Do seed and microsite limitation interact with seed size in determining invasion patterns in flooding Pampa grasslands?

Additive influences of the invasion ability of species (invasiveness) and the characteristics of the habitat which make it invasible (invasibility) cannot fully explain grassland invasion patterns. We tested the hypothesis that different species assemblages of grassland communities may partly result from interactive influences between the relative invasiveness of available species and community invasibility. During 10 months, we evaluated seed and microsite limitation of seedling emergence, survival, and recruitment of plants belonging to species with different seed size (large-seeded species vs. small-seeded species), with in a two-phase community mosaic typical of semi-natural grasslands in the southern flooding Pampa of Argentina. Seeds of large versus small-seeded species were sown either in species-poor patches dominated by a tall tussock grass (“pajonal”) or in species-rich patches dominated by short grasses (“matrix”), subjected to different levels of canopy disturbance (cut vs. uncut). Seed addition promoted seedling emergence for 7 out of the 10 species sown, and this effect was higher for large than for small-seeded species. After seed limitations were removed, interactive effects among seed size, community state and canopy disturbance reflected a strong positive influence of seed size on plant recruitment only in cut pajonal patches. Therefore, according to the stage of invasion process, relative species success may depend on non-interactive (seed-size effect on seed limitation to seedling emergence) or interactive influences among species invasibility and community invasiveness (from seedling emergence to plant recruitment). As a general conclusion, different assemblages of species are expected to successfully colonize spatially close grassland patches, according to both the available invasible species (seed size) and the community state and stage (species composition and canopy disturbance).     

Population dynamics of the perennial Plantago media in semi‐natural grasslands

Abstract. In this study we investigated population dynamics of a perennial forb, Plantago media, in semi‐natural grasslands in southeastern Sweden. Plantago media is a rather common inhabitant of grasslands in this region, but it has been shown to experience dispersal limitations both among managed grassland sites and at potential dispersal routes along road verges. The demography of P. media was studied in 22 permanent plots at two sites over five years. A seed sowing experiment was also performed at each site. The life‐cycle of P. media includes seedlings, juveniles, small vegetative rosettes, large vegetative rosettes and flowering rosettes. The population growth rates (λ‐values) were negative for both populations, with one exception: the first year transition matrix at one site. The projected life span for individuals surviving from seedlings to flowering rosettes varied between 19.6 yr and 227.8 yr. Elasticity analysis showed that remaining in the large rosette stages, both vegetative and flowering, was the most important factor influencing population growth rate. LTRE analysis gave similar results, indicating that spatial and temporal variation have similar effects on the population growth rate. The expected time to extinction in populations with an initial size of between 100 and 1000 individuals varied between 60 and 200 yr. The seed sowing experiment showed that seedling emergence was enhanced by both seed addition and disturbance, suggesting that recruitment is limited by seed and microsite availability. Recruitment was not found to have much impact on population growth rate. However, long‐term population maintenance may depend on occasional small‐scale disturbances that enhance pulses of recruitment. Fragmented landscape is likely to effect plant populations including rare and endangered species as well as more common species which have limited dispersal mechanisms.     

Factors Affecting Seed Germination and Seedling Establishment of Fen-Meadow Species

Availability of seeds and provision of “safe sites” for seedling recruitment are essential for successful restoration of seminatural grassland communities. Inability to provide species‐specific conditions for seedling recruitment appears to be a major factor limiting establishment of fen‐meadow species on restoration sites. This contention was tested in the field for both germination and establishment conditions for a selection of fen‐meadow species. A Cirsio‐Molinietum fen meadow and an agriculturally semi‐improved species‐poor grass dominated rush pasture were used. Seeds of Carex ovalis, Cirsium dissectum, Molinia caerulea, Succisa pratensis, and Holcus lanatus were sown onto treatments comprising either irrigation or no irrigation, presence or absence of existing vegetation canopy, and presence or absence of soil disturbance. Germination of all except H. lanatus was higher in the fen meadow than in the rush pasture. The fen‐meadow site was less susceptible to drought, provided more light to the seed environment, and showed a stronger day–night variation in relative humidity compared with the rush pasture. All the fen‐meadow species responded strongly to the experimental treatments, whereas H. lanatus showed only a small response. Soil disturbance was the major factor that increased germination. Removal of the vegetation canopy improved germination only in S. pratensis. Conditions affecting survival of seedlings were different from those affecting seed germination. Seedling survival was greater on the fen‐meadow site than on the rush pasture. Canopy presence was the major factor that reduced seedling survival. Few seedlings survived in the presence of the rush pasture canopy. Irrigation and soil disturbance were of minor importance for seedling survival on both sites. Safe sites for seed germination and seedling establishment of fen‐meadow species existed on the fen meadow even without soil disturbance and gap creation. Safe sites for seedling recruitment were not present in the rush pasture. The need for species‐specific definition of safe site characteristics at the two stages of seedling recruitment (i.e., for seed germination and for seedling survival) was demonstrated. The implications of these findings for restoration of seminatural grasslands are discussed.     

