Ecological application of biotic resistance to control the invasion of an invasive plant,Ageratina altissima

Biotic resistance is the ability of species in a community to limit the invasion of other species. However, biotic resistance is not widely used to control invasive plants. Experimental, functional, and modeling approaches were combined to investigate the processes of invasion by Ageratina altissima (white snakeroot), a model invasive species in South Korea. We hypothesized that (1) functional group identity would be a good predictor of biotic resistance to A. altissima, whereas a species identity effect would be redundant within a functional group, and (2) mixtures of species would be more resistant to invasion than monocultures. We classified 37 species of native plants into three functional groups based on seven functional traits. The classification of functional groups was based primarily on differences in life longevity and woodiness. A competition experiment was conducted based on an additive competition design with A. altissima and monocultures or mixtures of resident plants. As an indicator of biotic resistance, we calculated a relative competition index (RCI_avg) based on the average performance of A. altissima in a competition treatment compared with that of the control where only seeds of A. altissima were sown. To further explain the effect of diversity, we tested several diversity–interaction models. In monoculture treatments, RCI_avg of resident plants was significantly different among functional groups but not within each functional group. Fast‐growing annuals (FG1) had the highest RCI_avg, suggesting priority effects (niche pre‐emption). RCI_avg of resident plants was significantly greater in a mixture than in a monoculture. According to the diversity–interaction models, species interaction patterns in mixtures were best described by interactions between functional groups, which implied niche partitioning. Functional group identity and diversity of resident plant communities were good indicators of biotic resistance to invasion by introduced A. altissima, with the underlying mechanisms likely niche pre‐emption and niche partitioning. This method has most potential in assisted restoration contexts, where there is a desire to reintroduce natives or boost their population size due to some previous level of degradation.     

Giant Ragweed Invasion is Not Well Controlled by Biotic Resistance

The effect of native plant restoration on invasion by giant ragweed (Ambrosia trifida), an invasive species, is currently unknown. We hypothesized that (1) functional group identity would be a good predictor of biotic resistance to A. trifida, and (2) mixtures of species would be more resistant to invasion than monocultures. Using seven functional traits, 37 native and non-native plants were divided into three functional groups that differed primarily in longevity and woodiness. We conducted a competition experiment using an additive competition design with A. trifida and monocultures or mixtures of 14 species. Biotic resistance was evaluated by calculating a relative competition index (RCI_avg) based on the average performance of A. trifida in treatments compared with that in control. In monocultures, RCI_avg of resident plants did not significantly differ among the three functional groups or within each functional group. The highest RCI_avg (40%) was observed for some fast-growing annuals (FG1) such as Zea mays and Secale cereal, which were strong competitors. RCI_avg of resident plants was not significantly greater in mixtures than in monocultures. Taken together, the results show that plant diversity did not control invasion by A. trifida and that giant ragweed invasion cannot be well controlled by biotic resistance.     

Seasonality and facilitation drive tree establishment in a semi-arid floodplain savanna

A popular hypothesis for tree and grass coexistence in savannas is that tree seedlings are limited by competition from grasses. However, competition may be important in favourable climatic conditions when abiotic stress is low, whereas facilitation may be more important under stressful conditions. Seasonal and inter-annual fluctuations in abiotic conditions may alter the outcome of tree–grass interactions in savanna systems and contribute to coexistence. We investigated interactions between coolibah (Eucalyptus coolabah) tree seedlings and perennial C₄ grasses in semi-arid savannas in eastern Australia in contrasting seasonal conditions. In glasshouse and field experiments, we measured survival and growth of tree seedlings with different densities of C₄ grasses across seasons. In warm glasshouse conditions, where water was not limiting, competition from grasses reduced tree seedling growth but did not affect tree survival. In the field, all tree seedlings died in hot dry summer conditions irrespective of grass or shade cover, whereas in winter, facilitation from grasses significantly increased tree seedling survival by ameliorating heat stress and protecting seedlings from herbivory. We demonstrated that interactions between tree seedlings and perennial grasses vary seasonally, and timing of tree germination may determine the importance of facilitation or competition in structuring savanna vegetation because of fluctuations in abiotic stress. Our finding that trees can grow and survive in a dense C₄ grass sward contrasts with the common perception that grass competition limits woody plant recruitment in savannas.     

