A 16th-century biodiversity and crop inventory.

Documenting and understanding long-term biodiversity change is limited by the availability of historical data, particularly from periods preceding major anthropogenic changes associated with the Industrial Revolution. We mined the information of a standardized historical survey developed in 628 localities across Spain between 1574 and 1582 (Relaciones Topográficas de Felipe II) with the goal of producing a general characterization of Spanish settlements, including, though not limited to, natural resources. From this survey, we were able to gather 7309 records for 75 wild plant taxa, 89 wild animal taxa, and 60 crop and domestic animal taxa. These data can be used to reconstruct historical land use and habitat cover, and model historical distributions of many species, including emblematic ones such as wolf and bear, which can establish reference distributions to assess range and niche expansion, contraction, and shifts. Data are provided under a Creative Commons Attribution Non Commercial (CC-BY-NC) 4.0 License. Please cite this data paper when the data are used in publications. The authors are open to collaborate in projects based on this dataset.

Eurasian otter (Lutra lutra) as sentinel species for the long-term biomonitoring of the Guadiamar River after the Aznalcóllar mine spill.

Accidents at mines involving stored tailings have produced catastrophic environmental damage. In April 1998 the dam of the Aznalcóllar mine tailings pond in the surroundings of the Doñana National Park (southwestern Spain) broke, discharging into the Guadiamar River more than 6 million m3 of toxic mud and acidic water with high concentrations of heavy metals and arsenic. We used the Eurasian otter (Lutra lutra) as sentinel species to assess the potential impact of the toxic spill on the river ecosystems and their recovery with time by studying the spatial and temporal variation (1999-2003, 2006) of selected trace element (Cu, Zn, Cd, Pb and As) concentrations in feces. Throughout the sampling period, the highest heavy metal and As levels were found in the most spill-affected reaches of the Guadiamar River (i.e., the Middle and, to a lesser extent, the Lower reaches), pointing out the mining accident as the main origin of the contamination. Overall, levels of trace elements decreased with the time elapsed since the toxic spill, except for Cd (F1,352 = 0.29, P = 0.59). However, rebounds for some elements (Pb, As, and Cu) were also observed, especially in the Middle and Lower reaches of the river, which might be attributed to the residual contamination in abiotic compartments and/or new inputs from industrial and agricultural activities in the nearby areas. Concentrations were relatively high when compared to those reported for both our reference area (Guadalete River) and other metal-polluted zones. We found that the estimated amounts of Pb and As ingested during the first years after the spill in the Guadiamar Middle reach would be high enough to cause reproductive issues. This could affect the local population recovery, although evidence on distribution range and numbers suggests otherwise, with thriving populations at regional scale. Our results support the role of otters as sentinel species for biomonitoring contaminants and thus to evaluate fluvial ecosystem health.

Genetic rescue by distant trees mitigates qualitative pollen limitation imposed by fine-scale spatial genetic structure.

Restricted seed dispersal frequently leads to fine-scale spatial genetic structure (i.e., FSGS) within plant populations. Depending on its spatial extent and the mobility of pollinators, this inflated kinship at the immediate neighbourhood can critically impoverish pollen quality. Despite the common occurrence of positive FSGS within plant populations, our knowledge regarding the role of long-distance pollination preventing reproductive failure is still limited. Using microsatellite markers, we examined the existence of positive FSGS in two low-density populations of the tree Pyrus bourgaeana. We also designed controlled crosses among trees differing in their kinship to investigate the effects of increased local kinship on plant reproduction. We used six pollination treatments and fully monitored fruit production, fruit and seed weight, proportion of mature seeds per fruit, and seed germination. Our results revealed positive FSGS in both study populations and lower fruit initiation in flowers pollinated with pollen from highly-genetically related individuals within the neighbourhood, with this trend intensifying as the fruit development progressed. Besides, open-pollinated flowers exhibited lower performance compared to those pollinated by distant pollen donors, suggesting intense qualitative pollen limitation in natural populations. We found positive fine-scale spatial genetic structure is translated into impoverished pollen quality from nearby pollen donors which negatively impacts the reproductive success of trees in low-density populations. Under this scenario of intrapopulation genetic rescue by distant pollen donors, the relevance of highly-mobile pollinators for connecting spatially and genetically distant patches of trees may be crucial to safeguarding population recruitment.

