Arnan X., Comas L., Gracia M., Retana J. (2014) Composition and habitat use of small mammals in old-growth mountain forests. Journal of Natural History. 48: 481-494.LinkDoi: 10.1080/00222933.2013.800611
Old-growth mountain forests in the Pyrenees have natural gap dynamics, a well-developed shrub layer and a large amount of dead wood. Small mammal communities in two types of old-growth forests, silver fir and mountain pine, were studied in July and September in 2006 and 2007. Four species were trapped: bank vole (Myodes glareolus), wood mouse (Apodemus sylvaticus), garden dormouse (Eliomys quercinus) and common shrew (Sorex araneus). Bank voles and wood mice were most commonly trapped (78% of all captures). There were no differences in community composition in the two forest types, although the bank vole was more often captured in the silver fir than in the mountain pine forest. Mammals were more frequently captured at trap stations with high shrub cover, high tree regeneration cover and low herbaceous cover. Our results show that forest structure and, to a lesser extent, forest type determine small mammal community structure, and specifically fine-scale occurrence patterns, in these old-growth forests. © 2013 Taylor & Francis.
Arnan X., Escola A., Rodrigo A., Bosch J. (2014) Female reproductive success in gynodioecious Thymus vulgaris: Pollen versus nutrient limitation and pollinator foraging behaviour. Botanical Journal of the Linnean Society. 175: 395-408.LinkDoi: 10.1111/boj.12173
Gynodioecy is a dimorphic breeding system in which female individuals coexist with hermaphroditic individuals in the same population. Females only contribute to the next generation via ovules, and many studies have shown that they are usually less attractive than hermaphrodites to pollinators. Several mechanisms have been proposed to explain how females manage to persist in populations despite these disadvantages. The 'resource reallocation hypothesis' (RRH) states that females channel resources not invested in pollen production and floral advertisement towards the production of more and/or larger seeds. We investigated pollination patterns and tested the RRH in a population of Thymus vulgaris. We measured flower display, flower size, nectar production, visitation rates, pollinator constancy and flower lifespan in the two morphs. In addition, we measured experimentally the effects of pollen and resource addition on female reproductive success (fruit set, seed set, seed weight) of the two morphs. Despite lower investment in floral advertisement, female individuals were no less attractive to pollinators than hermaphrodites on a per flower basis. Other measures of pollinator behaviour (number of flowers visited per plant, morph preference and morph constancy) also showed that pollinators did not discriminate against female flowers. In addition, stigma receptivity was longer in female flowers. Accordingly, and contrary to most studies on gynodioecious species, reproductive success of females was not pollen limited. Instead, seed production was pollen limited in hermaphrodites, suggesting low levels of cross-pollination in hermaphrodites. Seed production was resource limited in hermaphrodites, but not in females, thus providing support for the RRH. © 2014 The Linnean Society of London.
Caut S., Jowers M.J., Arnan X., Pearce-Duvet J., Rodrigo A., Cerda X., Boulay R.R. (2014) The effects of fire on ant trophic assemblage and sex allocation. Ecology and Evolution. 4: 35-49.LinkDoi: 10.1002/ece3.714
Fire plays a key role in ecosystem dynamics worldwide, altering energy flows and species community structure and composition. However, the functional mechanisms underlying these effects are not well understood. Many ground-dwelling animal species can shelter themselves from exposure to heat and therefore rarely suffer direct mortality. However, fire-induced alterations to the environment may change a species' relative trophic level within a food web and its mode of foraging. We assessed how fire could affect ant resource utilization at different scales in a Mediterranean forest. First, we conducted isotopic analyses on entire ant species assemblages and their potential food resources, which included plants and other arthropods, in burned and unburned plots 1 year postfire. Second, we measured the production of males and females by nests of a fire-resilient species, Aphaenogaster gibbosa, and analyzed the differences in isotopic values among workers, males, and females to test whether fire constrained resource allocation. We found that, in spite of major modifications in biotic and abiotic conditions, fire had little impact on the relative trophic position of ant species. The studied assemblage was composed of species with a wide array of diets. They ranged from being mostly herbivorous to completely omnivorous, and a given species' trophic level was the same in burned and unburned plots. In A. gibbosa nests, sexuals had greater δ15N values than workers in both burned and unburned plots, which suggests that the former had a more protein-rich diet than the latter. Fire also appeared to have a major effect on A. gibbosa sex allocation: The proportion of nests that produced male brood was greater on burned zones, as was the mean number of males produced per nest with the same reproductive investment. Our results show that generalist ants with relatively broad diets maintained a constant trophic position, even following a major disturbance like fire. However, the dramatically reduced production of females on burned zones compared to unburned zones 1 year postfire may result in considerably reduced recruitment of new colonies in the mid to long term, which could yield genetic bottlenecks and founder effects. Our study paves the way for future functional analyses of fire-induced modifications in ant populations and communities. © 2013 The Authors.
