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- Unravelling the small‐island effect through phylogenetic community ecologyPublication . Matthews, Thomas J.; Rigal, François; Kougioumoutzis, Kostas; Trigas, Panayiotis; Triantis, KonstantinosAIM: The small-island effect (SIE) describes a different relationship between island area and species richness on smaller compared to larger islands. The pattern has recently gained widespread support. However, few studies have attempted to identify the actual mechanisms driving the SIE. Here, we use a phylogenetic community framework to study the SIE, based on the assumption that if the dominant assembly processes differ between small and large islands, patterns of phylogenetic community structure should shift across the area and habitat diversity gradient. LOCATION: The Aegean Archipelago, Greece. TAXON: Plants. METHODS: We used a large dataset of 3,262 vascular plant species distributed across 173 islands, in combination with a species-level phylogeny. The phylogenetic community structure of each island was calculated using a null modelling framework and was quantified using effect sizes (ES); negative values indicating phylogenetic clustering and positive values overdispersion. Habitat diversity, species richness, phylogenetic diversity (PD) and ES values were regressed against log10-transformed area and we tested for a SIE using piecewise regression models. We also assessed differences in taxonomic and phylogenetic composition between small and large islands using a beta diversity framework. RESULTS: We found evidence of a SIE using species richness, PD and phylogenetic community structure (ES values). Small islands displayed low variation in habitat diversity and tended to be more phylogenetically clustered, while large islands shifted from phylogenetic clustering towards phylogenetic overdispersion with increasing area and habitat diversity. In addition, we showed that phylogenetic composition tended to be more similar between small islands than expected. MAIN CONCLUSION: Overall, our results provide an example of a SIE in the analysis of island phylogenetic community structure, and point to a role of habitat diversity in driving the SIE more generally.
- Increase of insular exotic arthropod diversity is a fundamental dimension of the current biodiversity crisisPublication . Borges, Paulo A. V.; Rigal, François; Ros‐Prieto, A.; Cardoso, PedroA dramatic insect decline has been documented on the grasslands and forests of European or North American mainland. Yet, other parts of the world and other ecosystems remain much less studied with unknown patterns. Using a unique time-series dataset, we investigate recent trends on abundance and richness of arthropods sampled in Azorean native forest over 6 years (2013–2018). We test the hypothesis that biodiversity erosion drivers are changing the diversity and relative species abundance structure (species abundance distribution, SAD) of endemics, native non‐endemics and exotic species over time. We also examine temporal trends in abundance for each individual species. In contrast with mainland studies, we observed no decline in overall arthropod diversity, but a clear increase in the diversity of exotic arthropods and some evidence of a tendency for decreasing abundance for some endemic species. We also document stronger species turnover for exotic species, but no specific changes in the SAD. We argue that many changes, particularly in unique systems such as islands, will be noticed not at the richness but mostly at compositional level. Special attention should be given to exotic species which are known to be one of the major drivers of biodiversity erosion on islands.
- Medium-term effects of Ag supplied directly or via sewage sludge to an agricultural soil on Eisenia fetida earthworm and soil microbial communitiesPublication . Courtois, Pauline; Rorat, Agnieszka; Lemiere, Sébastien; Guyoneaud, Rémy; Attard, Eléonore; Longepierre, Manon; Rigal, François; Levard, Clément; Chaurand, Perrine; Grosser, Anna; Grobelak, Anna; Kacprzak, Malgorzata; Lors, Christine; Richaume, Agnès; Vandenbulcke, FranckThe widespread use of silver nanoparticles (AgNPs) in consumer products that release Ag throughout their life cycle has raised potential environmental concerns. AgNPs primarily accumulate in soil through the spreading of sewage sludge (SS). In this study, the effects of direct exposure to AgNPs or indirect exposure via SS contaminated with AgNPs on the earthworm Eisenia fetida and soil microbial communities were compared, through 3 scenarios offering increasing exposure concentrations. The effects of Ag speciation were analyzed by spiking SS with AgNPs or AgNO₃ before application to soil. SS treatment strongly impacted Ag speciation due to the formation of Ag₂S species that remained sulfided after mixing in the soil. The life traits and expression of lysenin, superoxide dismutase, cd-metallothionein genes in earthworms were not impacted by Ag after 5 weeks of exposure, but direct exposure to Ag without SS led to bioaccumulation of Ag, suggesting transfer in the food chain. Ag exposure led to a decrease in potential carbon respiration only when directly added to the soil. The addition of SS had a greater effect on soil microbial diversity than the form of Ag, and the formation of Ag sulfides in SS reduced the impact of AgNPs on E. fetida and soil microorganisms compared with direct addition.