Restoration of Araucaria Forest: The Role of Perches, Pioneer Vegetation, and Soil Fertility

The Araucaria forest is Brazil's highly threatened subtropical forest ecosystem that has been disappearing in recent decades. Restoration programs involving this forest type are scarce, and there is a lack of scientific information on how ecological processes such as competition, facilitation, and seed dispersal influence natural forest restoration. This work aims to investigate how use of perches to attract seed dispersers and the influence of pioneer vegetation and soil fertilization could affect the colonization of woody species in a degraded area. An experiment was conducted in an abandoned field where the natural establishment of seeds and seedlings of woody species was monitored under factorial combinations of the following treatments: (1) pioneer vegetation (presence and absence); (2) soil fertility (addition of NPK and control); and (3) perches (presence and absence). Seed and seedling abundance, seed and seedling species richness, and seedling mortality were recorded monthly during 12 months. Seed abundance and species richness were significantly greater in places with perches than in control plots. These results were consistent over the year and more pronounced when the surrounding forest produced a higher amount of fruit. Species richness and abundance of seedlings were significantly greater in places with perches than in control plots, and in places with vegetation than without. Soil fertility did not influence seedling establishment. Facilitation and seed dispersal are important factors affecting the colonization of woody species in this subtropical area. Nutrient availability neither regulates the facilitation process nor influences species replacement during the early stages of Araucaria forest succession.     

Are variations of direct and indirect plant interactions along a climatic gradient dependent on species’ strategies? An experiment on tree seedlings

Investigating how interactions among plants depend on environmental conditions is key to understand and predict plant communities’ response to climate change. However, while many studies have shown how direct interactions change along climatic gradients, indirect interactions have received far less attention. In this study, we aim at contributing to a more complete understanding of how biotic interactions are modulated by climatic conditions. We investigated both direct and indirect effects of adult tree canopy and ground vegetation on seedling growth and survival in five tree species in the French Alps. To explore the effect of environmental conditions, the experiment was carried out at 10 sites along a climatic gradient closely related to temperature. While seedling growth was little affected by direct and indirect interactions, seedling survival showed significant patterns across multiple species. Ground vegetation had a strong direct competitive effect on seedling survival under warmer conditions. This effect decreased or shifted to facilitation at lower temperatures. While the confidence intervals were wider for the effect of adult canopy, it displayed the same pattern. The monitoring of micro‐environmental conditions revealed that competition by ground vegetation in warmer sites could be related to reduced water availability; and weak facilitation by adult canopy in colder sites to protection against frost. For a cold‐intolerant and shade‐tolerant species (Fagus sylvatica), adult canopy indirectly facilitated seedling survival by suppressing ground vegetation at high temperature sites. The other more cold tolerant species did not show this indirect effect (Pinus uncinata, Larix decidua and Abies alba). Our results support the widely observed pattern of stronger direct competition in more productive climates. However, for shade tolerant species, the effect of direct competition may be buffered by tree canopies reducing the competition of ground vegetation, resulting in an opposite trend for indirect interactions across the climatic gradient.     

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.     

Limitations on tree seedling establishment across ecotones between abandoned fields and adjacent broad-leaved forests in eastern Japan

Field abandonment has increased over several decades under a long-term recession in Japanese agriculture. To support effective farmland management or reforestation, the patterns of tree recruitment in abandoned fields, which influence management costs, need to be clarified. We investigated tree seedling establishment and microsite variables along forest–field transects in 11 abandoned fields adjacent to secondary broad-leaved forests in the eastern Kanto region of Japan. Generalized linear mixed model analysis indicated that tree seedling establishment in abandoned fields was not correlated with the period of abandonment. Instead, it was positively correlated with vegetation openness. The dominance of a woody vine (Pueraria lobata), an annual liana (Humulus japonicus), and a dwarf bamboo (Pleioblastus chino) mainly explained low vegetation openness within the fields. In addition, the establishment of evergreen tree seedlings and tree seedlings dispersed by hoarding was negatively correlated with the distance from the forest edge. In the forest interior, taller understory vegetation at the edge correlated with lower seedling establishment. These results suggest that seedling establishment across forest–field ecotones is restricted by the dominance of competitive natives within abandoned fields and distance-dependent limitations in both abandoned fields and adjacent forest.     

