Edge Effects Act Differentially on Multiple Early Regeneration Stages of a Shade‐tolerant Tree Tapirira mexicana

Forest fragmentation is pervasive in tropical landscapes, and one pathway by which fragmentation may negatively impact populations is via edge effects. Early life‐stages are particularly important for species regeneration as they act as bottlenecks, but how edge effects may act differentially on different life‐stages is unknown. This study evaluated edge effects on multiple early life‐stages of a currently common animal‐dispersed, shade‐tolerant tree Tapirira mexicana (Anacardiaceae). The study was conducted in tropical premontane wet forest fragments in a highly deforested region of Costa Rica. The stages assessed were pre‐dispersal predation, primary dispersal, post‐dispersal predation, secondary dispersal, ex situ germination, in situ seed longevity, first and second year seedling abundance, second year seedling survivorship, and basal diameter growth. Results showed that impacts of edge effects were not equal across stages, but were limited to specific stages and times. One stage which may act as a bottleneck for species regeneration was pre‐dispersal predation. Over 60 percent of the seeds were predated by larvae, and predation was higher near the edge than interior habitat. Seeds lost viability within 10 d in the forest. Germination to first year seedling stage was also lower near edges, but such effect was eliminated within a year after that. Primary dispersal, seedling survivorship, and growth were not affected by proximity to edges, and both secondary dispersal and post‐dispersal predation were rare. This study demonstrates that current population abundance may not guarantee future species persistence and the importance of considering multiple life‐stages for a comprehensive assessment of forest fragmentation effects on species regeneration.     

Differential seedling regeneration patterns across forest–grassland ecotones in two tropical treeline species (Polylepis spp.)

Successful forest expansion into grassland can be limited by seed dispersal and adverse conditions for tree seedlings in the grassland environment. In the high‐elevation Andes, human‐induced fragmentation has exacerbated the patchy distribution of Polylepis forests, threatening their unique biological communities and spurring restoration interest. Studies of Polylepis forest extent in Peru suggest that forest borders have remained stable over the past century despite decreasing anthropogenic disturbance, suggesting that tree seedling recruitment is being limited in the open grassland habitat. We studied natural seedling dispersion patterns of Polylepis sericea and Polylepis weberbaueri (Rosaceae) at forest–grassland edges across a range of environmental conditions to examine seedling recruitment and colonization of grasslands in Huascaran National Park (Peru). Using data from 2367 seedlings found in 48 forest–grassland edge plots (15 m × 15 m) at forest patches between 3900–4500 masl, we employed generalized mixed modelling to identify the significant associations of seedling densities with environmental covariates. In addition, we compared these associations to patterns of adult presence on the landscape. Seedling densities were associated with a combination of variables varying within (distance to forest edge) and among (elevation and dry season solar irradiation) plots across the landscape. For both species, seedling densities decreased with increasing distance away from the forest in a manner consistent with short‐distance seed dispersal by wind. Our results suggest that such short‐distance dispersal may slow forest expansion, but that there also appear to be substantial post‐dispersal limitations to seedling establishment in the grassland. Polylepis sericea densities decreased with elevation, while P. weberbaueri increased with elevation and decreased with solar irradiation. Associations of adult presence with elevation and solar irradiation mirrored those of seedling densities. Management of areas with forest patches dominated by these species should consider these differences in their environmental tolerances, particularly during species selection and zonation for reforestation.     

