Population Status and Reproductive Success of an Endangered Epiphytic Orchid in a Fragmented Landscape

Habitat fragmentation and disturbance are two of the most significant drivers of species extinctions in plant populations. The degree of impact of fragmentation on plant populations depends on the level of specificity of plant–animal interactions, as well as on the availability of suitable sites for seedling recruitment. In this study, we describe the population density and structure, pollen limitation and reproductive success of the endangered tropical orchid Myrmecophila christinae, an epiphytic species with a specialized pollination system. We surveyed a total of 14 populations located in a fragmented landscape. Seedling density was related to habitat disturbance and host plant density; while density of juveniles was related to density of adults. Adult and total individual densities were related to habitat affectation. We also found that fragments <1 ha had significantly fewer seedlings, as well as an over‐representation of large adults. On the other hand, fruit production was higher in fragments >10 ha, and fruit set was significantly lower in highly disturbed fragments. Hand pollination experiments showed that M. christinae was pollen limited in all the studied populations, suggesting that pollen limitation is unrelated to habitat disturbance. Overall, our results suggest that fragmentation has affected key demographic features of M. christinae, including reproduction and recruitment.     

Cattle affect regeneration of the palm species Attalea princeps in a Bolivian forest–savanna mosaic

Attalea princeps is an important palm species that shapes the forest–savanna mosaic in Beni, Bolivia, as it dominates the two principal forest landscape elements (forest islands and gallery forest), and provides a vital microhabitat, food, and nesting source for numerous plant and animal species. The forest–savanna mosaic is used for extensive grazing, and the palm population is declining on the forest islands due to a low regeneration rate, which threatens the maintenance of this landscape. We therefore examined the (a)biotic factors that influence the population structure of Attalea in the centers and edges of forest islands and gallery forests. Ninety‐one 0.1‐ha plots were established, and 500 palm adults and 3,700 juveniles were measured for their size, health condition, and fire damage. For each plot, habitat characteristics, such as landscape position, grazing pressure, and soil conditions, were measured. Attalea population density was significantly lower on the forest islands than in the gallery forests, especially in the juvenile life stage. A structural equation model showed that juvenile density is positively related to the health condition of juveniles and amount of fruits present, where the amount of fruits is positively affected by the condition of adults. Juvenile density is negatively influenced by grazing, affecting the health condition of the juvenile, as well as organic matter and phosphate availability in the soil. Therefore, it is recommended to decrease the grazing pressure by decreasing livestock densities, fencing off vulnerable forest islands, or by rotating cattle.Abstract in Spanish is available with online material.     

Population structure and adult plant performance of forest herbs in three contrasting habitats

In Flanders, forests have been subjected to subsequent periods of reclamation and reforestation, leading to a mosaic of forest patches of various ages and with different abiotic conditions, depending on the duration and intensity of the past agricultural activities. While extensive research has been done on the relative importance of dispersal limitation and establishment limitation in explaining colonization patterns of so-called ancient forest plant species, little is known about the viability of populations of these species once they have colonized recent forests. Therefore, a comparative study on population structure and plant performance was carried out for three forest plant species with different colonization capacity ( Primula elatior , Geum urbanum and Arum maculatum ) in three different habitat types: ancient forests, recently established forests and small landscape elements (hedgerows). Contrary to the expectation, it was shown that adult plant performance – a combination of vegetative as well as reproductive characteristics – of the three species was generally higher in recent forests and small landscape elements. This also led to higher seedling densities and proportions in these habitat types. Adult densities, however, were extremely low in newly established forests, suggesting reduced seedling and juvenile survival under these circumstances, probably due to higher light availability and hence increased competition from species like Urtica dioica . Although there might be no immediate establishment limitation, installing a shrub layer in recent forests might enhance survival and result in a stable population structure. Hedgerows on the other hand seemed to contain viable populations of certain forest plant species and can therefore function as an important colonization source when incorporated in recently established forests.     

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.     

