximum sampled distances from mature rain forest margins at all sites. Lifeform and dispersal spectra data illustrated that: (1) the proportions of woodland trees, shrubs and graminoids declined with successional phase, with concomitant increases in rain forest primary trees and all other lifeform categories save rain forest trees; (2) the proportions of major dispersal syndromes did not vary between successional phases, neither for rain forest nor woodland taxa. Main conclusions Rain forest seedling distribution data for phases I and II sites illustrate three successional processes: margin extension – seedling density significantly negatively correlated with distance from mature rain forest margins at two sites; nucleation – seedling densities significantly positively correlated with tall trees at two sites; and irruption – seedling densities at two sites neither correlated with distance from mature rain forest margins, nor with measured vegetation structural features. The observation of irruptive rain forest regeneration at these sites, combined with decadal‐scale rain forest canopy development at the five remaining sites, illustrates that under conditions conducive to growth (moisture, substrate), low fire disturbance, and maintenance of diverse dispersal processes (high frugivore richness), rain forest can rapidly invade regional landscapes.     

Spatial distribution and regeneration of Quercus semecarpifolia and Quercus floribunda in a subalpine forest of western Himalaya, India

Studies were conducted on population status and regeneration of Quercus semecarpifolia and Quercus floribunda in a subalpine forest of western Himalaya. Saplings of Q. semecarpifolia belonging to the 51-100 cm size class and those of Q. floribunda belonging to the 101-150 cm size class were well represented in studied subalpine forests. Comparable seedling emergence and establishment of both species, either in the habitat dominated by same or different species seems to be an adaptational strategy of these species in subalpine forests. Seedling to tree ratio for Q. semecarpifolia was higher in the stand dominated by Q. floribunda, while comparable pattern was observed for Q. floribunda in the stand dominated by Q. semecarpifolia. Establishment of higher percent of seedlings of one species in the habitats dominated by another species of Quercus appears to indicate an oligarchic nature of oaks in subalpine forests. This study describes regeneration strategies, and spatial distribution of these species in a subalpine forest of Uttarakhand.     

Plant–plant interactions change during succession on nurse logs in a northern temperate rainforest

Plant–plant interactions change through succession from facilitative to competitive. At early stages of succession, early‐colonizing plants can increase the survival and reproductive output of other plants by ameliorating disturbance and stressful conditions. At later stages of succession, plant interactions are more competitive as plants put more energy toward growth and reproduction. In northern temperate rainforests, gap dynamics result in tree falls that facilitate tree regeneration (nurse logs) and bryophyte succession. How bryophyte‐tree seedling interactions vary through log succession remains unclear. We examined the relationships of tree seedlings, bryophyte community composition, bryophyte depth, and percent canopy cover in 166 1.0 m² plots on nurse logs and the forest floor in the Hoh rainforest in Washington, USA, to test the hypothesis that bryophyte‐tree seedling interactions change from facilitative to competitive as the log decays. Tree seedling density was highest on young logs with early‐colonizing bryophyte species (e.g., Rhizomnium glabrescens) and lowest on decayed logs with Hylocomium splendens, a long‐lived moss that reaches depths >20 cm. As a result, bryophyte depth increased with nurse log decay and was negatively associated with tree seedling density. Tree seedling density was 4.6× higher on nurse logs than on the forest floor, which was likely due to competitive exclusion by forest floor plants, such as H. splendens. Nurse logs had 17 species of bryophytes while the forest floor had six, indicating that nurse logs contribute to maintaining bryophyte diversity. Nurse logs enable both tree seedlings and smaller bryophyte species to avoid competition with forest floor plants, including the dominant bryophyte, H. splendens. H. splendens is likely a widespread driver of plant community structure given its dominance in northern temperate forests. Our findings indicate that plant–plant interactions shift with succession on nurse logs from facilitative to competitive and, thus, influence forest community structure and dynamics.     