Bryophyte Mats Inhibit Germination of Non-native Species in Burnt Temperate Native Grassland Remnants

Species-rich native grasslands in western Victoria, Australia, are often small, have a high perimeter to area ratio and are surrounded by non-native species. Few non-native species, however, have invaded them. A feature of species-rich grasslands is the presence of a bryophyte mat (composed of mosses and liverworts) that carpets the intertussock spaces. I assessed the role of these mats in plant invasions by sowing three non-native species (Briza maxima, Hypochoeris radicata, Plantago lanceolata) in replicated disturbed (mats removed) and undisturbed (mats intact) microsites at three grassland remnants (two recently burnt, one unburnt for 3 years) and followed seedling emergence, survival and growth for 5 months. Three native species were also sown for comparison. The rate of germination and total percent germination of non-native species were significantly enhanced at both burnt sites when the mat was disturbed. The large-seeded Briza maxima failed to germinate at both burnt sites in the absence of soil disturbance. The native species generally did not show a strong germination or growth response to soil disturbance in burnt areas. At the unburnt site, where monthly percent soil moisture was highest, final percent germination of the non-native and native species was greatest of any site in both microsites, and germination was not significantly affected by soil disturbance. Differences in the seed morphology of native and non-native species may play an important role in their ability to establish on bryophyte mats in moisture-limiting environments. Any activity that disrupts the mats in the frequently burnt, species-rich grassland remnants is likely to significantly enhance the germination and subsequent growth by non-natives. However, where burning is infrequent, germination of some non-native species may be expected, regardless of disturbance, although growth will likely be favoured in disturbed areas.     

子午岭三种生境下辽东栎幼苗定居限制

辽东栎(Quercus wutaishanica)是子午岭地区的乡土乔木树种,也是该地区气候顶极群落的建群种,其幼苗的补充更新影响着森林群落的结构及物种组成。在3种典型生境(辽东栎林、人工油松林、灌草丛)中,设置3因素(种子、干扰、遮荫)两水平的野外播种实验(随机区组设计),记录辽东栎幼苗出苗量,并监测幼苗同生群在3种生境中3a间的生长状况,以确定种子及微生境在辽东栎幼苗补充过程中的限制作用。播种实验样方大小30cm×30cm,共计216个样方。结果显示,在辽东栎林及油松林内,增加种子和干扰强度(去除枯落物),能引起出苗量和幼苗补充量的显著增加,且2种处理方式间存在交互作用,表明在郁闭林冠下,辽东栎幼苗的补充受到了种子和微生境的双重限制,枯落物是导致微生境限制的主要因素之一。在灌草丛生境,各种处理方式均不能增加幼苗的补充量,表明辽东栎无法在开阔生境(强光照、干旱)中完成实生幼苗的补充更新。3种生境中的幼苗同生群生存分析表明,辽东栎幼苗在森林群落中存活率显著高于灌草丛群落。根据幼苗生长指标判断,在3种生境中,人工油松林是辽东栎幼苗定居的最佳场所。与实验预期相反,灌木对辽东栎幼苗的补充无显著影响。     

Recruitment in species-rich grasslands: the effects of functional traits and propagule pressure

