Seasonal changes in mixed‐species bird flocks and antipredator information

Animals acquire information produced by other species to reduce uncertainty and avoid predators. Mixed‐species flocks (MSFs) of birds are ubiquitous in forest ecosystems and structured, in part, around interspecific information transfer, with “nuclear” species providing information that other species eavesdrop on. We hypothesized that in a seasonal tropical forest, the amount of information produced by birds about predation would be dynamic and particularly would decrease inside MSFs when the nuclear species leave MSFs to breed. We obtained baseline information on MSF encounter rate and species composition along established sampling routes over 9 months near the Sino‐Vietnamese border. We also conducted three experiments to quantify information produced by different species in response to typical predator encounters, including a moving predator stimulus presented inside of MSFs, and a stationary predator model presented both inside and outside of MSFs. MSFs were much less frequent in the breeding season with fewer individuals of the nuclear species, David's Fulvetta (Alcippe davidi), participating, though the diversity of other species remained stable. Fulvettas were the dominant producer of alarm‐related information both to the moving and stationary stimuli in MSFs and were also among the most active mobbers to stimuli presented outside of MSFs. In the breeding season, they tended to call less to the moving stimulus, and substantially fewer individuals responded to the in‐flock stationary stimulus. Other species increased their own information production at stationary predator stimuli (inside and outside of MSFs) during the breeding season, perhaps due to their increased investment in offspring during this time. Yet even during the breeding season, David's Fulvetta remained the highest producer of information about predators in MSFs. Hence, while we show that information production in MSFs can be somewhat dynamic, we describe a continually asymmetric communication system, in which a nuclear species is important to the whole community.     

Nest survival is influenced by parental behaviour and heterospecifics in a mixed‐species colony

Studies of avian nest success often focus on examining influences of variation in environmental and seasonal factors. However, in‐depth evaluations can also incorporate variation in individual incubation behaviour to further advance our understanding of avian reproductive ecology. We examined these relationships in colonially nesting Black‐crowned Night‐Herons Nycticorax nycticorax using intensive video‐monitoring methods to quantify incubation behaviours. We modelled nest survival as a function of both extrinsic factors and incubation behaviours over a 3‐year period (2010–12) on Alcatraz Island, USA. Model‐averaged parameter estimates indicated that nest survival increased as a function of greater incubation constancy (% of time spent incubating eggs within a 24‐h period), and average daily precipitation throughout the nesting stage. Common Ravens Corvus corax are the only known nest predator of Night‐Herons on Alcatraz Island, as on many other coastal Pacific islands. We also investigated the effects of heterospecific nesting of California Gulls Larus californicus and Western Gulls Larus occidentalis in a mixed‐species colony with Night‐Herons, based on nesting proximity data collected over a 2‐year period (2011–12). This second analysis indicated that, in addition to incubation behaviours, nesting heterospecifics are an important factor for explaining variation in Night‐Heron nest survival. However, contrary to our original expectation, we found that Night‐Herons experienced increased nest survival with increasing distance from gull colony boundaries. These results may apply to other areas with multiple colonial nesting species and similar predator communities and climatic patterns.     

Composition of mixed‐species flocks and shifts in foraging location of flocking species on Hainan Island,China

A total of 134 bird species were recorded at Jianfengling, Hainan Island, in China from May 2000 to September 2004, of which 44 participated in one or more of 134 mixed‐species flocks. These flocks averaged 3.8 ± 0.2 species and 20.3 ± 1.2 individuals. Flocking propensity in a given species ranged from 1.5 to 100%. For flocking species, frequency of flocking and number of individuals in flocks was positively correlated with frequency and number in point counts. Among all species pairs with flocking frequency above 5%, cluster and correlation analysis indicated there were two principal groups of flocking birds – canopy species and understorey species: associations were positive within a group, but negative between groups. Canopy birds had a higher flocking propensity than understorey birds. They also made significantly less use of inner branches and trunks and greater use of middle branches, and foraged at a significantly greater height when in mixed‐species flocks than when solitary. For understorey bird species, there were no significant differences in foraging locations between solitary and mixed‐species flocks. Higher flocking frequency occurred in the wet season for canopy birds, but in the dry season for understorey birds. Overall patterns were consistent with the explanation that flocking enables an expansion of foraging niche by reducing the risk of predation.     

