Species richness changes across two trophic levels simultaneously affect prey and consumer biomass

Increasing species richness of primary producers or consumers is proposed to increase primary and secondary production; however, the consequences of biodiversity change across trophic levels has been poorly investigated. We used a controlled marine microbial system to investigate the effects of simultaneous changes in biodiversity of consumer and prey species. Consumer (ciliates) and prey (algae) richness and identity were manipulated independently in a complete factorial design. The results showed clear biodiversity effects of both consumers and prey, within and across trophic levels. We found reduced prey and increased consumer biomass with increased consumer richness, with the most diverse prey assemblage supporting the highest biomass of consumers at the highest richness of consumers. Increasing prey richness did not increase resistance to consumption when consumers were present. Instead, our results indicated enhanced energy transfer with simultaneous increasing richness of consumers and prey.     

Trait means,trait plasticity and trait differences to other species jointly explain species performances in grasslands of varying diversity

Functional traits may help to explain the great variety of species performances in plant communities, but it is not clear whether the magnitude of trait values of a focal species or trait differences to co‐occurring species are key for trait‐based predictions. In addition, trait expression within species is often plastic, but this variation has been widely neglected in trait‐based analyses. We studied functional traits and plant biomass of 59 species in 66 experimental grassland mixtures of varying species richness (Jena Experiment). We related mean species performances (species biomass and relative yield RY) and their plasticities along the diversity gradient to trait‐based pedictors involving mean species traits (T_mean), trait plasticities along the diversity gradient (T_slope), extents of trait variation across communities (T_CV; coefficient of variation) and hierarchical differences (T_diff) and trait distances (absolute values of trait differences T_dist) between focal and co‐occurring species. T_mean (30–55%) and T_diff (30–33%) explained most variation in mean species performances and their plasticities, but T_slope (20–25%) was also important in explaining mean species performances. The mean species traits and the trait differences between focal species and neighbors with the greatest explanatory power were related to plant size and stature (shoot length, mass:height ratios) and leaf photosynthetic capacity (specific leaf area, stable carbon isotopes and leaf nitrogen concentration). The contribution of trait plasticities in explaining species performances varied in direction (positive or negative) and involved traits related to photosynthetic capacity, nitrogen acquisition (nitrogen concentrations and stable isotopes) as well as structural stability (shoot carbon concentrations). Our results suggest that incorporating plasticity in trait expression as well as trait differences to co‐occurring species is critical for extending trait‐based analyses to understand the assembly of plant communities and the contribution of individual species in structuring plant communities.     

Environmental predictors of global parrot (Aves: Psittaciformes) species richness and phylogenetic diversity

Aim Spatial patterns of phylogenetic diversity (PD) aid our ability to discern diversification rate mechanisms underlying hypotheses for the large‐scale distribution of biodiversity. We develop a predictive framework for the way in which spatial patterns of PD vary with those of species richness, depending on the balance between speciation and extinction rates. Within this framework, diversification processes thought to underlie the productive energy, ambient energy, topographic variability and habitat variety hypotheses predict that gradients of increase in species richness will be associated with: (1) decreasing extinction rates where driven by productive energy, hence increasing relative PD (i.e. PD controlling for species richness, or PD_rel); (2) a similar positive relationship between ambient energy and PD_rel; (3) increasing speciation rates where driven by topographic variability, hence decreasing PD_rel; and (4) no consistent relationship between PD_rel and habitat variety when driven by the latter. We test these predictions using distributional data on parrots. Location Neotropical, Afrotropical, Indo‐Malayan and Australasian realms. Methods Spatial models were used to test the predictions. Results Globally, a positive association between productive energy and PD_rel confirms prediction (1). However, within realms, hump‐shaped relationships suggest the importance of decreasing extinction rates up to a threshold level of productive energy, and the increasing importance of speciation rates thereafter. Ambient energy is positively associated with PD_rel in Australasia, Indo‐Malaya, and globally, supporting prediction (2). However, this is driven by the coincidence of highest PD_rel in areas of high ambient energy and intermediate productive energy (i.e. in seasonal tropical environments), which may be characterized by relatively low speciation and extinction rates. In the Neotropics, increasing topographic variability is associated with decreasing PD_rel and increasing species richness, suggesting an increasing gradient of speciation, supporting prediction (3). Elsewhere, the signal of this mechanism may be obscured by collinearities with energy gradients. The lack of an overall relationship between habitat diversity and PD_rel confirms prediction (4). Main conclusions Spatial patterns of PD_rel in relation to environmental gradients may be sensitive to collinearities among those gradients. Nevertheless, patterns emerge which have implications for the relative importance of speciation and extinction processes in generating latitudinal diversity gradients.     

