Spanish juniper gain expansion opportunities by counting on a functionally diverse dispersal assemblage community

Seed dispersal is typically performed by a diverse array of species assemblages with different behavioral and morphological traits which determine dispersal quality (DQ, defined as the probability of recruitment of a dispersed seed). Fate of ecosystems to ongoing environmental changes is critically dependent on dispersal and mainly on DQ in novel scenarios. We assess here the DQ, thus the multiplicative effect of germination and survival probability to the first 3 years of life, for seeds dispersed by several bird species (Turdus spp.) and carnivores (Vulpes vulpes, Martes foina) in mature woodland remnants of Spanish juniper (Juniperus thurifera) and old fields which are being colonized by this species. Results showed that DQ was similar in mature woodlands and old fields. Germination rate for seeds dispersed by carnivores (11.5%) and thrushes (9.12%) was similar, however, interacted with microhabitat suitability. Seeds dispersed by carnivores reach the maximum germination rate on shrubs (16%), whereas seeds dispersed by thrushes did on female juniper canopies (15.5) indicating that each group of dispersers performed a directed dispersal. This directional effect was diluted when survival probability was considered: thrushes selected smaller seeds which had higher mortality in the seedling stage (70%) in relation to seedlings dispersed by carnivores (40%). Overall, thrushes resulted low‐quality dispersers which provided a probability or recruitment of 2.5%, while a seed dispersed by carnivores had a probability of recruitment of 6.5%. Our findings show that generalist dispersers (i.e., carnivores) can provide a higher probability of recruitment than specialized dispersers (i.e., Turdus spp.). However, generalist species are usually opportunistic dispersers as their role as seed dispersers is dependent on the availability of trophic resources and species feeding preferences. As a result, J. thurifera dispersal community is composed by two functional groups of dispersers: specialized low‐quality but trustworthy dispersers and generalist high‐quality but opportunistic dispersers. The maintenance of both, generalist and specialist dispersers, in the dispersal assemblage community assures the dispersal services and increases the opportunities for regeneration and colonization of degraded areas under a land‐use change scenario.     

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.     

Validating the use of woodrat (Neotoma) middens for documenting natural invasions

Aim Plant macrofossils in fossil woodrat (Neotoma) middens are the primary source of information on late Quaternary biogeographical history of plants in arid and semi‐arid regions of North America. Macrofossil records from middens are playing particularly important roles in documenting spatial and temporal patterns of plant migrations and invasions since the last glacial maximum 21,000 years ago. However, relatively few actualistic studies comparing contents of modern middens with surrounding flora and vegetation have been carried out. The primary aim of this study is to assess the reliability of midden assemblages in detecting the presence of tree, shrub and several other plant species growing on the surrounding landscape. The secondary aims are to determine whether probability of occurrence of species in middens is related to abundance in vegetation, and whether representation of individual species in middens is contingent on presence/absence or abundance of other species. Location Our five study sites were bedrock escarpments or canyons in the central Rocky Mountains (north‐eastern Utah, central Wyoming and south‐central Montana). All sites were in conifer woodland or forest/woodland variously dominated by Juniperus osteosperma, J. scopulorum, Pinus edulis, P. flexilis and Pseudotsuga menziesii. Neotoma cinerea is the only woodrat species in the region. Methods Macrofossil assemblages from 59 modern middens (all showing clear signs of recent or ongoing woodrat activity) were compared with floristic composition of vegetation within 50 m of the middens, and with percent cover of vegetation within 30 m of the middens. Results Coniferous trees and shrubs were well‐represented in middens, occurring consistently even when abundance in the local vegetation was very low. Juniperus osteosperma and J. scopulorum were particularly well‐represented, occurring in middens regardless of local abundance. Other conifers (P. edulis, P. flexilis, P. ponderosa, Pseudotsuga menziesii) were occasionally absent from middens when abundance in vegetation was low (< 20% canopy cover). Occurrence of dicot shrubs and graminoids was less consistent. Main conclusions Middens constructed by N. cinerea are highly reliable sensors of presence/absence of J. osteosperma and J. scopulorum, and hence can be used to infer invasions and past biogeographical distributions of these species. The middens are also reliable in registering populations of other conifers, although presence in middens may be contingent on local abundance. Additional comparative studies are needed to develop a sound empirical basis for using middens of N. cinerea and other species to infer past presence/absence of plant species on the landscape, and to explore the vegetation‐sensing properties of midden assemblages.     