Influence of within‐patch habitat quality on high‐Andean Polylepis bird abundance

Habitat restoration strategies for fragmented high Andean forest landscapes must consider the influence of within‐patch habitat quality on bird abundance. I examined vegetation and bird abundance at three locations within a highly fragmented Polylepis forest landscape in the Cordillera Vilcanota, southern Peru. Across the landscape, there was significant variation in the vegetation structure of Polylepis forest patches of different size categories, especially in terms of tree girth, tree height, tree density, and canopy vegetation structure. Principal Component Analysis extracted five factors of habitat quality, which together accounted for 74.2% of the variability within 15 habitat variables. Polylepis bird species differed in their responses to habitat quality but, overall, variation in Polylepis bird abundance was not fully captured by the range of habitat quality variables. Tall, dense vegetation cover was clearly important for 11 conservation‐important species, a high density of large trees was important for 10 species and primary forest ground cover was important for eight species. Habitat quality exhibited no significant influence on the abundance of only one species –Asthenes urubambensis. The abundance of seven species was associated with lower elevation forest, but only one species was associated with higher elevation forest. Management of habitat quality in large and medium remnant forest patches throughout the Cordillera Vilcanota, particularly in the 3800–4200 m elevation range, will be a cornerstone in ensuring the persistence of the majority of conservation‐important bird species populations.     

Low plant density enhances gene dispersal in the Amazonian understory herb Heliconia acuminata

In theory, conservation genetics predicts that forest fragmentation will reduce gene dispersal, but in practice, genetic and ecological processes are also dependent on other population characteristics. We used Bayesian genetic analyses to characterize parentage and propagule dispersal in Heliconia acuminata L. C. Richard (Heliconiaceae), a common Amazonian understory plant that is pollinated and dispersed by birds. We studied these processes in two continuous forest sites and three 1‐ha fragments in Brazil's Biological Dynamics of Forest Fragments Project. These sites showed variation in the density of H. acuminata. Ten microsatellite markers were used to genotype flowering adults and seedling recruits and to quantify realized pollen and seed dispersal distances, immigration of propagules from outside populations, and reproductive dominance among parents. We tested whether gene dispersal is more dependent on fragmentation or density of reproductive plants. Low plant densities were associated with elevated immigration rates and greater propagule dispersal distances. Reproductive dominance among inside‐plot parents was higher for low‐density than for high‐density populations. Elevated local flower and fruit availability is probably leading to spatially more proximal bird foraging and propagule dispersal in areas with high density of reproductive plants. Nevertheless, genetic diversity, inbreeding coefficients and fine‐scale spatial genetic structure were similar across populations, despite differences in gene dispersal. This result may indicate that the opposing processes of longer dispersal events in low‐density populations vs. higher diversity of contributing parents in high‐density populations balance the resulting genetic outcomes and prevent genetic erosion in small populations and fragments.     

Effects of altitude and livestock on the regeneration of two tree line forming <Emphasis Type="Italic">Polylepis</Emphasis> species in Ecuador

Regeneration is known to be limited at many temperate tree lines, but very little data is available on the impacts of altitude and anthropogenic disturbance on regeneration patterns along tropical tree lines. The study focused on the reproductive traits of two Polylepis species in the Páramo de Papallacta in Ecuador along an altitudinal gradient, and involved different intensities of cattle trampling within subsequent altitudinal ranges. We analyzed flowering, fruit set, seed viability, germination, and seedling establishment as well as stand structure of Polylepis incana and P. pauta. The numbers of P. incana inflorescences and seedlings per m2 showed a marginally significant decrease with increasing altitude. Mean tree height was significantly lower at higher altitudes, while stem number increased. The number of P. pauta inflorescences also decreased significantly upslope. In both forest types, trampling was found to have a positive impact on seedling abundance, presumably due to the removal of the litter layer. Thus, there was no evidence of negative effects of moderate cattle grazing on both tree line species. However, sapling establishment was minimal inside the forest stands at all altitudes and grazing levels, and we consequently observed a low proportion of narrow stems within all investigated forests. Our results show that, along with vegetative growth limitations of adult trees, important regeneration traits such as seedling and inflorescence numbers are also influenced by altitude, which might contribute to the formation of the upper tree line. Nevertheless, recruitment in the forest interior was low overall indicating that further factors, such as light conditions, affect regeneration of the studied species.     

