Consumer pressure, seed versus safe-site limitation, and plant population dynamics

Plants often suffer reductions in fecundity due to insect herbivory. Whether this loss of seeds has population-level consequences is much debated and often unknown. For many plants, particularly those with long-lived seedbanks, it is frequently asserted that herbivores have minimal impacts on plant abundance because safe-site availability rather than absolute seed number determines the magnitude of future plant recruitment and hence population abundance. However, empirical tests of this assertion are generally lacking and the interplay between herbivory, spatio-temporal variability in seed- or safe-site-limited recruitment, and seedbank dynamics is likely to be complex. Here we use a stochastic simulation model to explore how changes in the spatial and temporal frequency of seed-limited recruitment, the strength of density-dependent seedling survival, and longevity of seeds in the soil influence the population response to herbivory. Model output reveals several surprising results. First, given a seedbank, herbivores can have substantial effects on mean population abundance even if recruitment is primarily safe-site-limited in either time or space. Second, increasing seedbank longevity increases the population effects of herbivory, because annual reductions in seed input due to herbivory are accumulated in the seedbank. Third, population impacts of herbivory are robust even in the face of moderately strong density-dependent seedling mortality. These results imply that the conditions under which herbivores influence plant population dynamics may be more widespread than heretofore expected. Experiments are now needed to test these predictions. Received: 3 November 1999 / Accepted: 15 February 2000     

Positive feedbacks between plant invasions and fire regimes: <Emphasis Type="Italic">Teline monspessulana</Emphasis> (L.) K. Koch (Fabaceae) in central Chile

Invasive species can increase fire frequency and intensity, generating favorable conditions for their self-perpetuation. Mediterranean south-central Chile may be especially prone to the effects of invasive species on fire regimes because it is less adapted to fire and it contains a highly endemic flora. Teline monspessulana (L.) K. Koch (syn. Cytisus monspessulanus L.; Genista monspessulana (L.) L.A.S. Johnson) is an introduced shrub that forms monotypic stands or is present as an understory species in native forests as well as in forestry plantations. Dense T. monspessulana stands are completely destroyed by fire, generating the conditions for it seeds to germinate and establish an abundant regeneration, with up to 900 plants/m². We report key evidence on abundance and biomass in adult stands, and patterns of seed bank and regeneration after fire in stands of T. monspessulana around the city of Concepción, Chile. We estimated living biomass in pure stands and underneath Eucalyptus plantations. In burned areas, we assessed T. monspessulana seed bank and studied regeneration patterns. We found that T. monspessulana densities reaches 52,778 plants/ha and 8.92 ton/ha in pure stands and 34,223 plants/ha and 2.31 ton/ha underneath Eucalyptus plantations. T. monspessulana generates small caliper fuel and acts as a ladder-fuel. Large soil seed banks allow for abundant regeneration after fire, with mean densities of 877,111 plants/ha, but an overall mortality of 37.2% in the first year after the fire. The high values of regeneration compared to final densities in adult stands suggest that density-dependent mortality. Our results indicate that T. monspessulana regeneration is not only favored by fires, but also that the species creates favorable conditions for intense and continuous fires, both under pure conditions, but also associated to exotic tree plantations. To understand the implications of positive feedbacks between invaders and fire, we recommend focusing in the mechanisms by which they increases fuel accumulation and fuel flammability, and how higher fire frequency and intensity favors invasive species recruitment over native species. Comprehension of this dynamics will allow for better management and control of these invasions which have major ecological, economical and social implications.     

Vegetation recovery patterns in early volcanic succession

Permanently plots were monitored from 1983 to the present on Mount Usu after the eruptions of 1977–78 which destroyed the pre-eruption vegetation by 1–3 m thick accumulations of ash and pumice in order to clarify the processes and mechanisms of succession. Until now, 163 species were recorded in the summit area. Most of these species were derived from vegetative reproduction throughout the volcanic deposits. Vegetative reproduction plays a major role on increases in cover. Although long-distance seed-dispersal species could immigrate to the crater basin, their cover increase was slow. Seedbank species only established in gullies where the original topsoil was exposed by erosion. Most annuais were supplied by the seedbank in the original topsoil and woody species originated via immigration, suggesting that the source greatly determines the species composition of establishing vegetation. Annual seedlings showed low survival, while overwintering perennial seedlings steadily established. Ground surface movements strongly restricted increases in plant cover and the distance from source vegetation was the principal determinant of plant density. Due to differences in disturbance intensity, successional rates were higher in the stable substrates outside gullies and lower on the exposed original topsoil in some gullies. Recipient of the Botanical Society Award of Young Scientists, 1994     