Competitor or facilitator? The ambiguous role of alpine grassland for the early establishment of tree seedlings at treeline

Alpine treelines are expected to move upslope with a warming climate. However, so far treelines have responded inconsistently and future shifts remain difficult to predict since many factors unrelated to temperature, such as biotic interactions, affect responses at the local scale. Especially during the earliest regeneration stages, trees can be strongly influenced by alpine vegetation via both competition and facilitation. We aimed to understand the relative importance of these two types of interaction in different vegetation structures for treeline regeneration dynamics. Effects of herbaceous alpine vegetation on seedling emergence and first‐year performance were studied in a field experiment in the French Alps (2100 m a.s.l.) with five important European treeline tree species: Larix decidua, Picea abies, Pinus cembra, Pinus uncinata and Sorbus aucuparia. Total emergence and locally‐germinated seedling survival were not affected, but for seedlings planted at two months of age, negative vegetation impacts dominated for all response parameters: first‐year survival, growth and carbohydrate accumulation. However, in the winter half‐year, evergreen tree seedlings increased carbohydrate reserves under the protection of senescent herbs. Also, responses of locally‐germinated seedlings suggest facilitative vegetation effects in the first two months after emergence. Thus, the interaction switched between competition and facilitation according to ontogenetic stage and seasons. Still, the net outcome after one year was negative, but species differed in their susceptibilities. Because initial establishment is the first bottleneck determining whether treelines remain stable or move upslope, understanding establishment, including site‐, life‐stage and species‐specific processes, is essential for understanding observed treeline spatial patterns and dynamics. When developing predictive models of treeline dynamics, all these ‘local’ aspects should be incorporated in addition to more global drivers like changes in temperature.     

Effects of vegetation canopy and climate on seedling establishment in Mediterranean shrubland

Abstract. Question: Does the influence of plant canopy on seedling establishment interact with climate conditions, and particularly, do intensified drought conditions, enhance a positive effect of the vegetation canopy on seedlings in Mediterranean‐type ecosystems. Location: Mediterranean shrubland near Barcelona, Spain at 210 m a.s.l. Methods: Over the course of four years we recorded seedling emergence and survival in open areas and below vegetation under control, drier and warmer experimental climatic conditions. Results: Seedling emergence is more sensitive to climate conditions than later stages of growth. When considering the whole set of species, the total number of established seedlings at the end of the experiment was lower in the drought and warming stands than in control ones, and vegetation canopy increased the number of these seedlings in the drought stands. Drought reduced seedling emergence but not warming, while the interaction between climate treatments and vegetation canopy was not significant. Seedling survival was lower in the warming treatment than in the control. Under drought conditions, vegetation canopy increased seedling emergence of the dominant Globularia alypum. In control stands, vegetation canopy reduced their survival. Vegetation canopy increased the survival of the dominant Erica multiflora in warming stands, and it reduced the survival of G alypum in drought stands. No significant effects of drought and warming were observed in the seed rain of these two species. Conclusions: The balance of the facilitation‐competition interactions between vegetation canopy and seedling establishment in Mediterranean‐type ecosystems determined by water availability, and drought conditions enhance the positive effect of vegetation canopy. This interaction is species‐specific and shows important between‐year variability.     

Plantation age,understory vegetation,and species‐specific traits of target seedlings alter the competition and facilitation role of Eucalyptus in South China

The Chinese government is considering ways to reconstruct Eucalyptus plantation monocultures in order to improve their ecosystem services. However, the effects of competition or facilitation of Eucalyptus on native species during the reconstruction have not been identified, especially during the formation of novel ecosystems. To better understand these relationships, in this study we tested the effects of competition and facilitation of Eucalyptus on three target species (Schima superba, Michelia macclurei, and Castanopsis fissa), in Eucalyptus plantations with different ages and understory vegetation. The results showed that the survival and growth of S. superba and M. macclurei were not sensitive to plantation age and understory removal treatment. The survival of C. fissa in understory removal was significantly higher and the growth of C. fissa increased gradually with increasing plantation age. The relative neighbor effect (RNE) index for growth indicated that the relationship between Eucalyptus and target species switched from competition to facilitation with an increase in plantation age. A redundancy analysis (RDA) showed that survival was positively correlated with soil water content and soil phosphorus. Growth was positively correlated with soil organic matter and soil nitrogen and negatively correlated with soil bulk density. Our results indicate that plantation age, understory vegetation, and species‐specific traits of target seedlings could change the competition and facilitation roles of Eucalyptus. Higher survival and growth rates indicated that C. fissa was an appropriate species for the reconstruction of mature Eucalyptus plantations and understory removal will benefit the process.     

