Population structure of the threatened perennial Serratula tinctoria in relation to vegetation and management

Question: How does vegetation and management affect the population stage structure of Serratula tinctoria, a species indicative of highly endangered and species‐rich flood‐plain meadows? Are different management regimes suitable to support viable populations of S. tinctoria? Location: Flood‐plain of the northern Upper Rhine, Germany. Methods: We analysed the population structure of S. tinctoria at 24 meadow sites which differed in vegetation and management. In this comparative study the investigated meadows were either (1) late mown in September; (2) early mown in June or (3) mown in June and then grazed by sheep. Results: The structure of the surrounding vegetation had a clear effect on the population structure of S. tinctoria. The percentage of bare soil, as well as the cover of bryophytes, were positively related, whereas the Ellenberg N‐value of the established vegetation was negatively related to the density and proportion of seedlings. Generally, we found only slight differences between the population structure of S. tinctoria in early and late mown meadows. Both management regimes had high densities of seedlings. In contrast, the meadow pastures supported significantly lower densities of seedlings and generative adults but relatively high densities of juveniles and vegetative adults, indicating a greater importance of clonal propagation under grazing pressure. Conclusions: Our results clearly showed that viable populations of S. tinctoria may occur in all of the studied management regimes. Although S. tinctoria was considered to be highly sensitive to early mowing, our findings suggest a facilitation of the species at more nutrient‐rich sites by mowing in June, which is also a benefit for the integration of management in farming systems.     

The effect of cattle and sheep grazing on salt-marsh vegetation at Skallingen,Denmark

The aggregated effect of cattle and sheep grazing on Puccinellion maritimae and other salt-marsh vegetation has been studied together with changes in species composition, the percentage cover of each species, total cover and the percentage of bare ground, six years after grazing had been prevented by construction of experimental exclosures. The results of these experiments are discussed in relation to the natural development of the vegetation that occurred in a permanent plot in the ungrazed part of the salt marsh. Six years without grazing caused a marked increase in total cover and a dramatic reduction in the amount of bare ground. The species composition of the Puccinellia maritima community did not change during these six years, but all species increased in cover. Whereas Artemisia maritima migrated into the Festuca rubra community and only two species Festuca and Halimione portulacoides, increased in cover, all other species showed reduced cover and Salicornia europaea disappeared from the plot after six years without grazing. During the same period of time, three species, Salicornia europaea, Suaeda maritima, and Glaux maritima, disappeared from the plot in the ungrazed marsh as a result of natural development. During thirty-five years the vegetation originally dominated by Puccinellia maritima and Salicornia europaea has changed into a community dominated by Halimione portulacoides, whereas the grazed salt marsh is still dominated by Puccinellia maritima and Salicornia europaea. These results are discussed in relation to the literature on the effect of cattle and sheep grazing on salt-marsh vegetation.     

Planting Seedlings in Tree Islands Versus Plantations as a Large‐Scale Tropical Forest Restoration Strategy

Planting tree seedlings in small patches (islands) has been proposed as a method to facilitate forest recovery that is less expensive than planting large areas and better simulates the nucleation process of recovery. We planted seedlings of four tree species at 12 formerly agricultural sites in southern Costa Rica in two designs: plantation (entire 50 × 50 m area planted) and island (six patches of three sizes). We monitored seedling survival, height, and canopy area over 3 years. To elucidate mechanisms influencing survival and growth, we measured soil and foliar nutrients, soil compaction, and photosynthesis. Survival of all species was similar in the two planting designs. Seedling height and canopy area were greater in plantations than islands at most sites, and more seedlings in islands decreased in height due to damage incurred during plot maintenance. Survival, height, and canopy area were both site‐ and species‐specific with the two N‐fixing species (Inga edulis and Erythrina poeppigiana) greater than the other species (Terminalia amazonia and Vochysia guatemalensis). Foliar N was higher in Terminalia and Vochysia in sites where Inga growth was greater. Soil nutrients, however, explained a small amount of the large differences in growth across sites. Leaf mass per area was higher in islands, and P use efficiency was higher in plantations. Our results show advantages (good seedling survival, cheaper) and disadvantages (more seedling damage, slightly lower growth) to the island planting design. Our study highlights the importance of replicating restoration strategies at several sites to make widespread management recommendations.     

