Arnan X., Andersen A.N., Gibb H., Parr C.L., Sanders N.J., Dunn R.R., Angulo E., Baccaro F.B., Bishop T.R., Boulay R., Castracani C., Cerdá X., Toro I.D., Delsinne T., Donoso D.A., Elten E.K., Fayle T.M., Fitzpatrick M.C., Gómez C., Grasso D.A., Grossman B.F., Guénard B., Gunawardene N., Heterick B., Hoffmann B.D., Janda M., Jenkins C.N., Klimes P., Lach L., Laeger T., Leponce M., Lucky A., Majer J., Menke S., Mezger D., Mori A., Moses J., Munyai T.C., Paknia O., Pfeiffer M., Philpott S.M., Souza J.L.P., Tista M., Vasconcelos H.L., Retana J. (2018) Dominance–diversity relationships in ant communities differ with invasion. Global Change Biology. 24: 4614-4625.EnlaceDoi: 10.1111/gcb.14331
The relationship between levels of dominance and species richness is highly contentious, especially in ant communities. The dominance-impoverishment rule states that high levels of dominance only occur in species-poor communities, but there appear to be many cases of high levels of dominance in highly diverse communities. The extent to which dominant species limit local richness through competitive exclusion remains unclear, but such exclusion appears more apparent for non-native rather than native dominant species. Here we perform the first global analysis of the relationship between behavioral dominance and species richness. We used data from 1,293 local assemblages of ground-dwelling ants distributed across five continents to document the generality of the dominance-impoverishment rule, and to identify the biotic and abiotic conditions under which it does and does not apply. We found that the behavioral dominance–diversity relationship varies greatly, and depends on whether dominant species are native or non-native, whether dominance is considered as occurrence or relative abundance, and on variation in mean annual temperature. There were declines in diversity with increasing dominance in invaded communities, but diversity increased with increasing dominance in native communities. These patterns occur along the global temperature gradient. However, positive and negative relationships are strongest in the hottest sites. We also found that climate regulates the degree of behavioral dominance, but differently from how it shapes species richness. Our findings imply that, despite strong competitive interactions among ants, competitive exclusion is not a major driver of local richness in native ant communities. Although the dominance-impoverishment rule applies to invaded communities, we propose an alternative dominance-diversification rule for native communities. © 2018 John Wiley & Sons Ltd
Gibb H., Sanders N.J., Dunn R.R., Arnan X., Vasconcelos H.L., Donoso D.A., Andersen A.N., Silva R.R., Bishop T.R., Gomez C., Grossman B.F., Yusah K.M., Luke S.H., Pacheco R., Pearce-Duvet J., Retana J., Tista M., Parr C.L. (2018) Habitat disturbance selects against both small and large species across varying climates. Ecography. 41: 1184-1193.EnlaceDoi: 10.1111/ecog.03244
Global extinction drivers, including habitat disturbance and climate change, are thought to affect larger species more than smaller species. However, it is unclear if such drivers interact to affect assemblage body size distributions. We asked how these two key global change drivers differentially affect the interspecific size distributions of ants, one of the most abundant and ubiquitous animal groups on earth. We also asked whether there is evidence of synergistic interactions and whether effects are related to species’ trophic roles. We generated a global dataset on ant body size from 333 local ant assemblages collected by the authors across a broad range of climates and in disturbed and undisturbed habitats. We used head length (range: 0.22–4.55 mm) as a surrogate of body size and classified species to trophic groups. We used generalized linear models to test whether body size distributions changed with climate and disturbance, independent of species richness. Our analysis yielded three key results: 1) climate and disturbance showed independent associations with body size; 2) assemblages included more small species in warmer climates and fewer large species in wet climates; and 3) both the largest and smallest species were absent from disturbed ecosystems, with predators most affected in both cases. Our results indicate that temperature, precipitation and disturbance have differing effects on the body size distributions of local communities, with no evidence of synergistic interactions. Further, both large and small predators may be vulnerable to global change, particularly through habitat disturbance. © 2017 The Authors
Doblas-Miranda, E., Alonso, R., Arnan, X., Bermejo, V., Brotons, L., de las Heras, J., Estiarte, M., Hódar, J.A., Llorens, P., Lloret, F., López-Serrano, F.R., Martínez-Vilalta, J., Moya, D., Peñuelas, J., Pino, J., Rodrigo, A., Roura-Pascual, N., Valladares, F., Vilà, M., Zamora, R., Retana, J. (2017) A review of the combination among global change factors in forests, shrublands and pastures of the Mediterranean Region: Beyond drought effects. Global and Planetary Change. 148: 42-54.EnlaceDoi: 10.1016/j.gloplacha.2016.11.012
Arnan X., Cerdá X., Retana J. (2016) Relationships among taxonomic, functional, and phylogenetic ant diversity across the biogeographic regions of Europe. Ecography. : 0-0.EnlaceDoi: 10.1111/ecog.01938
Understanding how different biodiversity components are related across different environmental conditions is a major goal in macroecology and conservation biogeography. We investigated correlations among alpha and beta taxonomic (TD), phylogenetic (PD), and functional diversity (FD) in ant communities in the five biogeographic regions most representative of western Europe; we also examined the degree of niche conservatism. We combined data from 349 ant communities composed of 154 total species, which were characterized by 10 functional traits and by phylogenetic relatedness. We computed TD, PD, and FD using the Rao quadratic entropy index, which allows each biodiversity component to be partitioned into α and β diversity within the same mathematical framework. We ran generalized least squares and multiple matrix regressions with randomization to investigate relationships among the diversity components. We used Pagel's λ test to explore niche conservatism in each biogeographic region. At the alpha scale, TD was consistently, positively related to PD and FD, although the strength and scatter of this relationship changed among the biogeographic regions. Meanwhile, PD and FD consistently matched up across regions. Accordingly, we found similar degrees of niche conservatism across regions. Nonetheless, these alpha-scale relationships had low coefficients of determination. At the beta scale, the three diversity components were highly correlated across all regions (especially TD and FD, as well as PD and FD). Our results imply that the different diversity components, and especially PD and FD, are consistently related across biogeographic regions and analytical scale. However, the alpha-scale relationships were quite weak, suggesting environmental factors might influence the degree of association among diversity components at the alpha level. In conclusion, conservation programs should seek to preserve functional and phylogenetic diversity in addition to species richness, and this approach should be applied universally, regardless of the biogeographic locations of the sites to be protected. © 2016 Nordic Society Oikos.