Floristic composition and structure of vegetation under isolated trees in neotropical pastures

Abstract. Large isolated trees are a common feature of the agricultural landscape in humid tropical regions originally covered by rain forest. These isolated trees are primarily used as a source of shade for cattle and people. 13 pastures (totalling ca. 80 ha) currently used as cattle pasture were studied. In them, we registered 265 isolated trees belonging to 57 species. 50 trees of the most frequent species (Ficus spp. n = 30 and Nectandra ambigens n = 20) were selected to examine the influence of isolated trees on floristic composition and vegetation structure in the pastures. At each tree, three 4–m² quadrats were sampled: under the canopy, directly under the canopy perimeter, and beyond the canopy in the open pasture. Under-canopy vegetation was structurally and floristically different from the other two sampling sites. Mean species richness per quadrat was significantly higher under the canopy (17.8 ± 4.3 SD) than at the canopy perimeter (11.2 ± 3.4) and in the open pasture (10.6 ± 3.6) sites. Stem density was higher at under-canopy sites, where greater proportions of endozoochorous and rain-forest species were found. Isolated trees function as nursery plants for rain-forest species by facilitating the establishment of zoochorous species whose seeds are deposited under the tree canopies by frugivorous birds or bats. Our results imply that isolated trees may play a major role in seed dispersal and establishment of native species, which is of consequence for the preservation of rain-forest species in these fragmented landscapes.     

Understorey gaps influence regeneration dynamics in subtropical coastal dune forest

Dominant understorey species influence forest dynamics by preventing tree regeneration at the seedling stage. We examined factors driving the spatial distribution of the monocarpic species Isoglossa woodii, a dominant understorey herb in coastal dune forests, and the effect that its cover has on forest regeneration. We used line transects to quantify the area of the forest understorey with I. woodii cover and with gaps in the cover. Paired experimental plots were established in semi-permanent understorey gaps with I. woodii naturally absent and in adjacent areas with I. woodii present to compare plant community composition, soil, and light availability between the two habitats. Isoglossa woodii was widespread, covering 65–95% of the understorey, while gaps covered the remaining 5–35% of the area. The spatial distribution of this species was strongly related to tree canopy structure, with I.␣woodii excluded from sites with dense tree cover. Seedling establishment was inhibited by low light availability (<1% of PAR) beneath I.␣woodii. When present, I. woodii reduced the density and species richness of tree seedlings. The tree seedling community beneath I. woodii represented a subset of the seedling community in gaps. Some species that were found in gaps did not occur beneath I. woodii at all. There were no significant differences between the sapling and canopy tree communities in areas with I. woodii gaps and cover. In the coastal dune forest system, seedling survival under I. woodii is dependent on a species’ shade tolerance, its ability to grow quickly during I. woodii dieback, and/or the capacity to regenerate by re-sprouting and multi-stemming. We propose a general conceptual model of forest regeneration dynamics in which the abundant understorey species, I. woodii, limits local tree seedling establishment and survival but gaps in the understorey maintain tree species diversity on a landscape scale.     

Density-dependent shift from facilitation to competition in a dwarf <Emphasis Type="Italic">Avicennia germinans</Emphasis> forest

The global effort to rehabilitate and restore destroyed mangrove forests is unable to keep up with the high mangrove deforestation rates, which exceed the average pace of global deforestation. Although facilitation theory presents new possibilities for the restoration of heavily degraded mangrove sites, knowledge of tree–tree interactions in stressed mangrove forest ecosystems is too limited to utilize facilitation appropriately. The aim was to determine the mode of local interaction among stressed mangrove trees by investigating the effect of clustering on tree size and crown morphology under contrasting stand densities. The study was conducted in a dwarf Avicennia germinans forest in Northern Brazil, in which tree growth is limited by infrequent inundation and high pore-water salinity. Autoregressive regression, Voronoi tessellation and spatial point pattern statistics were used to address the spatial processes underlying tree interaction. Under low stand density (1.2 trees m^?2) dwarf trees which grew in clustered cohorts of A. germinans had a less stunted crown morphology revealing the dominance of a positive neighborhood influence among plants. In contrast, dwarf trees in the denser forest stand (2.7 trees m^?2) were interacting competitively as indicated by the more negative effect of neighbors on crown morphology and size. The shift from facilitative to competitive interactions is an important feature of mangrove forest regeneration under harsh environmental conditions. If mangrove trees are unable to regenerate naturally on severely degraded sites, intraspecific facilitation could be used to assist regeneration by planting seedlings in clusters and not evenly spaced.     

