Vertical seed distribution in the soil constrains regeneration of Bromus pictus in a Patagonian steppe

Question: What is the effect of vertical seed distribution (i.e. seeds on soil surface or buried) and the relative importance of vertical and horizontal seed distribution (i.e. seeds near established vegetation or near bare soil) for seedling emergence? Location: Patagonian arid steppe. Methods: To evaluate the effect of vertical seed distribution on seedling emergence, four sowing treatments were used in field and greenhouse experiments. Vertical vs. horizontal seed distribution and the occurrence of seedlings were analysed in the field. Effects of trampling and mechanical drilling on seedling emergence were investigated experimentally in the field. Results: In field and glasshouse experiments buried seeds showed a higher emergence and larger seedling size than seeds lying on the soil surface. The observational field study indicated that natural emergence was not associated with particular microsites or plant cover (horizontal distribution), but that most seedlings emerged from buried positions (vertical distribution). Buried seeds represented less than 10% of the sampled seed cohort but account for almost 60% of the seedlings. The trampling/drilling experiment showed that sheep trampling and mechanical drilling increased seedling emergence compared to non‐treated controls. Conclusions: The population dynamics of Bromus pictus is strongly constrained by the vertical position of seeds, which largely affects emergence, and seed horizontal distribution, which largely affects seedling survival. Our results showed that sheep trampling after seed dispersal may be a low‐input technique for increasing grass recruitment.     

Interaction between large herbivore activities,vegetation structure,and flooding affects tree seedling emergence

Tree establishment in grazed vegetation mosaics involves a series of early bottlenecks, including seed dispersal, germination, seedling emergence, survival and growth. In a field experiment, we studied seedling emergence of two species with contrasting recruitment strategies, Fraxinus excelsior and Quercus robur, in five structurally different vegetations: grazed and ungrazed grassland, ruderal pioneer vegetation, soft rush tussocks, tall sedge mats and bramble scrub. In a simulation experiment, we studied the interaction effects of pre-emergence flooding (3 weeks of inundation), trampling and grazing (simulated by clipping) of grassland vegetation on the emergence and early growth of both tree species in grass swards. Seedling emergence was enhanced in low swards and sparse vegetation types. Despite different recruitment strategies, the interaction of flooding and trampling of swards enhanced seedling emergence of both species. Grazing of soft rush and tall sedges enhanced emergence of F. excelsior. Clipping grass swards increased early growth of emerging Q. robur. Our results support the hypothesis that natural disturbances of soil and vegetation create microsites for seedling emergence and reduce above-ground competition. In grazed systems however, these results suggest a discordant relationship between successful seedling emergence and subsequent seedling growth/survival during the establishment process in structurally different vegetations.     

Epizoochorous and post-dispersal processes in a rare plant species: Jurinea cyanoides (L.) Rchb. (Asteraceae)

We used Jurinea cyanoides as a model plant species to ask the question whether sheep play a dual role in enabling the establishment of a rare species (1) by epizoochorous transport of diaspores and (2) by influencing post-dispersal processes. We carried out two field experiments in sand grassland (Jurineo–Koelerietum glaucae) in the northern upper Rhine valley, Germany. In the first one, we attached diaspores to the fleeces of sheep and assessed retention time, seed shadow, seedling emergence and establishment. In the second one, diaspore displacement by sheep trampling was investigated.It could be shown that establishment of J. cyanoides is possible after sheep-epizoochorous dispersal, but is severely limited by several factors. First, retention time is mostly short (only 18% of the diaspores remained in the fleeces >2 h). Thus, the seed shadow is concentrated around the place of attachment (mostly <5 m distance, maximum 17 m). Second, seedling emergence of dispersed achenes is severely limited by above-ground granivory (99%). Third, establishment is limited by seedling mortality (68%) due to summer drought. The seedlings germinated from achenes that had been incorporated into the soil (or into deeper parts of the bryophyte layer). The second experiment showed that sheep trampling significantly enhanced the incorporation of achenes into the soil compared with controls. Our study gives evidence that sheep can play a dual role: diaspore transport and facilitation of establishment by trampling.     