Patterns of brown bear damages on apiaries and management recommendations in the Cantabrian Mountains, Spain.

Large carnivores are often persecuted due to conflict with human activities, making their conservation in human-modified landscapes very challenging. Conflict-related scenarios are increasing worldwide, due to the expansion of human activities or to the recovery of carnivore populations. In general, brown bears Ursus arctos avoid humans and their settlements, but they may use some areas close to people or human infrastructures. Bear damages in human-modified landscapes may be related to the availability of food resources of human origin, such as beehives. However, the association of damage events with factors that may predispose bears to cause damages has rarely been investigated. We investigated bear damages to apiaries in the Cantabrian Mountains (Spain), an area with relatively high density of bears. We included spatial, temporal and environmental factors and damage prevention measures in our analyses, as factors that may influence the occurrence and intensity of damages. In 2006-2008, we located 61 apiaries, which included 435 beehives damaged in the study area (346 km2). The probability of an apiary being attacked was positively related to both the intensity of the damage suffered the year before and the distance to the nearest damaged apiary, and negatively related to the number of prevention measures employed as well as the intensity of the damage suffered by the nearest damage apiary. The intensity of damage to apiaries was positively related to the size of the apiary and to vegetation cover in the surroundings, and negatively related to the number of human settlements. Minimizing the occurrence of bear damages to apiaries seems feasible by applying and maintaining proper prevention measures, especially before an attack occurs and selecting appropriate locations for beehives (e.g. away from forest areas). This applies to areas currently occupied by bears, and to neighbouring areas where dispersing individuals may expand their range.

Strong among population variation in frugivory strength by functional diverse frugivores: a 'reciprocal translocation' experiment.

Fruit-frugivore interactions are critical for the dynamics and evolution of many plant communities. The strength of the interactions between a given plant species and different frugivore guilds (e.g., seed dispersers, seed predators) often vary in space due to changes in plant extrinsic factors (e.g., frugivore abundances) and plant traits (e.g., fruit size and reward). By reciprocally translocating Pyrus bourgaeana ripe fruits representative of five Mediterranean localities during 2 consecutive years, we experimentally quantified guild-specific variations among populations in frugivory strength, while accounted for plant-intrinsic and- extrinsic factors. Though overall fruit removal did not differ among localities, there were strong guild-specific differences in fruit removal strength. Fruit removal by pulp feeders, seed dispersers, and fruit predators varied among populations up to 8.5-, 5.6-, and 4.0-folds, respectively. These strong variations seemed mediated by changes in frugivore relative abundances rather than on availability of alternative fruits. As expected, all fruit traits considered (e.g., fruit size, pulp amount) markedly varied among tree populations. However, no frugivore guild showed preference for fruits from any locality, suggesting that fruit traits did not contribute much to differences in frugivory strength among populations. Since the functional diverse frugivore guilds played contrasting roles in P. bourgaeana dynamics (e.g., seed dispersal vs. seed predation), our study highlights the importance of accounting for functional diversity in frugivore guilds when estimating spatial variations in the strenght of seed dispersal. This investigation also illustrates a neglected but widely applicable experimental approach to identify the relative importance of extrinsic factors and fruit traits in mediating fruit-frugivore interactions.

Spatio-temporal arrangement of Chamaerops humilis inflorescences and occupancy patterns by its nursery pollinator, Derelomus chamaeropsis.

Background and Aims: Nursery pollination is a highly specialized interaction in which pollinators breed inside plant reproductive structures. Pollinator occupancy of host plants often depends on plant location, flowering synchrony and sex. The nursery pollination system between the dioecious dwarf palm Chamaerops humilis (Arecaceae) and the host-specific palm flower weevil Derelomus chamaeropsis was investigated. For the first time, sex, flowering synchrony and spatial distribution of plants was related to the occupancy probability and the abundance of D. chamaeropsis larvae, important traits influencing both pollinator and plant fitness. Methods: During the flowering season, all inflorescences in anthesis were counted every 12 d and a flowering synchrony index was calculated taking into account all possible correlations with generalized linear mixed models. To analyse the spatial structure of plants, larva occupancy and abundance, different techniques of spatial point pattern analysis were used. Key results: In total, 5986 larvae in 1063 C. humilis inflorescences were recorded over three consecutive seasons. Male inflorescences showed a higher presence and abundance of weevil larvae than females, but interestingly approx. 30 % of the females held larvae. Also, larvae occurred mainly in highly synchronous plants with a low number of inflorescences, perhaps because those plants did not lead to a resource dilution effect. There was no evidence of spatial patterns in larva occupancy or abundance at any spatial scale, suggesting high dispersal ability of adult weevil. Conclusions: The results in a nursery-pollinated dioecious palm demonstrate that plant sex, flowering display and flowering synchrony act as additive forces influencing the presence and abundance of the specialized pollinator larvae. Contradicting previous results, clear evidence that female dwarf palms also provide rewarding oviposition sites was found, and thus the plant 'pays' for the pollination services. The findings highlight that plant local aggregation is not always the main determinant of pollinator attraction, whereas flower traits and phenology could be critical in specialized plant-pollinator interactions.