Population Dynamics and Potential for Biological Control of an Exotic Invasive Shrub in Hawaiian Rainforests

Introduction of biological control agents to reduce the abundance of exotic invasive plant species is often considered necessary but risky. I used matrix projection models to investigate the current population dynamics of Clidemia hirta (Melastomataceae), an invasive shrub, in two rainforest stands on the island of Hawaii and to predict the efficacy of hypothetical biological control agents in reducing population growth rates. Stage-structured matrix models were parameterized with field data collected over 3 years from 2906 C. hirta plants in a recently invaded forest with an open overstory (Laupahoehoe) and 600 plants in a less recently invaded forest with a closed canopy (Waiakea). Asymptotic population growth rates (λ) for both populations in all years were greater than one, demonstrating that both populations were growing. Composite elasticities were high for the seedling life-history stage and fecundity, and near-term demographic elasticities suggested that changes in seedling survival would have the largest effect on population size in the short term. However, simulations showed that almost 100% of seedlings or new recruits produced per reproductive adult would have to be destroyed to cause populations to go locally extinct under current environmental conditions. Herbivores or pathogens that decrease survival across all vegetative stages by 12% at Waiakea and 64% at Laupahoehoe were projected to cause the populations to decline. Thus, biocontrol agents that reduce survival of multiple life-history stages rather than seed production should be pursued to control C. hirta in Hawaiian rainforests.     

Population stage structure,survival and recruitment in the endangered East African forest herb <Emphasis Type="Italic">Saintpaulia</Emphasis>

Climatic seasonality, local habitat quality, and edge effects created by forest fragmentation due to human activity may affect the performance of endangered rain forest understory herbs. Viability and seasonal dynamics of the populations of the endangered Saintpaulia confusa, S. difficilis, and S. grotei were studied in a protected seasonal submontane forest in NE Tanzania by examining plant life-history traits, population stage structure, and the effects of habitat quality on plant performance. The population stage structures were of the dynamic type. There was a lower frequency of seedlings (57.9%) and higher frequency of juvenile (13.3%) and adult plants (28.8%) in S. confusa than in S. difficilis (74%, 10.7% and 15.3%, respectively). Seedling recruitment occurred from May to August with an average of 54, 103 and 38 emerged seedlings per 1 m² study plot (S. confusa, S. difficilis, and S. grotei, respectively). Presence of the seed bank was also an indication of the regeneration potential of the populations. Mortality was high during the dry and hot season from December to March. Survival was lowest in juvenile plants, higher in sterile adult plants and the highest in fertile adult plants. Because survival was the lowest on dry substrates under open canopy, our data suggest that forest fragmentation, by reducing shade and humidity, will increase mortality in Saintpaulia. Furthermore, since the mortality was highest in seedlings and juveniles, forest fragmentation is likely to impede the regeneration of the Saintpaulia populations.     

Influence of Patch Factors and Connectivity on the Avifauna of Fragmented Polylepis Forest in the Ecuadorian Andes

Human‐induced alteration of habitat is a major threat to biodiversity worldwide, especially in areas of high biological diversity and endemism. Polylepis (Rosaceae) forest, a unique forest habitat in the high Andes of South America, presently occurs as small and isolated patches in grassland dominated landscapes. We examine how the avian community is likely influenced by patch characteristics (i.e., area, plant species composition) and connectivity in a landscape composed of patches of Polylepis forest surrounded by páramo grasslands in Cajas National Park in the Andes of southern Ecuador. We used generalized linear mixed models and an information‐theoretic approach to identify the most important variables probably influencing birds inhabiting 26 forest patches. Our results indicated that species richness was associated with area of a patch and floristic composition, particularly the presence of Gynoxys (Asteraceae). However, connectivity of patches probably influenced the abundance of forest and generalists species. Elsewhere, it has been proposed that effective management plans for birds using Polylepis should promote the conservation of mature Polylepis patches. Our results not only suggest this but also show that there are additional factors, such as the presence of Gynoxys plants, which will probably play a role in conservation of birds. More generally, these findings show that while easily measured attributes of the patch and landscape may provide some insights into what influences patch use by birds, knowledge of other factors, such as plant species composition, is essential for better understanding the distribution of birds in fragmented landscapes.     

