Limiting factors for seedling emergence and establishment of the invasive non-native Rosa rugosa in a coastal dune system

The relative importance of seed, habitat and microhabitat limitation has rarely been investigated for invasive non-native species, although this is critical for their effective management and for predicting future range expansion. Rosa rugosa is an abundant non-native shrub in coastal habitats of NW Europe; it is common along the Baltic coast but more scarce in exposed dunes of the North Sea. To investigate whether invasion of exposed dunes is limited by seed, habitat and microhabitat limitation, seedling emergence and establishment were examined in a factorial sowing, transplant and disturbance experiment. Twenty plots were randomly placed in each of five dune habitats (white dune, Empetrum dune, grey dune, outer dune heath, inner dune heath), and studied over 2 years. Seedling emergence in control subplots was zero in all habitats, whereas 96% and 98% of the undisturbed and disturbed seeded subplots produced seedlings. Disturbance had a positive effect on emergence and subsequent survival in white dune, outer and inner dune heath. Seedling survival and growth, and growth of transplanted seedlings, were markedly lower in grey dune. These findings indicate that establishment of R. rugosa is seed-limited in coastal dune habitats, and that the species is able to establish in both active and fixed dunes once seeds have arrived. Although differential seedling emergence and growth indicate that habitats differ in their degree of invasion susceptibility by R. rugosa, the positive influence of small-scale disturbance suggests microhabitat limitation in some dune habitats as well. Dune management should aim to reduce seed production and dispersal of R. rugosa in near-natural sites, and anthropogenic changes of habitat dynamics should be prevented.     

地中海沿岸沙丘微生境对幼苗出现时空格局的影响

实验样地设在地中海沿岸沙丘,选择了3个不同的微生境代表：(1)稳定沙丘上的开阔地片段,(2)稳定沙丘上的灌丛下区域,(3)不稳定沙丘的路径区域;从2001年11月至2002年4月的整个生长季节,每一个微生境出现的幼苗在4个日期被监测,并在每一个取样日把每一株幼苗鉴定、计数后,用剪刀把地上部分移去;研究调查了3种微生境幼苗出现的时空分布格局,并分析了雨量与幼苗出现数量的关系。结果发现：在地中海沿岸沙丘生态系统,幼苗出现在时间上具有明显的分布特征,大多数幼苗出现在第一观测期,整个生长季幼苗都不断出现,但幼苗出现的数目却逐渐下降;各功能群的幼苗占幼苗总数的比例分别为：1年生阔叶草47．4％,多年生阔叶草2．5％,豆科植物17,0％,菊科植物14．5％,1年生禾草为11．7％,多年生禾草为1．9％,十字花科植物3．7％,伞形科植物1,4％。在空间上,总幼苗密度、物种丰富度和物种多样性等显示出重要的微生境差异,开阔地区域具有最大的幼苗密度、物种丰富度和物种多样性;3个微生境的幼苗出现不是同步的,微生境影响种子萌发的时间分布格局,灌丛下种子萌发具有滞后现象;大多数功能群的幼苗密度分布基本上具有显著的微生境差异,主要物种的幼苗分布也具有显著的微生境差异。雨量和萌发的幼苗数量间未发现显著的关系。     

Ontogenetic shifts in interactions of two dominant shrub species in a semi‐arid coastal sand dune system

Question: How does the interaction between two dominant shrub species in a coastal sand dune community change during their life history? Does this interaction influence their population dynamics? Location: A semiarid coastal sand dune system in southeast Spain. Methods: For 3 years we monitored physiological status, growth and reproductive effort of Juniperus phoenicea and Pistacia lentiscus, the dominant shrub species, growing either alone or in close spatial association. We also recorded adult mortality patterns and characterized seedling survival, soil properties and microclimate conditions beneath canopies and in bare ground. Results and conclusions: There was a strong bi‐directional interaction between the two studied species, with a net balance that changed in sign with increasing plant development. While mature individuals facilitated the establishment of seedlings of both species, adult mortality patterns suggested asymmetric competition at later life stages. The interaction with Pistacia negatively affected growth of juniper and contributed to its high mortality rates, while juniper had almost no effect on mature Pistacia individuals. Physiological data suggested that Pistacia had a competitive advantage over juniper, most likely because of differences in rooting patterns and tolerance to salinity, which may determine the source of water available for each species. Community dynamics are governed by facilitation at the seedling stage and shaped by differences in physiological traits in adult plants. Plant‐plant interactions, which are strongly affected by environmental gradients, are important drivers of community dynamics in this system.     