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.     

Absence of Howlers (<Emphasis Type="Italic">Alouatta palliata</Emphasis>) Influences Tree Seedling Densities in Tropical Rain Forest Fragments in Southern Mexico

The disappearance of frugivorous primates in fragmented forests can potentially lower the rates of seed dispersal and recruitment of endozoochorous tree species, thus altering plant community structure. We quantified seedling density for 7 tree species that are common in the feces of mantled howlers (Alouatta palliata) in 6 rain forest fragments in northern Chiapas, Mexico. Howlers were present in 3 of the fragments and absent in the other 3. We compared seedling density in primate sleeping sites in inhabited fragments with control sites, which were structurally similar to sleeping sites but where we did not find monkey feces, in both inhabited and uninhabited fragments. For each tree species, we determined the relationship between seedling density and the local density of seeds and adult trees. In fragments where howlers were present, seedling density for 4 of the focal tree species (Brosimum alicastrum, Dialium guianense, Manilkara zapota, and Nectandra ambigens) was greater in sleeping sites than in control sites found in the same fragments. Moreover, seedling density of Dialium guianense was greater in the control sites of fragments inhabited by howlers than in fragments where this primate is absent. Seedling density of these 4 species correlates positively with seed density on the forest floor; however, we observed no correlations between seedling density and the density of adult trees. Our results suggest that the diversity of the seedling community of tree species dispersed by howlers may decline in fragments where this seed disperser is absent. These findings, together with the fact that only 5% of the study region is currently covered by forest and 81% of the forest remnants are uninhabited by mantled howlers, suggest that the potential long-term recovery of associated populations of tropical tree species dispersed by this primate species is highly uncertain. Conservation and restoration efforts should be aimed at restoring or replacing the ecological role played by this important seed disperser in the region.     

Variation in structural diversity and regeneration potential of tree species in different tropical forest types of Similipal Biosphere Reserve,Eastern India

Understanding the regeneration potential of tree species in natural forest ecosystems is crucial to deliver suitable management practices for conservation of biodiversity. We studied the variation in structural diversity and regeneration potential of tree species in three different tropical forest types, namely: Dry Deciduous forest (DDF), Moist Deciduous forest (MDF) and Semi-evergreen forest (SEF) of Similipal Biosphere Reserve (SBR), Eastern India. Random sample plots were laid for studying the diversity and distribution pattern of tree, sapling, and seedling stages of the tree species. A total of 84 species belong to 73 genera and 35 families were recorded from the study area. The highest species richness was reported for tree (54 species) in DDF, sapling (24 species) in MDF and seedling (22 species each) in SEF and DDF. The overall density of trees with GBH (Girth at Breast Height) ≥ 10 cm was 881 individuals/ha. The regeneration potential of tree species was poor in DDF (39%) where as it was fair in SEF (43%) and MDF (49%). Most of the dominant tree species at each forest type performed good regeneration. The species such as Ehretia laevis Roxb., Bridelia retusa (L.)A.Juss., Mitragyna parviflora (Roxb.) Korth., Terminalia tomentosa Wight & Arn., Terminalia chebula Retz., Terminalia bellirica (Gaertn.) Roxb.etc. had either no regeneration or poor regeneration potential need immediate attention for conservation measures. The diversity of standing trees did not correlate with seedling or sapling diversity in all the cases but there was significant correlation among seedling and sapling diversity found in DDF (r = 0.67, p ≤ 0.05) and SEF (r = 0.83, p ≤ 0.05). Further, the diversity of tree species increased with their age (trees > saplings > seedlings) and the stem density decreased with their age (trees < saplings < seedlings) in all three forest types. The results of our study would be helpful in understanding the structural attributes, diversity and regeneration potential of different tropical forest types of India for their better conservation and management.     