Establishment and survival of Pinus brutia Ten. seedlings over the first growing season in abandoned fields in central Cyprus

Abstract

The present study was carried out in abandoned fields in central Cyprus. The main objective was to examine the impact of the regeneration environment on the establishment and survival of Pinus brutia seedlings. Sixty-four permanent plots of 16 m² were randomly established in two distinct sites. Four regeneration environments were recorded: (a) bare soil under the crown of a P. brutia tree, (b) soil under the canopy of a P. brutia tree and low shrubs, (c) bare soil in open areas, and (d) soil under the canopy of low shrubs in open areas. All P. brutia seedlings were classified in categories according to their regeneration environment. In all plots, the density of the P. brutia seedlings was measured in three different seasons (spring, summer, autumn). Soil temperatures were recorded, samples of surface soil were taken and the percentage of soil organic matter was measured. The main conclusions drawn from this research were the following: (1) the mature P. brutia trees and low shrubs facilitate the establishment and especially the survival of P. brutia seedlings, as all seedlings in bare vegetation ground had died by the end of the growing season, and (2) the importance of facilitation increases as abiotic stress rises.     

The distribution and ecology of coastal species on roadsides

The invasion of roadsides by coastal species is a response to the use of de-icing salt. The most widespread species on Britain's roadsides, Puccinellia distans, occurs in the north and east of England and Wales. This distribution is due to the varying use and effects of de-icing salt which are related in turn to regional differences in climate and traffic density. On roadsides, seed dispersal is greater in the direction of traffic flow and the smaller seeds of Puccinellia distans are carried further than the larger seeds of Plantago maritima. Species on roadsides occur in distinct zones with the coastal species confined to the saline road margin. The causes of this zonation were investigated by examining seedling establishment in experimental plots. Seedlings were only able to establish in grass plots when the salinity was sufficiently high to kill the competing glycophytic grasses. On bare soil, the halophytes established and grew satisfactorily when salt was added but grew very poorly in the absence of salt. After two years Plantago maritima overcame this apparent requirement for salt on bare soil but Puccinellia distans and Spergularia marina did not.Acknowledgements: N. E. Scott was in receipt of a N.E.R.C. Studentship. The authors wish to thank the members of the B.S.B.I. and the Biological Records Office for information on species distribution.     

Factors affecting the population dynamics of Suaeda maritima at initial stages of development

This study describes life history and population dynamics of the annual halophyte Suaeda maritima, at initial stages of seedling development, in salt-marshes of the Mont-Saint-Michel Bay. Effects of salinity on seed germination and seedling development were studied both in the laboratory and in the field. Salinity was measured as sediment electrical conductivity. Experiments in the laboratory showed that Suaeda maritima was relatively tolerant to salinity during the germination stage. In the field, germination occurred when soil salinity was minimal. Almost all seeds germinated in the field; this may explain the absence of a persistent seed bank. Salinity did not seem to affect the growth of seedlings either. Mortality in the field was attributed to physical factors such as anoxia during temporary immersion, burial by sediment and tidal uprooting. The intensity of these effects varied according to the location of Suaeda maritima seedlings: in hollows, along the edge of tidal creeks, in gaps among dense vegetation cover, or in pattern with Puccinellia maritima. Perennial vegetation did not restrict Suaeda maritima germination. In contrast, perennials such as Puccinellia maritima and Halimione portulacoides limited the development of Suaeda maritima seedlings since only a small number of this annual emerged beneath their canopies. The fate of seedlings depended to a great extent on the permeability of the perennial canopy to light. Thus, disturbance resulting in bare patches inside the perennial vegetation seemed essential for the development of this annual species. Suaeda maritima can grow in the presence of Puccinellia maritima so long as the latter present as an open matrix. Suaeda maritima may also benefit from protection against desiccation and tidal action where this occurs.     