- A global analysis of avian island diversity–area relationships in the AnthropocenePublication . Matthews, Thomas J.; Wayman, Joseph P.; Whittaker, Robert J.; Cardoso, Pedro; Hume, Julian P.; Sayol, Ferran; Proios, Konstantinos; Martin, Thomas E.; Baiser, Benjamin; Borges, Paulo A. V.; Kubota, Yasuhiro; dos Anjos, Luiz; Tobias, Joseph; Soares, Filipa C.; Si, Xingfeng; Ding, Ping; Mendenhall, Chase D.; Sin, Yong Chee Keita; Rheindt, Frank E.; Triantis, Kostas; Guilhaumon, François; Watson, David M.; Brotons, Lluís; Battisti, Corrado; Chu, Osanna; Rigal, FrançoisResearch on island species–area relationships (ISAR) has expanded to incorporate functional (IFDAR) and phylogenetic (IPDAR) diversity. However, relative to the ISAR, we know little about IFDARs and IPDARs, and lack synthetic global analyses of variation in form of these three categories of island diversity–area relationship (IDAR). Here, we undertake the first comparative evaluation of IDARs at the global scale using 51 avian archipelagic data sets representing true and habitat islands. Using null models, we explore how richness-corrected functional and phylogenetic diversity scale with island area. We also provide the largest global assessment of the impacts of species introductions and extinctions on the IDAR. Results show that increasing richness with area is the primary driver of the (non-richness corrected) IPDAR and IFDAR for many data sets. However, for several archipelagos, richness-corrected functional and phylogenetic diversity changes linearly with island area, suggesting that the dominant community assembly processes shift along the island area gradient. We also find that archipelagos with the steepest ISARs exhibit the biggest differences in slope between IDARs, indicating increased functional and phylogenetic redundancy on larger islands in these archipelagos. In several cases introduced species seem to have ‘re-calibrated’ the IDARs such that they resemble the historic period prior to recent extinctions.
- Automated Discovery of Relationships, Models, and Principles in EcologyPublication . Cardoso, Pedro; Veiga Branco, Vasco; Borges, Paulo A.V.; Carvalho, José Carlos; Rigal, François; Gabriel, Rosalina; Mammola, Stefano; Cascalho, José Manuel; Correia, LuísEcological systems are the quintessential complex systems, involving numerous high-order interactions and non-linear relationships. The most used statistical modeling techniques can hardly accommodate the complexity of ecological patterns and processes. Finding hidden relationships in complex data is now possible using massive computational power, particularly by means of artificial intelligence and machine learning methods. Here we explored the potential of symbolic regression (SR), commonly used in other areas, in the field of ecology. Symbolic regression searches for both the formal structure of equations and the fitting parameters simultaneously, hence providing the required flexibility to characterize complex ecological systems. Although the method here presented is automated, it is part of a collaborative human–machine effort and we demonstrate ways to do it. First, we test the robustness of SR to extreme levels of noise when searching for the species-area relationship. Second, we demonstrate how SR can model species richness and spatial distributions. Third, we illustrate how SR can be used to find general models in ecology, namely new formulas for species richness estimators and the general dynamic model of oceanic island biogeography. We propose that evolving free-form equations purely from data, often without prior human inference or hypotheses, may represent a very powerful tool for ecologists and biogeographers to become aware of hidden relationships and suggest general theoretical models and principles.
- The role of ecological specialization in shaping patterns of insular communitiesPublication . Sfenthourakis, Spyros; Triantis, Kostas A.; Proios, Konstantinos; Rigal, FrançoisAIM: Research on the response of species richness to area and environmental heterogeneity so far has not addressed possible effects of species’ differences in ecological specialization. Herein we provide a new metric, ‘ecorichness’, in an attempt to fill this gap. LOCATION: Aegean islands (Greece). TAXON: Terrestrial isopods. METHODS: ‘Ecorichness’ estimates an island's biodiversity by integrating species richness and the specialists-generalists spectrum. We calculated ‘ecorichness’ for terrestrial isopods from 43 Aegean islands based on the habitats they exploit. ‘Ecorichness’ then was regressed on area, habitat diversity and the K-parameter of the Choros model using linear and quadratic models, compared based on AICc. A reduced data set, without halophilous species and coastal habitats, as well as an alternative description of habitat diversity, also was explored. The small island effect (SIE) thresholds identified using both a path analysis approach and piecewise continuous linear models were compared to the area of maximum ‘ecorichness’. RESULTS: ‘Ecorichness’ response to area and habitat heterogeneity was best fitted by quadratic models with peaks located at an area similar to the SIE threshold identified by path analysis. Different measures of habitat diversity produced similar patterns. Exclusion of coastal species and habitats shows that the response of ‘ecorichness’ to area is mostly shaped by the increasing contribution of specialists in the assemblages of larger islands. MAIN CONCLUSIONS: ‘Ecorichness’ facilitates exploration of the role of ecological specialization in shaping community patterns. It can be applied to different community data sets, whenever habitat range exploitation can be quantified. Results from the case study accord with previous suggestions that the relative contribution of generalists and specialists differs between small and large island communities.