Soil heterogeneity effects on canopy structure and composition during early succession

Early succession from annuals to herbaceous and woody perennials on abandoned fields and grasslands is marked by vegetation heterogeneity; i.e., variation in canopy composition and structure over the field through time. Soil resource heterogeneity could promote vegetation heterogeneity. We created soil resource patchiness on two newly ploughed fields by establishing two types (fertilized, unfertilized) of checkerboard plots with two, four, or eight subplots (trenched, untrenched) to test effects of soil fertility heterogeneity scale on vegetation heterogeneity during early succession. Canopy composition and structure were censused in years 2, 4 and 6. On both fields, soil resource heterogeneity did not affect canopy composition or structure. Differential abundance of dominant species, Ambrosia trifida on one field and Solidago altissima on the other, between fertilized and unfertilized plots led to decreased vegetation heterogeneity among subplots within plots and increased vegetation heterogeneity among plots. Soil enrichment promoted taller and layered, but species-poor, canopy. In general, soil fertility effects on abundance, expansion, and dominance of individuals of dominant species promoted variation in canopy composition and structure, or vegetation heterogeneity, among plots on the two fields.     

Facilitation and sand burial affect plant survival during restoration of a tropical coastal sand dune degraded by tourist cars

Coastal sand dunes support various ecosystem services, including storm protection and tourism. Restoration programs are often critical to preserve this ecosystem due to its fragility and high degree of degradation. Dune restoration still suffers from a general lack of knowledge of the ecological processes controlling tropical dune communities. We investigated if facilitation can increase restoration success by assisting plant survival in a Brazilian coastal dune degraded by buggies at the Environmental Protection Area of Jenipabu, RN, Brazil. We performed two field experiments on dune crest sites degraded by buggies. The first experiment tested how the presence of established vegetation and coconut mesh, mimicking soil‐stabilizing effect of vegetation, facilitates seedling survival and establishment of the early successional dominant dune plant Canavalia maritima (Fabaceae). The second experiment tested if coconut mesh and initial irrigation would allow the establishment of C. maritima outside the vegetation, using both seeds and transplants. We found that the presence of established vegetation positively affected seedling survival. Application of mesh positively affected microclimatic conditions and slightly increased survival, but only for seeds. Initial irrigation only had a small effect on transplanted seedlings survival. Nevertheless, there was extreme seedling mortality in the experiment in particular from sand burial. We conclude that restoration of these dune crests is very difficult. Facilitation by vegetation can reduce plant mortality by sand burial and desiccation, but successful restoration is likely to require a major effort involving a very high number of seedlings or seeds.     

A Comparative Study of Germination Biology and Plant Performance in Two Dry Grassland Species

Over the last 20 years, the species assemblage of the species-rich dry grassland communities of central Germany has changed due to the ongoing abandonment of traditional land-use practices. In our study we wanted to investigate the germination biology and the plant-plant interaction of the low-growing and declining forb Alyssum montanum and the increasingly dominant grass Festuca rupicola. To investigate the germination behaviour we simulated cold (8/4°C), warm (20/10°C) and hot (32/20°C) conditions under a light-dark regime (12/12 hours) as well as in constant darkness. Germination of both species was similar with almost all non-dormant seeds germinating under intermediate temperature conditions (20/10°C). Whereas F. rupicola (F) germinated equally well under light changing conditions and constant darkness, the germination of A. montanum (A) was clearly reduced in darkness, which implies a strong competitive advantage of F. rupicola in dense vegetation cover. However, A. montanum germinated early under all temperatures regimes and better under 20/10°C as well as with light changing conditions. The interactions between the two species were tested in a pod-experiment with a replacement design, within which the composition of individuals was changed in the following way: 4A/1F, 3A/2F, 2A/3F, 1A/4F (eight replications per each). To evaluate any influence of intraspecific competition we created monocultures in an additive design (1A, 2A, 3A, 4A, 5A, 1F, 5F; eight replications per each). Despite that A. montanum developed faster and flowered after only four months in the replacement design experiment, the herb responded negatively to the presence of F. rupicola in several parameters (number of flowers, number of fruits and biomass per individual). The calculation of the Relative Neighbour Effect (RNE) indicated facilitation for F. rupicola and competition on A. montanum, which increased with increasing number of individuals of the other species. Our data suggest that A. montanum seedling recruitment, growth and fitness are negatively affected by increasing vegetation density of the expanding grass species F. rupicola.     