Tree Regeneration in Church Forests of Ethiopia: Effects of Microsites and Management

Tree regeneration is severely hampered in the fragmented afromontane forests of northern Ethiopia. We explored how trees regenerate in remnant forests along the gradient from open field, forest edge to closed sites and canopy gaps inside the forest. We investigated the effects of seed sowing, litter removal, and weeding on the regeneration success along this gradient. Regeneration success was investigated for four indigenous tree species, and measured in terms of seedling establishment, growth, and survival. Species performed differently according to site conditions. Within the forest, local canopy openings facilitated seed germination ( Ekebergia ), seedling growth (all species except Olea ), or survival ( Ekebergia and Olea ), suggesting that all species benefited from local high light conditions in the forest. Outside the forest, germination (all species) and growth rates ( Juniperus and Olea ) were lower in the open field, most probably due to water stress in the dry season. Outer edge conditions favored growth for three of the four species. Natural seed germination was, however, zero at any site for Juniperus and Olea and low for Ekebergia and Prunus in the open field. Soil scarification influenced germination positively, while weeding did not have a positive effect. These results suggest that simple measures may improve seedling establishment, and that, for some species, forest edges are particularly useful for growth and survival after succesful establishment. Together with erecting fences, needed to protect seedlings against grazing, seed sowing, planting seedling, and soil scarification may contribute to maintain and restore church forests in the fragmented landscapes of northern Ethiopia.     

Oomycetes associated with Prunus serotina persist in soil after tree harvest

Soil-borne pathogens can shape forest communities by lowering seedling survivorship. Many soil pathogens can persist long-term as survival spores, but how long pathogens outlive tree hosts in gap soils and whether they continue to affect seedling survival is uncertain. We studied the presence of oomycetes and evaluated seedling performance in soils near live Prunus serotina trees, and 0.5 and 1.5-y-old stumps. We isolated five species of oomycetes from soils, two of which were pathogenic (Pythium intermedium and Pythium irregulare) to Prunus serotina. There was a non-significant ~10.5% increase in conspecific seedling survival in stumps versus live trees, and pathogens were present in soils of all stump ages. The continued presence of pathogens of Prunus serotina in gap soils demonstrates the potential for impacts on conspecific regeneration after tree death, though the slight improvement in survival suggests that these effects may weaken with time.     

Nucleation-driven regeneration promotes post-fire recovery in a Chilean temperate forest

Nucleation is a successional process in which extant vegetation facilitates seed dispersal and recruitment of other individuals and species around focal points in the landscape, leading to ecosystem recovery. This is an important process in disturbed sites where regeneration is limited by abiotic conditions or restrictive seed dispersal. We investigated forest recovery in a large burned area of evergreen temperate rainforest in southern Chile subjected to seasonal soil waterlogging, and assessed the relevance of nucleation processes in overcoming biotic and physical barriers for tree species regeneration. We measured richness and abundance of woody species in relation to patch size, as well as abiotic factors such as light and soil moisture within and outside patches. We found higher tree regeneration in existing patches than in open areas. We recorded an increase of patch size over time, associated with the increase in number of individuals and tree species. Soils in open areas were waterlogged, especially in winter, while patches were not. Trees in patches also acted as perches, enhancing bird-mediated seed rain. Seeds of fleshy-fruited tree species arrived first at patches and seedlings were more frequent in smaller, younger patches, while the number of seedlings of trees with wind-dispersed seeds increased in larger, older patches. Our study shows that woody species seem incapable of recruiting in open and waterlogged soils and depend strongly on extant vegetation patches to establish. In this fire-disturbed evergreen temperate forest regeneration occurs via nucleation, where new individuals contribute to a centrifugal kind of patch growth.     

Interspecific variation in clonality in temperate lianas revealed by genetic analysis: Do clonal proliferation processes differ among lianas?

Lianas are important components of forest communities with a considerable impact on overall forest structure and function. Lianas are characterized by extensive clonal reproduction on the forest floor, which is important for their establishment and growth. Previous studies have suggested that clonal reproduction strategies vary substantially among liana species; however, few studies have quantitatively evaluated the clonality strategy in multiple co-occurring liana species. The primary objective of the present study was to evaluate the relative contribution of clonal reproduction and to understand the clonal proliferation processes in co-occurring liana species by assessing both small stems on the forest floor and mature stems that climbed trees. The clonal reproduction strategy in four common liana species (stem twiner: Wisteria floribunda; root climbers: Schizophragma hydrangeoides, Euonymus fortunei and Rhus ambigua) in a 6-ha plot and a belt transect within an old-growth temperate forest in central Japan was evaluated using genetic analysis. The contribution of clonal reproduction was smaller in root climbers than in W. floribunda. All W. floribunda genets with small ramets in the understory had genetically identical ramets that climbed trees, whereas few such ramets were found in root climbers. This indicates that W. floribunda mature ramets laterally produce small ramets via clonal proliferation, whereas seedlings of root climbers grow horizontally to climb trees. The results indicate that the clonal reproduction processes differ greatly among lianas and the clonal growth in lianas plays a contrasting role in their life-history strategy.     