Quantifying climate–growth relationships at the stand level in a mature mixed‐species conifer forest

A range of environmental factors regulate tree growth; however, climate is generally thought to most strongly influence year‐to‐year variability in growth. Numerous dendrochronological (tree‐ring) studies have identified climate factors that influence year‐to‐year variability in growth for given tree species and location. However, traditional dendrochronology methods have limitations that prevent them from adequately assessing stand‐level (as opposed to species‐level) growth. We argue that stand‐level growth analyses provide a more meaningful assessment of forest response to climate fluctuations, as well as the management options that may be employed to sustain forest productivity. Working in a mature, mixed‐species stand at the Howland Research Forest of central Maine, USA, we used two alternatives to traditional dendrochronological analyses by (1) selecting trees for coring using a stratified (by size and species), random sampling method that ensures a representative sample of the stand, and (2) converting ring widths to biomass increments, which once summed, produced a representation of stand‐level growth, while maintaining species identities or canopy position if needed. We then tested the relative influence of seasonal climate variables on year‐to‐year variability in the biomass increment using generalized least squares regression, while accounting for temporal autocorrelation. Our results indicate that stand‐level growth responded most strongly to previous summer and current spring climate variables, resulting from a combination of individualistic climate responses occurring at the species‐ and canopy‐position level. Our climate models were better fit to stand‐level biomass increment than to species‐level or canopy‐position summaries. The relative growth responses (i.e., percent change) predicted from the most influential climate variables indicate stand‐level growth varies less from to year‐to‐year than species‐level or canopy‐position growth responses. By assessing stand‐level growth response to climate, we provide an alternative perspective on climate–growth relationships of forests, improving our understanding of forest growth dynamics under a fluctuating climate.     

Characterization and cross‐species amplification of microsatellite loci in a Cyprinodon species flock

In this study, two new microsatellite loci from a library of Cyprinodon beltrani from Laguna Chichancanab, Mexico, were characterized. Additionally, cross‐species amplification with primer pairs developed for other Cyprinodon species was performed. The 11 markers show moderate to high levels of polymorphism (two to 33 alleles) in six species of the Cyprinodon flock from Laguna Chichancanab and in their sister species Cyprinodon artifrons. These loci were characterized for population genetic study to detect gene flow between the different endemic pupfish species.     

Reforestation with native mixed‐species plantings in a temperate continental climate effectively sequesters and stabilizes carbon within decades

Reforestation has large potential for mitigating climate change through carbon sequestration. Native mixed‐species plantings have a higher potential to reverse biodiversity loss than do plantations of production species, but there are few data on their capacity to store carbon. A chronosequence (5–45 years) of 36 native mixed‐species plantings, paired with adjacent pastures, was measured to investigate changes to stocks among C pools following reforestation of agricultural land in the medium rainfall zone (400–800 mm yr^?1) of temperate Australia. These mixed‐species plantings accumulated 3.09 ± 0.85 t C ha^?1 yr^?1 in aboveground biomass and 0.18 ± 0.05 t C ha^?1 yr^?1 in plant litter, reaching amounts comparable to those measured in remnant woodlands by 20 years and 36 years after reforestation respectively. Soil C was slower to increase, with increases seen only after 45 years, at which time stocks had not reached the amounts found in remnant woodlands. The amount of trees (tree density and basal area) was positively associated with the accumulation of carbon in aboveground biomass and litter. In contrast, changes to soil C were most strongly related to the productivity of the location (a forest productivity index and soil N content in the adjacent pasture). At 30 years, native mixed‐species plantings had increased the stability of soil C stocks, with higher amounts of recalcitrant C and higher C : N ratios than their adjacent pastures. Reforestation with native mixed‐species plantings did not significantly change the availability of macronutrients (N, K, Ca, Mg, P, and S) or micronutrients (Fe, B, Mn, Zn, and Cu), content of plant toxins (Al, Si), acidity, or salinity (Na, electrical conductivity) in the soil. In this medium rainfall area, native mixed‐species plantings provided comparable rates of C sequestration to local production species, with the probable additional benefit of providing better quality habitat for native biota. These results demonstrate that reforestation using native mixed‐species plantings is an effective alternative for carbon sequestration to standard monocultures of production species in medium rainfall areas of temperate continental climates, where they can effectively store C, convert C into stable pools and provide greater benefits for biodiversity.     