Competitive abilities of introduced and native grasses

Differencesin competitive ability may explain the maintenance of existing plantpopulationsand the invasion of new areas by plant species. We used field experiments toexamine the competitive responses of Agropyron cristatum(L.) Gaertn., an introduced C₃ grass, and Boutelouagracilis (HBK.) Lag., a native C₄ grass, and thecompetitive effects of Agropyron-dominated vegetation andsuccessional prairie. We also tested whether the outcome of competitiveinteractions varied with water availability. In each vegetation type,transplants of each species were grown under two levels of competition(presenceor absence of neighboring vegetation) and three levels of water availability(high, medium, or low). Transplant survival, growth, and biomass allocationpatterns were measured. Water availability had no effect on the measuredvariables, suggesting that both species were limited by another resource.Growthrates were affected more by competition, while survival and root: shoot ratiowere affected more by transplant species identity. In the successional prairie,neighboring vegetation suppressed the growth of Agropyrontransplants less than that of Bouteloua transplants,suggesting that Agropyron has a stronger ability to resistcompetitive suppression in that vegetation type. The spread ofAgropyron into surrounding vegetation may relate to itsability to resist competitive suppression. In theAgropyron-dominated vegetation, neighboring vegetationsuppressed the growth of both species by the same extent. However, competitionaccounted for more variation in transplant growth inAgropyron-dominated vegetation than in successionalprairie, suggesting that Agropyron has strong competitiveeffects which hinder plant growth and prevent other species from establishinginAgropyron fields.     

Intraspecific facilitation: a missing process along increasing stress gradients – insights from simulated shrub populations

In recent years many field studies have been conducted to assess the relative importance of facilitation and competition in structuring vegetation communities in different environments. Herein, we present a simulation model which systematically explores the relative importance of intra‐specific facilitation and competition between adult shrubs and seedlings for spatial pattern formation. A grid‐based simulation model was constructed and calibrated using data collected in the field from Sarcopoterium spinosum populations in Israel to simulate population dynamics along a rainfall gradient. A series of simulation experiments was conducted in which manipulations of seedling survival probabilities were carried out to assess the relative importance of these processes in generating spatial patterns. Increased survival probabilities of first‐year shrubs in open areas were used to simulate competition effects, while increased survival probabilities in the vicinity of shrubs were used to simulate facilitation effects. Simulation results were then compared to shrub spatial patterns observed in the field. The results indicate that facilitation is not an important process in generating intra‐specific spatial patterns. Rather, in mesic environments with high precipitation, competition is the dominant process generating spatial patterns, resulting in regular spacing of shrubs, similarly to the patterns observed in the field (L(h) values<0). In arid sites, where precipitation values are lower, and stress conditions are higher, the dominant process generating spatial patterns was random mortality due to drought conditions. The resulting spatial patterns in this case are random (L(h)～0), whereas observed field populations exhibited clumped patterns (L(h)>0). We conclude that as stress conditions increase, the importance of intraspecific neighborhood interactions decrease whereas the importance of environmental factors increase in dictating intra‐specific spatial pattern formation. Consequently in mesic environments intra‐specific competition among adults determines the emerging patterns, while intraspecific facilitation is a negligible process.     