Resource stoichiometry and consumers control the biodiversity-productivity relationship in pelagic metacommunities

Recent theory suggests that both biodiversity and productivity are constrained by resource supply rates and ratios and that resource stoichiometry is the key to understanding the relationship between biodiversity and productivity. We experimentally tested this theory using pelagic metacommunities. We amended existing predictions by explicitly considering evenness as an aspect of biodiversity and including control of algal biomass by consumption in addition to competition. The metacommunities received a different phosphorus (P) supply and the three patches within each metacommunity differed in their nitrogen (N) supply, which created different N∶P ratios (2, 16, and 128). All patches were inoculated with a phytoplankton assemblage consisting of five species, and half of the metacommunities received two ciliate species as consumers. At the level of the entire metacommunity, algal biomass increased with increasing P supply, whereas species richness and evenness decreased with increasing P supply. Without consumers, resource use efficiency (RUE; realized biomass per unit of P) increased with increasing richness and evenness. Consumer presence reduced overall biomass and richness and precluded a correlation between RUE and biodiversity. At the patch level, local evenness correlated with higher RUE at both imbalanced N∶P ratios (2 and 128) but not at a balanced N∶P ratio. In conclusion, overall P supply constrained realized biomass and altered diversity, whereas resource stoichiometry shaped the relationship between biodiversity and RUE.     

Effects of temporal and spatial heterogeneities created by consumer-driven nutrient recycling on algal diversity

A spatially explicit plant-herbivore model composed of planktonic herbivores, algal preys and nutrients was constructed to examine the effects of consumer-driven nutrient recycling (CNR) on the algal species richness with and without spatial structure. The model assumed that either of two essential nutrients (N and P) limited growth of algal populations and that consumer individuals moved randomly in the lattice and grazed all the algal species with the same efficiency. The results showed that when there was no CNR, the number of persistent algal species was affected by neither supply rates of external nutrients nor spatial structure and was consistently low. When consumers recycled nutrients according to their stoichiometry, the algal species richness changed with supply rates of external nutrients depending on spatial structure: the algal species richness decreased with increasing nutrient loadings when there were no spatial structure because CNR increased the probability of stochastic extinction of algal species by amplifying the oscillation of algae-consumer dynamics. However, when spatial structures were created by the migration of consumers, CNR increased the algal species richness in a range of nutrient loadings because spatial variation of grazing pressure functioned to stabilize the algal-consumer dynamics. The present study suggests that through grazing and nutrient recycling, consumer individuals can create ephemeral heterogeneity in growth environments for algal species and that this ephemerality is one of the keys to understanding algal species in nature.     

Population and community variability in randomly fluctuating environments

The prediction that environmental fluctuations may destabilise populations and yet stabilise aggregate community properties has remained largely untested. We examined population and community stability under constant and fluctuating temperatures in simple planktonic assemblages of differing algal richness. Temperature dependent resource competition produced a highly asymmetric community structure where algal community biomass was dominated by one species. For a given level of species richness, temperature fluctuations induced lower community covariance and thus stabilised community biomass. However, increasing algal species richness increased the variability of population abundance and growth rates, as well as population and community variability. Consumer dynamics were directly destabilised by environmental fluctuations. These results confirm recent theoretical studies suggesting a stabilising effect of environmental fluctuations at the community level. However, they also support the theoretical prediction that increasing species richness may be of limited value for community stability, most especially in asymmetric communities, when competition directly affects population variability.     

Temperature mediates competitive exclusion and diversity in benthic microalgae under different N:P stoichiometry

The dependence of competitive interactions on abiotic conditions is attracting increasing interest in the face of globally rising temperatures and altered biogeochemical cycles of major nutrients. In a microcosm experiment involving a natural inoculum of benthic microalgae, temperature and nutrient supply ratios were manipulated in order to test three main hypotheses: (1) temperature and nutrient supply ratios determine species composition and diversity of the assemblage, (2) the identity of the dominating species depends on nutrient supply and temperature, and (3) higher temperature leads to faster competitive exclusion and thus more rapid decline in species richness. Over a period of 7 weeks, algal biomass reached an equilibrium carrying capacity, with was higher at colder temperatures and intermediate N:P supply ratios (N:P = 16). Initial growth rate increased with temperature and under high P-supply. Species richness in the stationary phase of the experiment decreased with increasing temperature, reflecting a higher extinction rate in the warmer treatments, which were also characterized by higher dominance of single species. Thus, increasing temperature both altered the identity of the dominating species and accelerated competitive displacement. This experiment thus indicates that warming might influence outcome and temporal dynamics in species interactions, and thereby eventually local diversity.     