Aims The assembly of plant communities is a complex process which combines impacts from the species pool, dispersal and propagule pressure, niche requirements of colonizing species and the niche structure of the community. Recent theory development has incorporated all these aspects, e.g. in 'stochastic niche theory'. We investigated recruitment into a species-rich grassland community, using an experimental approach where we manipulated the trait composition of the community and examined the success of colonizing species entering with various propagule pressure. Specifically, we examined two predictions: (i) colonization success increases with increasing difference between traits of the colonizing species and the trait profile of the community and (ii) colonization success increases with increasing propagule pressure.Methods The examined communities were species-rich semi-natural grasslands located in southern Sweden. After a careful documentation of the composition of the plant communities at the experimental sites, we manipulated the trait profile of species-rich grassland plots based on the plant functional trait specific leaf area (SLA), which is correlated with several key life history functions. In addition to SLA, seed mass was also used to describe the trait profile of grassland plots. Seeds of 12 plant species from the regional species pool, varying in SLA and seed mass, were sown into plots using four different levels of propagule pressure. Recruitment was examined after 1 year. We also planted juvenile 'plug plants' of the same species which allowed us to examine survivorship and growth beyond the seedling stage.Important findings Overall we found very limited evidence for relationships between the traits of the colonizing species and the trait profile of the community and for recruitment after sowing these relationships were contrary to the prediction. Survival of plug plants after two seasons of growth was high irrespective of the trait profile of the community, but growth of plug plants was affected by the trait profile of the surrounding community. For four of the species there was a positive effect of increased propagule pressure on colonization. The results suggest that species assembly in species-rich grasslands is not strongly dependent on the niche structure of the community. However, the finding that colonization of only a third of the species responded positively to increased propagule pressure indicates that there might be niche-related effects that were not captured by our treatments. Overall, our results indicate that the factors determining colonization in this community are species specific. Some species are able to colonize irrespective of niche relationships, provided that the propagule pressure is sufficiently high to overcome stochastic mortality after seed arrival. For other species, however, we cannot exclude that niche assembly occurred, but we failed to identify the relevant niche factor.     

Species diversity and life history traits in calcareous grasslands vary along an urbanization gradient

Calcareous grasslands are among the most species-rich plant communities in Europe with a particularly high nature conservation value. During the past centuries their distribution has markedly decreased, at least partly due to urbanization. Thus we investigated the effects of urbanization on species diversity along a spatio-temporal urbanization gradient from traditionally managed grassland to areas affected by urban developments, which was situated in the plains northwest of Munich, Germany. Both a RLQ analysis linking species and environmental traits, and a redundancy analysis of the plant community features showed that soil disturbance, soil sealing and mean temperature explained most of the environmental variation along the gradient. The species in urban habitats showed increased insect pollination, earlier flowering and prolonged seed longevity. While urbanization favored short-lived species with dysochorous dispersal, the reference grasslands harbored more wind-pollinated perennials with effective vegetative spread and relatively large, short-lived seeds. Compared to the urban sites, traditionally used grasslands had a higher species diversity, more threatened species and a lower proportion of non-natives. We conclude that even under conservation management, urban habitats are not capable of maintaining the original biodiversity. However, we also found threatened species occurring exclusively in urban sites. Hence, urbanization decreased the area and diversity of traditional calcareous grasslands, but it also established niches for endangered species which are not adapted to the living conditions in calcareous grasslands.     

Differential effects of interspecific interactions and water availability on survival, growth and fecundity of three congeneric grassland herbs

Fitness of individual plants and of populations depends on the rates of survival, growth and fecundity. This study tested whether vital rates were differentially affected by biotic interactions and water availability. The effects of manipulations of above-ground competition (through clipping) and water availability (through water addition) on the vital rates of seedlings of three species (Viola elatior Fries, Viola pumila Chaix and Viola stagnina Kit.) were analysed in dry, mesic and wet grasslands. Water addition and grassland type had the largest effects on survival (accounting for 41 and 24% of total variation, respectively) across species. Height growth rate was positively affected by grassland type (19%) and water addition (12%) and varied among species (8%), while leaf accumulation rates and reproduction were affected by grassland type and clipping. The data suggested facilitative effects of the canopy on seedling survival in the dry grassland. This study presents evidence that environmental conditions and biotic interactions may have differential effects on seedling survival, growth and reproduction. The findings highlight the complex interplay between spatial and temporal environmental variation and biotic interactions in structuring plant communities.     