Where temperate meets tropical: multi-factorial effects of elevated CO2, nitrogen enrichment, and competition on a mangrove-salt marsh community

Our understanding of how elevated CO₂ and interactions with other factors will affect coastal plant communities is limited. Such information is particularly needed for transitional communities where major vegetation types converge. Tropical mangroves (Avicennia germinans) intergrade with temperate salt marshes (Spartina alterniflora) in the northern Gulf of Mexico, and this transitional community represents an important experimental system to test hypotheses about global change impacts on critical ecosystems. We examined the responses of A. germinans (C₃) and S. alterniflora (C₄), grown in monoculture and mixture in mesocosms for 18 months, to interactive effects of atmospheric CO₂ and pore water nitrogen (N) concentrations typical of these marshes. A. germinans, grown without competition from S. alterniflora, increased final biomass (35%) under elevated CO₂ treatment and higher N availability. Growth of A. germinans was severely curtailed, however, when grown in mixture with S. alterniflora, and enrichment with CO₂ and N could not reverse this growth suppression. A field experiment using mangrove seedlings produced by CO₂‐ and N‐enriched trees confirmed that competition from S. alterniflora suppressed growth under natural conditions and further showed that herbivory greatly reduced survival of all seedlings. Thus, mangroves will not supplant marsh vegetation due to elevated CO₂ alone, but instead will require changes in climate, environmental stress, or disturbance to alter the competitive balance between these species. However, where competition and herbivory are low, elevated CO₂ may accelerate mangrove transition from the seedling to sapling stage and also increase above‐ and belowground production of existing mangrove stands, particularly in combination with higher soil N.     

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.     

Facilitation and competition influence succession in a subtropical old field

The effects of neighboring vegetation and soil fertility on the establishment, survival and growth of tree species were studied in a subtropical old-field area in south Brazil. Seed damage, germination and seedling establishment of four tree species plus growth and survival of two transplanted tree species were monitored under factorial combinations of the following treatments: (1) pioneer vegetation (presence and absence); (2) soil fertility (addition of NPK and control). Facilitation was the main process affecting plant performance. The presence of pioneer vegetation significantly improved germination, establishment, growth and survival of most study species. Around 90% of sown seeds were damaged and the removal of pioneer vegetation significantly increased seed damage for all species studied, decreased germination in three out of four species, and decreased establishment in one species. Moreover a significantly higher seedling growth rate of Inga virescens was found after the first year of the experiment in plots where vegetation was present. The presence of vegetation significantly increased seedling survival of I. virescens by protecting seedlings from leaf loss due to winter frosts. Competition was detected by the second year when a higher growth of transplanted seedlings of the species Araucaria angustifolia occurred in plots where vegetation was present and fertilizer were applied. A lower growth rate was detected in plots where vegetation was present but fertilizer was not applied. These results indicate a balance between competition for soil nutrients and protection by neighbor vegetation. Damage of seedlings by leaf cutter ants was an important barrier for plant survival. Damage occurred in 80% of the A. angustifolia seedlings and 58% of these damaged seedlings died. The presence of neighboring vegetation tended to protect seedlings from ant damage. Although competition occurred, facilitation seems to be the main process driving early successional changes in this subtropical old field. This was mainly due to the improvement of local microclimatic conditions and protection against herbivores by neighboring vegetation. Facilitation occurred during establishment and growth phases in a subtropical area that is considered a productive, low stress environment. Our results indicate that facilitation may be more frequent in productive environments than previously thought.     

Effects of mixed cropping on population densities and parasitism rates of the diamondback moth, <Emphasis Type="Italic">Plutella xylostella</Emphasis> (Lepidoptera: Plutellidae)

Effects of mixed cropping and barrier crops on the population density and parasitism of the diamondback moth, Plutella xylostella (L.) (Lepidoptera: Plutellidae), were evaluated in field plots of cabbage grown in Bali, Indonesia. The densities of P. xylostella at larval and pupal stages, as well as the overall density at larval plus pupal stages, were significantly lower in cabbage/coriander mixed cropping subplots than in cabbage monoculture subplots. Parasitism of P. xylostella by the larval parasitoid Diadegma semiclausum (Hellen) (Hymenoptera: Ichneumonidae) was not significantly different between the mixed and monocultural cropping systems. These results do not support the so-called enemies hypothesis, but suggest that disruption of the host searching behavior of female moths by neighboring non-host plants is the mechanism behind the associational resistance observed in the coriander mixed cropping system. The inclusion of a Napier grass barrier between mixed crop and monoculture subplots did not affect the influence of mixed cropping on larval and pupal densities. Therefore, Napier grass, which is used locally as a fence for preventing livestock invasion of fields, would not obstruct the pest-reducing effect of coriander/cabbage mixed cropping.     

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

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