Salvador Rebollo

European rabbits as ecosystem engineers: warrens increase lizard density and diversity

Mammals that build extensive open burrow systems are often classified as ecosystem engineers, since they have the potential to modulate the availability of resources for themselves and other organisms. Lizards may benefit from the heterogeneity created by these structures, especially if coupled with an increased offer of sites for refuge and thermoregulation. However, information about these engineering effects by burrowing animals is scarce. We investigated the influence of European rabbit burrows on several parameters of a Mediterranean lizard community (abundance, density, diversity and body condition) in three different habitats (open pastures, holm oak and scrub patches). We found that lizards were positively associated with burrows, and that burrows determined lizard presence at otherwise unfavourable habitats. Moreover, community parameters such as density and species richness were higher in sites with burrows. Burrows influenced lizard species in different ways, and were also relevant for other Mediterranean vertebrates, as revealed by questionnaires to experts. We also explored the possible resources provided by burrows for lizards. Warrens offer relatively abundant prey and appropriate retreat sites for refuge and thermoregulation. Warrens may have further implications within the ecosystem, acting as stepping stones, allowing lizards to reach otherwise inaccessible habitat patches. This study shows that European rabbit warrens have a positive influence on lizard density and diversity, and confirms the role of rabbits as ecosystem engineers. This reinforces the need for appropriate conservation measures for rabbits, especially given their threatened status in the Iberian Peninsula. Furthermore, our study highlights that taking into account the influence of engineering activities increases our awareness of species interactions, and may translate into more adequate conservation measures for the preservation of biodiversity.

Role of disturbance in maintaining a savanna-like pattern in Mediterranean Retama sphaerocarpa shrubland

Can the interaction of episodic and chronic distur- bances explain the maintenance of savanna-like patterns? We explored the morphological and spatial patterns of the leguminous shrub Retama sphaerocarpa in a Mediterranean environment in relation to disturbance. Various morphological variables of R. sphaerocarpa shrubland were found to be deter- mined by a combination of two types of disturbance: (1) mechanical cutting: an episodic, heavy, short-term distur- bance of anthropogenic origin for management purposes, and (2) herbivore activity primarily by rabbits: generally a chronic, more lenient, long-term disturbance. The intensities of these two types of disturbance were not correlated. Me- chanical cutting effects on R. sphaerocarpa shrubland mor- phology predominated quantitatively over herbivore effects. Herbivores generally produced open shrubland with fewer, more scattered, thicker branched, l arger R. sphaerocarpa shrubs. In contrast, intense sprouting after cutting produced a higher density of smaller R. sphaerocarpa shrubs with denser aerial biomass. However, heavy herbivory in abnormally dry peri- ods produced some effects similar to those of mechanical cutting. The size of R. sphaerocarpa shrubs was positively related to seed production. Thus, the means of propagation depended upon the type of disturbance: episodic disturbances resulted in intense sprouting, whereas chronic herbivore activ- ity resulted in the formation of thick branches that produced a large number of seeds. The combination of these two distur- bances determine, in part, space occupancy patterns of domi- nant woody species in this Mediterranean landscape and simi- lar savanna-type ecosystems. Investigations of environmental constraints on vegetation distribution and abundance should take into account the historical role of herbivores in shaping present systems.

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.

Effects of Land use on Nocturnal Birds in a Mediterranean Agricultural Landscape

Abstract. Knowledge on the effects of land use on community composition and species abundance is crucial for designing realistic conservation strategies, particularly in highly dynamic systems such as Mediterranean agricultural mosaics that are subjected to intensive cultivation. We investigated these effects on the nocturnal bird species occurring in the study area (Stone Curlew Burhinus oedicnemus, Red-necked Nightjar Caprimulgus ruflcollis, Barn Owl Tyto alba, Eurasian Scops Owl Otus scops, Little Owl Athene noctua, Tawny Owl Strix aluco, Long-eared Owl Asio otus, Short-eared Owl Asio flammeus and Eagle Owl Bubo bubo) across an agricultural-natural habitat mosaic in Central Spain for three consecutive years. Shares of vineyards, scrubland, herbaceous cropland, water bodies, and roads significantly affected the composition of the nocturnal bird community. Herbaceous cropland and olive groves, which covered 50% of the study area, proved to be neutral for all species. Remnant patches of natural and semi-natural scrubland (around 10% of the study area) and water bodies (only 1.5% of the study area) showed a positive effect on Eagle Owls, Eurasian Scops Owls, Long-eared Owls, and Red-necked Nightjars. Vineyard (35% of the study area) had a negative influence on Eagle Owls, Long-eared Owls, and Eurasian Scops Owls. Our results indicate, first, that the relative extent of land use types was apparently not related with the presence of nocturnal bird species and, second, that natural scrublands and water bodies are key habitats for assuring the persistence of nocturnal birds in agricultural Mediterranean landscapes. Current land planning focused toward land use intensification will likely increase the areas of habitats that are neutral or have adverse effects on nocturnal birds.