Facilitation and sand burial affect plant survival during restoration of a tropical coastal sand dune degraded by tourist cars

Coastal sand dunes support various ecosystem services, including storm protection and tourism. Restoration programs are often critical to preserve this ecosystem due to its fragility and high degree of degradation. Dune restoration still suffers from a general lack of knowledge of the ecological processes controlling tropical dune communities. We investigated if facilitation can increase restoration success by assisting plant survival in a Brazilian coastal dune degraded by buggies at the Environmental Protection Area of Jenipabu, RN, Brazil. We performed two field experiments on dune crest sites degraded by buggies. The first experiment tested how the presence of established vegetation and coconut mesh, mimicking soil‐stabilizing effect of vegetation, facilitates seedling survival and establishment of the early successional dominant dune plant Canavalia maritima (Fabaceae). The second experiment tested if coconut mesh and initial irrigation would allow the establishment of C. maritima outside the vegetation, using both seeds and transplants. We found that the presence of established vegetation positively affected seedling survival. Application of mesh positively affected microclimatic conditions and slightly increased survival, but only for seeds. Initial irrigation only had a small effect on transplanted seedlings survival. Nevertheless, there was extreme seedling mortality in the experiment in particular from sand burial. We conclude that restoration of these dune crests is very difficult. Facilitation by vegetation can reduce plant mortality by sand burial and desiccation, but successful restoration is likely to require a major effort involving a very high number of seedlings or seeds.     

Adaptations enhancing survival and establishment of seedlings on coastal dune systems

Studies on the survival and establishment of seedlings in coastal and lacustrine sand dune systems suggest that nutrient deficiency, lack of moisture, sand accretion, salt spray and predation are probably the most important limiting factors. Seedlings employ both avoidance and tolerance strategies to with-stand the different stresses. For example, seedling recruitment of dune species coincided with periods of high moisture availability and occurred in years with high well distributed rainfall. The seedlings exhibited rapid vertical elongation of roots, avoided moisture stress by leaf rolling, dense hair on leaves, fleshy cotyledonary leaves, and by growing in the shade of nurse plants. A certain proportion of seedlings of all species survived partial burial and showed stimulation of leaf growth, total leaf area, number of tillers and total dry matter. Seedlings of most species could not survive complete burial, however, some species did grow through the sand deposit by an increase in the number of nodes and length of internodes. Plants responded to salt spray by inhibition of seed germination, developing resistance, increase in leaf thickness and reduced uptake of Na and Cl ions. Several characters such as higher vigour of seedlings, larger seed size and seed polymorphism were also of adaptive significance to the species.     

Facilitation of seedling establishment by an endemic shrub in tropical coastal sand dunes

Facilitation is predicted to occur on coastal sand dunes as these ecosystems have the harsh physical conditions common during primary succession. In a tropical dune system on the Gulf of Mexico the spatial patterns of plant distribution were analyzed and the hypothesis that facilitation is the responsible mechanism was tested by monitoring changes in the micro-environment and performing a seedling planting experiment under natural conditions. Densities of seedlings and adults of late colonizer grasses were two to six times greater beneath the early colonizer shrub, Chamaecrista chamaecristoides, than in the exposed areas. Temperatures on the sand surface, wind speed and sand accretion were significantly reduced by the shade of Chamaecrista. Only phosphate contents during the dry season were significantly higher in sand in the shade than in exposed sand. In the field experiment, successful establishment of seedlings of two late colonizer grasses (Trachypogon plumosus and Schizachyrium scoparium) was low but was concentrated exclusively beneath the shrubs. The introduced plants were reproductive one year after onset of the experiment and remained vigorous after three years. The observed spatial aggregation among the target species may be a result of improved conditions in the shade of the shrub, which facilitated the survival and establishment of late colonizers. On top of the environmental amelioration, protection from recurrent disturbances such as substrate mobility is an additional beneficial effect of the shrub.     