Characterization of polymorphic microsatellite markers for a coniferous shrub Juniperus sabina (Cupressaceae)

Juniperus sabina L. is an evergreen coniferous shrub, with a widespread distribution in Asia and Europe. It is one of the key species for restoration of the ecosystem in desertification areas. In the present study, we isolated and characterized eight microsatellite loci of this species. The number of alleles per locus ranged from two to 27, with observed heterozygosity values ranging from 0.256 to 0.744 and expected heterozygosity from 0.276 to 0.939. The markers developed in this study could be useful for population genetics studies of J. sabina.     

Life history of Juniperus sabina L. adapted to the sand shifting environment in the Mu Us Sandy Land,China: A review

We have reviewed publications on the physiological and ecological features of the growth and regeneration processes of Juniperus sabina L. which grows in semiarid sandy land in the Ordos plateau in northern China where desertification has progressed over time. J. sabina is a key native plant species used for ecological restoration in this region. The life history of J. sabina in this sandy land that has been revealed through this review includes several unique features: (1) both vegetative and seed propagations are observed, but seed propagation is not successful in the location where the mature J. sabina stands. Instead, seed propagation can occur at a different place with different landscapes from the mature stands. (2) Nurse plants play an important role in providing the microclimatic environment necessary for the growth of J. sabina seedlings and young plants. (3) During the horizontal and vertical growth processes of the J. sabina patch, the root system was affected by burial in shifting sand and consequently acquired greater access to the water supply in deeper soil horizons, which could support larger growth. These characteristics suggested that the regeneration by seed propagation and growth strategy of J. sabina in this region was strongly affected by sand movement and the landscape that is generated by sand movement.

     

Growth delay by winter precipitation could hinder Juniperus sabina persistence under increasing summer drought

Plants in Mediterranean mountains are particularly vulnerable to climatic change. In these environments, low temperature is combined with water shortage during summer, and as a result, the positive effect of global warming theoretically expanding the growing season length may be counterbalanced by rising drought stress. These circumstances may be exacerbated in the rear edge of species distribution, where warmer conditions occur. Here, we examined the climate-growth relationships of Juniperus sabina, a major prostrate shrub above the treeline in Mediterranean mountains, to investigate climate sensitivity and long-term signals stability in four rear-edge populations from southern Spain. We demonstrate that, over recent decades, local climatic conditions have modulated the response of J. sabina secondary growth to the ongoing climate change. We observed a negative effect of winter-spring moisture on secondary growth that suggests a limitation for earlier growth activity at higher elevation, potentially hindering the ability of J. sabina to compensate forthcoming increases in summer drought. At the driest site, we also detected a positive effect of October precipitation, suggesting a second growth pulse by early autumn. Our results provide an example of how local climatic conditions may limit plastic responses of secondary growth to climatic variability. The implications of growth limitation in J. sabina exceed the species-scale level, since these prostrate shrubs play a critical role as nurse plants and local biodiversity foci in Mediterranean mountains.     

Geographical and interspecific variation and the nutrient‐enrichment hypothesis as an adaptive advantage of myrmecochory

In myrmecochory, the relocation of diaspores to ant nests may lead to the enhancement of plant fitness because ant nests and their middens are often richer in essential nutrients than surrounding areas. This idea is the basis of the nutrient‐enrichment hypothesis (NEH), which suggests that nutrient enrichment may be a major selective influence in the evolution of myrmecochory. However, there is little evidence regarding whether the greater plant performance and fitness enhancement in ant nests is due to nutrient enrichment or other benefits of directed dispersal. Here, we present the results of a large‐scale seed‐sowing experiment that tests the NEH in the ant‐dispersed perennial herb Helleborus foetidus, exploring geographical and inter‐ant taxa variation. Experiments were conducted in three well‐separated regions of the Iberian Peninsula, targeting the nests of major and minor local ant dispersers (nine ant species in total) and the soil beneath maternal plants as seed destinations. Seedling emergence, survival and early establishment rates, as well as variation in soil characteristics, were obtained for each seed destination at each region. Our results do not fully support the NEH in our study system. Instead, we found that the advantage of ant nest soil for establishment in H. foetidus was conditional. Differences in soil fertility and concomitant differences in seedling establishment between ant nests and beneath the canopy of maternal plants were observed in some regions and for some ant species, but not in others. Thus, the conditional outcomes arise from inconsistencies among regions, between stages of seedling regeneration and among ant species in the advantages of being dispersed to nests. Because variation in the guilds of ant dispersers of myrmecochore plants across their ranges is common, this study illustrates the need to consider geographic and inter‐ant taxa variation for a complete evaluation of the NEH.     