Moms are better nurses than dads: gender biased self‐facilitation in a dioecious Juniperus tree

Questions: Can gender of nurse plants affect regeneration patterns and spatial population structure? Is there a seed‐seedling conflict in the regeneration process? What factors are responsible for the clumped spatial population structure observed for adult trees? Location: Mediterranean cold semi‐arid high mountains in Spain. Methods: The spatial pattern of adult Juniperus thurifera trees was studied by means of Ripley's K‐analysis. χ² analyses were used to test for natural seedling frequency in each of three main microhabitats: (1) under female and (2) male tree canopies and (3) in open interspaces. The observed pattern was explained experimentally by studying seed and seedling survival for two years. Survival probabilities were calculated across life stages for each of three main microhabitats. Results: Adult J. thurifera trees were aggregated in space. Most seedlings were found underneath female J. thurifera trees. Experimental studies demonstrated that from seed dispersal to seedling survival all life stages showed the same positive or negative trend within a given microhabitat, indicating stage coupling and no seed‐seedling conflicts. Attraction of frugivo‐rous birds by reproductive female junipers and improvement of environmental conditions beneath tree canopies were the main factors responsible for the variation in seedling density among microhabitats; highest underneath female trees and lowest in open interspaces. Conclusions: In dioecious species, the gender of nurse plants can significantly determine the spatial population structure. In J. thurifera forests, facilitation beneath female trees occurs among all life stages without any sign of seed‐seedling conflict. The most critical factors shaping the spatial population structure were directed seed dispersal and environmental amelioration beneath female conspecific trees.     

Insect Herbivory and Leaf Disease in Natural and Human Disturbed Habitats: Lessons from Early‐Successional Heliconia Herbs

The interaction of plants with insect herbivores and fungal pathogens can affect community dynamics, but there is little information on how this antagonistic interaction may be altered in human‐disturbed tropical systems. We examined whether the amount and quality of foliar damage on the pioneer herbs Heliconia latispatha and Heliconia collinsiana are distinct on road edges and secondary riparian vegetation compared with natural gaps in continuous forest (controls) in Mexico. We also investigated some physical and biological mechanisms that may jointly explain such differences. The overall insect damage in H. latispatha was similar between road edges and natural forest gaps (8.0% vs. 7.2% of leaf area). Damage by caterpillars, however, decreased from 4.2 percent in forest gaps to 0.5 percent on road edges, whereas damage by leaf‐cutting ants increased from 0 to 5.8 percent. In secondary riparian vegetation, where none of the leaves sampled were attacked by ants, overall herbivore damage in H. collinsiana was less than half that observed in forest gaps (3.0% vs. 6.7%), and driven mainly by differences in caterpillar damage (2.5% vs. 6.2%). By contrast, attack by leaf fungal pathogens was two to three times greater in both human‐disturbed habitats than in gaps (8.2–9.6% vs. 3.7–4.2%). Potential mechanisms underlying these differences involved human‐induced shifts in air and soil temperature driven by greater light availability, as well as changes in relative humidity, leaf toughness, foliar condensed tannins, and local abundance of herbivores. Our results indicate that human disturbance alters insect herbivory and may increase proliferation of leaf disease. Abstract in Spanish is available in the online version of this article.     