Contrasting effects of grazing on the early stages of woody encroachment in a Neotropical savanna

Woody encroachment in savannas represents an ecological process of current global interest given its negative impact on ecosystem functioning, particularly on forage production. Traditional savanna models propose competition and niche differentiation as the main mechanisms allowing tree-grass coexistence. Demographic models, instead, propose abiotic and biotic factors as bottlenecks controlling vital rates and transitions from seeds to adult trees. The role played by domestic grazing on woody encroachment is yet controversial. Here, using a multistage tree life approach, we combine both models and evaluate the role of grazing and herbaceous vegetation on woody recruitment in a Neotropical savanna dominated by Vachellia caven, a successful and widely spread encroacher tree species. We performed three experiments to evaluate seed predation, seedling emergence and survival of V. caven by manipulating cattle grazing (grazed and ungrazed areas) and herbaceous vegetation presence (vegetated and unvegetated). Finally, we combined the results of the three experiments to estimate the probability of plant recruitment across these experimental factors. Grazing decreased seed predation by half, did not modify seedling emergence and decreased seedling survival. Herbaceous vegetation did not affect seed predation nor seedling emergence rate, but increased seedling survival. Overall, the net effect of grazing on V. caven recruitment was neutral since the increase in seed availability due to the reduction in seed predation rate was compensated by the negative effect of grazing on seedling survival. Our analysis revealed that cattle grazing and herbaceous vegetation had contrasting effects on the seed and seedling life stages. We propose that in order to restrain the early stages of encroachment, cattle grazing pressure could be managed following the seasonality of demographic tree transitions. Through rotational grazing amongst paddocks, stocking rates could be relaxed during the primary dispersal stage to maximize granivory, and then increased to enhance the chance of seedling consumption and trampling.     

Effects of experimental and real land use on seedling recruitment of six fen species

Seedling emergence and early establishment of six fen species differing in seed mass and growth form were investigated under experimental land use with changed vegetation structure and under real land use in a calcareous fen. Seeds of all six species were sown in plots with different experimental land-use treatments: summer and autumn mowing with or without litter removal, trampling and abandonment. Additionally, emergence and survival of experimentally sown seeds was investigated under real land use on adjacent sites managed by mowing, grazing, intense trampling or abandonment.On abandoned plots and on plots without litter removal of the land-use experiment, emergence rates of all species were negatively affected either by high litter and moss cover or by tall canopy. No differences were found between autumn and summer mowing. Gap creation by experimental trampling did not increase germination rate. Under real land use, establishment of seedlings of most species was positively affected by litter cover and tall canopy. Trampling, in contrast, had a severe negative effect on seedling survival.The investigated species differed in their germination ability which was tested in the germination chamber and in their response to land use. Succisa pratensis with the highest seed mass germinated well in the chamber and in the field more or less regardless of land use. The low germination rate of Parnassia palustris in the germination chamber indicated a limitation of viable seeds. In the field, however, seedling emergence was additionally limited by microsite availability. Seeds of Serratula tinctoria and Primula farinosa germinated well in the germination chamber, but seedling recruitment in the field was hampered in the presence of a high litter or moss cover. Seeds of Tofieldia calyculata and Pinguicula vulgaris were strongly dependent on the availability of suitable microsites in the field. They hardly germinated under natural conditions, in spite of a high number of germinable seeds in the germination chamber.     

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.     

Is fire a selective force of seed size in pine species?

Fire is selectively shaping most of the traits of plants growing in fire-prone environments. However, seed size and other features related to seed production have not been studied in the light of the evolutionary role of fire. Our research tests the hypothesis that larger seeds have a higher chance of surviving wildfires and produce more vigorous seedlings with a lower death rate. To test this hypothesis the germination and early seedling growth of five Spanish pine species were studied. Weight, length and width of all seeds were measured. The biomass (fresh and dry weight) and length (root and total) of subsequent seedlings were also measured after 30 d from emergence. Seeds were submitted to elevated temperatures for periods in which the chance of survival was 50 % (calculated by means of a logistic model for each pine species). The differences observed among species suggests that fire may be adaptively shaping seed size in pines with larger seeds (Pinus canariensis and P. pinaster), because larger seeds are more likely to survive after heat shocks. Furthermore, in P. canariensis, seedlings after heat treatment are even larger than those submitted to control. In P. halepensis, despite being well adapted to fire, our results indicated no relationships between fire and seed characteristics. Finally, although heat treatment has a general adverse effect on seedling growth in the case of the two subalpine pines, we have detected a positive relationship between seed size and seedling growth but only in the largest seeds. This might also suggest the relevance of fire as a selective force for these pines which is outperformed by the relevance of dispersal and emergence time as adaptive traits in the post-fire scenario.     

