Testing the role of biotic stress in the stress gradient hypothesis. Processes and patterns in arid rangelands

Since many arid ecosystems are overstocked with domestic herbivores, biotic stress could have a stronger influence in modulating the balance of species interactions than expected from the stress gradient hypothesis (SGH). Here we tested a priori predictions about the effect of grazing on species interactions and fine scale spatial structure of grasses in water‐limited ecosystems. We used detailed vegetation mapping and spatial analysis, and performed a field experiment where the direct and indirect components of positive interactions were disentangled to provide evidence of links between process and pattern. We found associational resistance (biotic refuge) to be the dominant process in grazing situations, while competition, instead of direct facilitation, seemed to govern grass spatial patterns when herbivore pressure was relaxed. These results suggest that facilitation between grasses in arid communities may be related to herbivory rather than nurse plant effects. Associational resistance tends to have the strongest effect on spatial aggregation of species at intermediate grazing pressure. Results suggest that contrary to SGH, this physical clustering of species decreased when grazing pressure reached their maximum levels. Positive associations remained significant only when palatability differences between neighbours is large, suggesting that managing stocking rate is a key factor determining the persistence of herbivory refuges. These refuges are potential foci to initiate population recovery of high quality forage species in arid degraded areas.     

Responses of biotic interactions of dominant and subordinate species to decadal warming and simulated rotational grazing in Tibetan alpine meadow

Warming increases competition among plant species in alpine communities by ameliorating harsh environmental conditions, such as low temperatures. Grazing, as the main human activity, may mitigate the effect of warming, as previously reported. However, it is critical to refine the effects of warming on biotic interactions among species, for example, by taking the competitive ability of species into consideration. Based on a 10-year warming and grazing experiment in a Tibetan alpine meadow, we evaluated interspecific biotic interactions of dominant and subordinate species, using the approach of interspecific spatial associations. Warming significantly increased competition between subordinate and dominant species as well as among subordinate species, but not among dominant species. Moreover, facilitation of dominant-subordinate species also increased under warming. Simulated rotational grazing had similar effects to warming, with increasing interspecific competition. Our results show that, when studying the effects of warming on biotic interactions among species, it is necessary to characterize different species pairs relative to their competitive ability, and that simulated rotational grazing does not mitigate the effects of warming in the long term. Our results also provide evidence that the spatial pattern of species is a critical mechanism in species coexistence.     

Effects of human–livestock–wildlife interactions on habitat in an eastern Kenya rangeland

Human–livestock–wildlife interactions have increased in Kenyan rangelands in recent years, but few attempts have been made to evaluate their impact on the rangeland habitat. This study identified drivers of increased human–livestock–wildlife interactions in the Meru Conservation Area between 1980 and 2000 and their effects on the vegetation community structure. The drivers were habitat fragmentation, decline in pastoral grazing range, loss of wildlife dispersal areas and increase in livestock population density. Agricultural encroachment increased by over 76% in the western zone adjoining Nyambene ranges and the southern Tharaka area, substantially reducing the pastoral grazing range and wildlife dispersal areas. Livestock population increased by 41%, subjecting areas left for pastoral grazing in the northern dispersal area to prolonged heavy grazing that gave woody plant species a competitive edge over herbaceous life‐forms. Consequently, open wooded grassland, which was the dominant vegetation community in 1980, decreased by c. 40% as bushland vegetation increased by 42%. A substantial proportion of agro pastoralists were encountered around Kinna and Rapsu, areas that were predominantly occupied by pastoralists three decades ago, indicating a possible shift in land use in order to spread risks associated with habitat alterations.     

Grass-to-tree facilitation in an arid grazed environment (Aïr Mountains, Sahara)