Applying a continua landscape approach to evaluate plant response to fragmentation: Primula vulgaris in the Cantabrian mountains

Question: Continua landscape approaches conceptualize the effects of habitat fragmentation on the biota by considering fragmented landscapes as continuous gradients, departing from the view of habitat as either suitable (fragment) or unsuitable (matrix). They also consider the ecological gradients or the ‘Umwelt’ (species‐specific perception of the landscape) to represent the processes that ultimately limit organisms' ability to colonize and persist within habitat remnants. Are these approaches suitable for evaluating the response of plant species to fragmentation? Location: Fragmented mid‐elevation temperate forests, Cantabrian range, Spain. Methods: The presence, abundance and demographic structure of populations of the perennial herb Primula vulgaris were sampled across a continuous extent of 100 ha, subdivided into 400 50 m × 50 m sampling units. These variables were related to forest availability, forest subdivision and edge density, topography and the spatial clumpiness of populations (a measure of plant dispersal constraints and, hence, a major surrogate of plant Umwelt). Results: Fragmentation processes, especially habitat loss, negatively affect P. vulgaris, with a stronger effect on presence than on abundance and demography. Despite the importance of habitat availability, P. vulgaris does not occupy all potentially suitable forest habitat, mostly owing to dispersal constraints. A positive effect of slope on plant presence also suggests some effect of habitat quality in determining establishment and occupancy of forest landscape. Conclusions: Within‐habitat dispersal constraints are as important as forest fragmentation in determining the landscape‐scale distribution of P. vulgaris. By assessing the relative role of the diverse fragmentation processes, and of the species' landscape perception, a continua landscape approach proves to be a valuable tool for predicting plant response to landscape change.     

What explains landscape patterns of tree mortality caused by bark beetle outbreaks in Greater Yellowstone?

Aim Bark beetle outbreaks have recently affected extensive areas of western North American forests, and factors explaining landscape patterns of tree mortality are poorly understood. The objective of this study was to determine the relative importance of stand structure, topography, soil characteristics, landscape context (the characteristics of the landscape surrounding the focal stand) and beetle pressure (the abundance of local beetle population eruptions around the focal stand a few years before the outbreak) to explain landscape patterns of tree mortality during outbreaks of three species: the mountain pine beetle, which attacks lodgepole pine and whitebark pine; the spruce beetle, which feeds on Engelmann spruce; and the Douglas‐fir beetle, which attacks Douglas‐fir. A second objective was to identify common variables that explain tree mortality among beetle–tree host pairings during outbreaks. Location Greater Yellowstone ecosystem, Wyoming, USA. Methods We used field surveys to quantify stand structure, soil characteristics and topography at the plot level in susceptible stands of each forest type showing different severities of infestation (0–98% mortality; n= 129 plots). We then used forest cover and beetle infestation maps derived from remote sensing to develop landscape context and beetle pressure metrics at different spatial scales. Plot‐level and landscape‐level variables were used to explain outbreak severity. Results Engelmann spruce and Douglas‐fir mortality were best predicted using landscape‐level variables alone. Lodgepole pine mortality was best predicted by both landscape‐level and plot‐level variables. Whitebark pine mortality was best – although poorly – predicted by plot‐level variables. Models including landscape context and beetle pressure were much better at predicting outbreak severity than models that only included plot‐level measures, except for whitebark pine. Main conclusions Landscape‐level variables, particularly beetle pressure, were the most consistent predictors of subsequent outbreak severity within susceptible stands of all four host species. These results may help forest managers identify vulnerable locations during ongoing outbreaks.     