Seed dispersal effectiveness in a plant–lizard interaction and its consequences for plant regeneration after disperser loss

Mutualistic disruptions, such as those promoted by the loss of seed dispersers, can have negative effects on the plant regeneration of those species that strongly depend upon them. In order to adequately assess how plant communities are affected by such disruptions, we need to know the importance of the dispersal phase, both in its quantitative and qualitative components. We examined this in the narrow interaction between the shrub Daphne rodriguezii and its (only) disperser, the lizard Podarcis lilfordi. We quantified fruit removal and the effect of fruit/seed-size selection, seed treatment in the disperser’s guts and seed deposition patterns on seedling emergence and survival. In the only locality in which lizards persist, they removed most fruits and showed preference for larger ones in one of the two study years. Seed treatment in lizard’s guts had no effect on germination, although it tended to reduce the effect of seed size on germination (differences between large vs. small seeds in seed germination were higher for non-ingested seeds). Probability of seedling emergence, but not survival, was higher in the locality with lizards. Dispersed seeds under heterospecific shrubs showed higher seedling survival than those under conspecifics in all localities, especially the year with higher rainfall. Our findings support that the movement of seeds to nurse shrubs by lizards is the most important component of the seed dispersal process in the only remaining locality where both species coexist.     

Abiotic barriers limit tree invasion but do not hamper native shrub recruitment in invaded stands

The interplay between the invasion of alien plant species and re-colonization of native plant species is important for conservation. Sandy coastal plains (called restinga in Brazil) were used as a model system to explore the abiotic barriers that potentially limit the initial establishment of alien and native woody plants in invaded and non-invaded areas. The study tested the influence of light availability, soil type and litter layer on recruitment of a wind-dispersed alien tree (Casuarina equisetifolia) and two bird-dispersed native shrubs under a Casuarina stand and in the preserved restinga. The effect of soil type and the physical and allelopathic effects of Casuarina litter on seedling emergence of the three species were also evaluated under greenhouse conditions. Low dispersal associated with low seedling emergence and zero survival of young plants (mainly due to microhabitat conditions) apparently prevents the spread of Casuarina in the preserved restinga. The main cause of low recruitment of native species in the Casuarina stand was the physical barrier of the litter. However, if seeds overcome this physical barrier, the presence of litter improves seedling emergence and survival of young plants, mainly because the litter increases soil moisture. Sowing seeds below litter and planting young plants of native shrubs on litter can improve the re-colonization of native plants in invaded areas. In conclusion, Casuarina invasion on sandy coastal plains is strongly limited by abiotic barriers, but anthropogenic disturbances are altering the key processes that naturally make the restinga resistant to invasion.     

花楸树种子散布、萌发与种群天然更新的关系

花楸树是我国东北林区重要的非木质资源树种,其实生天然更新不良.本文通过研究花楸树种子散布、土壤种子库及种子萌发出土过程,分析花楸树实生天然更新的影响因素.结果表明：自然散落的花楸树果实96.1%分布于母株2 m范围内,凋落物层和土壤表层（0～2 cm）的种子数占土壤种子库总数的97.0%;不同季节花楸树土壤种子库种子数量差别很大,当年11月上旬种子数量最多,达(257.7±69.2)粒·m^-2;翌年7月下旬种子数量最少,仅为(2.9±2.9)粒·m^-2;温度不是花楸树种子萌发出土过程的限制因子,0 ℃～5 ℃时幼苗出苗率达(67.5±6.6)%,但对其出苗速率影响显著.土壤含水量为50%时,花楸树出苗率最高,达(74.7±4.2)%;含水量为60%时,幼苗死亡率最低,为(32.6±0.6)%.花楸树种子的散布格局和土壤种子库的时空分布格局影响种子的萌发出土过程,进而影响其种群的天然更新.     

Variation in dormancy and germination in three co-occurring perennial forest herbs