- Atmospheric iodine, selenium and caesium depositions in France : I. Spatial and seasonal variationsPublication . Roulier, Marine; Bueno, Maïté; Coppin, Frédéric; Nicolas, Manuel; Thiry, Yves; Rigal, François; Le Hécho, Isabelle; Pannier, FlorenceThe spatial distribution and seasonal variations of atmospheric iodine (I), selenium (Se) and caesium (Cs) depositions remain unclear and this precludes adequate inputs for biogeochemical models. We quantified total concentrations and fluxes of these elements in rainfalls from 27 monitoring sites in France with contrasted climatic conditions; monthly measurements were taken over one year (starting in 2016/09). Since speciation of I and Se can impact their behaviour in the environment, analysis of their inorganic compounds was also conducted. Our results showed that annual I concentrations in rainfall were much higher than those of Se and Cs (annual means = 1.56, 0.044 and 0.005 μg L-1, respectively). The annual iodine concentrations were highly positively correlated with those of marine elements (i.e. Na, Cl and Mg), involving higher I concentrations under oceanic climate than for transition, continental and mountainous ones. Furthermore, common patterns were found between Se concentrations and both marine and terrestrial components consistent with the various sources of Se in atmosphere. The association of Cs with two anthropogenic components (i.e. NH4+ and NO3-) used in agriculture supports the hypothesis of its terrestrial origin (i.e. from atmospheric dusts) in rainfall. We found higher rainfall concentrations of I during the warmest months for all climates. However, no specific seasonal trend occurred for Se and Cs. On annual average, rainfall contained mostly unidentified selenium compounds (inorganic Se proportions = 25-54%) and equal proportions of inorganic and unidentified I compounds. Concentrations of iodate were higher under oceanic climate consistent with an iodine marine-origin.
- Macaronesia as a Fruitful Arena for Ecology, Evolution, and Conservation BiologyPublication . Florencio, Margarita; Patiño, Jairo; Nogué, Sandra; Traveset, Anna; Borges, Paulo A. V.; Schaefer, Hanno; Amorim, Isabel R.; Arnedo, Miquel A.; Ávila, Sérgio P.; Cardoso, Pedro; Nascimento, Lea; Fernández-Palacios, José María; Gabriel, Sofia I.; Gil, Artur José Freire; Gonçalves, Vitor; Haroun, Ricardo J.; Illera, Juan Carlos; López-Darias, Marta; Martínez, Alejandro; Martins, Gustavo M.; Neto, Ana I.; Nogales, Manuel; Oromí, Pedro; Rando, Juan Carlos; Raposeiro, Pedro M.; Rigal, François; Romeiras, Maria M.; Silva, Luís; Valido, Alfredo; Vanderpoorten, Alain; Vasconcelos, Raquel; Santos, Ana M. C.Research in Macaronesia has led to substantial advances in ecology, evolution and conservation biology. We review the scientific developments achieved in this region, and outline promising research avenues enhancing conservation. Some of these discoveries indicate that the Macaronesian flora and fauna are composed of rather young lineages, not Tertiary relicts, predominantly of European origin. Macaronesia also seems to be an important source region for back-colonisation of continental fringe regions on both sides of the Atlantic. This group of archipelagos (Azores, Madeira, Selvagens, Canary Islands, and Cabo Verde) has been crucial to learn about the particularities of macroecological patterns and interaction networks on islands, providing evidence for the development of the General Dynamic Model of oceanic island biogeography and subsequent updates. However, in addition to exceptionally high richness of endemic species, Macaronesia is also home to a growing number of threatened species, along with invasive alien plants and animals. Several innovative conservation and management actions are in place to protect its biodiversity from these and other drivers of global change. The Macaronesian Islands are a well-suited field of study for island ecology and evolution research, mostly due to its special geological layout with 40 islands grouped within five archipelagos differing in geological age, climate and isolation. A large amount of data is now available for several groups of organisms on and around many of these islands. However, continued efforts should be made toward compiling new information on their biodiversity, to pursue various fruitful research avenues and develop appropriate conservation management tools.