Parr, C.L., Dunn, R.R., Sanders, N.J., Weiser, M.D., Photakis, M., Bishop, T.R., Fitzpatrick, M.C., Arnan, X., Baccaro, F., Brandão, C.R.F., Chick, L., Donoso, D.A., Fayle, T.M., Gómez, C., Grossman, B., Munyai, T.C., Pacheco, R., Retana, J., Robinson, A., Sagata, K., Silva, R.R., Tista, M., Vasconcelos, H., Yates, M., Gibb, H. (2016) GlobalAnts: A new database on the geography of ant traits (Hymenoptera: Formicidae). Insect Conservation and Diversity. : 0-0.EnlaceDoi: 10.1111/icad.12211
Arnan X., Cerdá X., Retana J. (2015) Partitioning the impact of environment and spatial structure on alpha and beta components of taxonomic, functional, and phylogenetic diversity in European ants. PeerJ. 2015: 0-0.EnlaceDoi: 10.7717/peerj.1241
We analyze the relative contribution of environmental and spatial variables to the alpha and beta components of taxonomic (TD), phylogenetic (PD), and functional (FD) diversity in ant communities found along different climate and anthropogenic disturbance gradients across western and central Europe, in order to assess the mechanisms structuring ant biodiversity. To this aim we calculated alpha and beta TD, PD, and FD for 349 ant communities, which included a total of 155 ant species; we examined 10 functional traits and phylogenetic relatedness. Variation partitioning was used to examine how much variation in ant diversity was explained by environmental and spatial variables. Autocorrelation in diversity measures and each trait's phylogenetic signal were also analyzed.We found strong autocorrelation in diversity measures. Both environmental and spatial variables significantly contributed to variation in TD, PD, and FD at both alpha and beta scales; spatial structure had the larger influence. The different facets of diversity showed similar patterns along environmental gradients. Environment explained a much larger percentage of variation in FD than in TD or PD. All traits demonstrated strong phylogenetic signals. Our results indicate that environmental filtering and dispersal limitations structure all types of diversity in ant communities. Strong dispersal limitations appear to have led to clustering of TD, PD, and FD in western and central Europe, probably because different historical and evolutionary processes generated different pools of species. Remarkably, these three facets of diversity showed parallel patterns along environmental gradients. Trait-mediated species sorting and niche conservatism appear to structure ant diversity, as evidenced by the fact that more variation was explained for FD and that all traits had strong phylogenetic signals. Since environmental variables explained much more variation in FD than in PD, functional diversity should be a better indicator of community assembly processes than phylogenetic diversity. © 2015 Arnan et al.
Retana J., Arnan X., Cerdá X. (2015) A multidimensional functional trait analysis of resource exploitation in European ants. Ecology. 96: 2781-2793.EnlaceDoi: 10.1890/14-2326.1
The major factors explaining ecological variation in plants have been widely discussed over the last decade thanks to numerous studies that have examined the covariation that exists between pairs of traits. However, multivariate relationships among traits remain poorly characterized in animals. In this study, we aimed to identify the main multivariate trait dimensions that explain variance in important functional traits related to resource exploitation in ants. To this end, we created a large ant trait database. This database includes information on 11 traits that are important in ant resource exploitation; data were obtained for 150 European species found in different biomes. First, we examined the pairwise correlations between the traits included in the database. Second, we used multivariate analyses to identify potential trait dimensions. Our study shows that, to a great extent, resource exploitation strategies align along two main trait dimensions. The first dimension emerged in both the overall and group-specific analyses, where it accounted for the same pairwise trait correlations. The second dimension was more variable, as species were grouped by levels of taxonomy, habitat, and climate. These two dimensions included most of the significant pairwise trait correlations, thus highlighting that complementarity, but also redundancy, exists among different pairs of traits. The first dimension was associated with behavioral dominance: dominance was associated with large colony size, presence of multiple nests per colony, worker polymorphism, and a collective foraging strategy. The second dimension was associated with resource partitioning along dietary and microhabitat lines: it ranged from species that consume liquid foods, engage in group foraging, and mainly nest in the vegetation to species that consume insects and seeds, engage in individual foraging, and demonstrate strictly diurnal activity. Our findings establish a proficient ecological trait-based animal research that minimizes the number of traits to be measured while maximizing the number of relevant trait dimensions. Overall, resource exploitation in animals might be framed by behavioral dominance, foraging strategy, diet, and nesting habitat; the position of animal species within this trait space could provide relevant information about their distribution and abundance, for today as well as under future global change scenarios. © 2015 by the Ecological Society of America.
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.EnlaceDoi: 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., Cerda X., Rodrigo A., Retana J. (2013) Response of ant functional composition to fire. Ecography. 36: 1182-1192.EnlaceDoi: 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.
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.EnlaceDoi: 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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