Differential seed dispersal systems of endemic junipers in two oceanic Macaronesian archipelagos: the influence of biogeographic and biological characteristics

This article evaluates the seed dispersal systems of two congeneric and endemic fleshy-fruited plants in the context of two relatively close oceanic archipelagos. For this purpose, representative populations of the endangered junipers Juniperus cedrus in the Canary Islands and Madeira, and Juniperus brevifolia in the Azores were studied. Despite both species sharing the same biogeographic region, we set out to test whether different conditions of the islands and biological characteristics of each juniper species determine the distinctive guilds of seed dispersers involved. We assessed the quantitative and qualitative role of the potential frugivores, showing that the wintering Turdus torquatus and the native Turdus merula were the main seed dispersers for J. cedrus and J. brevifolia, respectively (Frequency of occurrence: 74.9%, 80.2%; germination increase with respect to controls: 11.6%, 15.5%; for J. cedrus and J. brevifolia, respectively). The endemic lizard Gallotia galloti was quantitatively outstanding as seed disperser of J. cedrus, although its qualitative effect does not appear to be beneficial. The introduced rabbit Oryctolagus cuniculus acts as a disruptor in both natural seed dispersal systems, as inferred from the high percentage of damaged seeds found in their droppings. Our results indicate that J. cedrus and J. brevifolia are primarily adapted to ornithochory processes, T. torquatus and T. merula being their respective legitimate long-distance dispersers. Although these birds should be playing a key role in the connectivity of fragmented populations, the dependence of J. cedrus on a migrant bird involves a notable fragility of the system.     

Does sex matter? Gender-specificity and its influence on site-chronologies in the common dioecious shrub Juniperus communis

In recent years an increasing number of studies have shown shrubs to be reliable proxies of environmental conditions in regions where Trees − due to harsh climate conditions − are absent. Although many shrubs are monoecious, some are dioecious, which poses certain questions related to gender-specific growth as observed for trees in previous studies. Here, we address the questions whether dioecious shrubs, similar to trees, show growth differences between male and female plants, and − if so − whether this difference needs to be considered in terms of sample selection. We chose Juniperus communis. L., the most widely distributed woody plant, and a common and well-studied dioecious shrub species in the northern hemisphere, especially in the Boreal, Subarctic tundra and Alpine regions. Our samples were collected from four sites − three from the Ural Mountains and one site from Kirkenes in Norway. To see if there were differences in radial growth between sexes we performed four different analyses. First, we used multivariate explorative statistics to see if there were gender biased sub-populations and generally found no differences. Secondly, to compare growth over the lifetime of shrubs we computed cumulative annual increments of basal area which revealed no gender-specific growth patterns. Thirdly, to test if differences in radial growth between male and female shrubs affect the resulting site-chronology, we compared individual shrub chronologies with the site-chronology and found a significant differentiation between normalized correlations of gender-specific chronologies to the site-chronology. This significant difference was restricted to an overall comparison, but not evident at individual site-level. Lastly, we compared correlations of gender-specific chronologies and a mean site-chronology with monthly climate records to find only very few meaningful differences in their responses. In summary, we could not detect any clear gender-specific growth pattern in Juniperus communis but observed a trend towards more non-climatic signals in female junipers which may affect the resulting site-chronology.     