Staying cool or staying safe in a human-dominated landscape: which is more relevant for brown bears?

Pigeon et al. (2016) Staying cool in a changing landscape: the influence of maximum daily ambient temperature on grizzly bear habitat selection. Oecologia 181:1101. doi: 10.1007/s00442-016-3630-5 analyzed the effect of ambient temperature on the habitat selection of grizzly bears (Ursus arctos) in Alberta, Canada. They concluded that temperature played a significant role in bear habitat selection and that it was unlikely that human activity introduced biases to the habitat selection of bears. However, Pigeon et al. did not consider variables related to human activities in their analyses. They also misinterpreted previous research that has accounted for temperature in the habitat selection of brown bears. There is much literature published on the negative effects of human disturbance on wildlife in general and on bears in particular. Downplaying the role of human disturbance could have important negative consequences if, in fact, human disturbance were a more important factor than thermoregulation. Indeed, dismissing the importance of human influence, in the face of contradictory evidence, could tempt managers to disregard an important factor that is difficult and often unpopular to deal with in their conservation plans.

Spatiotemporal Dynamics of Genetic Variation in the Iberian Lynx along Its Path to Extinction Reconstructed with Ancient DNA.

There is the tendency to assume that endangered species have been both genetically and demographically healthier in the past, so that any genetic erosion observed today was caused by their recent decline. The Iberian lynx (Lynx pardinus) suffered a dramatic and continuous decline during the 20th century, and now shows extremely low genome- and species-wide genetic diversity among other signs of genomic erosion. We analyze ancient (N = 10), historical (N = 245), and contemporary (N = 172) samples with microsatellite and mitogenome data to reconstruct the species' demography and investigate patterns of genetic variation across space and time. Iberian lynx populations transitioned from low but significantly higher genetic diversity than today and shallow geographical differentiation millennia ago, through a structured metapopulation with varying levels of diversity during the last centuries, to two extremely genetically depauperate and differentiated remnant populations by 2002. The historical subpopulations show varying extents of genetic drift in relation to their recent size and time in isolation, but these do not predict whether the populations persisted or went finally extinct. In conclusion, current genetic patterns were mainly shaped by genetic drift, supporting the current admixture of the two genetic pools and calling for a comprehensive genetic management of the ongoing conservation program. This study illustrates how a retrospective analysis of demographic and genetic patterns of endangered species can shed light onto their evolutionary history and this, in turn, can inform conservation actions.

Historical citizen science to understand and predict climate-driven trout decline.

Historical species records offer an excellent opportunity to test the predictive ability of range forecasts under climate change, but researchers often consider that historical records are scarce and unreliable, besides the datasets collected by renowned naturalists. Here, we demonstrate the relevance of biodiversity records developed through citizen-science initiatives generated outside the natural sciences academia. We used a Spanish geographical dictionary from the mid-nineteenth century to compile over 10 000 freshwater fish records, including almost 4 000 brown trout (Salmo trutta) citations, and constructed a historical presence-absence dataset covering over 2 000 10 × 10 km cells, which is comparable to present-day data. There has been a clear reduction in trout range in the past 150 years, coinciding with a generalized warming. We show that current trout distribution can be accurately predicted based on historical records and past and present values of three air temperature variables. The models indicate a consistent decline of average suitability of around 25% between 1850s and 2000s, which is expected to surpass 40% by the 2050s. We stress the largely unexplored potential of historical species records from non-academic sources to open new pathways for long-term global change science.