Vole mound effects and disturbance rate in a mediterranean plant community under different grazing and irrigation regimes

A factorial field experiment was used to assess the influence of soil-disturber mammals in the structure of a 9-year-old Mediterranean annual plant community subjected to different sheep grazing and irrigation regimes. We estimated the disturbance rate (mound building activity) by Mediterranean voles, their effects on vegetation and the mechanisms of these effects during a period of vole outbreak. The effects on vegetation were analysed at the levels of species, functional groups and plant community. Disturbance rate was high and voles can disturb the entire soil surface once every four or five years. The availability of certain trophic resources (perennial plants) appeared to drive vole expansion in the experimental plots and it was independent of the irrigation and grazing treatments. Mound building activities largely affected vegetation but conserved plot differences. Total vegetation cover, absolute cover of all functional groups, mean vegetation height and species richness were less on mounds than on undisturbed ground. These effects did not change the relative abundance of annuals, perennials, grasses and forbs. Only the relative abundance of small-seeded species decreased on mounds. As the proportion of these seeds was similar in both types of patches, we suggest that small-seeded species had more difficulties for germinating or emerging when they are buried during mound formation. Irrigation and sheep grazing promoted large changes in the vegetation parameters but these effects were, in general, similar on mounds and undisturbed ground. Our results show that the availability of germinable seeds may be the major limitation for mound revegetation, probably due to the scarcity of seeds existing at the depths from which soils are excavated. Our results also suggested a resource limitation on mounds. The results provide additional evidence that soil disturbances by small herbivore mammals exert relevant ecological effects on abandoned Mediterranean croplands. We discuss the ecological implications of vole mound-building activities for plant succession, plant species conservation and forage resource availability for livestock.

Higher reproductive success of small males and greater recruitment of large females may explain strong reversed sexual dimorphism (RSD) in the northern goshawk

Reversed sexual dimorphism (RSD), which occurs when the female of a species is larger than the male, is the rule for most birds of prey but the exception among other bird and mammal species. The selective pressures that favour RSD are an intriguing issue in animal ecology. Despite the large number of hypotheses proposed to explain the evolution of RSD, there is still no consensus about the mechanisms involved and whether they act on one or both sexes, mainly because few intrapopulation studies have been undertaken and few raptor species have been investigated. Using the strongly size-dimorphic northern goshawk (Accipiter gentilis L.) as a model, we studied a population with one of the highest densities of breeding pairs reported in the literature in order to understand selective pressures that may favour RSD. We evaluated life-history processes, including recruitment of adult breeders and reproductive success, and we explored the mechanisms thought to act on each sex, including hunting efficiency, diet, body condition and mate choice. We found that smaller males produced more fledglings than larger ones, but there was no relationship between size and reproductive success for females. The mean body size of female breeders was larger than that of female fledglings, but male fledglings and breeders did not differ in size. Male body size was related to the type but not to the amount of prey captured during the nestling stage. We conclude that RSD may be favoured in this goshawk population because small males tend to enjoy higher reproductive success and large females greater recruitment. Our results do not support the hypotheses that evolutionary reduction in male size is driven by hunting efficiency, at least during the nestling stage, or the hypotheses that it is driven by greater recruitment. Our findings also suggest that increase in female size is driven by recruitment, rather than by reproductive success as previously postulated.

Evaluation of Trail-Cameras for Analyzing the Diet of Nesting Raptors Using the Northern Goshawk as a Model

Diet studies present numerous methodological challenges. We evaluated the usefulness of commercially available trail-cameras for analyzing the diet of Northern Goshawks (Accipiter gentilis) as a model for nesting raptors during the period 2007–2011. We compared diet estimates obtained by direct camera monitoring of 80 nests with four indirect analyses of prey remains collected from the nests and surroundings (pellets, bones, feather-and-hair remains, and feather-hair-and-bone remains combined). In addition, we evaluated the performance of the trail-cameras and whether camera monitoring affected Goshawk behavior. The sensitivity of each diet-analysis method depended on prey size and taxonomic group, with no method providing unbiased estimates for all prey sizes and types. The cameras registered the greatest number of prey items and were probably the least biased method for estimating diet composition. Nevertheless this direct method yielded the largest proportion of prey unidentified to species level, and it underestimated small prey. Our trail-camera system was able to operate without maintenance for longer periods than what has been reported in previous studies with other types of cameras. Initially Goshawks showed distrust toward the cameras but they usually became habituated to its presence within 1–2 days. The habituation period was shorter for breeding pairs that had previous experience with cameras. Using trail-cameras to monitor prey provisioning to nests is an effective tool for studying the diet of nesting raptors. However, the technique is limited by technical failures and difficulties in identifying certain prey types. Our study also shows that cameras can alter adult Goshawk behavior, an aspect that must be controlled to minimize potential negative impacts.