Mesic deciduous forest herbs often disperse seed with morphophysiological dormancy (MPD) that prevents germination during unfavorable periods for seedling survival. However, for seeds of some species with MPD, seasonal separation of root and shoot emergence and variation in dormancy levels can complicate interpretation of seedling emergence timing in the field. We tested whether dormancy-break and germination requirements differed among co-occurring perennial forest herbs, Actaea racemosa, Hydrastis canadensis, and Sanguinaria canadensis, which are wild-harvested for their medicinal properties and known to have MPD. Seeds of all species exhibited a summer → autumn → winter requirement for seedling emergence in spring. However, species differed in seed-bank persistence due to variation in primary dormancy levels and stratification requirement of seeds. A. racemosa and H. canadensis can form short-term persistent seed bank, whereas S. canadensis can form a long-term persistent seed-bank, regardless of whether elaiosomes were removed from seeds prior to burial. A. racemosa seeds are dispersed in autumn with weak physiological dormancy, as seeds germinated to high rates at 15/6°C after 8 weeks. In contrast, most seeds of the summer dispersed species, H. canadensis and S. canadensis, require summer temperatures to overcome physiological dormancy. Consequently, seedling emergence is reduced and delayed by 1 year if seeds are not sown immediately following the period of natural dispersal. Seedling emergence was much lower in the field than in controlled conditions for all species, especially in the small-seeded A. racemosa. Interspecific variation in dormancy levels and germination traits must be considered when establishing populations for conservation purposes and in understanding recruitment limitation in perennial forest herbs.     

<Emphasis Type="Italic">Pinus nigra</Emphasis> Arn. ssp <Emphasis Type="Italic">salzmannii</Emphasis> early recruitment and initial seedling growth in warmer and drier locations: the role of seed and soil provenance

Current decreasing precipitation and increasing temperatures promote the likelihood of extreme drought events and may alter the recruitment capacities of tree species. Spanish black pine (Pinus nigra ssp. salzmannii) initial recruitment is being one of the most affected pine species by changing conditions with alterations in the future species distribution. In this context, a cross-exchange experiment was implemented using an outdoor nursery located in a warmer and drier location for testing different Spanish Black pine seeds and soil provenance combinations in relation to early recruitment and initial seedling growth. Soil and seeds were collected at a high (HA, 1641 m.a.s.l.) and low (LA, 1099 m.a.s.l.) altitude in Cuenca Mountains (Spain). Then, a cross-sown experiment using HA and LA soils and seeds was set up in an outdoor nursery, which is located in Albacete (704 m.a.s.l.). Soil quality, seedling emergence, seedling survival, initial seedling growth and total seedling dry mass were measured after one year. We found higher seed emergence and seedling survival by combining LA soil with LA seeds or HA soil with HA seeds. Seedlings from LA seeds with both soil origins and seedlings from HA seeds with LA soils allocated more biomass to roots than seedlings from HA sites growing in HA soils under drier and warmer conditions. These results support the idea that autochthonous provenances have the potential to adapt to changing climatic conditions in their habitats.     

Seedlings of the introduced invasive shrub Eugenia uniflora (Myrtaceae) outperform those of its native and introduced non-invasive congeners in Florida

One particularly compelling explanation for the success of invasive species is the ability to outperform other species in characteristics affecting fitness. Past studies have compared native or introduced non-invasive species to their invasive counterparts, while a system incorporating both native and introduced non-invasive congeners provides an opportunity for multiple controls. We used such a system of Eugenia congeners in Florida to compare seedling performance. In order to determine if invasive Eugenia uniflora seedlings outperform those of its congeners, we sowed seeds in the field and a common garden and quantified seedling emergence, growth, and survival, as well as foliar damage by insect herbivores. We obtained similar results in the field and garden experiments. Although there were no differences in seedling emergence for E. uniflora seedlings when compared to some of its introduced congeners in certain years, emergence of E. uniflora seedlings was consistently high across years. However, emergence, growth, and survival rates of native species were consistently low. In addition, E. uniflora outperformed its introduced and native congeners in growth and survival traits in most comparisons, even when sustaining higher levels of herbivore damage by an introduced weevil, Myllocerus undatus Marshall. Our results support our predictions, indicating that invasive E. uniflora may possess a competitive advantage because its seedlings perform better than or equivalently to its congeners in all of the attributes quantified. Our study further suggests that measurements of such traits may be useful in determining the likelihood of invasion by newly introduced woody plant species.     

The Effects of Seed Ingestion by Livestock,Dung Fertilization,Trampling, Grass Competition and Fire on Seedling Establishment of Two Woody Plant Species