Role of mixed‐species flocks in the use of adjacent savannas by forest birds in the central Cerrado,Brazil

Abstract: We examined the role of mixed‐species flocks for forest birds during their breeding and non‐breeding seasons in the use of savannas adjacent to forests in central Cerrado, Brazil. Transect surveys (n = 64) were conducted in eight savanna patches. Distances of birds from forests were estimated. Recorded birds were classified as members or not of mixed‐species flocks. About half of the bird species recorded in savannas were found in at least one mixed‐species flock. As distance from the forest increased, the number of species in mixed‐species flocks tended not to vary, while the number of species foraging alone or in mono‐specific groups decreased. Thus, for some forest species, participation in mixed‐species flocks allowed a greater use of more distant savannas. This tendency of being in mixed‐species flocks at greater distances from forests also can be interpreted as a reluctance to forage alone or in mono‐specific groups due to higher predation risk in less protective vegetation distant from cover. There was strong seasonal variation in the participation of bird species in mixed‐species flocks. There were significantly more species in mixed‐species flocks than out of these associations in the non‐breeding season, while differences in the breeding season were not significant. These patterns occurred, in part because mixed‐species flocks tended to be more frequent, to have more species and to forage at greater distances from forests during the early non‐breeding season than in other periods. This study suggests that the formation of mixed‐species flocks plays an important role in promoting the use of adjacent savannas by forest birds at forest/savanna boundaries in Cerrado. It also pointed out a novel advantage gained by birds with participation in mixed‐species flocks – greater use of adjacent vegetation patches.     

Vocal Mimicry of Alarm‐Associated Sounds by a Drongo Elicits Flee and Mobbing Responses from Other Species that Participate in Mixed‐Species Bird Flocks

A growing number of studies have shown that vocal mimicry appears to be adaptive for some bird species, although the exact function of this behaviour varies among species. Previous work has looked at the function of the vocal mimicry of non‐alarm sounds by the Greater Racket‐tailed Drongo (Dicurus paradiseus). But drongos also imitate sounds associated with danger, such as predators' vocalisations or the mobbing‐specific vocalisations of other prey species, raising the question of whether the function of mimicry can vary even within a species. In a playback experiment, we compared the effect on other species of different drongo vocalisations including: (1) predator mimicry, (2) mobbing mimicry, (3) drongo species‐specific alarms, (4) drongo species‐specific non‐alarms and (5) a control (barbet) sound. Both mobbing mimicry and drongo species‐specific alarms elicited flee responses from the most numerous species in the flocks, the Orange‐billed Babbler (Turdoides rufescens). Mobbing mimicry also elicited mobbing responses from the Orange‐billed Babbler and from another gregarious babbler, the Ashy‐headed Laughingthrush (Garrulax cinereifrons); when responses from both species were considered together, they were elicited at a significantly higher level by mobbing mimicry than by the barbet control, and a level that tended to be higher (0.07 < p < 0.10) than the response to drongo‐specific alarms. Predator mimicry elicited flee and mobbing responses at an intermediary level. Our results support the hypotheses that mobbing mimicry is a specific category of mimicry that helps attract the aid of heterospecifics during mobbing and that alarm mimicry can in some cases be beneficial to the caller.     

Unchanged carbon balance driven by equivalent responses of production and respiration to climate change in a mixed‐grass prairie

Responses of grassland carbon (C) cycling to climate change and land use remain a major uncertainty in model prediction of future climate. To explore the impacts of global change on ecosystem C fluxes and the consequent changes in C storage, we have conducted a field experiment with warming (+3 °C), altered precipitation (doubled and halved), and annual clipping at the end of growing seasons in a mixed‐grass prairie in Oklahoma, USA, from 2009 to 2013. Results showed that although ecosystem respiration (ER) and gross primary production (GPP) negatively responded to warming, net ecosystem exchange of CO₂ (NEE) did not significantly change under warming. Doubled precipitation stimulated and halved precipitation suppressed ER and GPP equivalently, with the net outcome being unchanged in NEE. These results indicate that warming and altered precipitation do not necessarily have profound impacts on ecosystem C storage. In addition, we found that clipping enhanced NEE due to a stronger positive response of GPP compared to ER, indicating that clipping could potentially be an effective land practice that could increase C storage. No significant interactions between warming, altered precipitation, and clipping were observed. Meanwhile, we found that belowground net primary production (BNPP) in general was sensitive to climate change and land use though no significant changes were found in NPP across treatments. Moreover, negative correlations of the ER/GPP ratio with soil temperature and moisture did not differ across treatments, highlighting the roles of abiotic factors in mediating ecosystem C fluxes in this grassland. Importantly, our results suggest that belowground C cycling (e.g., BNPP) could respond to climate change with no alterations in ecosystem C storage in the same period.     