Spatial Scales of Pollen and Seed-Mediated Gene Flow in Tropical Rain Forest Trees

Gene flow via seed and pollen is a primary determinant of genetic and species diversity in plant communities at different spatial scales. This paper reviews studies of gene flow and population genetic structure in tropical rain forest trees and places them in ecological and biogeographic context. Although much pollination is among nearest neighbors, an increasing number of genetic studies report pollination ranging from 0.5–14 km for canopy tree species, resulting in extensive breeding areas in disturbed and undisturbed rain forest. Direct genetic measures of seed dispersal are still rare; however, studies of fine scale spatial genetic structure (SGS) indicate that the bulk of effective seed dispersal occurs at local scales, and we found no difference in SGS (Sp statistic) between temperate (N?=?24 species) and tropical forest trees (N?=?15). Our analysis did find significantly higher genetic differentiation in tropical trees (F _ST?=?0.177; N?=?42) than in temperate forest trees (F _ST?=?0.116; N?=?82). This may be due to the fact that tropical trees experience low but significant rates of self-fertilization and bi-parental inbreeding, whereas half of the temperate tree species in our survey are wind pollinated and are more strictly allogamous. Genetic drift may also be more pronounced in tropical trees due to the low population densities of most species.     

Seedling and Sapling Dynamics of Treefall Pits in Puerto Rico1

Seedling and sapling dynamics in a Puerto Rican rain forest were compared between forest understory and soil pits created by the uprooting of 27 trees during Hurricane Hugo. Soil N and P, organic matter, and soil moisture were lower and bulk densities were higher in the disturbed mineral soils of the pits than in undisturbed forest soils ten months after the hurricane. No differences in N and P levels were found in pit or forest soils under two trees with N–fixing symbionts (Inga laurina and Ormosia krugii) compared to soils under a tree species without N–fixing sym–bionts (Casearia arborea), but other soil variables (Al, Fe, K) did vary by tree species. Forest plots had greater species richness of seedlings (<10 cm tall) and saplings (10–100 cm tall) than plots in the soil pits (and greater sapling densities), but seedling densities were similar between plot types. Species richness and seedling densities did not vary among plots associated with the three tree species, but some saplings were more abundant under trees of the same species. Pit size did not affect species richness or seedling and sapling densities. Recruitment of young Cecropia schreberiana trees (>5 m tall) 45 months after the hurricane was entirely from the soil pits, with no tree recruitment from forest plots. Larger soil pits had more tree recruitment than smaller pits. Defoliation of the forest by the hurricane created a large but temporary increase in light availability. Recruitment of C. schreberiana to the canopy occurred in gaps created by the treefall pits that had lower soil nutrients but provided a longer–term increase in light availability. Treefall pits also significantly altered the recruitment and mortality of many understory species in the Puerto Rican rain forest but did not alter species richness.     