Spatial patterns of an endemic Mediterranean palm recolonizing old fields.

Throughout Europe, increased levels of land abandonment lead to (re)colonization of old lands by forests and shrublands. Very little is known about the spatial pattern of plants recolonizing such old fields. We mapped in two 21-22-ha plots, located in the Doñana National Park (Spain), all adult individuals of the endozoochorous dwarf palm Chamaerops humilisL. and determined their sex and sizes. We used techniques of spatial point pattern analysis (SPPA) to precisely quantify the spatial structure of these C. humilis populations. The objective was to identify potential processes generating the patterns and their likely consequences on palm reproductive success. We used (1) Thomas point process models to describe the clustering of the populations, (2) random labeling to test the sexual spatial segregation, and (3) mark correlation functions to assess spatial structure in plant sizes. Plants in both plots showed two critical scales of clustering, with small clusters of a radius of 2.8-4 m nested within large clusters with 38-44 m radius. Additional to the clustered individuals, 11% and 27% of all C. humilis individuals belonged to a random pattern that was independently superimposed to the clustered pattern. The complex spatial pattern of C. humilis could be explained by the effect of different seed-dispersers and predators' behavior and their relative abundances. Plant sexes had no spatial segregation. Plant sizes showed a spatial aggregation inside the clusters, with a decreasing correlation with distance. Clustering of C. humilis is strongly reliant on its seed dispersers and stressful environmental conditions. However, it seems that the spatial patterns and dispersal strategies of the dwarf palm make it a successful plant for new habitat colonization. Our results provide new information on the colonization ability of C. humilis and can help to develop management strategies to recover plant populations.

Variability in primary productivity determines metapopulation dynamics.

Temporal variability in primary productivity can change habitat quality for consumer species by affecting the energy levels available as food resources. However, it remains unclear how habitat-quality fluctuations may determine the dynamics of spatially structured populations, where the effects of habitat size, quality and isolation have been customarily assessed assuming static habitats. We present the first empirical evaluation on the effects of stochastic fluctuations in primary productivity--a major outcome of ecosystem functions--on the metapopulation dynamics of a primary consumer. A unique 13-year dataset from an herbivore rodent was used to test the hypothesis that inter-annual variations in primary productivity determine spatiotemporal habitat occupancy patterns and colonization and extinction processes. Inter-annual variability in productivity and in the growing season phenology significantly influenced habitat colonization patterns and occupancy dynamics. These effects lead to changes in connectivity to other potentially occupied habitat patches, which then feed back into occupancy dynamics. According to the results, the dynamics of primary productivity accounted for more than 50% of the variation in occupancy probability, depending on patch size and landscape configuration. Evidence connecting primary productivity dynamics and spatiotemporal population processes has broad implications for metapopulation persistence in fluctuating and changing environments.

Conflict Misleads Large Carnivore Management and Conservation: Brown Bears and Wolves in Spain.

Large carnivores inhabiting human-dominated landscapes often interact with people and their properties, leading to conflict scenarios that can mislead carnivore management and, ultimately, jeopardize conservation. In northwest Spain, brown bears Ursus arctos are strictly protected, whereas sympatric wolves Canis lupus are subject to lethal control. We explored ecological, economic and societal components of conflict scenarios involving large carnivores and damages to human properties. We analyzed the relation between complaints of depredations by bears and wolves on beehives and livestock, respectively, and bear and wolf abundance, livestock heads, number of culled wolves, amount of paid compensations, and media coverage. We also evaluated the efficiency of wolf culling to reduce depredations on livestock. Bear damages to beehives correlated positively to the number of female bears with cubs of the year. Complaints of wolf predation on livestock were unrelated to livestock numbers; instead, they correlated positively to the number of wild ungulates harvested during the previous season, the number of wolf packs, and to wolves culled during the previous season. Compensations for wolf complaints were fivefold higher than for bears, but media coverage of wolf damages was thirtyfold higher. Media coverage of wolf damages was unrelated to the actual costs of wolf damages, but the amount of news correlated positively to wolf culling. However, wolf culling was followed by an increase in compensated damages. Our results show that culling of the wolf population failed in its goal of reducing damages, and suggest that management decisions are at least partly mediated by press coverage. We suggest that our results provide insight to similar scenarios, where several species of large carnivores share the landscape with humans, and management may be reactive to perceived conflicts.