- Spatial Scaling Patterns of Functional DiversityPublication . Alirezazadeh, Saeid; Borges, Paulo A. V.; Cardoso, Pedro; Gabriel, Rosalina; Rigal, François; Borda-de-Água, LuísEcology, biogeography and conservation biology, among other disciplines, often rely on species identity, distribution and abundance to perceive and explain patterns in space and time. Yet, species are not independent units in the way they interact with their environment. Species often perform similar roles in networks and their ecosystems, and at least partial redundancy or difference of roles might explain co-existence, competitive exclusion or other patterns reflected at the community level. Therefore, considering species traits, that is, the organisms’ functional properties that interact with the environment, might be of utmost importance in the study of species relative abundances. Several descriptive measures of diversity, such as the species-area relationship (SAR) and the species abundance distribution (SAD), have been used extensively to characterize the communities and as a possible window to gain insight into underlying processes shaping and maintaining biodiversity. However, if the role of species in a community is better assessed by their functional attributes, then one should also study the SAR and the SAD by using trait-based approaches, and not only taxonomic species. Here we merged species according to their similarity in a number of traits, creating functional units, and used these new units to study the equivalent patterns of the SAR and of the SAD (functional units abundance distributions - FUADs), with emphasis on their spatial scaling characteristics. This idea was tested using data on arthropods collected in Terceira island, in the Azorean archipelago. Our results showed that diversity scales differently depending on whether we use species or functional units. If what determines species communities’ dynamics is their functional diversity, then our results suggest that we may need to revaluate the commonly assumed patterns of species diversity and, concomitantly, the role of the underlying processes.
- Habitat filtering and inferred dispersal ability condition across‐scale species turnover and rarity in Macaronesian island spider assemblagesPublication . Malumbres-Olarte, Jagoba; Rigal, François; Girardello, Marco; Cardoso, Pedro; Crespo, Luís C.; Amorim, Isabel R.; Arnedo, Miquel A.; Boieiro, Mário; Carvalho, José Carlos; Carvalho, Rui; Gabriel, Rosalina; Lamelas-López, Lucas; López, Heriberto; Paulo, Octávio S.; Pereira, Fernando; Pérez‐Delgado, Antonio J.; Rego, Carla; Romeiras, Maria M.; Ros-Prieto, Alejandra; Oromí, Pedro; Vieira, Ana; Emerson, Brent C.; Borges, Paulo A. V.AIM: Habitat diversity has been linked to the diversity and structure of island communities, however, little is known about patterns and processes within habitats. Here we aim to determine the contributions of habitat type and inferred dispersal frequency to the differences in taxonomic structure between assemblages in the same island habitat. LOCATION: The Macaronesian archipelagos (Azores, Madeira, the Canary Islands and Cabo Verde). TAXON: Spiders (Araneae). METHODS: We established forest and dry habitat sites (each with five plots) on two islands per archipelago. We collected spiders using standardised sampling protocols. We tested the differences in beta diversity separately for each habitat and for each inferred category of ballooning (an aerial dispersal strategy) frequency across geographic scales through nested non-parametric permutational multivariate analyses of variance. We then tested whether ballooning and habitat influenced heterogeneity in species composition (dispersion in beta diversity) in the two habitat types. We analysed the effects of habitat and ballooning on species abundance distribution (SAD) and rarity by fitting Gambin models and evaluating the contribution of ballooning categories to SAD. RESULTS: Communities of the same archipelago and habitat were taxonomically more similar, and beta diversity increased with geographic scale, being greater in dry habitats. There was greater species replacement among assemblages in dry habitats than in forests, with greater differences for rare ballooners. There were no differences in SAD between habitats although dry habitat sites seemed to harbour more species with low abundances (rare species) than forests. MAIN CONCLUSIONS: Habitat type does not only condition the differences between spider assemblages of the same habitat but also the scale at which they occur. These differences may be determined by the heterogeneity in the physical structure of each habitat as well as how much this structure facilitates aerial dispersal (ballooning), and should be considered in theories/hypotheses on island community assembly as well as in conservation strategies.
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