The increasing rate of woody plant encroachment in grasslands or savannas remains a challenge to livestock farmers. The causes and control measures of woody plant encroachment are of common interest, especially where it negatively affects the objectives of an agricultural enterprise. The objectives of this study were to determine the effects of gut passage (goats, cattle), dung (nutrients), fire, grass competition and trampling on establishment of A. nilotica and D. cinerea seedlings. Germination trials were subjected to the following treatments: 1) seed passage through the gut of cattle and goats and unpassed/ untreated seeds (i.e. not ingested), 2) dung and control (no dung), 3) grass and control (mowed grass), 4) fire and control (no fire), 5) trampling and control (no trampling). The interaction of animal species, grass and fire had an effect on seedling recruitment (P < 0.0052). Seeds retrieved from goats and planted with no grass and with fire (6.81% ± 0.33) had a significant effect on seedling recruitment than seeds retrieved from goats and planted with grass and no fire (2.98% ± 0.33). Significantly more D. cinerea and A. nilotica seeds germinated following seed ingestion by goats (3.59% ± 0.16) than cattle (1.93% ± 0.09) and control or untreated seeds (1.69% ± 0.11). Less dense grass cover, which resulted in reduced grass competition with tree seedlings for light, space and water, and improved seed scarification due to gut passage were vital for emergence and recruitment of Acacia seedlings. These results will contribute considerably to the understanding of the recruitment phase of woody plant encroachment.     

Linking seed dispersal,germination and seedling recruitment in the invasive species <Emphasis Type="Italic">Berberis darwinii</Emphasis> (Darwin’s barberry)

Seedling recruitment is a multi-phased process involving seed production, dispersal, germination, seedling establishment and subsequent survival. Understanding the factors that determine success at each stage of this process is of particular interest to scientists and managers seeking to understand how invasive species spread and persist, and identify critical stages for management. To understand the factors and processes influencing recruitment of the invasive species Berberis darwinii Hook. (Darwin’s barberry), temporal and spatial patterns of seed dispersal, germination and seedling establishment were examined. Seed dispersal from a large source population was measured over two fruiting seasons, and subsequent patterns of seedling emergence and survival within each cohort were measured. Seed longevity was tested under both natural and artificial conditions. Seeds were widely dispersed by birds, up to 450 m from the source population. Dispersal was essential to seedling establishment, as few seedlings survived beneath the parent canopy. Seeds were relatively short-lived in the soil under both field and glasshouse conditions, with few surviving for more than 1 year. Patterns of newly emerged seedlings largely reflected patterns of seed rain, but seedling survival was significantly affected by distance from source population, seedling density and light environment. These results suggest that recruitment of B. darwinii is dependent on dispersal of seeds to favourable microsites. Management priorities should include the removal of fruiting plants, and seedling control in highlight areas.     

Experimental Seed Predator Removal Reveals Shifting Importance of Predation and Dispersal Limitation in Early Life History Stages of Tropical Forest Trees

Recruitment limitation of trees in tropical forests can occur because of high rates of seed predation or low rates of seed dispersal, but the degree to which limitation is influenced by variation in seed predator abundance, and hence variation in seed predation and dispersal, is not well understood. We experimentally reduced the density of a granivorous small mammal (Heteromys desmarestianus) by 90 % to assess the degree to which its rates of seed predation and dispersal limit seed to seedling survival of nine species of trees in a Neotropical lowland forest. Overall, the proportion of seeds that germinated was influenced more by high rates of predation than by limited dispersal. Reduction in density of H. desmarestianus resulted in an order of magnitude decrease in fruit removal rates and an order of magnitude increase in both the absolute and relative numbers of seeds that germinated. However, the proportion of seeds that were cached remained relatively constant across all periods and between control grids and removal plots. In removal plots, H. desmarestianus dispersed and cached about 10 % of the fruits they handled, of which approximately 25 % germinated. This was 2 to 3 times greater than the germination rates of undispersed seeds, and for two species dispersed seeds were the only ones that germinated. The results suggest the simultaneous occurrence of both seed predation and dispersal limitation for trees with animal-dispersed seeds, but there may also be a hierarchy of importance in the relative strength of these two mechanisms that is determined by the dynamics of seed predator populations.     

Seedling emergence responds to both seed source and recruitment site climates: a climate change experiment combining transplant and gradient approaches