Regeneration of Salix arbuscula and Salix lapponum within a Large Mammal Exclosure: The Impacts of Microsite and Herbivory

Over‐grazing or browsing by large herbivores may result in the loss of individual plant species or entire plant communities. Restoration schemes often involve exclusion of large mammals, but the resulting changes in vegetation may alter other important ecological processes such as regeneration, via changes in microsite availability for seed germination or increases in populations of seedling predators. Working within a large fenced area from which large mammals were excluded, we experimentally tested the effects of microsite, small herbivores, and their interactions on post‐dispersal seed and early seedling mortality of one nationally scarce (Salix arbuscula) and one nationally rare (S. lapponum) species of montane willow. Seeds were sown in three different microsites: natural vegetation, mown vegetation (mimicking grazed sward), and bare ground. Small exclosures and slug pellets were used to examine the effects of small mammal and slug predation, respectively. Survival of seedlings was monitored during the summer following planting. The presence of bare ground, rather than the absence of herbivores, was of over‐riding importance for early seedling survival and establishment. Protecting seedlings from small mammals made no difference to the levels of survival; however, protecting seedlings from slugs (Arion spp.) resulted in approximately 45% of seedlings surviving until the end of the summer compared to only 30% when seedlings were available to slugs. Although excluding large herbivores may increase seed production of existing individuals, the impacts of changes to plant communities on processes such as regeneration need to be considered if restoration projects are to be fully successful.     

Utilisation of Wadden Sea salt marshes by geese in relation to livestock grazing

To arctic breeding geese, the salt marshes of the International Wadden Sea are important spring staging areas. Many of these marshes have always been grazed with livestock (mainly cattle and sheep). To evaluate the influence of livestock grazing on composition and structure of salt-marsh communities and its consequences for habitat use by geese, a total of 17 pairs of grazed and ungrazed marshes were visited both in April and May 1999, and the accumulated grazing pressure by geese was estimated using dropping counts. Observed grazing pressure was related to management status and to relevant vegetation parameters.The intensity of livestock grazing influences the vegetation on the marsh. Salt marshes that are not grazed by livestock are characterised by stands with a taller canopy, a lower cover of grasses preferred by geese, and a higher cover of plants that are not preferred.Overall goose-dropping densities are significantly lower in ungrazed marshes compared to marshes grazed by livestock. Some ungrazed marshes had comparatively high goose grazing pressure, and these were all natural marshes on a sandy soil, or artificial mainland marshes with a recent history of intensive livestock grazing. Goose grazing is associated with a short canopy. The plant communities with short canopy, dominated by Agrostis stolonifera, Festuca rubra and Puccinellia maritima, together account for 85% of all goose droppings in our data.The sites that were not visited by geese differed very little from those that were visited, in the parameters we measured. This might indicate that there was no shortage of available habitat for spring staging geese in the Wadden Sea, in the study period.     

Tree seedling development in tropical dry abandoned pasture and secondary forest in Costa Rica

Development of planted seedlings of four canopy tree species in recently abandoned pastures (mown and unmown) and in ca. 40-yr old secondary dry forest in Guana-caste National Park, Costa Rica, was studied from July 1989–June 1992. The species were the light-demanding Cedrela odorata and Swietenia macrophylla, and the shade-tolerant Hymenaea courbaril and Manilkara chicle. Seedling mortality was high and primarily correlated with dry season drought. After the first dry season, M. chicle and C. odorata showed < 5 % survival in the pasture and 20 % in the forest, but after 3 yr survival had decreased to 3% in all plots. Survival of S. macrophylla was highest in the mown pasture (45 % after 3 yr) and lowest in the forest (10 %). For H. courbaril, survival was 40–55 % under all three conditions. Growth rates were low, with a height increment of < 15 cm/yr, possibly due to low precipitation in 1990 and 1991. All species grew taller in the pasture than in the forest. Differences in soil depth, texture and drainage appeared to contribute to variation in the results both within and between plots. Herbivory contributed to the lower survival of C. odorata and S. macrophylla in the forest.     

Population size and habitat quality affect genetic diversity and fitness in the clonal herb <Emphasis Type="Italic">Cirsium dissectum</Emphasis>