Spatial variation of seedling distribution in an east Mediterranean pine woodland at the beginning of post-fire succession

Most of the area in pine woodlands is occupied by perennial seeders that regenerate from seeds in the first winter after the fire and by annuals. Control of the germination in the regenerating vegetation after wildfire is therefore a primary ecological component of the post-fire succession in this ecosystem. The aim of the study presented here was to determine the distribution of Pinus, Cistus and other plants seeds around burned Pinus halepensis trees, and to measure the conditions related to seed germination in the upper soil layers in the same locations. The study was carried out in a 50-year old planted Pinus halepensis woodland that was burned down by a wildfire in July 1995. The variation of seedbank density was determined by collecting samples under the canopies of burned trees and in a nearby open area. Pine seedbank density decreased and that of Cistus and annuals increased with increasing distance from the burned trunks. Most pine seeds were present in the ash layer while those of the other plants were in the soil. In situ germination experiments showed that seedling density decreased with distance from the burned trunks while the proportion of pines in the seedling population increased. This was a result of seedbank variation and germination inhibition by the high pH conditions caused by the ash. The establishment of sparse pine seedling under the dead tree canopies insured their rapid development without interference by other plants and played a key role in the regeneration and stability of the pine woodland community. The concomitant mass germination of the perennial seeders in the rest of the area prevented invasion by annuals.     

The invasion of Pteridium aquilinum and the impoverishment of the seed bank in fire prone areas of Brazilian Atlantic Forest

Several studies have documented that fires are widespread in the tropics. Because fire in the Atlantic Rain Forest is rare, fire events and their consequences at the community level have not been evaluated. This study describes the composition of different seed banks in areas of Atlantic Forest. The study was carried out in the National Park of Tijuca in the state of Rio de Janeiro. Four different areas were chosen considering their differences on fire history. Fifteen soil surface samples (23 cm (centimeter) diameter ×7 cm height at 5 cm deep) were obtained from random locations within each site. The dominant family at all study sites except in the most preserved one were Dennstaetiaceae, followed by Melastomataceae and Poaceae. The family Dennstaetiaceae was exclusively represented by Pteridium aquilinum (Klf.) Herter. There were no significant differences among areas when comparing the number of seedlings excluding P. aquilinum. However, the number of P. aquilinum was higher in the most disturbed area, while Melastomataceae and Rubiaceae were more frequent in a less impacted area. Our results suggest that the seed bank in disturbed areas of the Atlantic Forest possibly will not contribute for forest restoration after disturbance.     

Seedbank persistence and seedling emergence of seven weed species in autumn-sown crops following a single year's seeding

Seedbanks and seedling emergence, resulting from seeds shed in wheat crops and ploughed in, were monitored in two experiments for 3–4 years in successive autumn sown crops, in which herbicides were used to prevent re-seeding. Populations of Galium aparine, Papaver rhoeas, Lamium purpureum, Myosotis arvensis, Veronica persica, Viola arvensis and Veronica hederifolia were studied in separate plots. The numbers of viable seeds in the soil declined exponentially; G. aparine and V. hederifolia declined most, and P. rhoeas and V. arvensis least rapidly. Total weeds (all species) over the 3–4 years, (obtained by adding successive numbers of seedlings recorded in each autumn or winter), represented 3–4% of the first year seedbank for the combined species. Few seedlings emerged in the first year after seeding due to effective burial by the initial ploughing. The species showing the greatest emergence in the first year, relative to seedbank size, were G. aparine and V. hederifolia. Other, smaller seeded, species produced their main seedling flushes in the second and third years after seeding. It is concluded that poor weed control in one year is likely to result in large weed infestations for up to four years and probably longer for some species.     