Effects of the aqueous extract from<Emphasis Type="Italic">artemisia campestris</Emphasis> ssp.<Emphasis Type="Italic">caudata</Emphasis> on mycorrhizal fungi colonization and growth of sand dune grasses

We used the aqueous extract fromArtemisia campesttis ssp.caudata to investigate its effects on the colonization of sand dune grass roots by mycorrhizal fungi and seedling growth. The percent colonization decreased with higher extract concentrations, and growth of three grass species was inhibited. Colonization by mycorrhizal fungi was more sensitive to the extract than was seedling growth, and no significant differences in the latter were found between the mycorrhizal and non-mycorrhizal treatments.     

Positive responses of coastal dune plants to soil conditioning by the invasive Lupinus nootkatensis

Invasive nitrogen-fixing plants drive vegetation dynamics and may cause irreversible changes in nutrient-limited ecosystems through increased soil resources. We studied how soil conditioning by the invasive alien Lupinus nootkatensis affected the seedling growth of co-occurring native plant species in coastal dunes, and whether responses to lupin-conditioned soil could be explained by fertilisation effects interacting with specific ecological strategies of the native dune species. Seedling performance of dune species was compared in a greenhouse experiment using field-collected soil from within or outside coastal lupin stands. In associated experiments, we quantified the response to nutrient supply of each species and tested how addition of specific nutrients affected growth of the native grass Festuca arundinacea in control and lupin-conditioned soil. We found that lupin-conditioned soil increased seedling biomass in 30 out of 32 native species; the conditioned soil also had a positive effect on seedling biomass of the invasive lupin itself. Increased phosphorus mobilisation by lupins was the major factor driving these positive seedling responses, based both on growth responses to addition of specific elements and analyses of plant available soil nutrients. There were large differences in growth responses to lupin-conditioned soil among species, but they were unrelated to selected autecological indicators or plant strategies. We conclude that Lupinus nootkatensis removes the phosphorus limitation for growth of native plants in coastal dunes, and that it increases cycling of other nutrients, promoting the growth of its own seedlings and a wide range of dune species. Finally, our study indicates that there are no negative soil legacies that prevent re-establishment of native plant species after removal of lupins.     

种子萌发和幼苗生长对沙丘环境的适应机制

综述了植物沙生适应机制的研究及其进展.一些植物的种子在刚成熟时具有休眠特性.种子需要适度沙埋促进萌发并实现幼苗定居,但过度沙埋则会抑制种子萌发和出苗.在沙层深处,没有萌发的种子会进入休眠状态,形成土壤种子库.幼苗通过增加节数和延长节间来适应沙埋.沙埋深度超过植物的忍耐限度会抑制幼苗生长,甚至导致幼苗死亡.沙生植物必须适应其他环境因子,如盐风与土壤盐分、昆虫采食、土壤养分亏缺等,才能在沙丘上成功生长.沙蚀可导致幼苗根系暴露并干燥脱水.一些沙漠植物的幼苗在萌发后可忍耐一段时期的干燥,水分条件得到满足之后,幼苗能够恢复生长.     

地中海沿岸沙丘种子大小对植物及其种子多度的影响

分析了地中海沿岸沙丘3种微生境(灌丛下、灌丛之间的开阔地、路径)以及整个沙丘生态系统的种子大小与土壤种子库中休眠种子的数量、土壤种子库中总种子的数量、地上植被中各个植物种的个体数量和幼苗数量、每种植物在小样方中的出现频率等方面之间的关系,也分析了种子尺寸(长、宽、高之和的平均值)与土壤种子的长久性、土壤种子库中休眠种子数量之间的关系;还分析了具有不同种子大小的植物在沙丘生态系统和各微生境的分布比率,验证了生活周期短的植物的种子在土壤中更长久和被干扰的微生境具有更多的休眠种子这种假说。结果发现,在地中海沿岸沙丘生态系统中,具有特别大的种子和特别小种子的植物种类都很少,不同植物种子的大小呈现近正态分布,但绝大多数植物的种子重量都不超过10 mg;微生境影响种子尺寸与种子长久性的关系,在灌丛下、开阔地区域等两个微生境以及整个的沙丘生态系统都呈现显著的正向关系(p<0 .0 5 ) ,而在路径这种关系不很明显;种子多度与植被物种出现频率呈现强烈的正相关(p<0 .0 5 ) ,没有发现种子大小与土壤种子库中休眠种子数量、种子库中总的种子数量、植被物种出现频率、植被的物种多度、幼苗植物多度等方面有显著的关系;具有中度大小种子(0 .1～10 mg)的植物在总的土壤种子库、休     