Seedling regeneration,environment and management in a semi‐deciduous African tropical rain forest

Questions: How is seedling regeneration of woody species of semi‐deciduous rain forests affected by (a) historical management for combinations of logging, arboricide treatment or no treatment, (b) forest community type and (c) environmental gradients of topography, light and soil nutrients? Location: Budongo Forest Reserve, Uganda. Methods: Seedling regeneration patterns of trees and shrubs in relation to environmental factors and historical management types were studied using 32 0.5‐ha plots laid out in transects along a topographic gradient. We compared seedling species diversity, composition and distribution patterns along topographic gradients and within types of historical management regimes and forest communities to test whether environmental factors contributed to differences in species composition of seedlings. Results: A total of 85 624 woody seedlings representing 237 species and 46 families were recorded in this rain forest. Cynometra alexandri C.H. Wright and Lasiodiscus mildbraedii Engl. had high seedling densities and were widely distributed throughout the plots. The most species‐rich families were Euphorbiaceae, Fabaceae, Rubiaceae, Meliaceae, Moraceae and Rutaceae. Only total seedling density was significantly different between sites with different historical management, with densities highest in logged, intermediate in logged/arboricided and lowest in the nature reserve. Forest communities differed significantly in terms of seedling diversity and density. Seedling composition differed significantly between transects and forest communities, but not between topographic positions or historical management types. Both Chao‐Jaccard and Chao‐Sørensen abundance‐based similarity estimators were relatively high in the plot, forest community and in terms of historical management levels, corroborating the lack of significant differences in species richness within these groups. The measured environmental variables explained 59.4% of variance in seedling species distributions, with the three most important being soil organic matter, total soil titanium and leaf area index (LAI). Total seedling density was positively correlated with LAI. Differences in diversity of >2.0 cm dbh plants (juveniles and adults) also explained variations in seedling species diversity. Conclusions: The seedling bank is the major route for regeneration in this semi‐deciduous tropical rain forest, with the wide distribution of many species suggesting that these species regenerate continuously. Seedling diversity, density and distribution are largely a function of adult diversity, historical management type and environmental gradients in factors such as soil nutrient content and LAI. The species richness of seedlings was higher in soils both rich in titanium and with low exchangeable cations, as well as in logged areas that were more open and had a low LAI.     

Dispersal and recruitment dynamics in the fleshy‐fruited Persoonia lanceolata (Proteaceae)

Question: What is the role of dispersal, persistent soil seed banks and seedling recruitment in population persistence of fleshy‐fruited obligate seeding plant species in fire‐prone habitats? Location: Southeastern Australia. Methods: We used a long‐term study of a shrubby, fleshy‐fruited Persoonia species (Proteaceae) to examine (1) seed removal from beneath the canopy of adult plants; (2) seedling recruitment after fire; (3) the magnitude and location of the residual soil seed bank; and (4) the implications for fire management of obligate seeding species. We used demographic sampling techniques combined with Generalised Linear Modelling and regression to quantify population changes over time. Results: Most of the mature fruits (90%) on the ground below the canopy of plants were removed by Wallabia bicolor (Swamp wallaby) with 88% of seeds extracted from W. bicolor scats viable and dormant. Wallabies play an important role in moving seeds away from parent plants. Their role in occasional long distance dispersal events remains unknown. We detected almost no seed predation in situ under canopies (< 1%). Seedling recruitment was cued to fire, with post‐fire seedling densities 6‐7 times pre‐fire adult densities. After fire, a residual soil seed bank was present, as many seeds (77‐100%) remained dormant and viable at a soil depth where successful future seedling emergence is possible (0‐5 cm). Seedling survival was high (> 80%) with most mortality within 2 years of emergence. Plant growth averaged 17 cm per year. The primary juvenile period of plants was 7–8 years, within the period of likely return fire intervals in the study area. We predicted that the study population increased some five‐fold after the wildfire at the site. Conclusions: Residual soil seed banks are important, especially in species with long primary juvenile periods, to buffer the populations against the impact of a second fire occurring before the seed bank is replenished.     

