Aggregated seed dispersal by spider monkeys limits recruitment to clumped patterns in Virola calophylla

The initial spatial pattern of seed deposition influences plant population and community structure, particularly when that pattern persists through recruitment. In a vertebrate‐dispersed rain forest tree, Virola calophylla, we found that spatially aggregated seed deposition strongly influenced the spatial structure of later stages. Seed dispersion was clumped, and seed densities were highest underneath V. calophylla females and the sleeping sites of spider monkeys (Ateles paniscus), the key dispersal agent. Although these site types had the lowest per capita seed‐to‐seedling survival, they had the highest seedling/sapling densities. Conversely, seed and seedling/sapling densities were lowest, and seed survival was highest, at sites of diurnal seed dispersal by spider monkeys. Negative density‐dependent and positive distance‐dependent seed survival thinned seed clumps. Nonetheless, the clumped dispersion at sleeping and parental sites persisted to the seedling/sapling stage because differences in seed deposition were large enough to offset differences in seed survival among these site types.     

Does invasion by an alien plant species affect the soil seed bank?

Questions: How does invasion affect old‐field seed bank species richness, composition and density? How consistent are these effects across sites? Does the soil seed bank match vegetation structure in old‐fields? Location: Menorca, Balearic Islands, Spain, western Mediterranean basin. Methods: We monitored seed germination in soils from old‐fields that were both uninvaded and invaded (legacy effect) by the annual geophyte Oxalis pes‐caprae. We also added O. pes‐caprae bulbs to uninvaded soils to test O. pes‐caprae interference with seedling emergence (competitive effect). We compared species composition in the seed bank with that of the vegetation. Results: Species richness in the seed bank and in the vegetation was not significantly different between invaded and uninvaded areas. Uninvaded areas did not have larger seed banks than invaded areas. More seedlings, especially of geophytes, emerged when O. pes‐caprae bulbs were added to the soil. Species similarity between invaded and uninvaded areas was higher in the seed bank (74%) than in the vegetation (49%). Differences in species composition were as important as differences among sites. The degree of species similarity between the seed bank and the vegetation was very low (17%). Conclusions: Despite invasion by O. pes‐caprae not affecting species richness, the variation in the seed bank species composition in invaded and uninvaded areas, and the differences between the seed bank and the mature vegetation, highlights that even if the invader could be eradicated the vegetation could not be restored back to the exact composition as found in uninvaded areas.     

Population ecology of wild sunflowers: effects of seed density and post-dispersal vertebrate seed predators

Assessing the effects of seed density on the population dynamics of wild plant species with crop relatives will be vital in determining the potential effects of introducing traits into wild populations as a result of crop-to-wild gene flow. We examined experimental sunflower (Helianthus annuus) patches in eastern Kansas to determine the effects of seed density and predation on seedling recruitment and seed production in the next generation. High seed density treatment plots had significantly more seedlings and adult plants than did low seed density treatment plots. Overwinter vertebrate seed predator exclusion treatments resulted in increases in plant density compared to plots in which vertebrates were not excluded. Control patches (no seeds added) contained virtually no plants. Head production and estimated total seed production for a patch were not statistically different among treatments (excluding control plots). Although initial seed density and vertebrate post-dispersal seed predation do appear to have effects on seedling recruitment, neither appear to be limiting seed production of competing adult plants. Therefore, variation in seed densities (over the range examined) may have limited effects on local population dynamics. It is important to note that the choice of seed densities may affect the results obtained: the seed densities used in this study may, in retrospect, be higher than in the small roadside populations typical in eastern Kansas, yet other natural sites have much larger densities. Further, the effects of increased seed density at a local site may have other important effects such as altering metapopulation dynamics through increased long-distance dispersal or increased local seed bank size.     