Mass flowering and parental death in the regeneration of Cerberiopsis candelabra (Apocynaceae), a long-lived monocarpic tree in New Caledonia1

Monocarpy is rare among forest trees, and the selective conditions allowing this life history to evolve are poorly known. Here we examined the regeneration dynamics of a New Caledonian monocarpic tree species, Cerberiopsis candelabra, to better understand how postreproductive mortality and mass flowering might contribute to the success of this strategy. We investigated population size structures and seedling establishment patterns in 18 stands following the 2003 flowering event. We found little evidence of recent recruitment. Instead, population size structures suggested a history of one or more substantial recruitment events followed by recruitment failure, despite multiple flowering events in most populations. The canopy gap created by death of an individual parent appeared generally insufficient to allow seedling establishment, but mass flowering and consequent death of many adults appeared to enhance seedling recruitment by opening more of the canopy. Site means of seedling density below dead parents correlated strongly with light availability and with the proportional density of flowering trees. Therefore, we suggest that mass flowering and gregariousness may have been necessary preconditions for the evolution of monocarpy in this species. However, the degree to which regeneration relies on synchronous flowering and gregariousness vs. infrequent catastrophic disturbances remains uncertain.     

Seed Bank Effects on Recovery after Disturbance in Reconstructed Tallgrass Prairies

Litter‐removing disturbances such as fire in grasslands temporarily increase available resources for plants, opening a window of opportunity for new establishment as communities recover. At this time, new individuals or species could be added to the community as a result of germination from the local seed bank. In reconstructed grasslands this may be problematic, as the seed bank may contain a suite of undesired species reflective of prior and surrounding land uses. In two, 25‐year‐old, low‐diversity reconstructed grasslands, we tested the effect of local seed bank establishment following litter‐removing disturbance using seedling removal plots (1 m²) and plots where natural seedling establishment was allowed. Following disturbance, the vegetation was either left intact or hayed to enhance seedling establishment (a common practice following inter‐seeding efforts). Although the seed bank and seedling community were dominated by resident grasses (Andropogon gerardii and Poa pratensis), recruitment from the seed bank increased species richness and reduced evenness through the addition of forb species (including Cirsium arvense) in one of the study sites. Haying temporarily altered the abundances of the dominant grasses, but did not consistently affect seedling recruitment. Disturbances that facilitate seed bank recruitment may promote establishment of undesired species within reconstructed grassland communities, and we need to take steps to better manage the contributions into and recruitment from the seed bank to reconstruct sustainable grasslands.     

The role of species richness for recruitment in a seminatural grassland

I examined two aspects of how recruitment is influenced by species richness in a seminatural grassland: effects of species richness among colonisers in a seed mixture, and effects of species richness, ramet density, grass and moss cover in the colonised vegetation. The results suggest that recruitment is higher in more species-rich seed mixtures (for three of the four target species: Anthyllis vulneraria, Centaurea jacea, Filipendula vulgaris and Primula veris ). Average recruitment of the target species was negatively related to ramet density in the colonised vegetation, but not affected by grass or moss cover. The results imply that mechanisms other than interspecific competition among species are important in structuring the community during the recruitment phase. The negative effects of ramet density and the positive effects of species richness may be a result of strong intra-specific competition, or species-independent competition in combination with diversity effects enhancing microsite variability.     

Hybrid ecosystems can contribute to local biodiversity conservation

Calcareous grasslands have become severely threatened habitats in Europe. The aim of this study was to investigate the changes in plant species richness, and functional and phylogenetic diversity in northern Estonian calcareous (alvar) grasslands resampled after 90 years of land-use change. Functional traits characterizing species that have benefited most from decreased habitat area and altered environmental conditions, and additional species that can potentially inhabit the remaining grassland patches were identified. Also changes in the relative amount of habitat-specific species were studied to detect a possible decrease in habitat integrity. Although grasslands in the studied region had lost most of their original area (~90 %), species richness had substantially increased due to invasion by more competitive, nutrient-demanding native species. Functional diversity generally increased, whereas phylogenetic diversity showed no response to altered conditions. Overall, these grasslands have lost their integrity as calcareous grassland habitat type in the region, because the relative amount of habitat-specific characteristic species has declined significantly. However, although the grasslands have transformed to a ‘hybrid’ habitat type and restoration to their previous state is likely not reasonable, such degraded species-rich grassland fragments can still be recognized as important habitats to preserve high local biodiversity and several characteristic species of calcareous grasslands. As current landscapes consist of an increasing number of hybrid and novel communities, new tools to supplement traditional conservation or restoration practices are necessary to recognize and maintain regions and habitats of high local biodiversity.     