Abundant recent activity of retrovirus‐like retrotransposons within and among flycatcher species implies a rich source of structural variation in songbird genomes

Transposable elements (TEs) are genomic parasites capable of inserting virtually anywhere in the host genome, with manifold consequences for gene expression, DNA methylation and genomic stability. Notably, they can contribute to phenotypic variation and hence be associated with, for example, local adaptation and speciation. However, some organisms such as birds have been widely noted for the low densities of TEs in their genomes and this has been attributed to a potential dearth in transposition during their evolution. Here, we show that avian evolution witnessed diverse and abundant transposition on very recent timescales. First, we made an in‐depth repeat annotation of the collared flycatcher genome, including identification of 23 new, retrovirus‐like LTR retrotransposon families. Then, using whole‐genome resequencing data from 200 Ficedula flycatchers, we detected 11,888 polymorphic TE insertions (TE presence/absence variations, TEVs) that segregated within and among species. The density of TEVs was one every 1.5–2.5 Mb per individual, with heterozygosities of 0.12–0.16. The majority of TEVs belonged to some 10 different LTR families, most of which are specific to the flycatcher lineage. TEVs were validated by tracing the segregation of hundreds of TEVs across a three‐generation pedigree of collared flycatchers and also by their utility as markers recapitulating the phylogenetic relationships among flycatcher species. Our results suggest frequent germline invasions of songbird genomes by novel retroviruses as a rich source of structural variation, which may have had underappreciated phenotypic consequences for the diversification of this species‐rich group of birds.     

Ecosystem fluxes of hydrogen in a mid‐latitude forest driven by soil microorganisms and plants

Molecular hydrogen (H₂) is an atmospheric trace gas with a large microbe‐mediated soil sink, yet cycling of this compound throughout ecosystems is poorly understood. Measurements of the sources and sinks of H₂ in various ecosystems are sparse, resulting in large uncertainties in the global H₂ budget. Constraining the H₂ cycle is critical to understanding its role in atmospheric chemistry and climate. We measured H₂ fluxes at high frequency in a temperate mixed deciduous forest for 15 months using a tower‐based flux‐gradient approach to determine both the soil‐atmosphere and the net ecosystem flux of H₂. We found that Harvard Forest is a net H₂ sink (?1.4 ± 1.1 kg H₂ ha^?1) with soils as the dominant H₂ sink (?2.0 ± 1.0 kg H₂ ha^?1) and aboveground canopy emissions as the dominant H₂ source (+0.6 ± 0.8 kg H₂ ha^?1). Aboveground emissions of H₂ were an unexpected and substantial component of the ecosystem H₂ flux, reducing net ecosystem uptake by 30% of that calculated from soil uptake alone. Soil uptake was highly seasonal (July maximum, February minimum), positively correlated with soil temperature and negatively correlated with environmental variables relevant to diffusion into soils (i.e., soil moisture, snow depth, snow density). Soil microbial H₂ uptake was correlated with rhizosphere respiration rates (r = 0.8, P < 0.001), and H₂ metabolism yielded up to 2% of the energy gleaned by microbes from carbon substrate respiration. Here, we elucidate key processes controlling the biosphere–atmosphere exchange of H₂ and raise new questions regarding the role of aboveground biomass as a source of atmospheric H₂ and mechanisms linking soil H₂ and carbon cycling. Results from this study should be incorporated into modeling efforts to predict the response of the H₂ soil sink to changes in anthropogenic H₂ emissions and shifting soil conditions with climate and land‐use change.     

The shifting phenological landscape: Within‐ and between‐species variation in leaf emergence in a mixed‐deciduous woodland