Composition and diversity of herbaceous patches in woody vegetation: The effects of grazing,soil seed bank,patch spatial properties and scale

Herbaceous plants contribute much to plant diversity in Mediterranean-type ecosystems though mostly occupying relatively small patches within the dense woody vegetation. While studying species diversity in the herbaceous patches, we hypothesized that grazing, soil seed bank, and spatial properties of the patch affect plant diversity and composition at different spatial scales. The study site was in an LTER site located in the Mediterranean region in north Israel. We determined herbaceous species composition in: (1) randomly sampled quadrats in herbaceous patches in grazed and un-grazed plots; (2) soil seed bank samples taken from the same patches and germinated under optimal greenhouse conditions; (3) quadrats in the same patches sown with a homogenous mixture of local soil samples. Using GIS methods, we determined small-scale spatial characteristics of the herbaceous patches. Alpha and beta diversities were calculated at the patch and plot scales using Shannon's entropy H. Grazing increased alpha diversity of local untreated seed bank samples but decreased alpha diversity of the artificial homogenous soil seed bank mixture at both patch and plot scales. Positive relation between alpha diversity and patch area was detected only under grazing. Grazing increased beta diversity in all three treatments at the patch scale. Grazing decreased the similarity in species composition between above-ground vegetation and soil seed bank. The results indicate that moderate cattle-grazing affects species diversity in the herbaceous patches within the dense maquis. This effect is scale-dependent, and interacts with the effects of soil seed bank and patch spatial-properties: without grazing soil seed bank plays a more important role than patch spatial properties, but under grazing the size and the accessibility of the patch are more important in the determination of herbaceous species composition.     

Spatial patterns and associations in a Quercus‐Betula forest in northern China

Question: Are species‐specific regeneration strategies and competition the dominant processes facilitating species coexistence in a Quercus liaotungensis dominated temperate deciduous forest? Location: Dongling Mountains, North China, 1300 m a.s.l. Methods: Ripley's K‐function was used to characterize the spatial patterns and spatial associations of two dominant tree species, Quercus liaotungensis and Betula dahurica, and a common subcanopy species, Acer mono, at different growth stages (adult, sapling, seedling). Results: Seedlings, saplings and adults of all three species exhibited clumped distributions at most spatial scales. Quercus seedlings and saplings were positively associated with conspecific adult trees and spatially independent of dead trees suggesting that seed dispersal and vegetative regeneration influenced the spatial patterning of Quercus trees. Betula seedlings and saplings were positively associated with both live and dead trees of conspecific adults at small scales (<5 m) but negatively associated with live and dead trees of other species indicating sprouting as an important mechanism of reproduction. Saplings of Acer had a strong spatial dependence on the distribution of conspecific adult trees indicating its limited seed dispersal range. Negative associations between adult trees of Betula and Quercus demonstrated interspecific competition at local scales (<5 m). Conclusions: Different regeneration strategies among the three species play an important role in regulating their spatial distribution patterns, while competition between individuals of Betula and Quercus at the adult stage also contributes to spatial patterning of these communities. The recruitment limitations of Betula and Quercus may affect the persistence of these species and the long‐term dynamics of the forest.     

Assessing the impact of plant invasions on soil seed bank communities: use of univariate and multivariate statistical approaches

Questions: Are soil seed banks affected by invasions of alien plants? How can we rigorously assess alterations in seed bank communities associated with invasive species and account for the high spatial variability of seed bank data? How do multivariate approaches compare with more traditional approaches based on analysis of variance? Location: Three riparian sites, Ireland. Methods: A protocol based on a combination of multivariate techniques was used to characterize soil seed bank communities associated with the herbaceous invasive species Heracleum mantegazzianum in May and October. Permutational multivariate analysis of variance (PERMANOVA) was used to test the effects of the factors “invasion”, “site”, “plot” and “depth” on the soil seed bank, while multivariate analysis of dispersion (PERMDISP) provided a measure of the variability of seed bank data at different spatial scales. Similarity percentages analysis (SIMPER) was used to identify the species that contributed most to the differences between invaded and uninvaded communities. A comparison between the results of PERMANOVA and ANOVA analyses was also made. Results: The composition of seed bank communities invaded by H. mantegazzianum differed significantly from that of uninvaded seed banks. Invaded seed banks were less diverse and had reduced abundance, and were dominated by only a few species, such as Urtica dioica and Juncus effusus. Such patterns were recorded at each of three depth categories, indicating that invasive plants can affect both the transient and the more persistent component of the soil seed bank. Seed bank variability was significantly higher within uninvaded areas, supporting the notion that invasions tend to lead to more homogeneous communities. Conclusion: The analytical protocol used in this study was effective in quantifying the effect of plant invasions, at different spatial scales, providing a statistically robust analysis of alterations in soil seed bank communities. Compared to ANOVA, this protocol provided more biological information and was more appropriate for analysis of the data. This approach is therefore recommended in soil seed bank and invasion ecological studies.     