Isotopic niche variation in a higher trophic level ectotherm: highlighting the role of succulent plants in desert food webs.

Stable isotope analysis of animal tissues allows description of isotopic niches, whose axes in an n-dimensional space are the isotopic ratios, compared to a standard, of different isotope systems (e.g. δ(13)C, δ(15)N). Isotopic niches are informative about where an animal, population or species lives and about what it consumes. Here we describe inter- and intrapopulation isotopic niche (bidimensional δ(13)C-δ(15)N space) of the Orange-throated whiptail (Aspidoscelis hyperythra), an arthropodivorous small lizard, in ten localities of Baja California Sur (Mexico). These localities range from extreme arid to subtropical conditions. Between 13 and 20 individuals were sampled at each locality and 1 cm of tail-tip was collected for isotope analysis. As expected, interpopulation niche width variation was much larger than intrapopulation one. Besides, isotopic variation was not related to age, sex or individual size of lizards. This suggests geographic variation of the isotopic niche was related to changes in the basal resources that fuel the trophic web at each locality. The position of Bayesian isotope ellipses in the δ-space indicated that whiptails in more arid localities were enriched in 13C, suggesting most of the carbon they ingested came from CAM succulent plants (cacti, agaves) and in minor degree in C4 grasses. Contrarily, whiptails in subtropical areas were depleted in 13C, as they received more carbon from C3 scrubs and trees. Localities closer to sea-level tended to be enriched in 15N, but a clear influence of marine subsidies was detected only at individual level. The study contributes to identify the origin and pathways through which energy flows across the trophic webs of North American deserts.

Fecal nitrogen concentration as a nutritional quality indicator for European rabbit ecological studies.

Measuring the quality of the nutritional resources available to wild herbivores is critical to understanding trophic regulation processes. However, the direct assessment of dietary nutritional characteristics is usually difficult, which hampers monitoring nutritional constraints in natural populations. The feeding ecology of ruminant herbivores has been often assessed by analyzing fecal nitrogen (FN) concentrations, although this method has been less evaluated in other taxa. This study analyzed the suitability of FN as an indicator of ingesta quality in the European rabbit (Oryctolagus cuniculus), which is a keystone lagomorph species in Mediterranean ecosystems and of great conservation interest. Firstly, domestic O. cuniculus were used to evaluate under experimental conditions the accuracy of total FN and the metabolic FN as diet quality indicators of forages with characteristics similar to those available under natural conditions. Secondly, the accuracy of Near-Infrared Spectroscopy (NIRS) to calculate FN was tested using partial least squares regression. Thirdly, a pilot field study was conducted to monitor FN dynamics from wild O. cuniculus in three different habitats during wet and drought periods. A strong association was found between diet type and total FN and metabolic FN (Pseudo-R(2) ≥ 0.89). It was also found that NIRS calibrations were accurate for depicting nitrogen concentrations (R(2) > 0.98 between NIRS and chemical results). Finally, the seasonal FN dynamics measured in the field were consistent with current knowledge on vegetation dynamics and forage limitations in the three habitats. The results support the use of NIRS methods and FN indices as a reliable and affordable approach to monitoring the nutritional quality of rabbit habitats. Potential applications include the assessment of the mechanistic relationships between resource limitations and population abundance, e.g., in relation to natural drought cycles and to habitat interventions aimed at reinforcing rabbit populations.

Hierarchical levels of seed predation variation by introduced beetles on an endemic Mediterranean palm.

Seed predators can limit plant recruitment and thus profoundly impinge the dynamics of plant populations, especially when diverse seed predators (e.g., native and introduced) attack particular plant populations. Surprisingly, however, we know little concerning the potential hierarchy of spatial scales (e.g., region, population, patch) and coupled ecological correlates governing variation in the overall impact that native and introduced seed predators have on plant populations. We investigated several spatial scales and ecological correlates of pre-dispersal seed predation by invasive borer beetles in Chamaerops humilis (Arecaceae), a charismatic endemic palm of the Mediteranean basin. To this end, we considered 13 palm populations (115 palms) within four geographical regions of the Iberian Peninsula. The observed interregional differences in percentages of seed predation by invasive beetles were not significant likely because of considerable variation among populations within regions. Among population variation in seed predation was largely related to level of human impact. In general, levels of seed predation were several folds higher in human-altered populations than in natural populations. Within populations, seed predation declined significantly with the increase in amount of persisting fruit pulp, which acted as a barrier against seed predators. Our results revealed that a native species (a palm) is affected by the introduction of related species because of the concurrent introduction of seed predators that feed on both the introduced and native palms. We also show how the impact of invasive seed predators on plants can vary across a hierarchy of levels ranging from variation among individuals within local populations to large scale regional divergences.