Resprouting enhances seedling persistence in a subtropical coastal dune forest

In theory, resprouting enables species with low reproductive output (i.e., few seedlings) to persist. The advantage conferred by seedling sprouts on tree species persistence was evaluated in a subtropical coastal dune forest in South Africa. Species with a higher frequency of seedling sprouts demonstrated greater persistence as evidenced by a larger proportion of seedlings >1 year old and a larger seedling bank than species with few seedling sprouts. Resprouted seedlings had a larger basal diameter than true seedlings. Although resprouting resulted in the maintenance of multiple stems in some seedlings, the proportion of multi‐stemmed seedlings was low. Multi‐stemming was not a favoured form of seedling growth except in one species that occupied relatively open sites. Despite the apparent difference among species in resprouting ability, we found that seedling resprouting was not phylogenetically constrained. These results demonstrate that seedling sprouts form an important component of seedling banks in coastal dune forest.     

Effects of sand burial on purple sandgrass (Triplasis purpurea): the significance of seed heteromorphism

The effects of sand burial on seedling emergence, growth, and reproduction of the annual dune grass Triplasis purpurea (Walt.) Chapm. were examined. This species matures heteromorphic seeds on panicles enclosed by leaf sheaths along tiller nodes in a characteristic pattern: seeds at the lower nodes are heaviest, whilst those at the upper nodes are lightest. Field excavation of seedlings revealed that seedlings can emerge from seeds buried to 4 cm.In one experiment in an incubator, seeds collected from the upper and lower tiller nodes of glasshouse-grown parental plants were buried in sand at depths of 0, 3, and 6 cm. As expected, the effects of seed depth on seedling growth were due to delayed emergence for seedlings from buried seeds. However, seedlings from the larger seeds from the lower nodes of parental plants had significantly greater dry mass than those from the lighter seeds from upper nodes. Because lower node seeds are the most likely to become buried by sand in nature, the seed heteromorphism may be adaptive in the coastal environment.A second experiment involved the burial of seedlings to 0, 50, or 100% of their height. Although most seedlings did not survive complete burial (100% height), for partially buried seedlings (50% height) there was high survival and a marked stimulation of growth and eventual reproduction compared to unburied seedlings. This stimulation could be linked to increased root growth for seedlings that had been buried. The ability to respond positively to sand accumulation may represent an adaptation to the dynamic dune environment.     

Size‐related and diel variations in microhabitat use of three endangered small fishes in a Mediterranean coastal stream

This study analysed the microhabitat use of three endangered fish species, Andalusian toothcarp Aphanius baeticus , Iberian loach Cobitis paludica and sand smelt Atherina boyeri , in a coastal stream stretch. Plastic minnow traps were set both during daytime and at night on the bottom and at the surface. Depth and presence of effective refuge were recorded for each trap. To assess size‐related changes in microhabitat use individuals of each species were classified in three size classes. The three species preferentially used bottom positions in the water column, though this behaviour was more evident in the case of Iberian loach. While large Iberian loach remained active at night Andalusian toothcarp and sand smelt were strongly diurnal, especially larger individuals. The three species showed a clear ontogenetic change in microhabitat preferences towards deeper waters. Small Andalusian toothcarp and medium‐sized Iberian loach used deeper microhabitat in the presence of refuge. Large Andalusian toothcarp consistently preferred exposed microhabitat at any time. Andalusian toothcarp using refuge were smaller at any time, while Iberian loach followed this pattern only at night. The differential vulnerability of these species to different predators (aerial and aquatic; diurnal and nocturnal) could explain the observed patterns in microhabitat use. Fish tended to co‐occur in microhabitats either due to habitat characteristics independently of species or due to species independently of habitat. Andalusian toothcarp segregated spatio‐temporally from sand smelt and Iberian loach, but these species occurred independently of each other. According to these co‐occurrence patterns, Andalusian toothcarp would be more sensitive than Iberian loach or sand smelt to interspecific interactions.     