Biased dispersal of Metrioptera bicolor,a wing dimorphic bush‐cricket

In the highly fragmented landscape of central Europe, dispersal is of particular importance as it determines the long‐term survival of animal populations. Dispersal not only secures the recolonization of patches where populations went extinct, it may also rescue small populations and thus prevent local extinction events. As dispersal involves different individual fitness costs, the decision to disperse should not be random but context‐dependent and often will be biased toward a certain group of individuals (e.g., sex‐ and wing morph‐biased dispersal). Although biased dispersal has far‐reaching consequences for animal populations, immediate studies of sex‐ and wing morph‐biased dispersal in orthopterans are very rare. Here, we used a combined approach of morphological and genetic analyses to investigate biased dispersal of Metrioptera bicolor, a wing dimorphic bush‐cricket. Our results clearly show wing morph‐biased dispersal for both sexes of M. bicolor. In addition, we found sex‐biased dispersal for macropterous individuals, but not for micropters. Both, morphological and genetic data, favor macropterous males as dispersal unit of this bush‐cricket species. To get an idea of the flight ability of M. bicolor, we compared our morphological data with that of Locusta migratoria and Schistocerca gregaria, which are very good flyers. Based on our morphological data, we suggest a good flight ability for macropters of M. bicolor, although flying individuals of this species are seldom observed.     

Geographic variation in the adaptive landscape for seed size at dispersal in the myrmecochorous <Emphasis Type="Italic">Helleborus foetidus</Emphasis>

Literature on seed dispersal mutualisms suggests that plant populations should hardly adapt to their current dispersers. We address the predictions that selection pressures exerted by ants on dispersal-related diaspore traits of the ant-dispersed Helleborus foetidus are highly variable in space, and that geographic (inter-population) variation in these traits is unrelated to selection by current dispersers. To test these predictions we use the concept of the quantitative adaptive landscape for seed size at dispersal. Such landscape depicts the relationship between the population’s mean trait value (mean seed size in the present study) and the population’s mean fitness (mean dispersal probability in the present study). Adaptive landscapes make it possible to assess whether the mean population’s phenotype agrees with one favored by selection. We first analyse, in 12 populations of H. foetidus from southern Spain, the extent of divergence among populations in seed and elaiosome size, and the abundance, composition, and behavior of the ant communities. Seeds from a fixed set of five of these populations were offered to ants in all the study sites to fit the adaptive landscape for seed size. In addition, seeds from the local population were also offered in each site. Our results show that seed size has undergone a larger divergence among populations than elaiosome size. Despite geographic variation in ant assemblages, the adaptive landscapes for seed size at dispersal were remarkably similar among sites: ants create disruptive selection on seed size in 10 out of 12 study sites. As predicted, the basic features of these adaptive landscapes (curvature and location of the minimum) varied geographically in accordance with variation in the size of seed dispersers. Also as predicted, in most populations, the observed mean seed size does not agree with that expected from the adaptive landscapes at dispersal. However, the relevance of dispersers for seed size evolution should not be neglected since the agreement between observed and optimum seed size was stronger where dispersers were more abundant. Thus, against the general view, our results evidence that, in H. foetidus, the observed geographic variation in dispersal-related plant traits is partly linked to selection exerted by current dispersers. Geographic variation in ant assemblages determines both the existence of a selection mosaic and the degree of adjustment of populations to the patterns of selection in the mosaic.     

Using genetic evaluation to guide conservation of remnant Juniperus communis (Cupressaceae) populations

• Many critically endangered plant species exist in small, genetically depauperate or inbred populations, making assisted gene flow interventions necessary for long‐term population viability. However, before such interventions are implemented, conservation practitioners must consider the genetic and demographic status of extant populations, which are strongly affected by species’ life‐history traits. In northwestern Europe, Juniperus communis, a dioecious, wind‐pollinated and bird‐dispersed gymnosperm, has been declining for the past century and largely exists in small, isolated and senescent populations.
• To provide useful recommendations for a recovery plan involving translocation of plants, we investigated genetic diversity and structure of populations in Belgium using four microsatellite and five plastid single‐nucleotide polymorphism (SNP) markers.
• We detected no clonality in the populations, suggesting predominantly sexual reproduction. Populations exhibited high genetic diversity (H_e = 0.367–0.563) and low to moderate genetic differentiation (F_ST ≤ 0.133), with no clear geographic structure. Highly positive inbreeding coefficients (F_IS = 0.221–0.507) were explained by null alleles, population substructuring and biparental inbreeding. No isolation by distance was observed among distant populations, but isolation at close geographic proximity was found. Patterns were consistent with high historical gene flow through pollen and seed dispersal at both short and long distances. We also tested four pre‐germination treatments among populations to improve germination rates; however, germination rates remained low and only cold‐stratification treatments induced germination in some populations.
• To bolster population regeneration, introductions of cuttings from several source populations are recommended, in combination with in situ management practices that improve seedling survival and with ex situ propagation.