Limits to tree species invasion in pampean grassland and forest plant communities

Factors limiting tree invasion in the Inland Pampas of Argentina were studied by monitoring the establishment of four alien tree species in remnant grassland and cultivated forest stands. We tested whether disturbances facilitated tree seedling recruitment and survival once seeds of invaders were made available by hand sowing. Seed addition to grassland failed to produce seedlings of two study species, Ligustrum lucidum and Ulmus pumila, but did result in abundant recruitment of Gleditsia triacanthos and Prosopis caldenia. While emergence was sparse in intact grassland, seedling densities were significantly increased by canopy and soil disturbances. Longer-term surveys showed that only Gleditsia became successfully established in disturbed grassland. These results support the hypothesis that interference from herbaceous vegetation may play a significant role in slowing down tree invasion, whereas disturbances create microsites that can be exploited by invasive woody plants. Seed sowing in a Ligustrum forest promoted the emergence of all four study species in understorey and treefall gap conditions. Litter removal had species-specific effects on emergence and early seedling growth, but had little impact on survivorship. Seedlings emerging under the closed forest canopy died within a few months. In the treefall gap, recruits of Gleditsia and Prosopis survived the first year, but did not survive in the longer term after natural gap closure. The forest community thus appeared less susceptible to colonization by alien trees than the grassland. We conclude that tree invasion in this system is strongly limited by the availability of recruitment microsites and biotic interactions, as well as by dispersal from existing propagule sources.     

Gap disturbance triggers the recolonization of the clonal plant Ambrosia tenuifolia in a flooding grassland of Argentina

Abstract Flooding provokes the death of many dicotyledonous species in grazed grasslands of the Flooding Pampa in Argentina, including the clonal plant Ambrosia tenuifolia, which produce the opening of numerous gaps. The objective of this study was to investigate the recolonization of grassland by A. tenuifolia after this species disappeared due to the occurrence of prolonged flooding events. To this end, responses of seed germination to environmental factors associated with gaps, such as light quality and temperature regime, conditions related to seedling survival, and clonal growth of ramets outside the gaps were studied in two different experiments in the field. Environmental factors related to gaps promoted the recruitment of new genets. The combined effect of alternating temperatures and the high red : far‐red ratio set off germination from the soil seed bank; germination also was enhanced when signals were generated artificially under the intact canopy in the field. Higher resource availabilities and maximum seedling survival were recorded in canopy gaps, which were the focus of invasion. Grassland recolonization outside the gaps continued rapidly by clonal growth, from small gaps and large ones, even within the dense surrounding canopy. This provoked an intense competition with the other species. Gap opening by disturbances, seed germination in gaps and clonal growth were decisive for the recolonization of A. tenuifolia populations. This sequence of events triggered the recolonization of the plant community by this species, in sites where it had been eliminated by prolonged flooding. This process represents one of the most significant fluctuations in the vegetation dynamics of the Flooding Pampa Grasslands.     

Land use heterogeneity causes variation in demographic viability of a bioindicator of species-richness in protected fen grasslands

Land use change is one of the main drivers of species extinction. In Europe, grasslands are under active conflict between conservation efforts and increasing agricultural pressures. Here, we examine the demographic effects of differential land use on the herbaceous perennial Trollius europaeus L. (Ranunculaceae), a bioindicator of species-richness and ecosystem services in wet grasslands of Central Europe. Demographic data were collected in 2006–2009 from nine populations in seven protected sites of northeastern Germany representing four land use types. We constructed stage-based matrix population models to explore the effects of various land management on demographic viability of focal populations. We show that most studied populations are declining (λ < 1), although the estimates of local extinction vary between ≤15 years for grazed and woodland populations, and 20–99 years for mown and abandoned populations. The joint information from our elasticity analyses and life table response experiments revealed that reproduction, growth of small vegetative individuals and survival of reproductive stages are most important for population viability. Our study shows that the current land uses in protected areas where T. europaeus is found is incompatible with its long-term viability. We suggest that, when compatible with in situ practices, grasslands containing this species be mown after maturity in order to enhance seedling recruitment and to reduce competition for juveniles. Prolonged extinction times in abandoned populations offer a buffer to develop conservation schemes there. An improvement of conservation measures is urgently needed to maintain the populations of this important bioindicator and its associated community of moist species-rich fen grasslands.     