The scarcity of tree regeneration is a major concern for the long-term conservation of tree cover in the Aïr-Ténéré Nature Reserve (Sahara, Niger), and the causes of this shortage are poorly understood. We raised the hypothesis that livestock pressure is a key-factor in the mortality of tree seedlings and that facilitation processes among plants may enhance tree regeneration. We tested whether the tussock grass Panicum turgidum could facilitate the regeneration of a keystone tree species, Acacia tortilis var. raddiana, under a grazing regime. We transplanted 3-month-old seedlings of Acacia to plots representing different neighbourhood conditions (in areas without vegetation, amongst intact tussocks, among cropped tussocks, and under branches), which were either fenced or exposed to grazing by large herbivores. Survival and height of 160 transplants were recorded during one year. Acacia performance was much lower in the grazed plots, indicating that large herbivores had a strong negative impact on tree regeneration. Presence of Panicum tussocks had a positive effect on the height of seedlings, likely because they improved soil properties and water availability. No aboveground positive effect on Acacia was detected though the grass did reduce atmospheric aridity. We detected an indirect positive effect of Panicum on Acacia survival in the presence of herbivores when aridity and grazing pressure reached their maximum levels. We obtained evidence that indirect facilitation may occur in arid grazed areas, although it has been predicted to mainly exist in mesic competitive environments. Additionally, our results support the stress-gradient hypothesis against assumptions that competitive effects may dominate in extremely arid environments. Since most of the positive effect on Acacia survival is indirect (biotic grazing refuge) due to the presence of herbivores, transplantation of tree seedlings amongst Panicum may be a simple and low-cost technique for the restoration of Saharan ecosystems degraded by herbivore grazing.     

The roles of stochasticity and biotic interactions in the spatial patterning of plant species in alpine communities

Questions

Plant community composition can be influenced by multiple biotic, abiotic, and stochastic factors acting on the local species pool to determine their establishment success and abundance and subsequently the diversity of the community. We asked if the influences of biotic interactions on the composition of plant species in communities, as indicated by patterns of plant species spatial associations (independent, positive or negative), vary across a productivity gradient within a single ecosystem type. Do dominant species of communities show spatial patterning suggestive of competitive interactions with interspecific neighbors? Do species that span multiple community types exhibit the same heterospecific interactions with neighbours in each community?

Location

Three alpine communities in the southern Rocky Mountains.

Methods

We measured the occurrence of species in a 1‐cm spatial grid within 2 m × 2 m plots to determine the spatial patterns of species pairs in the three communities. A null model of independent species spatial arrangements was used to determine whether species pairs were positively, negatively or independently associated, and how these patterns differed among the communities across the gradient of resource supply and environmental stress.

Results

Positive associations, indicative of facilitation between species, were most common in the most resource‐poor and least productive community. However negative associations, suggestive of competitive interactions among species, were not more common in the two more resource‐rich, productive communities. The dominant species of these communities did exhibit higher negative than positive associations with neighbours relative to positive patterning. Independent interspecific patterning was equally common relative to positive and negative patterns in all communities. Species that previously were shown to either facilitate other species or compete with neighbours exhibited spatial patterning consistent with the earlier experimental work.

Conclusions

A large number of species exhibit a lack of net biotic interactions, and stochastic factors appear to be as important as competition and facilitation in shaping the structure of the three alpine plant communities we studied.

     

Impacts of simulated livestock grazing on Utah prairie dogs (Cynomys parvidens) in a low productivity ecosystem

Allometric foraging theory suggests that herbivores of greatly differing size should co-exist through niche segregation, but a few studies of large–small herbivore foraging relationships have reported competitive interactions. This study addresses the potential roles of habitat productivity and large herbivore grazing intensities on large–small herbivore foraging interactions. We examined effects of different intensity simulated grazing treatments on forage abundance and quality for Utah prairie dogs (Cynomys parvidens) in a low productivity ecosystem, and consequent effects on prairie dog individual growth rates, foraging preferences, and activity budgets. We hypothesized that simulated grazing would have predominantly facilitative impacts on Utah prairie dogs, as was found for black-tailed prairie dogs in higher productivity ecosystems. To test this hypothesis, we measured the effects of simulated grazing on forage nitrogen, digestibility, and biomass. Simulated grazing increased average forage nitrogen and digestibility while decreasing forage biomass. These effects were associated with reduced individual growth rates, increased juvenile foraging time, and reduced juvenile vigilance. Results suggest that the negative effects of reduced vegetation biomass greatly outweighed positive treatment effects in this study. However, prairie dogs in the moderate intensity defoliation treatment showed some preference for “grazed” plots over “ungrazed” plots, and this preference increased with time. Our study lends support to the idea that habitat productivity and herbivore densities may mediate shifts between facilitative and competitive interactions between different-sized herbivores.     