     

The influence of demography and local mating environment on sex ratios in a wind‐pollinated dioecious plant

Negative frequency‐dependent selection should result in equal sex ratios in large populations of dioecious flowering plants, but deviations from equality are commonly reported. A variety of ecological and genetic factors can explain biased sex ratios, although the mechanisms involved are not well understood. Most dioecious species are long‐lived and/or clonal complicating efforts to identify stages during the life cycle when biases develop. We investigated the demographic correlates of sex‐ratio variation in two chromosome races of Rumex hastatulus, an annual, wind‐pollinated colonizer of open habitats from the southern USA. We examined sex ratios in 46 populations and evaluated the hypothesis that the proximity of males in the local mating environment, through its influence on gametophytic selection, is the primary cause of female‐biased sex ratios. Female‐biased sex ratios characterized most populations of R.  hastatulus (mean sex ratio = 0.62), with significant female bias in 89% of populations. Large, high‐density populations had the highest proportion of females, whereas smaller, low‐density populations had sex ratios closer to equality. Progeny sex ratios were more female biased when males were in closer proximity to females, a result consistent with the gametophytic selection hypothesis. Our results suggest that interactions between demographic and genetic factors are probably the main cause of female‐biased sex ratios in R. hastatulus. The annual life cycle of this species may limit the scope for selection against males and may account for the weaker degree of bias in comparison with perennial Rumex species.     

Restoration of alvar vegetation on Öland,Sweden

Abstract. Alvars in the Baltic region, particularly on the Swedish islands of Öland and Gotland and in western Estonia, are well‐known for their plant species richness and extensive populations of rare species. Grasslands make up most of alvar vegetation. The extent of these grasslands decreases because of bush enchroachment which takes place in most alvars when traditional land use practice changes, notably when grazing is ceased. The main threat for the alvar grassland is formed by the rapid expansion of Juniperus communis and Potentilla fruticosa in dry and wet sites, respectively. Applied research has been carried out during several years in order to develop plans for the restoration of alvar grassland. Thus it was shown that species richness in areas with a mixture of grassland and shrubland decreases in relation to the increased cover by shrubs. In the case of Juniperus there is a distinct drop in species number as soon as the cover exceeds 75‐80%, while the decrease is more gradual with increased cover of Potentilla. The seed banks under dense stands of these shrubs only contain ca. 20% of the species found in dry and wet alvar meadows. This was confirmed by clearing experiments. Long‐term recordings have shown that establishment of juniper seedlings takes place mainly in half‐open areas between already existing junipers. Intermediate‐sized junipers expanded faster than old and big shrubs. Potentilla shrubs recover fast from cutting or burning. After 2‐4 yr they have almost attained their former size. This recovery can be prevented when cattle grazing is introduced. In the framework of a local alvar restoration project on Öland, covering 7000 ha, grazing regimes are re‐established, fences erected and large‐scale clearings carried out. Recently established juniper shrubs are being cleared, intermediate sized junipers (30‐50 yr old) are selectively removed while creating a mosaic landscape with high biological diversity. Older dense juniper stands are left alone or are only partly cleared. Potentilla stands in moist areas are cut to create moist meadows which are breeding grounds for waders, and to establish corridors between remaining open areas. Three items are discussed (1) the importance of the Stora Alvar area; (2) re‐introduction of grazing and (3) re‐introduction of species. The outstanding importance of the area regarding species richness and endemism should be recognized through a ‘golden list’ to be used along with red and blue lists.     