Seasonal leaf dynamics across a tree density gradient in a Brazilian savanna

Interactions between trees and grasses that influence leaf area index (LAI) have important consequences for savanna ecosystem processes through their controls on water, carbon, and energy fluxes as well as fire regimes. We measured LAI, of the groundlayer (herbaceous and woody plants <1-m tall) and shrub and tree layer (woody plants >1-m tall), in the Brazilian cerrado over a range of tree densities from open shrub savanna to closed woodland through the annual cycle. During the dry season, soil water potential was strongly and positively correlated with grass LAI, and less strongly with tree and shrub LAI. By the end of the dry season, LAI of grasses, groundlayer dicots and trees declined to 28, 60, and 68% of mean wet-season values, respectively. We compared the data to remotely sensed vegetation indices, finding that field measurements were more strongly correlated to the enhanced vegetation index (EVI, r ²=0.71) than to the normalized difference vegetation index (NDVI, r ²=0.49). Although the latter has been more widely used in quantifying leaf dynamics of tropical savannas, EVI appears better suited for this purpose. Our ground-based measurements demonstrate that groundlayer LAI declines with increasing tree density across sites, with savanna grasses being excluded at a tree LAI of approximately 3.3. LAI averaged 4.2 in nearby gallery (riparian) forest, so savanna grasses were absent, thereby greatly reducing fire risk and permitting survival of fire-sensitive forest tree species. Although edaphic conditions may partly explain the larger tree LAI of forests, relative to savanna, biological differences between savanna and forest tree species play an important role. Overall, forest tree species had 48% greater LAI than congeneric savanna trees under similar growing conditions. Savanna and forest species play distinct roles in the structure and dynamics of savanna–forest boundaries, contributing to the differences in fire regimes, microclimate, and nutrient cycling between savanna and forest ecosystems.     

Habitat fragmentation changes the adaptive value of seed mass for the establishment of a tropical canopy tree

To cope with the limiting light conditions in the rain forest understory, many tropical tree species have evolved large seeds that provide the emerging seedlings with nutritional reserves. Habitat fragmentation might change the adaptive value of seed size by modifying the biotic and physical conditions of the forest understory. We experimentally assessed the potential of fragmentation to alter how seed mass affects seedling survival, vigor, and attack by natural enemies of the tropical tree Nectandra ambigens. Seeds from different mother trees (families) were individually weighed and sown in experimental sites established in continuous forest and in forest fragments. Seedling survival, vigor, and damage by herbivores and pathogens were recorded periodically. While seedlings derived from larger seeds had higher survival rates in both habitats, seedling survival and vigor were significantly greater in forest fragments, and the seedlings also suffered fewer attacks by natural enemies. We found genetic variance for seed mass among families with a heritability value (h²) of 0.66, and we found evidence for selection on seed size. Average seed size differed between dead and living seedlings in three sites. In one fragment, seed size was selectively neutral in relation to survival. Overall, selection for seed size promoted survival and thus appears to affect the distribution of this trait. The maintenance of genetic variance could be related to the stochastic nature of the formation of light gaps. Our results highlight the importance of evaluating the adaptive value of traits susceptible to environmental changes for conservation purposes.     

A Neighborhood Analysis of the Consequences of Quercus suber Decline for Regeneration Dynamics in Mediterranean Forests

In forests, the vulnerable seedling stage is largely influenced by the canopy, which modifies the surrounding environment. Consequently, any alteration in the characteristics of the canopy, such as those promoted by forest dieback, might impact regeneration dynamics. Our work analyzes the interaction between canopy neighbors and seedlings in Mediterranean forests affected by the decline of their dominant species (Quercus suber). Our objective was to understand how the impacts of neighbor trees and shrubs on recruitment could affect future dynamics of these declining forests. Seeds of the three dominant tree species (Quercus suber, Olea europaea and Quercus canariensis) were sown in six sites during two consecutive years. Using a spatially-explicit, neighborhood approach we developed models that explained the observed spatial variation in seedling emergence, survival, growth and photochemical efficiency as a function of the size, identity, health, abundance and distribution of adult trees and shrubs in the neighborhood. We found strong neighborhood effects for all the performance estimators, particularly seedling emergence and survival. Tree neighbors positively affected emergence, independently of species identity or health. Alternatively, seedling survival was much lower in neighborhoods dominated by defoliated and dead Q. suber trees than in neighborhoods dominated by healthy trees. For the two oak species, these negative effects were consistent over the three years of the experimental seedlings. These results indicate that ongoing changes in species’ relative abundance and canopy trees’ health might alter the successional trajectories of Mediterranean oak-forests through neighbor-specific impacts on seedlings. The recruitment failure of dominant late-successional oaks in the gaps opened after Q. suber death would indirectly favor the establishment of other coexisting woody species, such as drought-tolerant shrubs. This could lead current forests to shift into open systems with lower tree cover. Adult canopy decline would therefore represent an additional factor threatening the recruitment of Quercus forests worldwide.