Differential seed and seedling predation by crabs: impacts on tropical coastal forest composition

Recently, the importance of seed predation by crabs on mangrove species distributions and densities has been established by several studies. In a tropical coastal terrestrial forest in Costa Rica, we investigated the relative importance of predation by land crabs, Gecarcinus quadratus, and hermit crabs, Coenobita compressus, on measured forest composition through a series of seed removal and seedling establishment experiments. We also used natural light-gaps and adjacent non-gap sites to test how canopy cover affects crab predation (seed removal) and seedling establishment. We found fewer tree species (S=18) and lower densities (seedlings, saplings, and adults) in the coastal zone within 100 m of coastline, than in the inland zone (S=59). Land crab densities were higher in the coastal zone (3.03±1.44 crabs m^–2) than in the inland zone (0.76±0.78 crabs m^–2), and hermit crabs were not present in the inland zone. Seed removal and seedling mortality also were higher in the coastal zone than in the inland zone, and in the open controls than in the crab exclosures. Mortality of seeds and seedlings was two to six times higher in the controls than exclosures for four of the five experiments. Crabs preferred seeds and younger seedlings over older seedlings but showed no species preferences in the seed (Anacardium excelsum, Enterolobium cyclocarpum, and Terminalia oblonga) and seedling (Pachira quinata and E. cyclocarpum) stages. We conclude that the observed differences in tree densities were caused by differential crab predation pressure along the coastal gradient, while the differences in species composition were due to predator escape (satiation) by seed quantity. Canopy cover did not affect seed removal rates, but did affect seedling survival with higher mortality in the non-gap versus gap environments. In summary, crab predation of seeds and seedlings, and secondarily canopy cover, are important factors affecting tree establishment in terrestrial coastal forests.     

Predator satiation and recruitment in a mast fruiting monocarpic forest herb

Background and Aims

Cross-pollination and satiation of seed predators are often invoked to explain synchronous mast reproduction in long-lived plants. However, explanations for the synchronous death of parent plants are elusive. The roles of synchronous seeding and post-reproductive mortality were investigated in a perennial monocarpic herb (Isoglossa woodii) in coastal dune forest in South Africa.

Methods

Pre-dispersal seed predation and seed production were assessed by measuring fruit and seed set of inflorescences sprayed with insecticide or water and with no spray treatments. Seed predation was measured at different densities of I. woodii plants by monitoring removal rates of seed from the forest floor. The influence of adult plants on establishment of I. woodii seedlings was assessed by monitoring growth and survivorship of seedlings in caged and uncaged 1 × 1 m plots in understorey gaps and thickets.

Key Results

Fruit and seed set were similar between spray treatments. An I. woodii stem produced 767·8 ± 160·8 seeds (mean ± s.e.) on dune crests and 1359·0 ± 234·4 seeds on the foredune. Seed rain was greater on the foredune than in other topographic locations. Seed predation rates were 32 and 54 % on dune crests and in dune slacks, respectively, and decreased with seed abundance, number of inflorescences per stem and plant height. Seedling recruitment was greater beneath synchronously dying adult plants than in natural understorey gaps (no I. woodii). However, seedling growth rate beneath I. woodii mid-way through its life-cycle was less than in gaps, although survivorship was similar.

Conclusions

The selective advantage of masting in I. woodii derives from satiation of both pre- and post-dispersal seed predators. In addition, post-seeding mortality of adult plants facilitates seedling establishment. Satiation of seed predators and the benefits of seedling establishment are strong drivers of the evolution of synchronous monocarpy in I. woodii.     

Subtle differences in two non‐native congeneric beach grasses significantly affect their colonization,spread, and impact