Facilitation by herbivore-mediated nurse plants in a threatened tree, Taxus baccata: local effects and landscape level consistency

We investigated the regeneration of a threatened tree, the yew Taxus baccata, in relation to the presence of fleshy‐fruited woody plants acting as seed dispersal foci as well as protecting yew recruits against ungulate herbivores. We seek to determine if local facilitative effects are consistent across landscape in the Cantabrian range (NW Spain). Yew seed rain by birds mostly concentrated under yew trees and beneath hollies Ilex aquifolium. Seedling emergence distributed similarly to seed rain, but first‐year seedling survival was higher beneath hollies. In one site where woody vegetation was structured as nucleation centres (multispecific patches of fleshy‐fruited plants acting as foci for seed rain) yew recruits mostly occurred in yew‐dominated centres, suggesting dispersers‐mediated facilitation. However, holly was the main nurse plant for most of these recruits, considering the nurse as the species whose canopy covered directly the yew recruit. Living beneath nurse plants reduced herbivore damage on saplings and enhanced seedling survival. A planting experiment with yew rooted‐cuttings beneath different spiny shrubs corroborated this effect. Additional evidence on yew recruitment limitation by herbivory emerged from one population where ungulates were fence‐excluded. Our results suggest that nurse plants mitigate the negative effect of herbivores on yew regeneration, by providing defence against browsing and trampling. Shelter ability related to nurse structure, cone‐shaped shrubs with branches at their bases acting better as a barrier. Paradoxically, this structure resulted from heavy browsing on nurse plants. The study of yew regeneration and habitat structure in seven sites provided evidence for the consistency of facilitation by holly at the landscape level, since local values of yew recruitment positively related to nurse ground cover. Range‐scale yew management must consider the local functioning of the interaction among avian seed‐dispersers, nurse fleshy‐fruited plants and ungulate herbivores, in combination with regional measures, targeting the habitats where facilitation emerges.     

Effect of shrubs and seasonal variability of rainfall on the establishment of Aspidosperma quebracho‐blanco in two edaphically contrasting environments

Abstract Three experiments were conducted to verify if an increase in environmental stress level would affect the interactions between two species of nurse shrubs and seedlings of Aspidosperma quebracho‐blanco. This is a mesic species with a generalist distribution over an extensive environments gradient. The relationship between Larrea divaricata and seedlings of A. quebracho‐blanco was studied in two contrasting soils, a silty loam soil with higher surface clay content and a sandy loam soil. The effect of seasonal variability of rainfall on the initial establishment of seedlings under the shade of L. divaricata was evaluated in three consecutive years. The effect of nurse plant shade was tested comparing two shrub species with different types of leaf life span (sclerophyllous‐evergreen and leguminous‐deciduous). The natural establishment of A. quebracho‐blanco depended on shaded microsites, but not on the type of shade provided by different nurse shrubs. Emergence and initial establishment depended on interactions of soil type and seasonal rainfall variation with nurse plants. The importance of facilitation increased with clay soil (CS). Sandy soil was ‘less humid’ than CS under shrub shade. However, establishment success depends on opportune even rainfall distribution in interaction with nurse plant presence.     

Pinus monophylla establishment in an expanding Pinus‐Juniperus woodland: Environmental conditions,facilitation and interacting factors

Abstract. The tree species comprising Pinus‐Juniperus woodlands are rapidly expanding into shrub‐grasslands throughout their range. Observational studies indicate that establishment is facilitated by nurse plants, but little information exists on the mechanisms involved. I examined both abiotic and biotic factors influencing Pinus monophylla establishment in Artemisia tridentata steppe with expanding populations of P. monophylla and Juniperus osteosperma. I determined soil water contents, temperatures, and nutrient characteristics for the primary establishment microhabitats, i.e. under Pinus, under Juniperus, under Artemisia, tree interspace and sage interspace, and evaluated the emergence and survival response of two seedling cohorts over a 3‐yr period for the different microhabitats. I also examined the effects of seed burial and predation on seedling establishment. Microhabitats under trees and shrubs had higher extractable P and K, higher organic matter, total nitrogen and cation exchange capacity than interspace microhabitats. Soil water contents (0–15 cm) were lower in interspaces than under shrubs or trees due to dry surface (0–5 cm) soils. Soil temperatures (at 1 and 15 cm) were lowest under trees, intermediate under shrubs, and highest in interspaces. Timing and rate of seedling emergence were temperature dependent with the order of emergence paralleling mean growing season temperatures: tree interspace = shrub interspace > under shrub > under Juniperus under Pinus. Seed burial was required for rooting and the highest emergence occurred from depths of 1 and 3 cm indicating that caching by birds and rodents is essential and that animals bury seeds at adequate if not optimal depths for emergence. Seedlings required microenvironmental modification for survival; all seedlings, including those that emerged from seeds and transplants, died within the first year in interspace microhabitats. Survival in under‐tree or under‐shrub microhabitats depended on soil water availability and corresponded closely to soil water contents over the 3‐yr study. Under‐shrub microhabitats had more favourable soil and micro‐environmental characteristics than under‐tree microhabitats and had the highest seedling life spans for the first‐year seedling cohort. Predation of Pinus seedlings by rodents was a significant cause of mortality with caged transplants exhibiting life spans that were 74% longer overall than uncaged transplants. Emergence and survival of P. monophylla within the expanding woodland were dependent upon a complex set of interacting factors including growing season conditions, microhabitat characteristics, and animal species.     