Quevedo L., Arnan X., Boet O., Rodrigo A. (2014) Post-fire selective thinning of Arbutus unedo L. coppices keeps animal diversity unchanged: The case of ants. Annals of Forest Science. 71: 897-905.LinkDoi: 10.1007/s13595-014-0381-5
Context: In the Mediterranean area, different post-fire management strategies are used for coppices of resprouting species to promote a more regular forest structure, enhance plant growth, and reduce fire risk. However, the effects of these management treatments on forest-associated fauna are unknown, which in turn could be limiting their beneficial effects.Aims: The aim of this work was to determine whether forest management of a recently burned area dominated by a vigorous resprouting tree species (Arbutus unedo L.) affects ant communities.Methods: Ant communities, sampled using pitfall traps, were examined from unmanaged and selective thinning coppices of A. unedo. Ants are here used as bioindicators of ecosystem health and surrogates for other animal groups.Results: Very limited effects of these post-fire management strategies on the structure and composition of ant communities were found. The lack of effects could be due to the reported small changes in physical conditions among treatments; or either, the most sensitive ant species to these post-fire management treatments might be the same ones affected by fire and, consequently, the ant species that would potentially be affected most were no longer in the study area.Conclusion: The lack of any significant effects caused by these post-fire management practices on the associated fauna of A. unedo coppices points out the suitability of these treatments in these circumstances. © 2014, INRA and Springer-Verlag France.
Torne-Noguera A., Rodrigo A., Arnan X., Osorio S., Barril-Graells H., Da Rocha-Filho L.C., Bosch J. (2014) Determinants of spatial distribution in a bee community: Nesting resources, flower resources, and body size. PLoS ONE. 9: 0-0.LinkDoi: 10.1371/journal.pone.0097255
Understanding biodiversity distribution is a primary goal of community ecology. At a landscape scale, bee communities are affected by habitat composition, anthropogenic land use, and fragmentation. However, little information is available on local-scale spatial distribution of bee communities within habitats that are uniform at the landscape scale. We studied a bee community along with floral and nesting resources over a 32 km2 area of uninterrupted Mediterranean scrubland. Our objectives were (i) to analyze floral and nesting resource composition at the habitat scale. We ask whether these resources follow a geographical pattern across the scrubland at bee-foraging relevant distances; (ii) to analyze the distribution of bee composition across the scrubland. Bees being highly mobile organisms, we ask whether bee composition shows a homogeneous distribution or else varies spatially. If so, we ask whether this variation is irregular or follows a geographical pattern and whether bees respond primarily to flower or to nesting resources; and (iii) to establish whether body size influences the response to local resource availability and ultimately spatial distribution. We obtained 6580 specimens belonging to 98 species. Despite bee mobility and the absence of environmental barriers, our bee community shows a clear geographical pattern. This pattern is mostly attributable to heterogeneous distribution of small (<55 mg) species (with presumed smaller foraging ranges), and is mostly explained by flower resources rather than nesting substrates. Even then, a large proportion (54.8%) of spatial variability remains unexplained by flower or nesting resources. We conclude that bee communities are strongly conditioned by local effects and may exhibit spatial heterogeneity patterns at a scale as low as 500-1000 m in patches of homogeneous habitat. These results have important implications for local pollination dynamics and spatial variation of plant-pollinator networks. © 2014 Torné-Noguera et al.