Local‐scale spatial dynamics of ants in a temperate agroecosystem

At the local scale, spatial aggregations in ant distribution are often thought to be driven by competitive interactions among dominant ant species, although niche preferences and habitat heterogeneity might also lead to patchiness. Nevertheless, competitive interactions might be particularly important in agroecosystems that are structurally more homogeneous than natural habitats. The spatial patterns of ants in two Australian vineyards were investigated by intensive pitfall trapping to examine if non‐random patterns occur and whether these might be the result of competitive species interactions as well as the influence of woody vegetation adjacent to the vineyards. Null model analyses suggested competitive species interactions within ant assemblages that might have been driven by numerically dominant species, even though both positive and negative associations between these were found. Consistent spatial aggregations indicated significant spatial overlap in distributions of some species. Such overlap suggests that potential coexistence might be attributed to temporal partitioning or differences in foraging strategies. The presence of woody vegetation had a marked influence on ant assemblage structure and competitive interactions, and might facilitate coexistence by increasing resource heterogeneity. The implications of these findings for sampling strategies and ecological processes within vineyards are discussed.     

Nurse plants,tree saplings and grazing pressure: changes in facilitation along a biotic environmental gradient

Current conceptual models predict that an increase in stress shifts interactions between plants from competitive to facilitative; hence, facilitation is expected to gain in ecological importance with increasing stress. Little is known about how facilitative interactions between plants change with increasing biotic stress, such as that incurred by consumer pressure or herbivory (i.e. disturbance sensu Grime). In grazed ecosystems, the presence of unpalatable plants is reported to protect tree saplings against cattle grazing and enhance tree establishment. In accordance with current conceptual facilitation-stress models, we hypothesised a positive relationship between facilitation and grazing pressure. We tested this hypothesis in a field experiment in which tree saplings of four different species (deciduous Fagus sylvatica, Acer pseudoplatanus and coniferous Abies alba, Picea abies) were planted either inside or outside of the canopy of the spiny nurse shrub Rosa rubiginosa in enclosures differing in grazing pressure (low and high) and in exclosures. During one grazing season we followed the survival of the different tree saplings and the level of browsing on these; we also estimated browsing damage to the nurse shrubs. Shrub damage was highest at the higher grazing pressure. Correspondingly, browsing increased and survival decreased in saplings located inside the canopy of the shrubs at the high grazing pressure compared to the low grazing pressure. Saplings of both deciduous species showed a higher survival than the evergreens, while sapling browsing did not differ between species. The relative facilitation of sapling browsing and sapling survival – i.e. the difference between saplings inside and outside the shrub canopy – decreased at high grazing pressure as the facilitative species became less protective. Interestingly, these findings do not agree with current conceptual facilitation-stress models predicting increasing facilitation with abiotic stress. We used our results to design a conceptual model of facilitation along a biotic environmental gradient. Empirical studies are needed to test the applicability of this model. In conclusion, we suggest that current conceptual facilitation models should at least consider the possibility of decreasing facilitation at high levels of stress.     

Ecotypic differentiation determines the outcome of positive interactions in a dryland annual plant species

Positive interactions among plants have rarely been investigated with respect to their evolutionary consequences and vice versa. The outcome of facilitative interactions depends on the competitive ability and stress tolerance of the species. We tested whether this also applies to populations of conspecifics that are locally adapted to different environments and thereby differ in these traits. We hypothesised that ecotypes from less stressful environments experience a greater effect of facilitation when grown in stressful environments compared to populations adapted to these conditions.Seeds of two ecotypes of the annual grass species, Brachypodium distachyon, were collected from Mediterranean and arid origins and transplanted at an arid environment within the species’ distribution range. To examine the effect of biotic interactions on these ecotypes, we transplanted the individuals with and without the presence of the shrub Gymnocarpos decander (underneath or away from the shrub), and with and without the presence of annual vegetation (removal experiment). We examined the effect of these interactions on the two B. distachyon ecotypes by comparison of emergence success, biomass, and survival to reproduction.The presence of shrubs had a positive effect on all three variables in both ecotypes. Facilitation by shrubs enabled individuals from Mediterranean origin to grow and reproduce in arid conditions. Unlike the locals, they failed to survive to reproduction away from the shrubs, because of the markedly shorter growing season in open areas. The annual vegetation did not affect emergence or survival to reproduction in either ecotype; however, the positive effect of shrubs on biomass was reduced in the presence annual vegetation in the Mediterranean ecotype.This demonstrates that ecotypes adapted to arid conditions respond differently to these biotic interactions compared to Mediterranean populations. We argue that facilitation may have important evolutionary consequences by enabling maladapted ecotypes to invade and colonize stressful habitats.     