Recovery from clearing,cyclone and fire in rain forests of Tonga,South Pacific: Vegetation dynamics 1995–2005

Abstract In late 2001 a category 3 cyclone impacted forest plots that were established in Tonga in 1995, and additionally, one plot was accidentally burned by an escaped land‐clearing fire. Subsequent surveys provide observations of 10 years of forest dynamics in this poorly studied region, and the first reported observations of large interannual variation in juvenile (seedling and sapling) abundance in the western tropical Pacific. The severely disturbed (burned) plot was initially colonized by a non‐native early pioneer, Carica papaya L., but 3.5 years later a native pioneer, Macaranga harveyana (Muell. Arg.) Muell. Arg., was the most abundant tree species. The seedling layer included some long‐lived pioneers and shade‐tolerant species. Two mature forest plots affected only by the cyclone changed very little over a decade. Late‐successional shade‐tolerant species that dominated the overstory were also abundant as seedlings and saplings. This is in contrast with a 30‐ to 40‐year‐old, formerly cultivated, secondary forest plot that still shows no recruitment of late‐successional dominants, in spite of the proximity of remnant forest patches. This study suggests differing pathways of succession following shifting cultivation versus cyclone and fire disturbances in Tonga. Land use legacies appear to have a long‐lasting effect on community composition.     

Comparative demography of two giant caulescent rosettes (Espeletia timotensis and E. spicata) from the high tropical Andes

Using field data from previous studies we built matrix models for two populations of giant rosettes, Espeletia timotensis Cuatrec. and E. spicata Sch. Bip. Wedd., from the Andes Cordillera in Mérida, Venezuela. We analysed the models and calculated population growth rate (λ), sensitivities, elasticities and the sensitivity of the elasticities to changes in the vital rates. The analysis showed that the two species behave alike in general demographic terms. In both models, population growth rate is positive and sensitivities of λ to changes in vital rates decrease markedly in this order: plant establishment, progression of juvenile–adult, germination and survival. The relative contributions of vital rates to λ (elasticities) are very similar to those of other woody plant species: a higher contribution of survival and a very low contribution of fecundity. Transition from seedling to juvenile is most important and the younger established stages (juveniles and young adults) play a predominant demographic role in both populations. Seed banks and older adults are playing a relatively minor role in the dynamics of both populations. However, they may be important in relation to unpredictable, favourable or detrimental events. Perturbation analysis of elasticities showed that increasing the rate of plant establishment will decrease the relative importance of stasis. We conclude that both species are demographically very close, and similar to other long‐lived woody plant species. However, the two species differ in the role of the seed bank, which seems more important in the demography of E. spicata than in E. timotensis.     

Contrasting demographics of tropical savanna and temperate forest eucalypts provide insight into how savannas and forests function. A case study using Corymbia clarksoniana from north‐eastern Australia

Eucalypts (Eucalyptus spp. and Corymbia spp.) dominate many communities across Australia, including frequently burnt tropical savannas and temperate forests, which receive less frequent but more intense fires. Understanding the demographic characteristics that allow related trees to persist in tropical savannas and temperate forest ecosystems can provide insight into how savannas and forests function, including grass–tree coexistence. This study reviews differences in critical stages in the life cycle of savanna and temperate forest eucalypts, especially in relation to fire. It adds to the limited data on tropical eucalypts, by evaluating the effect of fire regimes on the population biology of Corymbia clarksoniana, a tree that dominates some tropical savannas of north‐eastern Australia. Corymbia clarksoniana displays similar demographic characteristics to other tropical savanna species, except that seedling emergence is enhanced when seed falls onto recently burnt ground during a high rainfall period. In contrast to many temperate forest eucalypts, tropical savanna eucalypts lack canopy‐stored seed banks; time annual seed fall to coincide with the onset of predictable wet season rain; have very rare seedling emergence events, including a lack of mass germination after each fire; possess an abundant sapling bank; and every tropical eucalypt species has the ability to maintain canopy structure by epicormically resprouting after all but the most intense fires. The combination of poor seedling recruitment strategies, coupled with characteristics allowing long‐term persistence of established plants, indicate tropical savanna eucalypts function through the persistence niche rather than the regeneration niche. The high rainfall‐promoted seedling emergence of C. clarksoniana and the reduction of seedling survival and sapling growth by fire, support the predictions that grass–tree coexistence in savannas is governed by rainfall limiting tree seedling recruitment and regular fires limiting the growth of juvenile trees to the canopy.     