Andersen A.N., Arnan X., Sparks K. (2013) Limited niche differentiation within remarkable co-occurrences of congeneric species: Monomorium ants in the Australian seasonal tropics. Austral Ecology. 38: 557-567.LinkDoi: 10.1111/aec.12000
Niche theory predicts that few closely related species can co-occur because such species tend to be ecologically similar and niche differentiation is required to avoid competitive exclusion. We analyse the co-occurrence of a remarkable 10-15 species of the ant genus Monomorium occurring within single 10×10m plots in a tropical savanna of northern Australia. Most of the species are undescribed, so we use genetic analysis to validate our species demarcations. We document nest dispersion patterns, and investigate differentiation in the three primary niche dimensions: space, time and food. We also examine species differences in competitive abilities, by describing rates of foraging activity, foraging ranges, worker aggression, and levels of behavioural dominance. Analyses of nest and forager distributions showed very limited evidence of spatial segregation within plots. The great majority of species foraged either exclusively or primarily during daylight hours. Body size and isotopic analyses indicated very limited dietary differentiation. Such limited niche partitioning occurred despite the species differing markedly in their competitive abilities as measured by rates of resource discovery, recruitment and monopolization. Our findings defy the traditional assumption that multiple closely related and ecologically similar species of highly interactive taxa cannot co-occur. It seems very likely that species coexistence in our study system is determined to a very large degree by stochastic processes relating to dispersal and establishment, as predicted by neutral theory. However, neutral theory assumes competitive equivalence, whereas we found very marked differences in the competitive abilities of our co-occurring species. We suggest that competitive exclusion is prevented by the modular nature of ant colonies, with competition limiting colony performance but not preventing occurrence. We conclude that other factors that allow species persistence, and not just competitive equivalence, can allow dispersal and establishment processes to drive species coexistence. © 2012 Ecological Society of Australia.
Arnan X., Cerda X., Rodrigo A., Retana J. (2013) Response of ant functional composition to fire. Ecography. 36: 1182-1192.LinkDoi: 10.1111/j.1600-0587.2013.00155.x
Little is known about the impact of disturbances on functional diversity and the long-term provisioning of ecosystem services, especially in animals. In this work we analyze the effect of wildfire on the functional composition of Mediterranean ant communities. In particular, we asked whether a) fire changes functional composition (mean and dissimilarity of trait values) at the community level; and b) such fire-induced functional modification is driven by changes in the relative abundance-dominance of species or by a replacement of species with different traits. We sampled ant communities in burned and unburned plots along 22 sites in a western Mediterranean region, and we computed two complementary functional trait composition indices ('trait average' and 'trait dissimilarity') for 12 functional traits (related to resource exploitation, social structure and reproduction) and with two different datasets varying in the way species abundance is considered (i.e. abundance and occurrence data). Our results suggest a set of functional responses that seem to be related to direct mortality by fire as well as to indirect fire-induced modifications in environmental conditions relevant for ants. Trait average of colony size, worker size, worker polymorphism and the ratio between queen and worker size, as well as the trait dissimilarity of the proportion of behaviorally dominant species and of liquid food consumption, and overall functional diversity, were higher in burned than in unburned areas. Interestingly, different patterns arise when comparing results from abundance and occurrence data. While the response to fire in trait averages is quite similar, in the case of trait dissimilarity, the higher values in response to fire are much more marked when considering occurrence rather than abundance data. Our results suggest that changes in trait average are driven at the same time by replacement of species with different traits and by changes in the relative abundance-dominance of species, while fire promotes a higher diversity of functions that is primarily driven by rare species that are functionally unique. Overall, we observed major fire-induced changes in functional composition in Mediterranean ant communities that might have relevant consequences for ecosystem processes and services. © 2013 The Authors.