The potential role of seed banks in the recovery of dune ecosystems after removal of invasive plant species

Question: How resilient is the seed bank of an invaded dune system? Is that resilience dependent on duration of invasion? How does the accumulated litter layer contribute to the soil seed bank? Location: Coastal sand dunes invaded by Acacia longifolia, Portugal. Methods: Seedling emergence was used to quantify and compare soil seed banks in long‐invaded, recently invaded and non‐invaded areas. Changes in seed banks were also compared with areas where A. longifolia and the litter layer were removed. Results: Species richness, seedling density and diversity were higher in non‐invaded and recently‐invaded areas than in long‐invaded areas. Although there was an apparent similarity between non‐invaded and recently‐invaded areas, analyses of species traits revealed differences. Non‐invaded areas had a wider array of traits. Exotic/invasive species dominated invaded seed banks while native species dominated non‐invaded seed banks. Life forms, growth forms, longevity and dispersal mode showed differences between areas, with cleared plots of long‐invaded areas being apparently the most similar to non‐invaded plots. Acacia longifolia seeds were most abundant in long‐invaded areas, particularly where the litter layer remained. Removal of A. longifolia plus the litter had little effect on the seed bank composition of recently‐invaded areas but resulted in noticeable changes in seed banks of long‐invaded areas. Conclusions: Long‐invaded areas are less resilient and show a higher reinvasion potential, despite severe alteration of the seed banks of both areas. Seed bank studies can be a useful tool to guide management, but can give misleading results when invasion periods are protracted.     

Plant species richness and composition of heathland relics in north‐western Belgium: evidence for a rescue‐effect?

Aim To study the effects of the degree of fragmentation of heathland patches on their species richness and species composition, and to infer the ecological mechanisms behind the observed patterns. Location The heathland patches of the north‐western part of Flanders, Belgium. During the last 200 years, the heathland area in this study area has been reduced from c. 10,000 to c. 40 ha, yielding c. 150 small and highly isolated relic fragments. Methods Different isolation measures were calculated for each of the 153 inventoried heathland patches. The influence of isolation, area and habitat diversity on species richness was investigated using correlation techniques. Community composition of the patches was tested for nestedness, and the mechanisms potentially underlying this pattern were determined. Results and main conclusions Both the analyses at the species richness and community composition level yielded evidence for a positive species–area relation. This relation was not caused by higher habitat heterogeneity in larger patches. Patch isolation, measured in different ways, however seemed much more important in explaining species richness and community composition than fragment area. Our results indicate that area effects are overcompensated by the rescue‐effect: if a patch is close to other patches, species can disperse between them and prevent the species from going extinct. Species having a short living seed bank were also more sensitive to isolation than species with a long living seed bank, indicating that the latter most probably depend on the seed bank to survive periods when environmental conditions are harsh. Analogously to the spatial rescue‐effect, the existence of a persistent seed bank may lead to a sort of temporal rescue‐effect, where the extinction of a plant species is prevented through survival in the seed bank of a patch.     

Seed bank spatial pattern in a temperate secondary forest

Abstract. Seed bank spatial pattern was studied in a secondary forest dominated by Fagus sylvatica and Betula celtiberica in the Urkiola Natural Park (N Spain). Soil samples were taken every 2 m in a regular grid (196 points) and divided into two fractions (0–3 cm and 3–10 cm deep). The viable seed bank was studied by monitoring seedling emergence for ten months. The effect of different factors on seed bank composition and patterning was analysed using constrained ordination as a hypothesis testing tool. Furthermore, the existence of spatial autocorrelation was evaluated by geostatistical analysis. Seed density was high, 7057 seed.m^?2, with a few species dominating. Species composition in the various layers were significantly correlated. The seed bank showed significant spatial structure, which was partially explainable by the spatial structure of the canopy and understorey vegetation. Spatial clumping from 0–8 m was observed in seed bank density and composition, mainly due to the pattern of two abundant taxa Juncus effusus and Ericaceae. The Ericaceae seed bank was related to the spatial distribution of dead stumps of Erica arborea. J. effusus was not present in the above‐ground vegetation, which indicates that its seed bank was formed in the past. As expected, the seed bank of this forest reflects its history, which is characterized by complex man‐induced perturbations. The seed bank appears to be structured as a consequence of contrasting driving forces such as canopy structure, understorey composition and structural and microhabitat features.     

