Animal collective movements are a key example of a system that links two clearly defined levels of organization: the individual and the group. Most models investigating collective movements have generated coherent collective behaviours without the inclusion of individual variability. However, new individual-based models, together with emerging empirical information, emphasize that within-group heterogeneity may strongly influence collective movement behaviour. Here we (i) review the empirical evidence for individual variation in animal collective movements, (ii) explore how theoretical investigations have represented individual heterogeneity when modelling collective movements and (iii) present a model to show how within-group heterogeneity influences the collective properties of a group. Our review underscores the need to consider variability at the level of the individual to improve our understanding of how individual decision rules lead to emergent movement patterns, and also to yield better quantitative predictions of collective behaviour.This article is part of the theme issue 'Collective movement ecology'.

Summary1. Network analysis, a branch of discrete mathematics that quantifies the structure of links between a set of nodes, is emerging as a powerful methodology to approach complex ecological interactions. 2. There has been a rapidly increasing body of research targeting the topological description of trophic interactions. Here we categorize studies applying a topological approach to empirical trophic networks, with the aim of identifying recent trends and deficiencies in this approach to analyse trophic interactions.3. There were biases in the taxonomic groups addressed and in the methodological approaches used for data collection and analyses. Studies on antagonistic interactions were generally focused on a single kind of bipartite interaction, and most studies compared network attributes across time, ecosystems or treatments. 4. We encourage a broader use of both interlinked and unipartite networks that would allow for describing indirect effects of trophic interactions, as well as time series of networks across seasons and phenological states of organisms. We also noted shortages of studies identifying interaction structures across levels of organization, on the correspondence between network structure and function, and particularly of studies on the behaviour of trophic networks in the face of environmental perturbations, which could provide guidance for preserving ecological interactions.

a b s t r a c tA large body of studies have linked sexual segregation in dimorphic ungulates with differential resource selection and nutritional requirements of the sexes. However, detailed patterns of sexual dietary preferences have rarely been assessed as keystone mechanisms of sexual segregation. We compared diet and nutritional selection by 3 sex-age classes in the Iberian red deer (Cervus elaphus hispanicus Hilzheimer) across reproductive states and according to seasonal fluctuations in resource availability in a Mediterranean environment. Foraging segregation between sexes was found during rut when female/juvenile selection of shrubs was higher than that of males. Observed foraging patterns relate to a stronger selection of nitrogen, tannins and lignin by females and juveniles, and fibres by males. Our findings are associated to the Mediterranean climate, where rut and lactation concur with a shortage period, the particularly dry summer. Foraging segregation between sexes during the rut could be shaped by a conjunction of factors such as the low quality of resources and different fitness enhancement strategies. We highlight both the importance of including the dietary component and providing a temporal framework when documenting ungulate sexual segregation, and the interest of considering regional conditions when addressing management of ungulates with a wide distribution.

Ungulates select diets with high energy, protein, and sodium contents. However, it is scarcely known the influence of essential minerals other than Na in diet preferences. Moreover, almost no information is available about the possible influence of toxic levels of essential minerals on avoidance of certain plant species. The aim of this research was to test the relative importance of mineral content of plants in diet selection by red deer (Cervus elaphus) in an annual basis. We determined mineral, protein and ash content in 35 common Mediterranean plant species (the most common ones in the study area). These plant species were previously classified as preferred and non-preferred. We found that deer preferred plants with low contents of Ca, Mg, K, P, S, Cu, Sr and Zn. The model obtained was greatly accurate identifying the preferred plant species (91.3% of correct assignments). After a detailed analysis of these minerals (considering deficiencies and toxicity levels both in preferred and non-preferred plants) we suggest that the avoidance of excessive sulphur in diet (i.e., selection for plants with low sulphur content) seems to override the maximization for other nutrients. Low sulphur content seems to be a forgotten factor with certain relevance for explaining diet selection in deer. Recent studies in livestock support this conclusion, which is highlighted here for the first time in diet selection by a wild large herbivore. Our results suggest that future studies should also take into account the toxicity levels of minerals as potential drivers of preferences.