Testing the Home‐Site Advantage in Forest Trees on Disturbed and Undisturbed Sites

Restoration of plant populations is often undertaken using seed or plants from local sources because it is assumed they will be best adapted to the prevailing conditions. However, the effect of site disturbance on local adaptation has rarely been examined. We assessed local adaptation in three southwestern Australian forest tree species (Eucalyptus marginata, Corymbia calophylla, and Allocasuarina fraseriana) using reciprocal transplant trials at disturbed and undisturbed sites. Performance of plants within the trials was assessed over 2 years. Planting location accounted for the majority of the variation in most measures of performance, although significant variation of percent emergence among source populations was also detected. In all species, percent emergence and survival of plants sourced from Darling Range populations was significantly higher than that of plants from the Swan Coastal Plain, regions of contrasting edaphic and climatic environment. Survival of E. marginata over the first 18 months and emergence of C. calophylla were both higher in local plants, providing at least weak evidence for local adaptation. Where a local advantage was observed, the relative performance of local and nonlocal seed did not vary among disturbed and undisturbed sites. Evidence for enhanced establishment from local seed in at least one species leads us to recommend that where sufficient high‐quality seed supplies exist locally, these should be used in restoration. We also recommend longer‐term studies to include the possibility of local adaptation becoming evident at later life history stages.     

Contrasting Demographic Structure of Short‐ and Long‐lived Pioneer Tree Species on Amazonian Forest Edges

Although tropical forests have been rapidly converted into human‐modified landscapes, tree species response to forest edges remains poorly examined. In this study, we addressed four pioneer tree species to document demographic shifts experienced by this key ecological group and make inferences about pioneer response to forest edges. All individuals with dbh ≥ 1 cm of two short‐lived (Bellucia grossularioides and Cecropia sciadophylla) and two long‐lived species (Goupia glabra and Laetia procera) were sampled in 20 1‐ha forest edge plots and 20 1‐ha forest interior plots in Oiapoque and Manaus, Northeast and Central Amazon, respectively. As expected, pioneer stem density with dbh ≥ 1 cm increased by around 10–17‐fold along forest edges regardless of species, lifespan, and study site. Edge populations of long‐lived pioneers presented 84–94 percent of their individuals in sapling/subadult size classes, whereas edge populations of short‐lived pioneers showed 56–97 percent of their individuals in adult size classes. These demographic biases were associated with negative and positive net adult recruitment of long‐ and short‐lived pioneers, respectively. Our population‐level analyses support three general statements: (1) native pioneer tree species proliferate along forest edges (i.e., increased density), at least in terms of non‐reproductive individuals; (2) pioneer response to edge establishment is not homogeneous as species differ in terms of demographic structure and net adult recruitment; and (3) some pioneer species, particularly long‐lived ones, may experience population decline due to adult sensitivity to edge‐affected habitats.     

Accessibility predicts structural variation of Andean Polylepis forests

High Andean mountain forests, formed almost purely by trees of the genus Polylepis, occur nowadays as scattered remnant patches of a more continuous past distribution. Apparently, the destruction of Polylepis forests has mainly been caused by millennia of human disturbance, although forest distribution may also have fluctuated according to prevailing climatic conditions. Nowadays, the remaining Polylepis forest stands are still threatened by anthropogenic disturbance, which gradually degrades the forests. The aim of our study was to test if the structural variation of Polylepis forest patches, as an indication of forest degradation, can be predicted by accessibility to humans. The study was carried out in the Cordilleras Vilcanota and Vilcabamba, Cuzco, Peru. We used indices of forest biomass and proportion of vegetative regeneration as forest structural variables. First we examined the dependence of these variables on elevation with linear regressions. We did this separately for different Polylepis species and combining the species within humid and dry areas. Thereafter, we used the residual forest structural variation to assess possible relationships with accessibility, quantified as geographical distance to the nearest village, road or market centre. We found several significant relationships between the structural variables and accessibility, which may reflect different landscape related preferences in forest use. The results suggest accessibility can be used for rapid spatial prediction of Polylepis forest degradation, which facilitates identifying Polylepis forests that are potentially the most degraded and therefore in the most urgent need of restoration or conservation activities.     

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.     