Seed bank spatial structure in semi-arid environments: beyond the patch-bare area dichotomy

The prevalence of patchy structures in vegetation is a common feature in semi-arid ecosystems. Although the effect of patches on seed density is widely known, we still lack information on how patch features affect seed bank density and composition. Our aim was to answer two basic questions: (1) How do seed bank density and composition vary within and outside patch aboveground physical limits? and (2) Do patch characteristics affect soil seed bank density and composition? We sampled 50 shrub patches in a semi-arid gypsum system in Central Spain, measuring patch size, composition and structure, and seed bank at three locations per shrub (centre, edge and outside). We calculated the effect of interior patch location, patch composition and structure on seed density and composition. Patches acted both as seed sources, increasing seed density in neighbouring areas and as seed sinks by trapping seeds from bare areas. Patch structure (erect perennial cover) had the greatest effect on seed bank density, whereas patch size and microslope had the greatest influence on bare area density. Patch structure, composition and interior location explained the variation in seed bank composition. Patch effect extends to the surrounding bare matrix creating a seed bank gradient in density and composition. This effect is modulated by patch structure and composition and affects seed bank composition. Our results suggest that the spatial structure of gypsum community seed banks may act as a mechanism for a spatial storage effect contributing to the maintenance of high levels of diversity in semi-arid environments     

Avian seed dispersal and seedling distribution of the endangered tree species,Taxus chinensis,in patchy habitats

Background: There is limited understanding about bird dispersal behaviour and seedling distribution of endangered tree species in patchy environments, although these processes are important for plant species persistence.

Aims: We tested how patch features affected bird behaviour and seed dispersal, and thus seedling distribution of the endangered Chinese yew tree (Taxus chinensis).

Methods: In the present study, we combined field data of bird dispersal behaviour and GIS-based information to elucidate the influence of spatial features of habitat patches on bird dispersal behaviour, and the resulting effects on the seedling distribution of the endangered Chinese yew in two patchy habitats.

Results: Our results showed that the only seed source patch could attract eight bird species for dispersal at the two sites. Post-foraging movements of bird dispersers was strongly related to both topography and the relative locations of habitat patches. Yew seedlings aggregated only at the seed source and bamboo recruitment patches, which was affect by both the spatial distribution of recruitment patches and patch use by dispersers.

Conclusions: Our results emphasise that bamboo patches in both patchy environments provide the necessary conditions for germination of yew seeds, and the post-foraging behaviour of dispersers determines seed deposited in these patches. Our study highlights the importance of the dispersal behaviour of frugivorous birds in the successful regeneration and colonisation of yew populations in patchy habitats.     

Belowground bud banks and meristem limitation in tallgrass prairieplant populations

Rhizome meristem populations were sampled in tallgrass prairie to quantify the size, grass?:?forb composition, and temporal and spatial variability of the soil bud bank and to compare fire effects on bud bank and seed bank composition. Soil cores (10.5 cm diameter, 15 cm deep) were collected from replicate annually and infrequently burned tallgrass prairie sites, and intact rhizomes and rhizome buds were censused. Bud bank densities ranged from approximately 600 to 1800 meristems/m(2) among sites and had high spatial and seasonal variability. In annually burned prairie, the total bud bank density was two-fold greater and the grass?:?forb meristem ratio was more than 30-fold greater than that of infrequently burned prairie. These patterns are opposite those observed in soil seed banks at this site. The rhizome population in annually burned prairie was 34% larger than the established aboveground tiller population. By contrast, the bud bank density in unburned prairie was significantly lower than aboveground stem densities, indicating possible belowground meristem limitation of stem density and net primary production on infrequently burned prairie. The patterns observed in this study suggest that the densities and dynamics of tallgrass prairie plant populations, as well as their response to disturbance (e.g., fire and grazing) and climatic variability, may be mediated principally through effects on the demography of belowground bud populations. Patterns of seed reproduction and seed bank populations have little influence on short-term aboveground population dynamics of tallgrass prairie perennials.     