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The ecological effects of consumer guilds are strongly driven by the relative resource use of included species. Here we use stable isotopes in faecal samples from 6 cooccurring mammalian herbivores in an African savanna to identify species-specific trophic niches and detect patterns of interactions between herbivores and their feeding resources. Species-specific foraging strategies were reflected in the faecal  13 C values, with species aligning along both a browser-grazer gradient and a niche breadth gradient. Within the broad foraging strategies outlined by  13 C,  15 N indicated a seasonal shift in resource use for some herbivores. We expected that 13 C isotope niches would overlap in a nested fashion, with the niches of grazers and browsers being included in those of mixed feeders, and that 15 N niches would separate into discrete modules reflecting leguminous and protein content of respective diets. However, the observed structure was more complex, with combined modular and nested patterns of interactions between herbivores and 13 C isotopes. We identified an isolated module comprising eland and its exclusive δ 13C values, and a series of nested structures with a set of specialist herbivores (hartebeest and sable) which δ 13 C values were nested within those of more generalist herbivores (impala, waterbuck and zebra). Networks based on δ 15 N, however, reflected a higher level of overlap in resource use with random patterns in herbivore interactions with resources, and only a significant modular interaction pattern during the dry season. We suggest that the combined modular and nested pattern of δ 13 C interactions reflect the simultaneous mutualistic and antagonistic characteristics of plant-herbivore interactions. We argue that such interaction patterns could stabilize ecosystems by constraining effects of perturbations to specific modules and by increasing functional redundancy through nested interactions.

Context Supplemental feeding of large mammalian herbivores is a common management tool mainly aimed at promoting healthy populations and at increasing productivity and trophy sizes. Such management measure may indirectly affect herbivore effects on plant communities through altered foraging patterns. The quantification of the ecological effects of large herbivore management is important for designing holistic management and conservation programs. Aims Here we aimed at quantifying the ecological effects of supplemental feeding of Iberian red deer, Cervus elaphus hispanicus, on the composition of and on the browsing effects on Mediterranean woody plant community. Methods An experiment was set up in a hunting rangeland located in central Spain, where female deer were kept in enclosures with either exclusive access to natural forages or with additional ad libitum access to a nutritionally rich concentrate. The experiment also included a control area where deer were absent. Key results We observed significant differences in browsing impacts among the supplemented, non-supplemented and control areas, and such effect varied for the different plant species. Plant species which nutritional content complemented that of fodder were more highly consumed, for instance, Erica spp., which digestible fibre content is higher and N content lower than that of provided fodder. The presence of deer and the concentrate supplied, instead, did not influence the relative abundances of shrub species. Conclusions Artificial supplemental feeding provided to ungulates led to increased browsing on plant species which nutritional composition complemented that of the supplement provided. Implications So as to alleviate herbivory impact on all shrubs, we suggest that composition of supplemental feeding should adjust both to the natural forage availability and quality and to ungulate requirements across seasons.

We report evidence of hierarchical resource selection by large herbivores and plant neighbouring effects in a Mediterranean ecosystem. Plant palatability was assessed according to herbivore foraging decisions. We hypothesize that under natural conditions large herbivores follow a hierarchical foraging pattern, starting at the landscape scale, and then selecting patches and individual plants. A between- and within-patch selection study was carried out in an area formed by scrubland and pasture patches, connected by habitat edges. With regard to between-patch selection, quality-dependent resource selection is reported: herbivores mainly consume pasture in spring and woody plants in winter. Within-patch selection was also observed in scrub habitats, influenced by season, relative patch palatability and edge effect. We defined a Proximity Index (PI) between palatable and unpalatable plants, which allowed verification of neighbouring effects. In spring, when the preferred food resource (i.e. herbs) is abundant, we observed that in habitat edges large herbivores basically select the relatively scarce palatable shrubs, whereas inside scrubland, unpalatable shrub consumption was related to increasing PI. In winter, a very different picture was observed; there was low consumption of palatable species surrounded by unpalatable species in habitat edges, where the latter were more abundant. These outcomes could be explained though different plant associations described in the literature. We conclude that optimal foraging theory provides a conceptual framework behind the observed interactions between plants and large herbivores in Mediterranean ecosystems.