雪灾干扰后林冠开阔度对黄心树幼苗更新的影响

幼苗更新是森林更新的关键过程,而林窗对于植物幼苗更新、生长及最终存活具有重要影响。极端气候事件会使森林在短时间内形成大量林窗,从而显著影响林内幼苗更新和存活。本研究以2015年雪灾后云南省哀牢山中山湿性常绿阔叶林优势种黄心树(Machilus bombycina)幼苗为对象,对194个1 m×1 m样方中的幼苗生长和死亡动态进行为期4年的监测,利用线性混合模型(LMM)和广义线性混合模型(GLMM),分析林冠开阔度对幼苗高度相对生长率、叶片相对增长率、新增率和死亡率的影响,研究了雪灾干扰后的林冠开阔度对植物幼苗更新的影响。结果表明:(1)林冠开阔度随时间推移逐渐变小(20.07%～9.97%),且速度由快到慢;(2) 2015—2016年,已有幼苗高度相对生长率与林冠开阔度呈显著正相关(P<0.05),其余年份两者间无显著相关性; 2016—2017年已有幼苗叶片相对增长率与林冠开阔度呈显著正相关(P<0.05),其余年份两者间无显著相关性;(3)林冠开阔度与幼苗新增率和死亡率均无显著相关性。本研究表明,雪灾后造成的林冠开阔度增加有利于黄心树幼苗生长。     

Defining grassland fire events and the response of perennial plants to annual fire in temperate grasslands of south-eastern Australia

Many species-rich Themeda triandra grasslands in south-eastern Australia have been burnt annually in summer as a land management practice for decades. The characteristics of annual fires (maximum surface temperatures, maximum soil temperature changes at 10 mm depth, Byram fire intensity and duration of surface heating) were compared to fires that occur less frequently (2, 4 or 7-year inter-fire interval). The impacts of annual summer fires on seed survival, perennial plant dynamics and flowering were also documented at two sites over two years.Annual grassland fires differ principally from other grassland fire types in their duration of surface temperatures above 100°C (i.e., <1 min versus >2–3 min in grasslands burnt at 4–7-year intervals). This was correlated to fuel load, but not rate of spread or Byram fire intensity. Maximum surface temperatures were variable within annual fire events (98–458°C ) and did not differ significantly from those observed in other grassland fires. All sites experienced temperatures above 350°C but no site had its soil temperature increase at 10 mm depth by >10°C during fire. Byram fire intensity was lower in annually burnt sites, but positively correlated with rate of spread moreso than fuel load.Many perennial species (34–55%) and most perennial individuals (70–83%) avoid the annual summer fire event by being dormant at the time of burning. All perennial species regenerated rapidly by vegetative means after fire and seedlings were absent or rare for most species. As a result, turnover of species at the small-scale was low.Annual burning permits high perennial species richness (mean 18 species/0.25 m²) and high numbers of individual perennial plants (range 189–1036 plants/0.25 m²) to co-exist at the small-scale. Seasonal plant density maxima peaked in spring, 6–8 months after fire, and the number of plants in a species' population each year following fire depended on their seasonal response more than their direct fire response. The direction and magnitude of population change from one year to the next was species and site specific and did not correlate with life form or a plant's ability for vegetative spread. All perennial species have a short secondary juvenile period (i.e., 6–11 months) but relatively few individual plants flower in any one year. Seed that has made its way into the soil is completely protected from the direct effects of fire and hence, occasional post-fire seedling recruitment may be possible for all species.Maintaining annual burning in grasslands as a management regime is unlikely to lead to a decline in richness and plant density in the short-term. Rather, delayed burning (i.e., >3 year intervals), and the impact that this may have on interval dependent processes such as above-ground competition, are predicted to have more substantial long-term effects on the small-scale dynamic of this community.     

Effect of seed predation on seed bank size and seedling recruitment of bush lupine (Lupinus arboreus)