Recruitment limitations in Primula vulgaris in a fragmented landscape

Plant recruitment is limited by dispersal, if seeds cannot arrive at potential recruitment sites, and by establishment, due to a low availability of safe sites for recruitment. Seed-sowing experiments, scarcely applied along gradients of landscape alteration, are very useful to assess these limitations. Habitat loss and fragmentation may foster recruitment limitations by affecting all the processes from seed dispersal to seedling establishment. In this study, we perform a seed-sowing experiment to disentangle the importance of dispersal and establishment limitations in different stages of recruitment of the perennial herb Primula vulgaris in fragmented forests of the Cantabrian Range (Northwestern Spain). We evaluated the influence of ecological gradients resulting from habitat loss and fragmentation (modifications of habitat amount at the landscape and microhabitat scales, changes in the species’ population size, changes in seed predation and seedling herbivory) on seedling emergence, survival and early growth. We found strong evidence of dispersal limitation, as seedling emergence was very low in experimental replicates where no seeds were added. This limitation was independent of landscape alterations, as we found no relation with any of the ecological gradients studied. Establishment limitations at the germination phase were also unrelated to ecological gradients, probably because these limitations are more related to fine-scale environmental gradients. However, further monitoring revealed that seedling survival after summer and winter periods and seedling growth were conditioned by landscape alteration, as we found effects of habitat amount at the landscape and microhabitat scales, of presence of populations of P. vulgaris and of seedling herbivory. These effects were complex and sometimes opposite to what can be expected for adult plants, revealing the presence of different requirements between life stages.     

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.     

Bumble bee species exhibit divergent responses to urbanisation in a Southern California landscape

1. Bees are ecologically important pollinators that are threatened by disease, habitat degradation, and habitat loss. Bee species that share ecological traits (e.g. body size, diet breadth, and sociality) may often respond similarly to environmental disturbance; however, few studies have examined the extent to which closely related, ecologically similar species exhibit divergent responses to the same forms of disturbance. 2. In the present study, the responses of bumble bees (Apidae: genus Bombus) to urbanization are examined by combining field surveys with the quantification of local‐ and landscape‐level variables using Geographic Information Systems (GIS). During the spring and summer of 2012 and 2013, 57, bumble bees were surveyed in 1‐ha plots in San Diego County, California that spanned a gradient of urbanisation. Species‐level, occurrence (presence/absence) data and logistic regressions were then used to identify determinants of Bombus occurrence across this landscape. 3. Approximately 1000 Bombus individuals belonging to three species were observed: B. californicus Smith, B. melanopygus Cresson, and B. vosnesenskii Radoszkowski. The occurrence of all Bombus species was negatively related to impermeable surface cover within study plots, but species responded uniquely to other plot‐scale variables and exhibited distinctive patterns of scale‐dependency with respect to impermeable surface cover surrounding plots. 4. Urbanisation negatively affected all three focal bumble bee species in this study. Species‐specific responses to plot‐scale and landscape‐scale variables presumably reflect interspecific trait differences (e.g. body size, tongue length, and foraging behaviour). Unique responses to urbanisation caution against pooling species into functional groups based merely on taxonomic relationships or perceived ecological similarities.     