Many organisms rely on synchronizing the timing of their life‐history events with those of other trophic levels—known as phenological matching—for survival or successful reproduction. In temperate deciduous forests, the extent of matching with the budburst date of key tree species is of particular relevance for many herbivorous insects and, in turn, insectivorous birds. In order to understand the ecological and evolutionary forces operating in these systems, we require knowledge of the factors influencing leaf emergence of tree communities. However, little is known about how phenology at the level of individual trees varies across landscapes, or how consistent this spatial variation is between different tree species. Here, we use field observations, collected over 2 years, to characterize within‐ and between‐species differences in spring phenology for 825 trees of six species (Quercus robur, Fraxinus excelsior, Fagus sylvatica, Betula pendula, Corylus avellana, and Acer pseudoplatanus) in a 385‐ha woodland. We explore environmental predictors of individual variation in budburst date and bud development rate and establish how these phenological traits vary over space. Trees of all species showed markedly consistent individual differences in their budburst timing. Bud development rate also varied considerably between individuals and was repeatable in oak, beech, and sycamore. We identified multiple predictors of budburst date including altitude, local temperature, and soil type, but none were universal across species. Furthermore, we found no evidence for interspecific covariance of phenology over space within the woodland. These analyses suggest that phenological landscapes are highly complex, varying over small spatial scales both within and between species. Such spatial variation in vegetation phenology is likely to influence patterns of selection on phenology within populations of consumers. Knowledge of the factors shaping the phenological environments experienced by animals is therefore likely to be key in understanding how these evolutionary processes operate.     

Relationship between heatwave‐induced forest die‐off and climatic suitability in multiple tree species

In recent decades, many forest die‐off events have been reported in relation to climate‐change‐induced episodes, such as droughts and heat waves. To understand how these extreme climatic events induce forest die‐off, it is important to find a tool to standardize the climatic conditions experienced by different populations during a specific climatic event, taking into account the historic climatic conditions of the site where these populations live (bioclimatic niche). In this study, we used estimates of climatic suitability calculated from species distribution models (SDMs) for such purpose. We studied forest die‐off across France during the 2003 heatwave that affected Western Europe, using 2,943 forest inventory plots dominated by 14 single tree species. Die‐off severity was estimated by Normalized Difference Vegetation Index (NDVI) loss using Moderate‐resolution Imaging Spectroradiometer remote sensor imagery. Climatic suitability at the local level during the historical 1979–2002 period (HCS), the episode time (2003; ECS) and suitability deviance during the historical period (HCS‐SD) were calculated for each species by means of boosted regression tree models using the CHELSA climate database and occurrences extracted from European forest inventories. Low HCS‐SD and high mean annual temperature explained the overall regional pattern of vulnerability to die‐off across different monospecific forests. The combination of high historical and low episode climatic suitability also contributed significantly to overall forest die‐off. Furthermore, we observed different species‐specific relationships between die‐off vulnerability and climatic suitability: Sub‐Mediterranean and Mediterranean species tended to be vulnerable in historically more suitable localities (high HCS), whereas Euro‐Siberian species presented greater vulnerability when the hot drought episode was more intense. We demonstrated that at regional scale, past climatic legacy plays an important role in explaining NDVI loss during the episode. Moreover, we demonstrated that SDMs‐derived indexes, such as HCS, ECS and HCS‐SD, could constitute a tool for standardizing the ways that populations and species experience climatic variability across time and space.     

The adult body size variation of Dryas iulia (Lepidoptera,Nymphalidae, Heliconiinae) in different populations is more influenced by temperature variation than by host‐plant availability during the seasons

The body size of insects is affected by environmental conditions during development and can present considerable intraspecific variations, which can be seen as an ultimate consequence/adaptation to environmental conditions. This paper evaluated whether the body size of the butterfly Dryas iulia from subtropical populations was influenced by changing climate conditions and food source availability during the seasons. The likely reasons behind body size variation were also investigated. First, field data on body size variation, host‐plant availability and climate fluctuation throughout the seasons were recorded. Then, the effects of host‐plant species and temperature on body size were analyzed by controlled experiments. Field data revealed that body size and host‐plant availability varied significantly through the seasons. Populations had the smallest body size during the spring and the biggest size during summer, whereas host‐plant availability was lower during winter and higher during spring. The controlled experiments revealed that both temperature and host‐plant had significant effect on the plasticity of body size. Larvae subjected to winter temperature treatment led to smaller butterflies when compared to immatures reared under summer temperature treatment, and larvae fed with Passiflora misera produced bigger adults when compared to larvae reared on Passiflora suberosa. The combination of data gathered in the field and in the laboratory suggests that seasonal body size variation in D. iulia is related mainly to differences in the temperatures to which larvae are subjected during development, while host‐plant shifts caused by differential availability of food through the seasons had slightly effects on the variation observed.     