Revisiting food-based models of territoriality in solitary predators.

Food availability is considered a major factor determining spacing behaviour in territorial species, especially for females. Theoretically, spatial overlap (considered the opposite of territoriality) and food availability are related in a nonlinear manner (hypothesized inverted-U function), with high overlap levels at the extremes of a food availability gradient and low overlap at intermediate levels of this gradient. Similar patterns are expected for encounter frequencies owing to its expected correlation with spatial overlap. However, these predictions have rarely been tested in highly structured social systems on a broad gradient of food availability, which implicitly requires experimental manipulation. We test these predictions in a solitary, territorial and trophic specialist, the Iberian lynx Lynx pardinus, taking advantage of a three-decade data set of spatial behaviour in different scenarios of food availability (i.e. rabbit density). In contrast with expectations, home range overlap among resident females was low (median overlap index = 0.08, range 0-0.57) and core area overlap was nearly nil (median overlap index = 0, range 0-0.22) throughout the entire gradient of prey availability. Furthermore, spatial associations between pairs of females were negligible regardless marked variation in prey availability. Therefore, we did not find support for a model of flexible lynx territoriality driven by food availability. Our results suggest that the exclusive use of space in the Iberian lynx was not related to food. Lack of influence of prey availability on lynx territoriality may be adaptive to cope with the consequences of frequent drought-induced periods of prey scarcity or other disturbance typically affecting wild rabbit populations in Mediterranean environments. Thus, lynx would adopt an obstinate strategy of territoriality that consists in defending exclusive areas across a broad range of resource availability ensuring an exclusive access to the minimum amount of prey necessary for survival and eventually reproduction even during periods of prey scarcity. However, we found signs that territoriality was influenced by lynx density in a nonlinear fashion. Our results suggest the occurrence of population regulation through territoriality in this species.

Possible extinction vortex for a population of Iberian lynx on the verge of extirpation.

Theory suggests that demographic and genetic traits deteriorate (i.e., fitness and genetic diversity decrease) when populations become small, and that such deterioration could precipitate positive feedback loops called extinction vortices. We examined whether demographic attributes and genetic traits have changed over time in one of the 2 remaining small populations of the highly endangered Iberian lynx (Lynx pardinus) in Doñana, Spain. From 1983 to 2008, we recorded nontraumatic mortality rates, litter size, offspring survival, age at territory acquisition, and sex ratio. We combined these demographic attributes with measures of inbreeding and genetic diversity at neutral loci (microsatellites) and genes subjected to selection (major histocompatibility complex). Data on demographic traits were obtained through capture and radio tracking, checking dens during breeding, track surveys, and camera trapping. For genetic analyses, we obtained blood or tissue samples from captured or necropsied individuals or from museum specimens. Over time a female-biased sex ratio developed, age of territory acquisition decreased, mean litter size decreased, and rates of nontraumatic mortality increased, but there were no significant changes in overall mortality rates, standardized individual heterozygosity declined steadily, and allelic diversity of exon 2 of class II major histocompatibility complex DRB genes remained constant (2 allelic variants present in all individuals analyzed). Changes in sex ratio and age of territory acquisition may have resulted from demographic stochasticity, whereas changes in litter size and nontraumatic mortality may be related to observed increases in inbreeding. Concomitant deterioration of both demographic attributes and genetic traits is consistent with an extinction vortex. The co-occurrence, with or without interaction, of demographic and genetic deterioration may explain the lack of success of conservation efforts with the Doñana population of Iberian lynx.

Thieves or mutualists? Pulp feeders enhance endozoochore local recruitment.