Reinstatement of traditional mowing regimes counteracts population senescence in the rare perennial Primula vulgaris

Question: Traditional management of grassland verges or ditch banks included mowing as a way to provide additional harvesting of hay. Nowadays, such sites are often left unmanaged, as mowing verges is no longer profitable in modern agricultural systems. Are vulnerable plant species able to withstand competition with the surrounding vegetation and maintain viable populations under these circumstances? How do they respond to reinstatement of traditional mowing regimes? Location: Oedelem, northwestern Belgium. Methods: To investigate the effect of reinstatement of the rare perennial Primula vulgaris, demography and adult plant performance were monitored in a grassland verge between 1999 and 2003 under different mowing regimes. Year transitions between life stages were analysed with matrix population models. To disentangle the contributions of the deviations in different life stage transitions to the variation in overall population growth rate, life table response experiments were used. Results: Both management and year had a strong impact on demographic traits of P. vulgaris. If plots were left unmanaged, lower plant performance and declining population growth rates were observed. While population growth rates differed significantly between mowing regimes, mowing of plots only in July did not differ from mowing in July and October in terms of vegetative and reproductive output of adults. Mowing twice a year appeared to be most efficient in increasing population growth rate both by raising recruitment and growth of individuals into large reproductive adults. Conclusions: Large P. vulgaris populations show a good ability to recover from recent abandonment of traditional management regimes. By mowing twice a year, managers are able to target vital rates that are most influential: growth and flowering of adult individuals.     

Landscape context influences the abundance of amphibians and the strength of their food web interactions in small ponds

Many species, including most amphibians, undergo an ontogenetic niche shift (ONS) from an aquatic larval stage to a terrestrial adult stage. We use the ratio of aquatic to terrestrial habitat in a landscape as a tool to understand the influence of landscape context on the population growth of ONS species. The aquatic to terrestrial ratio (ATR) of habitats can be viewed as an analog to the influence of resource ratios on the population growth of consumers and depends on the degree to which each habitat type limits the growth of a given population. Population growth rates of shorter‐lived species tend to be more limited by demographic rates in early (aquatic) life stages. As a result, increasing the ATR should lead to a higher total population size in the landscape (and higher densities in the terrestrial habitat), but have little influence on the density of individuals in any given aquatic habitat. Alternatively, population growth rates of longer‐lived species tend to be more limited by demographic rates in later (terrestrial) life stages and increasing the ATR should have little influence on the total population size in the landscape, but decrease the density of individuals in any given aquatic habitat. We show that among‐landscape variation in the breeding‐pond densities of three widespread amphibians with contrasting life histories is consistent with this framework. Within‐pond densities of Pseudacris crucifer, a species with short‐lived adults, were not influenced by ATR, whereas within‐pond densities of Hyla versicolor, a longer‐lived member of the same family (Hylidae), declined as ATR increased. Ambystoma maculatum, a long‐lived salamander, also had lower densities in ponds with higher ATR. Because A. maculatum larvae are important predators in ponds, we use structural equation modeling to show that landscape context (ATR) can moderate community structure via direct (amphibian abundances) and indirect (prey species richness) effects.     

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.     

Bottlenecks for plant recruitment in woodland remnants: An ornithochorous shrub in a Mediterranean ‘relictual’ landscape

Habitat fragmentation may lead to declines in plant populations and ultimately to extinction through a disruption of demographic processes, which may result in bottlenecks or even a collapse in regeneration. Nevertheless, very few studies have assessed the net effects of habitat fragmentation on plant recruitment integrating its multiple demographic processes. Using data from a four-year field study, we analyze how habitat fragmentation affects particular demographic processes and the overall magnitude of plant recruitment. We use as a case study the bird-dispersed shrub Myrtus communis in Mediterranean woodland patches within an extremely fragmented landscape (～1% woodland cover). By means of observations and experiments, we quantified fecundity, fruit removal by frugivores, seed rain, post-dispersal seed predation by rodents and seedling emergence and survival. Within each patch, we quantified post-dispersal processes in different target microhabitats. We considered the life cycle to be a combination of consecutive life stages connected by transitional processes with specific probabilities. We calculated the overall probability of recruitment for each patch as the product of all of these probabilities. The demographic processes negatively affected by fragmentation were bird-generated seed rain and seedling emergence and survival, which were attributable, respectively, to lower fruit abundance and poorer habitat quality in the smaller patches. The negative effect of fragmentation on M. communis recruitment became stronger when all the demographic processes were integrated. Of all processes, seedling emergence and, above all, seedling survival were clearly bottlenecks for recruitment that were associated with habitat fragmentation. Results from our observations and experiments were consistent with natural patterns of regeneration given that we found higher seedling densities in larger patches and old population structures (with no saplings/juveniles) in some small patches. Our study shows that habitat fragmentation has serious negative effects on recruitment in M. communis due to demographic bottlenecks in seedling establishment. The available evidence (this and a companion study) suggests that the impoverishment of habitat quality associated with habitat fragmentation (edge effects, disturbances associated with management and microhabitat availability) can explain these results. Given that restoration at a landscape scale is likely to be extremely difficult, initial management actions should aim to improve habitat quality in the smallest woodland remnants.     