Seedling recruitment allows genetic recombination and production of dispersal units. Both the climate experienced by the source populations (seed source effect) and the weather experienced by the seeds during germination and seedling emergence (recruitment site effects) are important for seedling recruitment. Separating these effects in the field is essential to assess potential climate change impacts on plant population. We combine experimental seed transplant and gradient analyses to separate the effects of seed source and recruitment site temperature and precipitation for the seedling emergence of two alpine/lowland species pairs (Viola biflora/Viola palustris, Veronica alpina/Veronica officinalis). Combining these approaches allows us to compare local responses versus responses along environmental gradients, but also tests for local adaptation and/or pre-conditioning effects (adaptive seedling emergence responses). Veronica officinalis emergence increased with increasing seed source temperature in both the experimental and the gradient approaches, and showed adaptive seedling emergence. Viola biflora, Viola palustris and Veronica alpina emergence decreased with recruitment site temperature in both approaches. Both Violas emergences increased with recruitment site precipitation, in both approaches for the alpine violet, and in the gradient approach for lowland one. Emergence was primarily affected by the environment of the recruitment site, whereas seed source climate and adaptive seedling emergence impacted recruitment in only one of our species. The responses to recruitment site temperatures were negative, whereas the response to seed source temperature was positive. Ignoring the distinctions between these different mechanisms can lead to erroneous conclusions regarding potential climate change impacts on plant recruitment.     

Influence of elaiosome on postdispersal dynamics of an ant-dispersed plant

Specialized myrmecochorous plants bear an elaiosome attached to the seed. Ants benefit from food provided by this appendage and plants get their seeds eventually dispersed. Aside from the benefits conferred by seed dispersal, we hypothesize that the influence of the elaiosome might extend to later life history stages such as emergence or survival, but few studies have examined rigorously such delayed effects on seed and seedling dynamics. This paper tests this hypothesis for the ant-dispersed herb Helleborus foetidus. Presence or lack of the elaiosome had some interesting delayed influences. Specifically, our results show that elaiosome removal not only enhanced seedling emergence, but also reduced cumulative long-term seed predation. The combination of these two effects, together with the inherent positive effect of the elaiosome on seed dispersal, highlights the ecological and evolutionary importance of myrmecochorous systems.     

Effects of invasive rats and burrowing seabirds on seeds and seedlings on New Zealand islands

Rats (Rattus rattus, Rattus norvegicus, Rattus exulans) are important invaders on islands. They alter vegetation indirectly by preying on burrowing seabirds. These seabirds affect vegetation through nutrient inputs from sea to land and physical disturbance through trampling and burrowing. Rats also directly affect vegetation though consumption of seeds and seedlings. Seedling communities on northern New Zealand islands differ in composition and densities among islands which have never been invaded by rats, are currently invaded by rats, or from which rats have been eradicated. We conducted experimental investigations to determine the mechanisms driving these patterns. When the physical disturbance of seabirds was removed, in soils collected from islands and inside exclosures, seedling densities increased with seabird burrow density. For example, seedling densities inside exclosures were 10 times greater than those outside. Thus the negative effects of seabirds on seedlings, by trampling and uprooting, overwhelm the potentially beneficial effects of high levels of seed germination, seedling emergence, and possibly seed production, which result from seed burial and nutrient additions. Potential seedling density was reduced on an island where rats were present, germination of seeds from soils of this island was approximately half that found on other islands, but on this island seedling density inside exclosures was 7 times the density outside. Although the total negative effects of seabirds and rats on seedling densities are similar (reduced seedling density), the differences in mechanisms and life stages affected result in very different filters on the plant community.     

Effective nut dispersal by magpies (<Emphasis Type="Italic">Pica pica</Emphasis> L.) in a Mediterranean agroecosystem

Scatter-hoarding animals such as corvids play a crucial role in the dispersal of nut-producing tree species. This interaction is well known for some corvids, but remains elusive for other species such as the magpie (Pica pica), an abundant corvid in agroecosystems and open landscapes of the Palearctic region. In addition, the establishment of the individual dispersed seeds—a prerequisite for determining seed-dispersal effectiveness—has never before been documented for the interaction between corvids and nut-producing trees. We analyzed walnut dispersal by magpies in an agroecosystem in southern Spain. We used several complementary approaches, including video recording nut removal from feeders, measuring dispersal distance using radio tracking (with radio transmitters placed inside nuts), and monitoring the fate of dispersed nuts to the time of seedling emergence. Magpies were shown to be highly active nut dispersers. The dispersal distance averaged 39.6 ± 4.5 m and ranged from 4.1 to 158.5 m. Some 90% of the removed walnuts were cached later, and most of these (98%) were buried in the soil or hidden under plant material. By the time of seedling emergence, ca. 33% of nuts remained at the caching location. Finally, 12% of the cached nuts germinated and 4% yielded an emerged seedling, facilitating the transition to the next regeneration stage. The results demonstrate for the first time that magpies can be an effective scatter-hoarding disperser of a nut-producing tree species, suggesting that this bird species may play a key role in the regeneration and expansion of broadleaf forests in Eurasia.