Remaining populations of plant species in fragmented landscapes are threatened by declining habitat quality and reduced genetic diversity, but the interactions of these major factors are rarely studied together for species conservation. In this study, the interactions between population size, habitat quality, genetic diversity and fitness were investigated in 22 populations of the clonal herb Cirsium dissectum throughout the British Isles. Regression analysis was used to identify significant factors, and a structural equation model was developed to illustrate and integrate these interactions. It was found that smaller populations (measured as the total number of plants) had lower genetic diversity (proportion of polymorphic loci), and that reduced genetic diversity (allelic richness) had a negative impact on the survival of seedlings grown under standard conditions. Habitat quality also had a large effect on C. dissectum. Unmanaged sites with tall vegetation, no bare soil and higher nutrient levels had smaller populations of C. dissectum, but flowering was promoted. Flowering was suppressed in heavily grazed sites with short vegetation. Higher levels of bare soil and phosphorus both had a positive relationship with genetic diversity, but probably for distinctly different reasons: bare soil provides safe sites for establishment, whilst phosphorus may promote flowering and improve seed germination. In order to conserve C. dissectum, management needs to maintain site heterogeneity so that C. dissectum can flower and establishment gaps are still available for seedlings; when either component is reduced, negative feedbacks through reduced genetic diversity and individual fitness can be expected. This study therefore highlights the importance of considering both conservation genetics and habitat quality in the conservation of plant species.     

Lipid Composition of the Roots of Plantago Species: Response to Alteration of the Level of Mineral Nutrition and Ecological Significance

In a study of the adaptation of Plantago species to their specific environment the lipid composition of the roots of several species: Plantago major L. ssp. major, Plantago major L. ssp. pleiosperma Pilger, Plantago lanceolata L., Plantago media L., Plantago maritima L., Plantago coronopus L. was studied as well as the effect of the nutritional regime. Upon exposure to low-salt conditions Plantago major L. ssp. major L. and Plantago maritima L. maintained the level of free sterols in the roots, despite a depressed level of total sterols, and the root lipids were more saturated than under high-salt conditions. Both factors may reduce nutrient leakage from the roots to the low-salt condition. Upon exposure to low-salt conditions, all Plantago species showed a decreased level of galactolipid (exception: Plantago coronopus) and a decreased level of sitosterol (exception: Plantago maritima); the latter being compensated by an elevated level of cholesterol + tocopherol in Plantago major ssp. pleiosperma, Plantago lanceolata and Plantago media. Plantago coronopus was the only species which under low-salt conditions showed an increased level of free sterols, among which cholesterol was the most important; thus indicating a high degree of regulation of membrane permeability under alternating nutritional conditions. The level of sulfolipid was kept constant in all Plantago species, with the highest level observed in Plantago maritima. The role which various lipids may play in maintenance of membrane integrity under alternating nutritional conditions is discussed.     

Behaviour of male and female parasitoids in the field: influence of patch size,host density,and habitat complexity

1. Two field experiments were carried out to examine the role of patch size, host density, and complexity of the surrounding habitat, on the foraging behaviour of the parasitoid wasp Cotesia glomerata in the field. 2. First, released parasitoids were recaptured on patches of one or four Brassica nigra plants, each containing 10 hosts that were placed in a mown grassland area. Recaptures of females were higher than males, and males and females aggregated at patches with four plants. 3. In experiment 2, plants containing 0, 5 or 10 hosts were placed in unmown grassland plots that differed in plant species composition, on bare soil, and on mown grassland. Very low numbers of parasitoids were recaptured in the vegetated plots, while high numbers of parasitoids were recaptured on plants placed on bare soil or in mown grassland. Recaptures were higher on plants on bare soil than on mown grassland, and highest on plants containing 10 hosts. The host density effect was significantly more apparent in mown grassland than on bare soil. 4. Cotesia glomerata responds in an aggregative way to host density in the field. However, host location success is determined mostly by habitat characteristics, and stronger host or host‐plant cues are required when habitat complexity increases.     

Niche differentiation and preferences of Elaphrus cupreus Duftschmid, 1812 and Elaphrus uliginosus (Fabricius, 1792) (Coleoptera: Carabidae) as reason for their different endangerment in Central Europe

Ecological demands were studied in Elaphrus cupreus, a frequent species, and E. uliginosus, one of the most endangered carabids in Europe. Ecological experiments were performed in the laboratory and the field in northern Germany. Abundance was measured using the mark-and-recatch method. Both species prefer sites with water content changes lower than 25%. E. uliginosus prefers higher temperatures than E. cupreus and niche breadth was on average wider in E. cupreus than in E. uliginosus. E. uliginosus is restricted to grassland sites with a low fraction (>0 and 25%) of bare soil. E. cupreus was mainly found on sites having >75–100% bare soil. This preference of E. uliginosus can be referred to its high temperature demands using open regions for warming up and its predatory behaviour using hideouts for a rapid predatory push. The wider niche structure and lower temperature demand of E. cupreus allows the inhabitation of cooler alder woods. It prefers large fractions of bare soil because its predatory behaviour is to run randomly on the soil. Thus, E. uliginosus demands wet extensively grazed grassland where cattle produces low fractions of bare soil, while E. cupreus prefers intensively grazed sites in grassland or alder woods.     