Are irrigation and grazing effects transferred, accumulated, or counteracted during plant recruitment?

How do effects from perturbations such as irrigation and grazing that have an impact at one stage of the recruitment process (e.g., seedling) affect performance at later stages (e.g., adult)? Such effects may be transferred to later stages without any further change (transferred effects), reinforced by a similar effect so that their importance increases (accumulative effect), or counteracted at later stages by an opposite effect (counteractive effect). We analysed the predominance of transferred, accumulative, and counteractive effects depending on (1) organization level (community, functional group, and species levels), (2) life cycle stage (seeds, seedlings, and adults), (3) grass/forb functional group (forbs versus grasses), and (4) seed mass (small-seeded species, medium-seeded species, and large-seeded species). The study was conducted in Alcalá de Henares, central Spain. During one annual cycle, we counted the number of readily germinable seeds (3,909 seeds: 2,156 forbs and 1,753 grasses), emerged seedlings (2,126 seedlings: 1,238 forbs and 888 grasses), and surviving seedlings up to reproductive adult status (917 adults: 217 forbs and 700 grasses) of all species (74 species) in a factorial field experiment under three different simulated rainfall and three seasonal sheep grazing regimes in a natural Mediterranean annual plant community composed of forbs (dicotyledonous herbs) and grasses (monocotyledonous herbs). Our main results were: (1) the proportion of transferred and counteractive effects was similar between the three studied organization levels (community, functional group, and species levels); (2) we detected many more counteractive effects in the seedling stage than in the adult stage; (3) we did not find more counteractive effects in forbs versus grasses, but transferred effects were more frequent in grasses; and (4) we found more counteractive effects with decreasing seed mass and more transferred effects with increasing seed mass. Our main conclusions were that the prevalence of counteractive effects reduces the sensitivity of the whole recruitment process and enhances the relative importance of the final stage (adults). The prevalence of transferred effects makes the recruitment process more predictable and more dependent on the early stages (seeds and seedlings).

Assessing the ability of novel ecosystems to support animal wildlife through analysis of diurnal raptor territoriality

Novel ecosystems have emerged through human intervention and are rapidly expanding around the world. Whether they can support animal wildlife has generated considerable controversy. Here we developed a new approach to evaluate the ability of a novel forest ecosystem, dominated by the exotic tree species Eucalyptus globulus, to support animal wildlife in the medium and long term. To evaluate this ability, we took advantage of the fact that species territory size decreases with increasing habitat quality, and we used territoriality of a raptor guild composed of Northern Goshawk (Accipiter gentilis), Eurasian Sparrowhawk (A. nisus) and Common Buzzard (Buteo buteo) as indicator. We compared the territoriality of these species in the novel ecosystem with that in other ecosystems found in the literature. Average distances between con-specifics in the novel ecosystem were similar, or even shorter, than those in other ecosystems. Average distances between Goshawk con-specifics were among the shortest described in the literature. All three species nested preferably in mixed stands abundant in large exotic trees, with high structural complexity and abundance of native species within the stand. Key factors supporting this diverse and dense raptor community were the special forest management system implemented in the study area and the agricultural matrix located close to forest plantations that complements the supply of prey. Our results suggest that forest management that promotes a complex and suitable forest structure can increase the ability of novel forest ecosystems to support wildlife biodiversity, particularly a diverse nesting community of forest-dwelling raptors and their preys. The results further suggest the suitability of territoriality for assessing this potential of novel ecosystems.

Prey preferences and recent changes in diet of a breeding population of the Northern Goshawk Accipiter gentilis in Southwestern Europe

ABSTRACT Capsule: Northern Goshawk Accipiter gentilis diet has changed significantly since the 1980s, probably due to changes in populations of preferred prey species. Aims and methods: To assess changes to the breeding season diet of the Northern Goshawk in southwest Europe over three decades. We examined prey remains at and around nests and assessed avian prey availability using point count and line transect surveys. Results: During 2008–11, Goshawks mainly ate birds, with Feral Pigeons Columba livia f. domestica being the most important prey species. Goshawks preferred prey of 100−400 g and forest prey species to non-forest species. Goshawk diet has changed significantly over recent decades: 22% of current prey items belong to species that were not part of the diet in the 1980s. We suggest that these dietary changes reflect changes in the abundance of prey species of the preferred size caused by changes in land use leading to an increase in forest cover, new prey species colonization and changes in the abundance and management of domestic prey. Conclusion: This study emphasizes that major transformations occurring in agroforestry systems are affecting the main preferred prey of important forest predators, which may have consequences for conservation of both the predators and their prey.