Colonization of Abandoned Land by Juniperus thurifera Is Mediated by the Interaction of a Diverse Dispersal Assemblage and Environmental Heterogeneity

Land abandonment is one of the most powerful global change drivers in developed countries where recent rural exodus has been the norm. Abandonment of traditional land use practices has permitted the colonization of these areas by shrub and tree species. For fleshy fruited species the colonization of new areas is determined by the dispersal assemblage composition and abundance. In this study we showed how the relative contribution to the dispersal process by each animal species is modulated by the environmental heterogeneity and ecosystem structure. This complex interaction caused differential patterns on the seed dispersal in both, landscape patches in which the process of colonization is acting nowadays and mature woodlands of Juniperus thurifera, a relict tree distributed in the western Mediterranean Basin. Thrushes (Turdus spp) and carnivores (red fox and stone marten) dispersed a high amount of seeds while rabbits and sheeps only a tiny fraction. Thrushes dispersed a significant amount of seeds in new colonization areas, however they were limited by the presence of high perches with big crop size. While carnivores dispersed seeds to all studied habitats, even in those patches where no trees of J. thurifera were present, turning out to be critical for primary colonization. The presence of Pinus and Quercus was related to a reduced consumption of J. thurifera seeds while the presence of fleshy fruited shrubs was related with higher content of J. thurifera seeds in dispersers’ faeces. Therefore environmental heterogeneity and ecosystem structure had a great influence on dispersers feeding behaviour, and should be considered in order to accurately describe the role of seed dispersal in ecological process, such as regeneration and colonization. J. thurifera expansion is not seed limited thanks to its diverse dispersal community, hence the conservation of all dispersers in an ecosystem enhance ecosystems services and resilience.     

Selection against immigrants in wild seabird populations

Immigration is a major demographic parameter shaping population dynamics and is an important driver of eco‐evolutionary patterns, but the fitness consequences for individuals following their settlement to a new population (immigrants) remain poorly tested in wild animal populations, particularly among long‐lived species. Here we show that immigrants have a lower fitness than residents in three wild seabird populations (wandering albatross Diomedea exulans, southern fulmar Fulmarus glacialoides, snow petrel Pagodroma nivea). Across all species and during a 32‐year period, immigrants made on average ?9 to 29% fewer breeding attempts, had 5–31% fewer fledglings, had 2–16% lower breeding success and produced 6–46% fewer recruits. Female immigration and male residency were also favored through differences in breeding performance. We provide evidence for selection against immigrants in wild populations of long‐lived species and our results are consistent with female‐biased dispersal in birds being driven by asymmetric limiting resources and the competitive ability of dispersers vs. non‐dispersers.     

The role of avian ‘seed predators’ as seed dispersers

Seed dispersal is a central process in plant ecology with consequences for species composition and habitat structure. Some bird species are known to disperse the seeds they ingest, whereas others, termed ‘seed predators’, digest them and apparently play no part in dispersal, but it is not clear if these are discrete strategies or simply the ends of a continuum. We assessed dispersal effectiveness by combining analysis of faecal samples and bird density. The droppings of seed dispersers contained more entire seeds than those of typical seed predators, but over a quarter of the droppings of seed predators contained whole seeds. This effect was further magnified when bird density was taken into account, and was driven largely by one frequent interaction: the Chaffinch Fringilla coelebs, a typical seed predator and the most abundant bird species in the area and dispersed seeds of Leycesteria formosa, a non‐native plant with berry‐like fruits. These results suggest the existence of a continuum between seed predators and seed dispersers.