Community structure and positive interactions in constraining environments

Spatial patterns in plant communities are thought to be controlled by the interplay of species interactions and environmental constraints. To evaluate the role of plant–plant interactions in shaping these communities we quantified species co-occurrence and interaction in seven environmentally distinct communities. These included four different semiarid habitats in southeast Spain, one alpine system in the Sierra Nevada range (Spain), and two sites in Venezuela, a secondary savanna near Caracas (Altos de Pipe), and a sclerophyllous shrubland in the Gran Sabana plateau. We expected that facilitation would be stronger at sites with more spatial associations. The four semiarid sites in Spain and the shrubland in Gran Sabana showed a high degree of positive species associations. Of the other two communities, one showed both positive and negative associations while negative ones predominated in Altos de Pipe. The direct experimental measure of neighbors' effect showed that positive interactions among species prevailed in communities where positive species associations dominated. The appearance of benefactor species in patches increased species richness compared with the surrounding inter-shrub spaces. Our results provide a link between spatial patterns and species interactions, where aggregation points to positive interactions and segregation to competitive or interference effects. Facilitation appears as a relevant process shaping communities under environmental constraints.     

Pairwise null model analyses of temporal patterns of bird assemblages contradict the assumptions of competition theory

Compensatory dynamics assume inverse patterns of population dynamics of species with similar ecological resource requirements (temporal segregation). The objective of this study was to test this hypothesis on temporal samples (10–57 years) of 19 breeding bird assemblages of various habitats. We used presence/absence null model (SIM2) in combination with the C-score and Sørensen indices. The C-score index estimates the average number of checkerboards for two species, while the Sørensen index measures the qualitative similarity of co-occurrence between two species in a time series. We used pairwise null model analysis to select significant species pairs based on three selection criteria: the standard confidence interval criterion, conservative empirical Bayes mean based criterion and confidence limit based criterion. Altogether, 21 402 species pairs were analysed. The SIM2 algorithm detected from 157 to 7 segregated pairs depending on the selection criterion. The number of significant negative pairs with possible biological significance (foraging guild membership, predator–prey interactions) was far lower and represented approximately 0.0–0.3% (4–65) of pairs in a matrix. Indeed, the number of detected negative associations depended on the selection criterion. Moreover, the number of segregated pairs was negatively related to the area of the census plots and fill of the species matrix. Our results underline the minor importance of interspecific competitive interactions in temporal patterns of bird assemblages. Instead, we suggest that stochastic factors, climate or heterospecific social information may lead to more or less synchronous dynamics of bird pairs.     

Refuge effects of a cactus in grazed short‐grass steppe

Abstract. Question: Which factors influence the effectiveness of biotic refuges for harbouring grazing‐sensitive species in pastures with a long history of grazing by large herbivores? Previous research showed that spiny clumps of the cactus Opuntia polyacantha provided refuges from cattle grazing for plants and for inflorescence production on short‐grass steppe. In this paper, seven factors that may have a potential positive influence on the refuge effects of cactus at a landscape scale were assessed. Location: Short‐grass steppe of the Great Plains of North America. Methods: The study was conducted in eight long‐term grazed pastures and their respective ungrazed controls that were established 60 years ago. Results: Heavy grazing intensities were necessary for some positive effects of cactus to manifest, and some refuge effects changed to negative effects under lower grazing pressure. Refuge effects increased with plant community productivity due to greater abundances of grazing‐sensitive species, and greater grazing intensities in the more productive areas. Cover of cactus cladodes (spine‐covered pads) inside clumps appeared to be the main limiting factor for refuge effects, probably by limiting available space for grazing‐sensitive species in the clumps. Other factors such as size and density of cactus clumps, and the presence of large refuges in the proximity of clumps had minor influence on the effectiveness of cactus refuges. Conclusions: The effects of biotic refuges largely varied with ecological conditions and structural characteristics of the refuge. Refuge effects were mainly influenced by grazing intensity, plant community productivity, and structural characteristics of the biotic refuges. A conceptual model of factors influencing refuge effects at a local landscape scale in plant communities grazed by large herbivores is presented.     