Factors Limiting Post‐logging Seedling Regeneration by Big‐leaf Mahogany (Swietenia macrophylla) in Southeastern Amazonia,Brazil, and Implications for Sustainable Management1

Post‐logging seedling regeneration density by big‐leaf mahogany (Swietenia macrophylla), a nonpioneer light‐demanding timber species, is generally reported to be low to nonexistent. To investigate factors limiting seedling density following logging within the study region, we quantified seed production rates, germinability, dispersal patterns, and seed fates on the forest floor through germination and the first seedling growing season in southeastern Amazonia, Brazil. Fruit production rates were low by three logged and one unlogged populations compared to reports from other regions. Commercial‐sized trees (>60 cm diameter) were more fecund than noncommercial trees (30–60 cm diameter) at two sites, averaging 14.5 vs. 3.9 fruits/tree/year, respectively, at Marajoara, a logged site, over 8 yr. Fruit capsules contained an average of 60.3 seeds/fruit, 70 percent of which appeared viable by visual inspection. Sixty‐seven to 72 percent of apparently viable seeds germinated in nursery beds 2.5 mo after the dispersal period, when wet season rains began. Dry season winds blew most seeds west‐northwest of parent trees, with median dispersal distances of 28 and 9 m on west and east sides of parent trees, respectively. Nearly 100 percent of seeds fell within an area of 0.91 ha. On the forest floor beneath closed canopies, mammals, invertebrates, and fungal pathogens killed 40 percent of apparently viable seeds, while 36 percent germinated. Nine months after seedling establishment—midway through the first logging season following seed dispersal—14 percent of outplanted seeds survived as seedlings, representing 5.8 seeds/fruit. We conclude that seedlings are likely to survive in logging gaps at appreciable densities only in rare cases where previous year fruit production rates by logged trees were high (4–12.5% of commercial‐sized trees/year at Marajoara) and where tree crowns were felled in west or northwest directions.     

The Role of Natural Enemies in the Germination and Establishment of Pachira (Malvaceae) Trees in the Peruvian Amazon

Seeds from Pachira brevipes from white‐sand forest and Pachira insignis from clay forest were placed in a reciprocal transplant experiment that manipulated herbivore presence. Pachira insignis experienced 80 percent leaf loss in both habitats from herbivores, causing 79 percent mortality in white‐sand compared with 47 percent in clay in just 3 mo. Pachira brevipes suffered pathogen attack in both habitats that killed 90 percent of its establishing seedlings. Natural enemies thus strongly influence seedling recruitment of these two Pachira species in these forests and appear to have the largest impact within the seedlings' first few months.     

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).     

Demographic consequences of greater clonal than sexual reproduction in Dicentra canadensis

Clonality is a widespread life history trait in flowering plants that may be essential for population persistence, especially in environments where sexual reproduction is unpredictable. Frequent clonal reproduction, however, could hinder sexual reproduction by spatially aggregating ramets that compete with seedlings and reduce inter‐genet pollination. Nevertheless, the role of clonality in relation to variable sexual reproduction in population dynamics is often overlooked. We combined population matrix models and pollination experiments to compare the demographic contributions of clonal and sexual reproduction in three Dicentra canadensis populations, one in a well‐forested landscape and two in isolated forest remnants. We constructed stage‐based transition matrices from 3 years of census data to evaluate annual population growth rates, λ. We used loop analysis to evaluate the relative contribution of different reproductive pathways to λ. Despite strong temporal and spatial variation in seed set, populations generally showed stable growth rates. Although we detected some pollen limitation of seed set, manipulative pollination treatments did not affect population growth rates. Clonal reproduction contributed significantly more than sexual reproduction to population growth in the forest remnants. Only at the well‐forested site did sexual reproduction contribute as much as clonal reproduction to population growth. Flowering plants were more likely to transition to a smaller size class with reduced reproductive potential in the following year than similarly sized nonflowering plants, suggesting energy trade‐offs between sexual and clonal reproduction at the individual level. Seed production had negligible effects on growth and tuber production of individual plants. Our results demonstrate that clonal reproduction is vital for population persistence in a system where sexual reproduction is unpredictable. The bias toward clonality may be driven by low fitness returns for resource investment in sexual reproduction at the individual level. However, chronic failure in sexual reproduction may exacerbate the imbalance between sexual and clonal reproduction and eventually lead to irreversible loss of sex in the population.     