Development of ice biota in a temperate sea area (Gulf of Bothnia)

A study of sea ice biota was carried out in the Gulf of Bothnia (northern Baltic Sea) during the winter of 1989–1990. Samples (ice cores) were taken at a coastal station at regular time intervals during the ice season. Chlorophyll a concentration, algal species distribution, bacterial numbers, and primary and bacterial production were measured. Colonization of the ice began in January when daylight was low. As the available light increased, the algae started to grow exponentially. The vertical chlorophyll a distribution changed and algal species composition and biomass changed during the season. During the initial and middle phase of colonization, ice-specific diatoms, Nitzschia frigida and Navicula pelagica, dominated the algal biomass. Nutrients (PO₄ ^3– and NO_3j) were found to be depleted during the time of algal exponential growth. The maximum algal biomass exceeded 800 g C 1^–1. The primary production supplied food for heterotrophic organisms. The presence of heterotrophic organisms of different trophic levels (bacteria, flagellates, ciliates and rotifers) indicated an active microbial food web.     

中国蚂蚁丰富度地理分布格局及其与环境因子的关系

物种丰富度分布格局及其形成机制的研究对于生物多样性保护具有重要意义。为了了解中国蚂蚁物种丰富度分布格局,利用中国省级尺度蚂蚁物种分布数据和环境信息,结合GIS和数理统计方法,探讨蚂蚁物种丰富度的地理分布格局与环境因子之间的关系。研究结果表明:(1)蚂蚁丰富度随纬度增加呈逐渐递减趋势,但缺乏显著的经度梯度。丰富度最高的地区主要集中在南方省份,我国北方、西北干旱区和青藏高原北部地区丰富度较低;(2)简单线性回归分析表明,能量、水分和季节性因素中,影响蚂蚁物种丰富度最强的因子分别为最冷月均温(TEMmin)(R2adj=0.532)、年均降水量(PREC)(R2adj=0.376)和年温度变化范围(TEMvar)(R2adj=0.539),而单个生境异质性因子对蚂蚁物种丰富度的影响均不显著;(3)最优模型由年均温(TEM)、海拔变化范围(ELErange)和年温度变化范围(TEMvar)组成,能够解释68.4%的蚂蚁丰富度地理分异。鉴于海拔变化范围更多地反映与温度相关的生境异质性,因此温度是限制中国蚂蚁分布的最重要因素。另外,分析结果还表明,海南、贵州、江西、四川、安徽和山西等6省蚂蚁区系调查最不充分,是未来发现蚂蚁新分布的热点地区。     

物种丰富度对集群生物量的影响及抽样、互补效应的检验

通过8种甲藻构建不同物种丰富度梯度的集群进行室内一次性培养实验,探讨了集群生物量随物种丰富度的增长模式,运用多因素方差分析检验集群内是否存在抽样效应. 运用超产分析、相对产量之和分析和子集分析方法,检验了集群内的生态位互补效应,并运用分离加性方程法定量分离了抽样与互补效应.结果表明: 集群生物量随物种丰富度呈饱和型上升趋势;集群内存在较强的互补效应;在指数生长期,某些特定物种对集群生物量有显著影响,但在生长稳定期,集群生物量不受个别物种出现与否的影响;在生长稳定期,抽样效应总体上为负,而互补效应与净生物多样性效应则总体为正.     

A Compositional and Structural Analysis of Phytoplankton in Lake Khurpatal, (U.P.), India

The phytoplankton community in Lake Khurpatal (Kumaun Himalya), India, was analyzed in relation to physico-chemical variables. Phytoplankton biomass ranged from 2.7 to 20.0 g m⁻³ in the euphotic zone of the lake. Dinoflagellates monopolized the algal community with a mean annual contribution of 94.5 % to the total phytoplankton biomass. The community coefficient used to quantify the seasonal changes in algal population reflected the fact that successional episodes were not very significant, the percentage similarity among the consecutive algal samples ranging from 47.8 to 89.9 %. The phytoplankton community was characterized by low species richness, low equitability and species diversity. Various ecological characteristics of the community are discussed and the phytoplankton biomass is also compared with that of some tropical and temperate lakes.     