The role of below-ground competition during early stages of secondary succession: the case of 3-year-old Scots pine (Pinus sylvestris L.) seedlings in an abandoned grassland

In abandoned or extensively managed grasslands, the mechanisms involved in pioneer tree species success are not fully explained. Resource competition among plants and microclimate modifications have been emphasised as possible mechanisms to explain variation of survivorship and growth. In this study, we evaluated a number of mechanisms that may lead to successful survival and growth of seedlings of a pioneer tree species (Pinus sylvestris) in a grass-dominated grassland. Three-year-old Scots pines were planted in an extensively managed grassland of the French Massif Central and for 2 years were either maintained in bare soil or subjected to aerial and below-ground interactions induced by grass vegetation. Soil temperatures were slightly higher in bare soil than under the grass vegetation, but not to an extent explaining pine growth differences. The tall grass canopy reduced light transmission by 77% at ground level and by 20% in the upper part of Scots pine seedlings. Grass vegetation presence also significantly decreased soil volumetric water content (Hv) and soil nitrate in spring and in summer. In these conditions, the average tree height was reduced by 5% compared to trees grown in bare soil, and plant biomass was reduced by 85%. Scots pine intrinsic water-use efficiency (A/g), measured by leaf gas-exchange, increased when Hv decreased owing to a rapid decline of stomatal conductance (g). This result was also confirmed by δ ¹³C analyses of needles. A summer ¹⁵N labelling of seedlings and grass vegetation confirmed the higher NO₃ capture capacity of grass vegetation in comparison with Scots pine seedlings. Our results provide evidence that the seedlings' success was linked to tolerance of below-ground resource depletion (particularly water) induced by grass vegetation based on morphological and physiological plasticity as well as to resource conservation.     

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.     

Effects of Sward Height, Gap Size, and Slug Grazing on Emergence and Establishment of Trollius europaeus (Globeflower)

Abstract To determine the effects of sward height and gap size on the emergence and subsequent development of Trollius europaeus, seeds were introduced into artificially created gaps in mown and unmown wet meadow grassland in Ayr, Scotland. Emergence and establishment of nondormant seed (previously leached in running tap water and soaked in gibberellic acid) placed in a range of circular gaps (25 mm, 50 mm with and without root barrier, 100 and 200 mm diameter) were monitored over 305 days. Trollius demonstrated high field emergence (mean, 46% of viable seed sown) in this experiment. Emergence was not significantly affected by either sward height or gap size, although emergence was greater in mown swards. The critical factor determining subsequent survival of seedlings was slug predation rather than sward height or gap size. However, losses due to slug predation were significantly greater in unmown swards, resulting in more seedlings in mown swards. One hundred eleven days after sowing (September) seedling numbers had declined substantially in both unmown and mown swards, and by day 305 (April) seedlings were extinct in both sward types. The implications of the research for the restoration and management of T. europaeus are discussed.     

The effects of sheep-grazing on the subterranean termite fauna (Isoptera) of the Western Australian wheatbelt

Abstract The majority of existing remnants of wandoo Eucalyptus capillosa woodland in the Western Australian wheatbelt have been grazed by sheep for several decades and are often visibly degraded. A pilot survey was conducted into the effects of sheep on vegetation and soil variables, and the abundance, diversity and species frequency of occurrence of subterranean termite communities. Ten 1/4 ha study plots were used for paired grazed/ungrazed comparisons. Ungrazed plots had more litter mass (dry weight), leaf and woody litter, canopy cover (%) and soil moisture (moisture content <1.2% across study plots); grazed plots had a higher percentage of bare ground. Termites were as abundant, and as diverse, in grazed as in ungrazed plots, and were equally often sampled in the soil and surface wood. Termite species eating sound wood, decayed wood/debris and grass were sampled equally often, and were of equal diversity in sheep-grazed as in ungrazed plots. The mounds of Drepanotermes tamminensis were more abundant in grazed plots. These findings indicate that prolonged sheep grazing in remnants of wandoo woodland of the Western Australian wheatbelt has had no detrimental or beneficial effect on its subterranean termites.     