Arnan X., Quevedo L., Rodrigo A. (2013) Forest fire occurrence increases the distribution of a scarce forest type in the Mediterranean Basin. Acta Oecologica. 46: 39-47.LinkDoi: 10.1016/j.actao.2012.10.005
Here we report how fire recurrence increases the distribution of a scarce forest type in NE Spain that is dominated by the resprouter tree species Arbutus unedo. We used a combination of GIS and field surveys to determine the effect of fire and pre-fire vegetation on the appearance of A. unedo forests. In the field, we also analyzed the factors that promote fire and lead to the appearance of A. unedo forests. Our results reveal an increased occurrence of A. unedo forests in NE Spain in recent years; this phenomenon was strongly related to fire recurrence and the vegetation type present prior to fire. Most Pinus halepensis forests that burned more than once gave rise to A. unedo forests. Our results indicate that these conversions were related to a reduction in pine density coupled with increases in the density and size of A. unedo trees due to recurrent fires. Given that fires are increasing in number and magnitude in the Mediterranean, we predict a major change in landscape structure and composition at the regional scale. © 2012 Elsevier Masson SAS.
Lazaro-Gonzalez A., Arnan X., Boulay R., Cerda X., Rodrigo A. (2013) Short-term ecological and behavioural responses of Mediterranean ant species Aphaenogaster gibbosa (Latr. 1798) to wildfire. Insect Conservation and Diversity. 6: 627-638.LinkDoi: 10.1111/icad.12018
Fire greatly affects plant and animal biodiversity. There is an extensive body of literature on the effects of fire on insect communities, in which a large variability of responses has been observed. Very few studies, however, have addressed functional responses at the species level, information that would greatly enhance our understanding of the impact of fire at higher organisational levels. The aim of this study is to analyse the short-term ecological and behavioural responses of the Mediterranean ant Aphaenogaster gibbosa to fire-induced environmental changes. We compared aspects of the abiotic and biotic environment relevant to this species, as well as differences in colony foraging behaviour, on unburnt and burnt plots in a Mediterranean area that was affected by a wildfire. Our results showed that fire modified plant cover around nests and daily cycles of soil temperature close to the nest. Although there were no significant differences in food quantity, food quality (particularly seed composition) was different between unburnt and burnt plots. In accordance with these environmental changes, we found significant differences in the daily activity rhythms and diet composition of A. gibbosa between unburnt and burnt plots. Overall, these differences did not result in significant changes in overall foraging activity and efficiency, allowing ant colonies to maintain the same food intake regardless of the habitat they occupied. We conclude that A. gibbosa uses behavioural plasticity to modify its foraging strategy in recently burnt environments and thus survive post-fire conditions. © 2013 The Royal Entomological Society.
Pino J., Arnan X., Rodrigo A., Retana J. (2013) Post-fire invasion and subsequent extinction of Conyza spp. in Mediterranean forests is mostly explained by local factors. Weed Research. 53: 470-478.LinkDoi: 10.1111/wre.12040
This work explored the invasion patterns of Conyza species in Mediterranean pine forests after fire and identified their main correlates through a temporal study approach. We hypothesised that wildfires might favour Conyza spp. invasion in these forests, but only transiently and depending on fire regime. We recorded Conyza spp. invasion and subsequent extinction in plots from species' occurrence and cover in vegetation surveys. We also explored the association of Conyza spp. presence and cover with a set of climatic, landscape and local (plot) factors using GLZ and GLM. We assessed changes in significant factors over time with a Wilcoxon test for paired samples. Evidence for Conyza spp. establishment was found in two-thirds of the study plots, with an invasion peak 2 years after fire. Local factors related to resource availability, including high fire severity, low soil stoniness and total vegetation cover and high herbaceous cover, were significantly correlated with Conyza occurrence in plots at the invasion peak. However, Conyza cover was always low (≤6%) and populations did not persist more than several years, thus becoming rarer as plant cover increased. Landscape and climatic factors showed no association with Conyza occurrence. In conclusion, wildfires favour transient invasion of European Mediterranean pine forests by Conyza spp. Invasion is mostly enhanced by local fire severity and constrained by subsequent vegetation recovery, while it is poorly explained by climate and landscape, either current or historical. © 2013 European Weed Research Society.
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