浮游植物生物量增长模式及抽样、互补效应检验

选取我国近海18种常见浮游植物,以不同物种构建集群,试验研究不同浮游植物集群生物量随物种丰富度的增长模式,运用多因素方差分析检验集群的抽样效应,并分别采用超产分析、相对产量之和分析、子集分析法,检验集群内的生态位互补效应.结果表明:集群生物量随物种丰富度的增长模式不是单一的,当物种数<5时,集群生物量随物种丰富度增加而增大,当物种数>5时,集群生物量与物种丰富度之间无明显规律性;集群内存在较强的互补效应,随物种丰富度的增加呈近似"钟型"曲线变化;集群在生长稳定期存在抽样效应.     

Prey diversity and prey identity affect herbivore performance on different time scales in a long term aquatic food‐web experiment

The question whether and how diversity‐mediated productivity at the base of food‐webs influences adjacent trophic levels is still unclear. Experiments revealed negative effects on consumers due to the increasing dominance of inedible species under grazing pressure, and positive effects due to a greater variety of prey resources. We experimentally investigate two more hypotheses, which have not been addressed in detail so far: first, more diverse primary producer communities potentially use limiting resources more efficiently, and are, therefore, more productive. This effect can be considered functionally similar to a direct enrichment with limiting resources, potentially resulting in a higher stochastic risk of herbivore extinction (‘paradox of enrichment’). Second, in a stable environment, enclosed primary producer communities should evolve towards a ‘climax state’, eventually dominated by one or few prey species. Therefore, long‐term diversity effects in producer communities should more likely result from the specific traits of the dominating species, than from complementarity. To address these hypotheses, we conducted long‐term laboratory experiments, exposing the freshwater grazer Daphnia magna to a gradient of algal species richness (1, 2, 4 or 8 edible chlorophyte species). The experiments were run in batch cultures, without exchange of growth medium after the start of the experiment. Six parameters related to Daphnia population demography, biomass accrual, and stability were followed and determined over a period of up to 263 days. Producer diversity exhibited strong positive effects on the short‐term performance of grazers (first reproduction, first population peak), and on grazer mean standing stocks. However, herbivore long‐term dynamics (day of extinction and temporal stability) depended on prey species identity, namely the presence of Chlamydomonas reinhardtii. Our experiments suggest that both prey diversity and identity can have positive effects on consumer performance, but act on different time scales.     

Low temperature constrains growth rates but not short-term ingestion rates of Antarctic ciliates

Low environmental temperature is a major factor affecting the feeding activities, growth rates, and growth efficiencies of metazooplankton, but these features are poorly characterized for most protistan species. Laboratory experiments were conducted to examine the growth and ingestion rates of cultured herbivorous Antarctic ciliates. Three ciliates fed several algal species individually at 0 °C exhibited uniformly low growth rates (<0.26 day^?1), but the algae varied substantially in their ability to support ciliate growth. Specific ingestion rate (prey biomass consumed per unit ciliate biomass per unit time) was strongly affected by ciliate physiological state (starved vs. actively growing). Starved cells ingested many more prey than cells in balanced growth during short-term (minutes-to-hours) experiment but did not grow faster, indicating temperature compensation of ingestion rate but not growth rate. Field experiments were also conducted in the Ross Sea, Antarctica, to characterize the feeding rates of ciliates in natural plankton assemblages. Specific ingestion rates of two dominant ciliates were an order of magnitude lower than rates reported for temperate ciliates, but estimated rates were strongly affected by prey abundance. Our data indicate that short-term ingestion rates of Antarctic ciliates were not constrained by low environmental temperature although overall growth rates were, indicating the need for caution when designing experiments to measure the ingestion rates of these species at low environmental temperature. We present evidence that artifacts arising from estimating ingestion in short-term experiments may lead to errors in estimating feeding impact and growth efficiencies that are particularly large for polar protists.     

Does ambient substrate composition influence consumer diversity effects on algal removal?