The persistence of mutualisms despite the strong incidence of exploiters could be explained if exploiters deny one or more services (i.e., cheat) while eventually supplying some subtler but critical services. Pulp feeders usually ingest fruit reward without dispersing seeds and thus are considered to be mainly cheaters or thieves of seed-disperser mutualisms. By consuming the fruit pulp, however, they could release seeds from pulp inhibitory effect, enhancing germination and, potentially, subsequent seedling emergence, growth, survival, and thus local recruitment. We evaluated such a largely neglected hypothesis by considering the interaction between Pyrus bourgaeana and its pulp feeders. We experimentally showed that pericarp removal had a consistent strong positive effect on seed performance (e.g., lower rotting and higher germination percentages) and seedling fate (greater emergence, growth, and survival to two years old). Interestingly, these relatively large positive effects of depulpation on plant fitness persisted for a surprisingly long time. Though seedlings experienced higher mortality under fruiting conspecifics, the benefits of depulpation were not overridden by high propagule mortality beneath fruiting trees or in adverse microhabitats after two years of monitoring. Specifically, the cumulative probability of establishment for depulped seeds was 4-25 times higher than for seeds in whole ripe fruits. Thus, under some circumstances, pulp feeders can provide essential services to endozoochorous plants. Our study contributes to clarifying the apparent paradox of plant-frugivore mutualisms that persist in the face of exploitation by pulp feeders. Because "thieves" and "mutualists" refer to the extremes of a complex continuum, and because organisms displaying concurrent cheating and honest behaviors during different host stages are likely prevalent, the persistent language of mutualists vs. thieves, cheaters, or exploiters might be misleading.

Microgeographical, inter-individual, and intra-individual variation in the flower characters of Iberian pear Pyrus bourgaeana (Rosaceae).

Flower characteristics have been traditionally considered relatively constant within species. However, there are an increasing number of examples of variation in flower characteristics. In this study, we examined the variation in attracting and rewarding flower characters at several ecological levels in a metapopulation of Pyrus bourgaeana in the Doñana area (SW Spain). We answered the following questions: what are the variances of morphological and nectar characters of flowers? How important are intra-individual and inter-individual variance in flower characters? Are there microgeographical differences in flower characters? And if so, are they consistent between years? In 2008 and 2009, we sampled flowers of 72 trees from five localities. For six flower morphological and two nectar characteristics, we calculated coefficients of variation (CV). The partitioning of total variation among-localities, among-individuals, and within-individuals was estimated. To analyze differences among localities and their consistency between years, we conducted generalized linear mixed models. The CVs of nectar characters were always higher than those of morphological characters. As expected, inter-individual variation was the main source of variation of flower morphology, but nectar characters had significant variation at both intra- and inter-individual levels. For most floral traits, there were no differences among localities. Our study documents that variation is a scale-dependent phenomenon and that it is essential to consider intra- and inter-individual variance when investigating the causes and consequences of variation. It also shows that single year studies of floral characters should be viewed with caution.

Prisoners in their habitat? Generalist dispersal by habitat specialists: a case study in southern water vole (Arvicola sapidus).

Habitat specialists inhabiting scarce and scattered habitat patches pose interesting questions related to dispersal such as how specialized terrestrial mammals do to colonize distant patches crossing hostile matrices. We assess dispersal patterns of the southern water vole (Arvicola sapidus), a habitat specialist whose habitat patches are distributed through less than 2% of the study area (overall 600 km²) and whose populations form a dynamic metapopulational network. We predict that individuals will require a high ability to move through the inhospitable matrix in order to avoid genetic and demographic isolations. Genotypes (N = 142) for 10 microsatellites and sequences of the whole mitochondrial Control Region (N = 47) from seven localities revealed a weak but significant genetic structure partially explained by geographic distance. None of the landscape models had a significant effect on genetic structure over that of the Euclidean distance alone and no evidence for efficient barriers to dispersal was found. Contemporary gene flow was not severely limited for A. sapidus as shown by high migration rates estimates (>10%) between non-neighbouring areas. Sex-biased dispersal tests did not support differences in dispersal rates, as shown by similar average axial parent-offspring distances, in close agreement with capture-mark-recapture estimates. As predicted, our results do not support any preferences of the species for specific landscape attributes on their dispersal pathways. Here, we combine field and molecular data to illustrate how a habitat specialist mammal might disperse like a habitat generalist, acquiring specific long-distance dispersal strategies as an adaptation to patchy, naturally fragmented, heterogeneous and unstable habitats.