Sexual vs. vegetative reproduction in relation to forest dynamics in the understorey shrub,Hydrangea paniculata (Saxifragaceae)

In the forest understorey, shrubs usually reproduce vegetatively rather than sexually, but the relative contribution of these two reproductive modes may vary with temporal changes in environmental conditions (e.g., light, substrates of forest floor) that are closely related to forest dynamics. To evaluate the occasional changes in reproductive modes of the clonal understorey shrub Hydrangea paniculata, the reproductive characteristics (i.e., flowering, seedling establishment, vegetative propagation, reproductive size and age) were investigated in four different developmental stages in a Japanese beech forest (early gap, EG; late gap, LG; building, BU; mature, MA). Flowering individuals occurred only in EG and LG, and a much greater number of seedlings was observed in both EG and LG than in BU and MA, mainly because canopy gaps provided sufficient light for flowering and suitable substrates (fallen trees and exposed mineral-soil) for seedling establishment. The flowering individuals originated from plants that had persisted in the shaded understorey until gap formation. In contrast to sexual reproduction, clonal fragmentation increased with forest development in the order LG, EG, BU, MA. This is because, in later stages of forest dynamics (BU and MA), large individuals, some of which had previously flowered in gap stages, were layered on the forest floor and subsequently produced a substantial number of clonal fragments by separation of the buried branches from the main stems which had decayed. This resulted in a large number of clonal fragments originating from a single mother plant. These results suggest that the reproductive modes of H. paniculata is strongly influenced by the changes in environmental conditions with respect to the dynamics of canopy trees. Reproduction from seed in gap stages, despite their short period, provide new genets in established populations, whereas vegetative propagation via fragmentation in closed-canopy stages enhance the stability of the population until the next disturbance.     

Spatial patterns of tree recruitment in a relict population of Pinus uncinata: forest expansion through stratified diffusion

Aim To infer future changes in the distribution of isolated relict tree populations at the limit of a species’ geographical range, a deep understanding of the regeneration niche and the spatial pattern of tree recruitment is needed. Location A relict Pinus uncinata population located at the south‐western limit of distribution of the species in the Iberian System of north‐eastern Spain. Methods Pinus uncinata individuals were mapped within a 50 × 40‐m plot, and their size, age and reproductive status were estimated. Data on seed dispersal were obtained from a seed‐release experiment. The regeneration niche of the species was assessed based on the associations of seedling density with substrate and understorey cover. The spatial pattern of seedlings was described using point‐pattern (Ripley's K) and surface‐pattern (correlograms, Moran's I) analyses. Statistical and inverse modelling were used to characterize seedling clustering. Results Pine seedlings appeared aggregated in 6‐m patches. Inverse modelling estimated a longer mean dispersal distance (27 m), which corresponded to the size of a large cluster along the north to north‐eastward direction paralleled by an eastward trend of increasing seedling age. The two spatial scales of recruitment were related to two dispersal processes. The small‐scale clustering of seedlings was due to local seed dispersal in open areas near the edge of Calluna vulgaris mats: the regeneration niche. The long‐range expansion might be caused by less frequent medium‐distance dispersal events due to the dominant north‐westerly winds. Main conclusions To understand future range shifts of marginal tree populations, data on seed dispersal, regeneration niche and spatial pattern of recruitment at local scales should be obtained. The monitoring of understorey communities should be a priority in order to predict correctly shifts in tree species range in response to global warming.