Benthic substrates constitute an important habitat template for aquatic communities and may affect the contributions of benthic organisms to ecological processes. To test the effects of ambient substrate composition on the process of algae accrual and removal, we conducted an experiment to examine how substrate type influenced consumer richness effects. We hypothesized that algal removal from focal substrates (ceramic tiles) would be influenced by the surrounding ambient substrate through its effect on nutrient cycling and subsequent algal growth. We manipulated consumer richness in mesocosms at one or three species while holding consumer biomass constant. Aquatic consumers were an amphipod, a snail, and a water boatman, and ambient substrates were either sand or gravel. After 21 days, ambient substrate influenced epilithic algal accrual on tiles, affected physio-chemical parameters within mesocosms, and modified consumer behavior. Chlorophyll a was approximately 2× greater on control tiles surrounded by sand, and FPOM and turbidity were greater on sand than gravel when consumers were present. Substrate modified consumer behavior such that consumers congregated around focal substrates in sand, but dispersed around them in gravel. Consumers also had substrate-specific influences on epilithic chlorophyll, causing a decrease in sand and an increase in gravel. Algal assemblages on focal tiles were dominated by diatoms, and their composition responded to consumer richness and identity, but not substrate. Our data suggest that direct effects (e.g., consumptive removal of epilithon from focal tiles) were more pronounced in sand, whereas indirect effects (e.g., bioturbation and enhanced mixing) promoted algal accrual in gravel. These results show that algae production on exposed surfaces may change as underlying substrate composition changes, and that substrate type can alter consumer diversity effects on algal removal.     

Consumer and resource diversity effects on marine macroalgal decomposition

Few manipulative experiments that explicitly test the relationship between biodiversity and ecosystem function have focussed on regenerative processes such as decomposition and nutrient cycling. Of the studies that have taken place, most have concentrated on the effects of leaf litter diversity rather than the effects of consumer diversity on decomposition. In the present study, we established an in‐situ mesocosm experiment on an intertidal mudflat in the Ythan Estuary, Scotland, to investigate the interactive effects of consumer diversity, resource diversity and microbial activity on algal consumption and decomposition. We assembled communities of three commonly occurring macrofaunal species (Hediste diversicolor, Hydrobia ulvae and Littorina littorea) in monoculture and all possible combinations of two and three species mixtures and supplied them with single vs two‐species mixtures of the algae Fucus spiralis and Ulva intestinalis. Further, we also investigated whether algal decomposition changes nutrient remineralisation within the sediment by determining the C:N ratio of the surficial sediment. Data were analysed using extended linear regression with generalized least squares estimation to characterise the variance structure. We found that consumer species diversity effects are best explained by compositional effects and that species richness per se may not be the single most important determinant of resource use and decomposition in this community. Algal identity and invertebrate identity effects underpin the observed response and reflect species‐specific traits associated with algal consumption and processing. The role of the microbial community is comparatively weak, but strongly linked to faunal activities and behaviour. The C:N ratio of the sediment increased with consumer species richness, indicating increased mineralisation in more diverse communities. Overall, our results suggest that although consumer species richness effects per se are weak, decomposition and subsequent incorporation of resources is nevertheless dependent on the composition of the decomposer community, which, in turn, has important implications for biogeochemical nutrient cycling in marine coastal habitats.     

Effects of stream predator richness on the prey community and ecosystem attributes

It is important to understand the role that different predators can have to be able to predict how changes in the predator assemblage may affect the prey community and ecosystem attributes. We tested the effects of different stream predators on macroinvertebrates and ecosystem attributes, in terms of benthic algal biomass and accumulation of detritus, in artificial stream channels. Predator richness was manipulated from zero to three predators, using two fish and one crayfish species, while density was kept equal (n = 6) in all treatments with predators. Predators differed in their foraging strategies (benthic vs. drift feeding fish and omnivorous crayfish) but had overlapping food preferences. We found effects of both predator species richness and identity, but the direction of effects differed depending on the response variable. While there was no effect on macroinvertebrate biomass, diversity of predatory macroinvertebrates decreased with increasing predator species richness, which suggests complementarity between predators for this functional feeding group. Moreover, the accumulation of detritus was affected by both predator species richness and predator identity. Increasing predator species richness decreased detritus accumulation and presence of the benthic fish resulted in the lowest amounts of detritus. Predator identity (the benthic fish), but not predator species richness had a positive effect on benthic algal biomass. Furthermore, the results indicate indirect negative effects between the two ecosystem attributes, with a negative correlation between the amount of detritus and algal biomass. Hence, interactions between different predators directly affected stream community structure, while predator identity had the strongest impact on ecosystem attributes.     