Litter legacy after spruce plantation removal hampers initial vegetation establishment

Restoration of areas used for intensive even-aged Norway spruce (Picea abies Karst.) plantations often involves felling and subsequent spontaneous vegetation succession. However, the accumulated litter layer may hamper vegetation development, and thereby postpone recovery or even change the outcome.We studied effects of the litter layer on vegetation establishment during two seasons following a clear-cut of Norway spruce in Denmark. We experimentally assessed the response of multiple vegetation properties to litter removal, with and without wildlife exclusion by fencing, and in combination with sowing of trees, while fencing. Burning was tested as an alternative way to remove the litter layer.Vegetation establishment was poor, when the litter layer was intact, and cover developed slowly remaining below 10% after two years, irrespective of fencing. In contrast, litter removal and fencing together gave significantly faster recovery and reached nearly 60% mean cover. Vegetation cover was driven by few dominant species, especially the sedge Carex pilulifera. Species richness was similar in all treatments, but increased with sowing of trees. Fencing resulted in taller birch seedlings independently of litter removal, but enhanced by seedling density. Litter removal seemed to favor species with lighter seeds, lower specific leaf area (SLA) and lower Ellenberg N value, i.e. associated with relative infertile conditions. Disturbing the litter by burning seemed to have an effect comparable to mechanical removal, and could be a management alternative.Our results showed that a persistent litter layer after spruce plantation removal may hamper the initial vegetation establishment. Actively removing litter may serve as an additional restoration intervention to overcome this legacy. However, as grazing can keep this potential in check, wildlife exclusion may be necessary as well. To speed up recovery and diversify vegetation structure after spruce plantation removal, we suggest patchy disturbance of the litter, essentially combined with wildlife exclusion.     

Variable effects of endophytic fungus on seedling establishment of fine fescues

Seedborne systemic endophytic fungi of grasses are thought to be plant mutualists, because they have been shown to improve their host’s resistance against biotic and abiotic stresses. The interactions in plant–endophyte associations vary from mutualistic to parasitic with environmental conditions and the genotypes of interacting species. The possible pros and cons of endophytic fungi are expected to be most evident during the seedling establishment, where host fitness is most directly affected. If this holds true, endophytes may play a focal role in local adaptation of hosts to different environments. We examined if endophyte-infected and uninfected seeds and seedlings of two native grass species, Festuca rubra and F. ovina, differ in seed germination and seedling growth rates under greenhouse conditions. The germination of F. rubra seeds was also studied in the field. This is the first time that the effects of Epichloë endophyte on seedling establishment of fine fescues from natural populations have been experimentally evaluated. Mother plant (seed family) had a marked effect on many response variables in both grass species. Length and mean biomass of tillers of endophyte-infected (E+) F. ovina seedlings were lower, but root:shoot ratios were higher than in endophyte-free (E?) seedlings. In F. rubra, the effects of the endophyte were dependent on the habitat where the seeds were collected. The E+ seeds from river banks germinated faster than E+ seeds from meadows, and E+ seedlings from the river banks produced fewer but taller and heavier tillers than the other seedlings. Our data suggest that the effects of the endophyte infection on the seedling stage of fine fescues are dependent the species of grass, host genetic background and mother plant habitat. The germination strategy and growth form of E+ red fescue seedlings from river banks may be beneficial to surviving in the harsh conditions of that habitat.     

The germination process in vernal pools: sensitivity to environmental conditions and effects on community structure

Variation in timing and amount of rainfall in California has been noted to cause strong year-to-year differences in the composition of vernal pool communities. We explored the effects of possible variation of the “rainy season” in a seedbank germination study conducted with three factors: monthly timing of first soil moistening, length of moist period before inundation, and length of inundation. Monthly timing was the most important factor in determining the number of plants and number of species that germinated in the pots. Many species, mostly generalist wetland and pool-endemic species, showed sharp decreases in number in later trials. Five species did not germinate in the final trial, and two of these were abundant only in the first trial. Inundation was required for the germination of two species. Treatments with a combination of inundated and moist conditions resulted in higher numbers of species and numbers of plants compared to treatments with only moistness or only inundation. Our results indicate that vernal pool species have mechanisms that keep them from emerging under unfavorable conditions. The sensitivity of vernal pool species to out-of-season germination suggests that they could be negatively affected by climatic change, with generalist species like non-native Lythrum hyssopifolium having a potential advantage. Received: 15 July 1996 / Accepted: 14 July 1997