American marten and fisher do not segregate in space and time during winter in a mixed‐forest system

Understanding the mechanisms of coexistence between ecologically similar species is an important issue in ecology. Carnivore coexistence may be facilitated by spatial segregation, temporal avoidance, and differential habitat selection. American martens Martes americana and fishers Pekania pennanti are medium‐sized mustelids that occur sympatrically across portions of North America, yet mechanisms of coexistence between the two species are not fully understood. We assessed spatial and temporal partitioning in martens and fishers in the Upper Peninsula of Michigan, USA, using camera trap data collected during winter 2013–2015. To investigate spatial segregation, we used a dynamic occupancy model to estimate species’ occupancy probabilities and probabilities of persistence and colonization as a function of covariates and yearly occupancy probability for the other species. Temporal segregation was assessed by estimating diel activity overlap between species. We found weak evidence of spatial or temporal niche partitioning of martens and fishers. There was high overlap in forest cover selection, and both marten and fisher occupancy were positively correlated with deciduous forests (excluding aspen [Populus tremuloides]). There was strong temporal overlap (; CI = 0.79–0.82) with both species exhibiting largely crepuscular activity patterns. Co‐occurrence of martens and fishers appears to be facilitated by mechanisms not investigated in this study, such as partitioning of snow features or diet. Our results add additional insights into resource partitioning of mesocarnivores, but further research is required to enhance our understanding of mechanisms that facilitate marten and fisher coexistence.     

Seed survival on experimental dishes in a central European old‐growth mixed‐species forest – effects of predator guilds,tree masting and small mammal population dynamics

Predation of tree seeds can be a major factor structuring plant communities. We present a three year study on tree seed survival on experimental dishes in an old‐growth forest in central Europe in Austria. We addressed species specific, spatial and temporal aspects of post‐dispersal seed predation. Seeds of Norway spruce Picea abies, European beech Fagus sylvatica, and silver fir Abies alba were exposed on dishes in different types of exclosures which allowed access only to specific guilds of seed predators. Removal experiments were carried out in two old‐growth forests and a managed forest (macro‐sites), including micro‐sites with and without cover of ground vegetation. We conducted the experiment in three consecutive years with a mast year of beech and spruce before the first year of the study. The seed removal experiments were combined with live trapping of small mammals being potential seed predators. Our experiments showed a distinctly different impact of different predator guilds on seed survival on the dishes with highest removal rates of seeds from dishes accessible for small mammals. We observed differing preferences of small mammals for the different tree species. Seed survival in different macro‐ and micro‐habitats were highly variable with lower seed survival in old growth forests. In contrast to our assumption, and in contrast to the satiation hypothesis which assumes higher seed survival in and directly after mast years, seed survival was lower in the year following the mast year of beech when a population peak of small mammals occurred and higher in intermast periods when subsequently small mammal population crashed. This suggests a higher importance of sporadic masting shortly after mast years in intermast periods for establishment of forest trees provided that pollination efficiency is high enough in such years. Combined with the high seed mortality observed after the mast year, this corroborates the important role of seed predation for forest dynamics. An altered synchrony or asynchrony of masting of different tree species and changed masting frequencies through climate change may thus lead to strong and non‐linear effects on forest dynamics.     

Nutrient limitation and nutrient‐driven shifts in plant species composition in a species‐rich fen meadow

Question: We studied the development and persistence of the effects of nutrient pulses on biomass production and species composition in a fen meadow. Location: Nature reserve, central Netherlands, 5 m a.s.l. Methods: Single pulse fertilization with N and P in a factorial design on an undrained central and a drained margin site in a species‐rich fen meadow (Cirsio dissecti‐Molinietum). Biomass production and species composition were monitored during four years. Results: At the central site, N addition boosted biomass production, but only during one year. The species composition was not changed. P fertilization increased the biomass production and changed the species composition from a vegetation dominated by Carex panicea to a grassland community with abundant Holcus lanatus, but not before the second year. At the margin site, P fertilization changed the species composition in a similar way, but biomass production was not increased. N fertilization had no effect. At both sites the P induced shift in species composition persisted for four years although the P effect declined during the experiment. Conclusions: The biomass responses show that N was limiting in the central site. Another nutrient, besides N and P (probably K) must have been limiting in the marginal site. The fast decline of the N effect on biomass is ascribed to increased denitrification and biomass removal. The delay in the P effect on biomass and species composition and the persistence of the P effect on species composition are ascribed to fast immobilisation and subsequent slow release of fertilizer P in the peat soil. Recurrence of the P pulses is expected to cause permanent changes in species composition.