Depth distribution and composition of seed‐banks in alien‐invaded and uninvaded fynbos vegetation

South African fynbos vegetation is threatened on a large scale by invasive woody plants. A major task facing nature conservation managers is to restore invaded areas. The aim of this study was to determine the restoration potential of fynbos following dense invasion by the Australian tree Acacia saligna. The impacts of dense invasion on seed‐bank composition and depth distribution were investigated to determine which fynbos guilds and species have the most persistent seed‐banks. Soil samples were excavated at three different depths for invaded and uninvaded vegetation at two sand plain and mountain fynbos sites. Seed‐banks were determined using the seedling emergence approach. Invasion caused a significant reduction in seed‐bank density and richness at all sites. There was a significant, but smaller, reduction in seed‐bank density and richness with soil depth at three sites. Seed‐bank composition and guild structure changed following invasion. Low persistence of long‐lived obligate seeders in sand plain fynbos seed‐banks indicates that this vegetation type will be difficult to restore from the seed‐bank alone following alien clearance. The dominance of short‐lived species, especially graminoids, forbs and ephemeral geophytes, suggests that regenerating vegetation will develop into a herbland rather than a shrubland. It is recommended that seed collecting and sowing form part of the restoration plan for densely invaded sand plain sites. As seed density remained higher towards the soil surface following invasion, there is no general advantage in applying a mechanical soil disturbance treatment. However, if the shallow soil seed‐bank becomes depleted, for example following a hot fire through dense alien slash, a soil disturbance treatment should be given to exhume the deeper viable seed‐bank and promote recruitment.     

Topsoil properties and seedling recruitment in Lavandula latifolia: stage-dependence and spatial decoupling of influential parameters

This paper investigates the influence of topsoil texture and chemistry on long‐term patterns of seedling recruitment in Lavandula latifolia (Labiatae), an evergreen Mediterranean shrub. The questions addressed are: (1) Are spatial patterns of seedling emergence and survival probabilities related to spatial variation in soil properties? (2) Are the two components of recruitment similarly influenced by the same soil parameters or, alternatively, are they influenced in different ways and/or by different soil parameters? (3) If the latter applies, then do the sets of soil parameters influencing emergence and survival show congruent patterns of spatial variation? Seed progenies from fourteen maternal families were sown at ten widely spaced field sites (between‐site distances in the scale of a few kilometers). Sites differed in average soil characteristics. There was also considerable variation in some soil parameters within sowing sites (distances between soil sampling points in the scale of meters). The seedling recruitment process was monitored for six years. Both the probability of one seed of L. latifolia giving rise to an emerged seedling, and the probability of an emerged seedling remaining alive six years past emergence and entering the juvenile age class, were significantly related to topsoil properties. Seedling emergence and survival tended to be influenced by largely non‐overlapping sets of soil properties that were related, respectively, to texture‐ and fertility‐related gradients. These two soil gradients tended to be uncorrelated at the scale of the study, and differed in the apportionment of variance among small (within sites) and large (between sites) spatial scales. While most variance in the emergence‐influencing, texture‐related gradient occurred at the between‐site scale, a considerably greater fraction of variance in the survival‐influencing, fertility‐related gradient occurred at the within‐site scale. It is suggested that, because of fine scale variability in soil fertility, local adaptations of L. latifolia to soil textural properties enhancing seedling emergence will be easier to evolve than adaptations to fertility‐related features enhancing seedling survival.     

Blue fan palm distribution and seed removal patterns in three desert oases of northern Baja California, Mexico

Information on the current processes controlling the distribution patterns of relict palm populations at the limit of their northwestern distribution in America are poorly known. We explored the importance of post-dispersal seed removal by vertebrates, recruitment, and distribution patterns of the blue fan palm, Brahea armata, in the Northern Baja California peninsula, by evaluating (i) the levels of blue fan palm seed removal by vertebrates at two spatial scales and the initial fate of dispersed seeds, (ii) the spatial distribution and association of seedlings and adults at two spatial scales, (iii) seed removal levels and seedling densities based on density and distance to adult palm trees, and (iv) the population age structure. Overall, seed removal levels were low at all sites, varied at regional but not at local scales, with small rodents apparently being responsible of most removals. Adult density and distribution differed among oases, but seedling densities did differ locally. Whereas seed removal did not vary with distance from seed sources or parent trees, a weak positive association between seedlings and adults at the whole patch level indicated that establishment tended to occur in or near those grid cells where adults had established successfully. However, the analysis showed a negative association between seedling and adult densities within the patches, indicating that within the grid cells where growth was most successful, seedlings established preferentially in relatively open spaces. All life-history categories were adequately represented within each site and populations seemed to be in a fairly good conservation state. We did not find evidence to establish that post-dispersal seed removal by rodents is a main factor defining the recruitment patterns in these oases. In contrast, the effect of flood pulses seems to be significant and may have strong and conclusive effects on palm seedling distributions. We suggest that local biotic (nurse plants) and/or abiotic (canyon physiography, nurse objects) factors at each particular canyon have the potential to affect post-dispersal seed removal activity patterns by rodents, as well as to provide vital protection for palm seedling establishment from the extreme floods. This baseline information will help in further investigations on the possible mechanisms that sustain palm populations at their distribution limits and in stressful environments.     