Non-native competitive perennial grass impedes the spread of an invasive annual grass

Invasive plants are degrading wildlands around the globe by displacing native species, reducing biodiversity, and altering ecological functions. The current approach of applying herbicides to invasive plants in wildlands has not been effective at curtailing their expansion and, in certain circumstances, may do more harm than good. Preventing the spread of invasive species has been identified as an important strategy to protect wildlands. However, few prevention strategies have actually been tested. We hypothesized that establishing competitive vegetation next to infestations would increase the biotic resistance of the plant community to invasion and decrease the invasive species propagule pressure beyond the competitive vegetation. To evaluate this, we established twelve competitive vegetation barriers in front of invasive annual grass, Taeniatherum caput-medusae (L.) Nevski, infestations. The non-native perennial grass Agropyron desertorum (Fisch. ex Link) Schult. was seeded into plant communities adjacent to the infestations to create the competitive vegetation barriers. Soil nutrient concentrations and the spread of T. caput-medusae were compared between where A. desertorum was seeded and not seeded (control treatment) 3 years after treatment. Less T. caput-medusae and lower soil ammonium and potassium concentrations in the competitive vegetation barrier than control treatment (P ≤ 0.01) suggest that establishing competitive vegetation increased the biotic resistance of the plant communities to invasion. Taeniatherum caput-medusae cover and density in the plant communities protected by the competitive vegetation barrier (locales across the barriers from the infestations) were ~42- and 47-fold less, respectively, than unprotected plant communities (P < 0.01). This suggests that invasive plant propagule pressure was decreased in the plant communities protected by competitive vegetation barriers. The establishment of competitive vegetation around infestations may be an effective strategy to prevent or at least reduce the spread of invasive plant species.     

Do associations between native and invasive plants provide signals of invasive impacts?

Do invasive plant species act more as “passengers” or drivers of ecological change in native plant communities? Snapshot studies based on correlations at the site scale ignore longer-term dynamics and variation in how particular invaders affect particular native species. We analyzed patterns of co-occurrence between three invading species (Alliaria petiolata, Lonicera x bella, and Rhamnus cathartica) and 70 native plant species in 94 southern Wisconsin forests at two scales to test four hypotheses. Surveys at these sites in the 1950s and again in the 2000s allowed us to assess how initial plant diversity and site conditions affected subsequent patterns of invasion. Sites with more native species in the 1950s experienced fewer invasions of Lonicera and Rhamnus. However, this result may reflect the fact that more fragmented habitat patches supported both fewer species in the 1950s and more invasions. At the site-level, few negative correlations exist between invasive and native species’ abundances. Sites with higher Alliaria densities in the 2000s, however, support fewer native species and lower populations of several declining natives. Rhamnus-invaded sites support lower populations of two increasing species. Association (C-score) analyses detect more associations and more negative associations at the 1 m² scale than at the site scale. Most strong associations between invasive and increasing native species are positive while those with declining natives are often negative. Species restricted to specialized habitats rarely co-occur with invaders. Alliaria has more negative associations at fragmented sites where it is more abundant and invasions may be older. Fine-scale invasive-native associations were stronger, easier to detect, and less consistent than those detectable at the site-level. Thus, screening large numbers of local associations using observational data may allow us to identify particular invasive-native interactions worth further investigation. Although invading plants sometimes act as passive passengers, increasing in tandem with certain native plants in response to disturbed fragmented habitats, they may also contribute to the declines we observe in many native species. Monitoring invasions would allow us to assess whether local associations serve to predict subsequent invasive species impacts.     

Disentangling the roles of chance,abiotic factors and biotic interactions among epiphytic bryophyte communities in a tropical rainforest (Yunnan,China)

• Epiphytes offer an appealing framework to disentangle the contributions of chance, biotic and abiotic drivers of species distributions. In the context of the stress-gradient theory, we test the hypotheses that (i) deterministic (i.e., non-random) factors play an increasing role in communities from young to old trees, (ii) negative biotic interactions increase on older trees and towards the tree base, and (iii) positive interactions show the reverse pattern.
• Bryophyte species distributions and abiotic conditions were recorded on a 1.1 ha tropical rainforest canopy crane site. We analysed co-occurrence patterns in a niche modelling framework to disentangle the roles of chance, abiotic factors and putative biotic interactions among species pairs.
• 76% of species pairs resulted from chance. Abiotic factors explained 78% of non-randomly associated species pairs, and co-occurrences prevailed over non-coincidences in the remaining species pairs. Positive and negative interactions mostly involved species pairs from the same versus different communities (mosses versus liverworts) and life forms, respectively. There was an increase in randomly associated pairs from large to small trees. No increase in negative interactions from young to old trees or from the canopy to the base was observed.
• Our results suggest that epiphytic bryophyte community composition is primarily driven by environmental filtering, whose importance increases with niche complexity and diversity. Biotic interactions play a secondary role, with a very marginal contribution of competitive exclusion. Biotic interactions vary among communities (mosses versus liverworts) and life forms, facilitation prevailing among species from the same community and life form, and competition among species from different communities and life forms.