Whether seed consumers affect plant establishment is an important unresolved question in plant population biology. Seed consumption is ubiquitous; at issue is whether seedling recruitment is limited by safe-sites or seeds. If most seeds inhabit sites unsuitable for germination, post-dispersal seed consumption primarily removes seeds that would otherwise never contribute to the population and granivory has minimal impacts on plant abundance. Alternatively, if most seeds ultimately germinate before they lose viability, there is greater potential for seed consumption to affect plant recruitment. Of the many studies on seed consumption, few ask how seed loss affects seedling recruitment for species with long-lived seed banks. We examined post-dispersal seed predation and seedling emergence in bush lupine (Lupinus arboreus), a woody leguminous shrub of coastal grasslands and dunes in California. We followed the fate of seeds in paired experimental seed plots that were either protected or exposed to rodent granivores in grassland and dune habitats. Significantly more seeds were removed by rodents in dunes than grasslands. In dunes, where rodent granivory was greatest (65% and 86% of seeds removed from plots by rodents in two successive years), there is a sparse seed bank (6.6 seeds m⁻²), and granivory significantly reduced seedling emergence (in the same two years, 18% and 19.4% fewer seedlings emerged from exposed versus protected plots), suggesting seed rather than safe-site limited seedling recruitment. In contrast, rodents removed an average of 6% and 56% of seeds from grassland plots during the same two years, and the grassland seed bank is 43-fold that of the dunes (288 seeds m⁻²). Even high seed consumption in the second year of the study only marginally influenced recruitment because seeds that escaped predation remained dormant. Burial of seeds in both habitats significantly reduced the percentage of seeds removed by rodents. Results suggest that granivores exert strong but habitat-dependent effects on lupine seed survival and seedling emergence. Received: 24 October 1996 / Accepted: 4 February 1997     

Experimental demography of the rare Gentianella germanica: seed bank formation and microsite effects on seedling establishment

The persistence of populations of short-lived species requires regular reproduction and seedling establishment. A persistent seed bank can buffer populations against extinction in unfavourable years. We experimentally investigated seed fate in Gentianella germanica, an endangered biennial species characteristic for species-rich nutrient-poor calcareous grasslands in central Europe. We studied the effect of experimental gaps on seedling establishment from sown seeds and the fate of seeds buried in bags over two years. In December 1993 experiments were established at seven calcareous grassland sites in the Swiss Jura mountains. In spring 1994 seedlings emerged in all plots where seeds had been sown, including previously unoccupied patches. This suggests that limited dispersal within sites contributes to small population sizes. Significantly more seedlings emerged at sites with current populations of G. germanica than at unoccupied sites (5.95% vs 3.40%). Because this difference was not explained by germinations from the natural seed bank it indicates differences in habitat quality. Clipping of the vegetation and disturbance of the soil reduced vegetation cover in the following spring and enhanced seedling emergence. In undisturbed plots 4.5% of seeds sown produced a seedling in spring 1994, whereas in plots with clipped vegetation 9.9% and in disturbed plots 12.7% produced seedlings (p>0.01). This suggests that management measures which create gaps in the vegetation (e.g. grazing) could positively influence population size and persistence of G. geymanica. On average, we recovered 7.55% viable seeds after one year of burial in bags, and 4.05% after two years, indicating that G. geymanica has a persistent seed bank. The demographic data indicate that the number of viable seeds in the seed bank exceeds the number of established plants in a population at least by a factor of 20. Restoration of extinct populations of the species from the seed bank may thus be possible if appropriate management measures are taken within a few years.     

Marginal holly (Ilex aquifolium L.) populations in Mediterranean central Spain are constrained by a low-seedling recruitment

Seedling density and survival was studied in holly woodland located in the mountains of the Sistema Central, Madrid (Spain) in 1998. Ten permanent quadrats of 50×50 cm were randomly placed in each of the following habitats: under closed holly canopy, in canopy gaps, at the edge of the holly woodland, in the adjacent open grassland, and under fleshy- and dry-fruited shrubs located in this grassland. Emerged seedlings were periodically monitored and their survival was checked during the following 2 years. Additionally, a set of 20 seedlings were planted in each of the cited habitats and their survival was followed in parallel with the naturally emerged seedlings. The influence of light on holly survival was tested by experimental planting of holly seedlings under six different controlled light intensities.Seedling emergence was strongly habitat dependent, with the highest seedling densities in the closed holly woodland and in canopy gaps, and a complete absence in the grassland. Survival to the first year was nil. No naturally emerged seedling reached 1 year of age, independent of the habitat studied. Only one of the planted seedlings in the field survived to the first year. Despite this, mean seedling lifespan was different between habitats, with canopy gaps, woodland edge, and under fleshy-fruited shrubs being the habitats with highest average lifespan. Seedling response to the distinct light intensities is largely similar to that seen in the different field habitats, but other factors in addition to light, mainly summer drought and disturbances are proposed to explain the 100% mortality observed.Seedling incorporation is the recruitment dynamic bottleneck of this holly population, being more severe here than in other places studied further north. This may explain the increasing rarity of the species in the southern half of the Iberian Peninsula, where Ilex is close to its southern limit of its distribution.