Role of the soil Seed Bank in the restoration of a native pinewood at Glen Garry,Inverness-shire

Summary

The potential recovery of ground vegetation in a pinewood at Glen Garry, from which introduced tree species had been removed, was studied by sampling the upper soil seed bank and comparing the species composition of germinating seedlings with extant vegetation. Mean numbers and species of seedlings emerging differed between a control site (not under-planted) and two formerly under-planted sites cleared at different times. Germinating seedling numbers also differed with depth (0–10 cm). Calluna vulgaris and Juncus spp. dominated fifteen species germinating from the seed bank, while some other pinewood species were not found. Although seedling pine occurred, their survival will be affected by competition from Betula spp. and Deschampsia flexuosa. Other elements of the vegetation will recover from the seed bank or by migration, their distribution being influenced by topographic variations and the nature of the developing tree stand. Timing of clearance of exotics in relation to pine seed production and soil scarification might accelerate recovery of the pinewood flora generally.     

Woody plant invasion at a semi‐arid/arid transition zone: importance of ecosystem type to colonization and patch expansion

Question: How do patterns in colonization and patch expansion of an invasive woody plant (Larrea tridentata, Zygo‐phyllaceae) differ between two grassland ecosystems at a biome transition zone? Location: Semi‐arid/arid transition zone in central New Mexico. Methods: Frequency of occurrence, height, and surface area of saplings (n= 134) and patches of adult plants (n= 247) of the invasive shrub, L. tridentata, were measured within a mosaic of ecosystems dominated either by the Chihuahuan Desert species, Bouteloua eriopoda (Poaceae), or the shortgrass steppe species, B. gracilis, located within 1 km of the L. tridentata‐dominated ecosystem. Distances between L. tridentata patches and patch area were used to estimate connectivity as a measure of propagule pressure. Sapling age (estimated from height using previously established relationships) and distance to the L. tridentata‐dominated ecosystem was used to evaluate patterns in dispersal. Cover by species or functional group inside each L. tridentata patch was compared with surrounding vegetation to estimate changes in species composition with patch expansion. Results: L. tridentata saplings (< 1%) and adult patches (15%) occurred less frequently in B. gracilis‐dominated ecosystems than expected based on areal extent of this ecosystem type. Propagule pressure did not differ with distance from the core ecosystem dominated by L. tridentata. Evidence for both local and long‐distance dispersal events was found. Similar relationships between number of plants and patch area in both grassland types indicate similar patterns in patch expansion. Cover of perennial forbs was higher and cover of dominant grasses was lower in L. tridentata patches compared with the surrounding vegetation for both ecosystem types. Conclusions Spatial variation in L. tridentata saplings and patches at this biome transition zone is related to the different susceptibilities to invasion by two grassland ecosystems. The persistence of grasslands at this site despite region‐wide expansion by L. tridentata may be related to the spatial distribution of B. gracilis‐dominated ecosystems that resist or deter invasion by this woody plant.     