Disentangling the roles of spatial and environmental variables in shaping benthic algal assemblages in rivers of central and northern China

Benthic algae were collected from central and northern Chinese rivers to test the hypothesis that geographic location has significant contributions in shaping algal assemblages. We used Moran’s eigenvector maps (MEM) to model spatial components and variation partitioning to quantify the influences of spatial and environmental variables on regional patterns of algal richness and community composition, respectively. We found that variation in algal richness was attributed to MEM component 2, 8, and 9 and the quadratic term of N–NO₃. Regarding abundance data, latitude, longitude, and MEM component 1, 2, and 7 were important spatial variables. Although P–PO₄, pH, and annual mean temperature were significant environmental variables influencing algal community composition, they were all spatially structured. Among the total explained variance in both algal metrics, spatial proportions were higher than that of environmental variables. We also found that abundant species of Achnanthidium minutissimum, Cocconeis placentula, Cymbella delicatula, Cymbella affinis, Cymbella turgidula, and Synedra ulna displayed clear spatially related patterns. In conclusion, the contributions of spatial and environmental variables to regional variation of algal assemblages are scale-dependent. As for our study scale (~1,000 km), spatial control may be more important. Since spatial effects could obscure local environmental impacts on algal communities, appropriate study scale and statistical methods should be taken into account in algal bioassessment. We recommend inclusion of both algal richness and community composition in study of algal biogeography, due to their different relationships with spatial and environmental variables.     

Effects of additional illumination and fertilization on seasonal changes in fine-scale grassland community structure

Abstract. Fine-scale structure of a species-rich grassland was examined for seasonal changes caused by manipulated changes in the availability of above and below-ground resources (additional illumination with the help of mirrors and fertilization) in a field experiment. If changes induced by fertilization — which are expected to lead to a reduction in small-scale diversity —are due to intensified light competition, they should be compensated for by additional light input. Permanent plots of 40 cm × 40 cm were sampled by the point quadrat method at three angles (60°, 90° and 120° from the horizontal North-South direction), using a laser beam to position the quadrats, in early July and early September. The applied treatments did not cause apparent changes in plant leaf orientation. The degree of spatial aggregation of biomass increased seasonally in fertilized, non-illuminated plots: greater productivity at a constant light supply led to a faster growth rate of potentially dominant species, as compared to the subordinate ones. Additional illumination mitigated this effect of fertilization, indicating that the observed changes in biomass aggregation were due to increased light competition. There was a considerable seasonal decrease of variance ratio (ratio of observed variance of richness at a point and variance expected at random) in fertilized only and in illuminated only plots. In fertilized plots this was due to the positive relationship between biomass aggregation and expected variance of richness. Biomass constancy occurs to be inversely related to deficit in variance of richness. In illuminated plots, in contrast, only the observed variance of richness decreased seasonally, indicating a more uniform use of space by different species. Evidently, a deficit in variance of richness can be caused by drastically different processes, showing that the variance ratio statistic may not have a significant explanatory value in fine-scale community studies.     

Abundance, species richness and energy availability in the North American avifauna

Aim To determine how species richness, abundance, biomass, energy use and mean number of individuals per species scale with environmental energy availability in wintering and breeding avian assemblages, and to contrast assemblages of (i) common and rare species and (ii) breeding residents and migrants. To assess whether such patterns are compatible with the ‘more individuals hypothesis’ (MIH) that high‐energy areas are species‐rich because they support larger populations that are buffered against extinction. Location The North American continent (latitudinal range 23.4 °?48.1 °N; longitudinal range 124.2°?68.7° W). Methods Avian species richness, abundance, biomass and energy use were calculated for 295 Resident Bird Count plots. Environmental energy availability was measured using ambient temperature and the Normalized Difference Vegetation Index (NDVI), a close correlate of plant productivity. Analyses took plot area into account, and were conducted (with and without taking habitat type into account) using general linear models and spatial mixed models. Results Positive species–energy relationships were exhibited by both wintering and breeding assemblages, but were stronger in the former. The structure of winter assemblages responded more strongly to temperature than NDVI, while breeding assemblages tended to respond more strongly to NDVI. Breeding residents responded to annual measures of energy availability while breeding migrants and the winter assemblage responded more strongly to seasonal measures. In the winter assemblage, rare and common species exhibited species–energy relationships of a similar strength, but common breeding species exhibited a much stronger relationship than rare breeding species. In both breeding and wintering assemblages, abundance, biomass and energy use increased with energy availability and species richness. Energy availability was a poor predictor of the mean number of individuals per species. Main conclusions The nature of the species–energy relationship varies seasonally and with the manner in which energy availability is measured. Our data suggest that residents are less able to respond to seasonal fluxes in resource availability than long‐distance migrants. Increasing species richness and energy availability is associated with increasing numbers of individuals, biomass and energy use. While these observations are compatible with the MIH our data provide only equivocal support for this hypothesis, as the rarest species do not exhibit the strongest species–energy relationships.