Comparisons of congeneric species have provided unique insights into invasion ecology. Most often, non‐native species are compared to native ones to look for traits predicting invasion success. In this study, we examine a different facet of congeneric comparisons in which both species are non‐native. Ecological variability among non‐native congeners might 1) lead to the inhibition or facilitation of either species’ ability to colonize and spread, 2) result in larger cumulative impacts due to synergies between species, and 3) depend on the physical context of the invaded habitat. To explore these possibilities, we studied the distribution and abundance of two non‐native beach grasses: European beach grass Ammophila arenaria and American beach grass Ammophila breviligulata, their interaction with one another, and their biotic and physical impacts on dune ecosystems of the Pacific coast of North America. We found that over a two‐decade period, A. breviligulata has increased its dominance over A. arenaria on dunes where it was originally planted in 1935 and has actively spread to new sites formerly dominated by A. arenaria. Our results also show that dune plant species richness was lower at A. breviligulata sites, although there was an increase in the native beach grass Elymus mollis. More significantly, we found that the two grass species are associated with significantly different foredune shapes that are likely controlled by a combination of variability in sand supply along the coast and subtle differences in the congeners’ morphology and growth form. These differences have significant implications for the coastal protection services of dunes to humans and the conservation of native species. They provide a cautionary tale on the impacts of introducing novel species based purely on analogy with closely related species.     

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.     

Factors influencing the natural regeneration of the pioneering shrub Calligonum mongolicum in sand dune stabilization plantations in arid deserts of northwest China

Calligonum mongolicum is a successful pioneer shrub to combat desertification, which is widely used for vegetation restoration in the desert regions of northwest China. In order to reveal the limitations to natural regeneration of C. mongolicum by asexual and sexual reproduction, following the process of sand dune stabilization, we assessed clonal shoots, seedling emergence, soil seed bank density, and soil physical characteristics in mobile and stabilized sand dunes. Controlled field and pot experiments were also conducted to assess germination and seedling emergence in different dune soil types and seed burial depths. The population density of mature C. mongolicum was significantly different after sand dune stabilization. Juvenile density of C. mongolicm was much lower in stabilized sand dunes than mobile sand dune. There was no significant difference in soil seed bank density at three soil depths between mobile and stabilized sand dunes, while the emergence of seedlings in stabilized dunes was much lower than emergence in mobile dunes. There was no clonal propagation found in stabilized dunes, and very few C. mongolicum seedlings were established on stabilized sand dunes. Soil clay and silt content, air‐filled porosity, and soil surface compaction were significantly changed from mobile sand dune to stabilized dunes. Seedling emergence of C. mongolicm was highly dependent on soil physical condition. These results indicated that changes in soil physical condition limited clonal propagation and seedling emergence of C. mongolicum in stabilized sand dunes. Seed bank density was not a limiting factor; however, poor seedling establishment limited C. mongolicum's further natural regeneration in stabilized sand dunes. Therefore, clonal propagation may be the most important mode for population expansion in mobile sand dunes. As a pioneer species C. mongolicum is well adapted to propagate in mobile sand dune conditions, it appears unlikely to survive naturally in stabilized sand dune plantations.     

Spatial concordance at a regional scale in the regeneration process of a circum-Mediterranean relict (Buxus balearica): connecting seed dispersal to seedling establishment

Both microhabitat and temporal scales have significant effects on the regeneration process of plant species. However, the consistency of those patterns at large scales is largely unknown, despite of being essential from both an ecological and a conservation viewpoint. In this study, we examined the spatio‐temporal variation in the process of regeneration through seeds of the relict Mediterranean shrub Buxus balearica. By means of observations and experiments, we quantified the losses at different recruitment phases (seed rain, seed predation, seedling emergence and establishment) both at a small spatial scale (distance and microhabitat effects) and at a temporal scale (four years), and compared them at a regional scale (10 different localities). We additionally studied transition probabilities between phases in two populations and measured the size structure of 18 localities encompassing all the distributional range of the species. The results show that seed shadow follows a leptokurtic pattern which is consistently maintained until the seedling phase (i.e. there is spatial concordance). Despite post‐dispersal factors are spatially uncoupled, they do not remove the gradient established in the seed shadow. Although the importance of different seed predators varied between regions, post‐dispersal seed predation (ca 80%), together with seedling emergence (usually lower than 10%, some years even nil), and seedling mortality during the first summer (ca 70%) were – consistently in all populations – the most critical bottlenecks in the regeneration of the species. Moreover, an absence of seed bank prevents the buffering of dramatic losses during such phases. Watering of sown seeds increased seedling emergence and survival, and number of leaves per seedling, and survival was consistently higher under vegetation coverage. These results suggest that hydric stress notably contributes to limit the regeneration of this once widespread species evolved under a subtropical climate, which nowadays appears in old‐structured populations throughout its distributional area.