     

Does Extensive Grazing Benefit Butterflies in Coastal Dunes?

Grazing at low stocking rates has become a common management practice in nature restoration projects in the Netherlands. However, detailed knowledge of grazing impact is often poor, in particular for invertebrates. This study addressed the impact of extensive grazing on butterflies. Butterflies are critical indicators of habitat quality for many plant and animal species. We compared monitoring data from 1992 to 1996 for calcareous coastal dune areas in the Netherlands with different management: 11 grazed areas, 7 ungrazed areas and 4 areas managed by annual cutting. Grazing typically concerned year‐round grazing by cattle and/or ponies, at low stocking rates (0.05–0.26 head ha^?1 yr^?1). Butterfly abundance was related to species composition and structure of the vegetation. Changes in butterfly abundance were positive in grazed and ungrazed areas compared to cut areas. Species richness was not affected by management, but individual species differed in their response. Species from open grassland benefited most from grazing, particularly Issoria lathonia (Queen of Spain Fritillary) and Lycaena phlaeas (Small Copper). No clear negative effects of grazing were observed, but species occurrence was not always positively related to the environmental characteristics associated with grazing. In the long run, even lower stocking rates might prove more beneficial to the butterfly community as a whole. Four of the more frequently observed species, I. lathonia, Hipparchia semele (Grayling), Pyrgus malvae (Grizzled Skipper) and Aricia agestis (Brown Argus), are listed as threatened to susceptible in the Netherlands. All were apparently favored by grazing. It is concluded that extensive grazing has good potential to enhance butterfly diversity in restoration projects.     

Different patterns in root and soil fungal diversity drive plant productivity of the desert truffle Terfezia claveryi in plantation

The desert truffle Terfezia claveryi is one of the few mycorrhizal fungi currently in cultivation in semiarid and arid areas. Agroclimatic parameters seem to affect its annual yield, but there is no information on the influence of biotic factors. In this study, fungal diversity was analysed by high-throughput sequencing of the ITS2 rDNA region from soil and root samples to compare productive and non-productive mycorrhizal plants in a 4-years old plantation (Murcia, Spain). The fungal metaprofile was dominated by Ascomycota phylum. Desert truffle productivity was driven by different patterns of fungal species composition in soil (species replacement) and root (species richness differences). Moreover, positive associations for ectomycorrhizal and negative for arbuscular mycorrhizal guilds were found in productive roots, and positive associations for fungal parasite-plant pathogen guild in non-productive ones. Soil samples were dominated by pathotroph and saprotroph trophic modes, showing positive associations for Aureobasidium pullulans and Alternaria sp. in productive areas, and positive associations for Fusarium sp. and Mortierella sp. were found in non-productive soils. Finally, some significant OTUs were identified and associated to ascocarp producing patches, which could serve as predictive and location markers of desert truffle production.     

Floodplain ecosystem dynamics under extreme dry and wet phases in semi‐arid Australia

相似文献   

Small-scale species dynamics in semi-arid steppe vegetation in Inner Mongolia

Abstract. Small-scale species dynamics in semi-arid steppe vegetation under grazing and recovering from long-term grazing were followed during five years. Most species successively increased their frequency during the study period, whereas some early successional species decreased in frequency. Perennial Artemisia spp. dominated the vegetation and showed low mobility, whereas small annual or short-lived perennial species had a lower frequency and higher mobility. Significant associations between species were few.     

Switch between competition and facilitation within a seasonal scale at colony level in bryophytes

The relative importance of positive and negative interaction in species assemblages is thought to be dependent on the harshness of the physical environment. I studied the consistency of this prediction in a field experiment using growth of the target species Warnstorfia exannulata as influenced by the presence or absence of two adjacent species, Sphagnum warnstorfii and Scapania undulata. In particular, I focused on the mechanism by which colony–colony interactions occur, elucidating how the balance of positive and negative interactions changes along a water gradient. Because the natural fluctuations of the environment modify the water gradient, it was expected that the competitive hierarchies of the species would not remain consistent over time. Results indicated that the different hydrological properties of the colonies, thought to be the necessary condition for the appearance of species interactions, were not sufficient to explain the outcome of the species interactions. The switch from competition to facilitation under more stressful conditions was not confirmed along a water stress gradient. In addition, natural climatic fluctuations, by affecting the length of the water gradient, changed the competitive hierarchies of the species on a seasonal scale.