Factors Influencing Seedling Establishment Rates in Zostera marina and Their Implications for Seagrass Restoration

Selection of strategies to efficiently utilize limited seed supplies in efforts to restore the seagrass Zostera marina (eelgrass) requires a better understanding of the processes that limit seedling establishment at potential restoration sites. We investigated the effect of seed distribution timing on seedling establishment and tested for interactive effects of seed burial and distribution timing. We also investigated the effect of habitat type on seedling establishment by distributing Z. marina seeds inside and outside of established Ruppia maritima (widgeongrass) patches and examined mechanisms causing habitat differences by manipulating seed position (buried or unburied) and vulnerability to seed predators (unprotected or protected in packets). Seeds distributed on the sediment surface in the summer (July or August) produced fewer seedlings than seeds distributed in fall (October) in five of six trials over 3 years. Seed burial increased success rates for seeds distributed in summer at one of two sites tested, eliminating the effect of season, but reduced success at the other site. Seeds placed in R. maritima generally produced fewer seedlings than seeds in bare sand, and although seed burial and protection in packets increased success in bare sand at three of four sites, the effect was less consistent in R. maritima. We conclude that seed predation and physical interactions were influential in reducing seedling establishment in R. maritima, contrary to hypotheses positing a nursery role for existing vegetation. Efficient restoration efforts with Z. marina seeds should target unvegetated areas after summertime sources of mortality have diminished. Direct seed burial may enhance seedling establishment rates.     

Effects of Planting Depth,Sediment Grain Size,and Nutrients on Ruppia maritima and Potamogeton perfoliatus Seedling Emergence and Growth

Protocols are now available for seed harvest, storage and germination of several mesohaline and polyhaline species; however, low seedling survival rates point to the need for an increased understanding of factors affecting seedling establishment. Depth of seed burial in sediments and initial seedling growth rates are shown to be limiting factors for photosynthetic competency of Ruppia maritima and Potamogeton perfoliatus. Seedling emergence is inversely proportional to planting depth on sediments ranging in grain size from coarse sands (850 μm) to silt (63 μm). Less than 6% of the seeds of either species emerged when buried to a depth of 3 cm in test sediments. Germination was greatest for seeds placed on the surface of sediments; however, these seedlings were subject to displacement because of the weak and fragile roots produced during early growth. Fine sediments may be more favorable for R. maritima seedling establishment, because seedling emergence and height decreased with increasing sediment grain size. Potamogeton perfoliatus seedlings seem to be more tolerant of a wider range of sediment grain sizes than R. maritima as indicated by the lack of an effect of sediment grain size on P. perfoliatus seed emergence, seedling height, and biomass. Increasing nutrients stimulated seedlings of both species; however, even at the highest concentrations tested, growth, as determined by shoot elongation and leaf and root formation, slowed within 7–10 days. This suggests factors other than mineral nutrients and light limit growth or that growth shifts from aboveground biomass production to belowground vegetative spread.     

Impacts of secondary seed dispersal and herbivory on seedling survival in Aesculus turbinata

Abstract. Aesculus turbinata is a tree species with large seeds (6.2 g mean dry weight). We studied the demography of its seeds and seedlings in a temperate deciduous forest in northern Japan to elucidate the ecological significance of large seeds with special reference to herbivory and secondary dispersal. Both seed and seedling stages suffered greatly from herbivores. Seedling herbivory was important judged from experiments with shoot clipping and hypogeal cotyledon removal. However, some seedlings survived through re-sprouting after herbivory. Survival rate and percentage resprouting seedlings were lower than those with remaining cotyledons, though seedling size was not affected. This suggests that stored resources in hypogeal cotyledons are working as a kind of ‘risk hedge’ against severe aboveground shoot clipping experienced by A. turbinata. The spatial distribution of seedlings was expanded via seed scatter-hoarding by rodents. Seedling survival rate was higher within canopy gaps than under closed canopy, indicating that canopy gaps are safe sites for establishment, and was negatively correlated with seedling density. Therefore, secondary seed dispersal in this species seems to be effective in ‘finding’ safe sites and in ‘escaping’ density-dependent mortality. The large seeds and seedlings of A. turbinata are attractive to herbivores, but the high resistance of seedlings to herbivory due to large reserves and the effective secondary dispersal appear to mitigate these disadvantages.