Year-to-year Oscillations in Demography of the Strictly Biennial <Emphasis Type="Italic">Pedicularis sylvatica</Emphasis> and Effects of Experimental Disturbances

In 1998–2001, I studied disturbance effects on the population structure and dynamics of a grassland strict biennial Pedicularis sylvatica, and on the species demography (monthly dynamics of seedling recruitment in 1998 and within- and between-year survival in 1998–2000). In two Czech populations, I established three experimental disturbance regimes: (1) a gap treatment, that simulated grazing by clipping vegetation and creating small gaps, (2) a mowing treatment, where I clipped the vegetation, and (3) a no management treatment, where I left the vegetation untreated. The number of recruiting seedlings varied greatly by year, and demographic structure of populations showed significant year-to-year oscillations in mean seedling numbers, from low (3 ± 0.7 s.e. per 0.25 m² plot) to high (103 ± 20). Inversely in the same years and plots, mean adult numbers in populations oscillated from high (12 ± 2) to low (0.7 ± 0.3). Disturbance effects were only important for seedling recruitment in early census dates in all years. In 1998, most seedlings recruited in April–May in gaps in both sites, but most died before winter. Within- and between-year survival was not affected by disturbance regimes but fluctuated significantly among years. Between-year survival increased with increasing size of the overwintering bud and was higher in disturbance treatments. Since the oscillations in population structure did not significantly vary in response to experimental disturbances, population dynamics may be driven endogenously rather than by disturbance events. The weak disturbance effects on species demography may also indicate population resilience to changes in habitat quality. However, since disturbances promoted seedling recruitment, grazing or mowing regimes are strongly recommended, as they create regeneration opportunities and maintain habitat quality, meeting the species long-term conservation goals.     

Density dependence in blowfly populations: experimental evaluation of non‐parametric time‐series modelling

Two approaches for describing density dependence in demographic rates of stage‐structured populations are compared in this study. Time‐series data from laboratory blowfly populations (Lucilia sericata) have been analysed in a separate study, with a statistical modelling approach that incorporated density dependences as unspecified (non‐parametric) functions. In this study, we assessed density‐dependent structures by manipulating densities of larvae and adults in cohorts of blowflies and measuring the demographic rates. We here compare the density‐dependent structures revealed by the cohort experiments with those estimated by the non‐parametric model. This model estimates the demographic rates to have the following density‐dependent structures: (i) larval survival was non‐linearly density‐dependent (a ‘humped’ function), (ii) adult survival was density‐independent, and (iii) reproductive rate decreased with adult density. In the cohort experiments reported here, (i) juvenile survival exhibited a positive density dependence in low densities (facilitation), which became negative at higher densities (competition). Pupal and adult size decreased with initial larval density. (ii) Adult survival was reduced by high initial larval density, but it was independent of adult density. (iii) Reproductive rate was reduced by high initial larval density, and by high adult density in populations of large individuals (from low larval density). Hence, the results from these experiments support the non‐parametric model estimates regarding density‐dependent structures of demographic rates in the blowfly populations. The mean demographic rates, however, were apparently underestimated by the model. We conclude that non‐parametric modelling is a useful first approach for exploratory analysis of ecological time‐series data.