The effects of plant dispersion and prey density on parasitism rates in a naturally patchy habitat

Despite extensive research on parasitoid-prey interactions and especially the effects of heterogeneity in parasitism on stability, sources of heterogeneity other than prey density have been little investigated. This research examines parasitism rates by three parasitoid species in relationship to prey density and habitat spatial pattern. The herbivore Itame andersoni (Geometridae) inhabits a subdivided habitat created by patches of its host plant, Dryas drummondii, in the Wrangell Mountains of Alaska. Dryas colonizes glacial moraines and spreads clonally to form distinct patches. Habitat subdivision occurs both on the patch scale and on the larger spatial scale of sites due to patchy successional patterns. Itame is attacked by three parasitoids: an ichneumonid wasp (Campoletis sp.), a braconid wasp (Aleiodes n. sp.), and the tachinid fly (Phyrxe pecosensis). I performed a large survey study at five distinct sites and censused Itame density and parasitism rates in 206 plant patches for 1–3 years. Parasitism rates varied with both plant patch size and isolation and also between sites, and the highest rates of overall parasitism were in the smallest patches. However, the effects of both small- and large-scale heterogeneity on parasitism differed for the three parasitoid species. There was weak evidence that Itame density was positively correlated with parasitism for the braconid and tachinid at the patch scale, but density effects differed for different patch sizes, patch isolations, and sites. At the site scale, there was no evidence of positive, but some indication of negative density-dependent parasitism. These patterns do not appear to be driven by negative interactions between the three parasitoid species, but reflect, rather, individual differences in habitat use and response to prey density. Finally, there was no evidence that parasitism strongly impacted the population dynamics of Itame. These results demonstrate the importance of considering habitat pattern when examining spatial heterogeneity of parasitism and the impacts of parasitoids. Received: 3 June 1999 / Accepted: 4 October 1999     

Moms are better nurses than dads: gender biased self‐facilitation in a dioecious Juniperus tree

Questions: Can gender of nurse plants affect regeneration patterns and spatial population structure? Is there a seed‐seedling conflict in the regeneration process? What factors are responsible for the clumped spatial population structure observed for adult trees? Location: Mediterranean cold semi‐arid high mountains in Spain. Methods: The spatial pattern of adult Juniperus thurifera trees was studied by means of Ripley's K‐analysis. χ² analyses were used to test for natural seedling frequency in each of three main microhabitats: (1) under female and (2) male tree canopies and (3) in open interspaces. The observed pattern was explained experimentally by studying seed and seedling survival for two years. Survival probabilities were calculated across life stages for each of three main microhabitats. Results: Adult J. thurifera trees were aggregated in space. Most seedlings were found underneath female J. thurifera trees. Experimental studies demonstrated that from seed dispersal to seedling survival all life stages showed the same positive or negative trend within a given microhabitat, indicating stage coupling and no seed‐seedling conflicts. Attraction of frugivo‐rous birds by reproductive female junipers and improvement of environmental conditions beneath tree canopies were the main factors responsible for the variation in seedling density among microhabitats; highest underneath female trees and lowest in open interspaces. Conclusions: In dioecious species, the gender of nurse plants can significantly determine the spatial population structure. In J. thurifera forests, facilitation beneath female trees occurs among all life stages without any sign of seed‐seedling conflict. The most critical factors shaping the spatial population structure were directed seed dispersal and environmental amelioration beneath female conspecific trees.     

The relationship between soil seed bank,above‐ground vegetation and disturbance intensity on old‐field successional permanent plots

Questions: How does disturbance and successional age influence richness, size and composition of the soil seed bank? What is the potential contribution of the soil seed bank to the plant community composition on sites differing in their successional age or disturbance intensity? Location: Experimental Botanical Garden of Göttingen University, central Germany. Methods: Above‐ground vegetation and soil seed bank were studied on formerly arable fields in a 36‐year‐old permanent plot study with five disturbance intensities, ranging from yearly ploughing via mowing to long‐term uninterrupted succession. We compared species compositions, seed densities and functional features of the seed bank and above‐ground vegetation by using several methods in parallel. Results: The seed bank was mainly composed of early successional species typical of strongly disturbed habitats. The difference between seed bank composition and above‐ground vegetation decreased with increasing disturbance intensity. The species of greatest quantitative importance in the seed bank was the non‐native forb Solidago canadensis. Conclusions: The ability of a plant community to regenerate from the soil seed bank dramatically decreases with increasing time since abandonment (successional age) and with decreasing disturbance intensity. The present study underlines that plant species typical of grasslands and woodlands are limited by dispersal capacity, owing to low capacity for accumulation of seeds in the soil and the fact that most species do not build up persistent seed banks. Rare and target species were almost absent from the seed bank and will, after local elimination, depend on reintroduction for continuation of their presence.