TerrANTALife 1.0 Biodiversity data checklist of known Antarctic terrestrial and freshwater life forms.

Background: Incomplete species inventories for Antarctica represent a key challenge for comprehensive ecological research and conservation in the region. Additionally, data required to understand population dynamics, rates of evolution, spatial ranges, functional traits, physiological tolerances and species interactions, all of which are fundamental to disentangle the different functional elements of Antarctic biodiversity, are mostly missing. However, much of the fauna, flora and microbiota in the emerged ice-free land of the continent have an uncertain presence and/or unresolved status, with entire biodiversity compendia of prokaryotic groups (e.g. bacteria) being missing. All the available biodiversity information requires consolidation, cross-validation, re-assessment and steady systematic inclusion in order to create a robust catalogue of biodiversity for the continent. New information: We compiled, completed and revised eukaryotic species inventories present in terrestrial and freshwater ecosystems in Antarctica in a new living database: terrANTALife (version 1.0). The database includes the first integration in a compendium for many groups of eukaryotic microorganisms. We also introduce a first catalogue of amplicon sequence variants (ASVs) of prokaryotic biodiversity. Available compendia and literature to date were searched for Antarctic terrestrial and freshwater species, integrated, taxonomically harmonised and curated by experts to create comprehensive checklists of Antarctic organisms. The final inventories comprises 470 animal species (including vertebrates, free-living invertebrates and parasites), 306 plants (including all Viridiplantae: embryophytes and green algae), 997 fungal species and 434 protists (sensu lato). We also provide a first account for many groups of microorganisms, including non-lichenised fungi and multiple groups of eukaryotic unicellular species (Stramenophila, Alveolata and Rhizaria (SAR), Chromists and Amoeba), jointly referred to as "protists". In addition, we identify 1753 bacterial (obtained from 348117 ASVs) and 34 archaeal genera (from 1848 ASVs), as well as, at least, 14 virus families. We formulate a basic tree of life in Antarctica with the main lineages listed in the region and their "known-accepted-species" numbers.

Dung removal increases under higher dung beetle functional diversity regardless of grazing intensification.

Dung removal by macrofauna such as dung beetles is an important process for nutrient cycling in pasturelands. Intensification of farming practices generally reduces species and functional diversity of terrestrial invertebrates, which may negatively affect ecosystem services. Here, we investigate the effects of cattle-grazing intensification on dung removal by dung beetles in field experiments replicated in 38 pastures around the world. Within each study site, we measured dung removal in pastures managed with low- and high-intensity regimes to assess between-regime differences in dung beetle diversity and dung removal, whilst also considering climate and regional variations. The impacts of intensification were heterogeneous, either diminishing or increasing dung beetle species richness, functional diversity, and dung removal rates. The effects of beetle diversity on dung removal were more variable across sites than within sites. Dung removal increased with species richness across sites, while functional diversity consistently enhanced dung removal within sites, independently of cattle grazing intensity or climate. Our findings indicate that, despite intensified cattle stocking rates, ecosystem services related to decomposition and nutrient cycling can be maintained when a functionally diverse dung beetle community inhabits the human-modified landscape.

Exploring the impact of data curation criteria on the observed geographical distribution of mosses.

Biodiversity data records contain inaccuracies and biases. To overcome this limitation and establish robust geographic patterns, ecologists often curate records keeping those that are most suitable for their analyses. Yet, this choice is not straightforward and the outcome of the analysis may vary due to a trade-off between data quality and volume. This problem is particularly recurrent for less-studied groups with patchy sampling effort. The latitudinal pattern of mosses richness remains inconsistent across studies and these may emerge purely from sampling artefacts. Our main objective here is to assess the effect of different curation criteria on this spatial pattern in the Temperate Northern Hemisphere (above 20° latitude). We contrasted the geographical distribution of moss species records and the latitude-species richness relation obtained under different data curation scenarios. These scenarios comprehend five sources of taxonomical standardisations and eight data cleaning filters. The analyses are based on the selection of well-surveyed cells at 100 km cell resolution. The application of some 'data curation scenarios' severely affects the number of records selected for analysis and substantially changes the proportion of richness per cell. The sensitivity to data curation becomes detectable at regional and at the cell scales showing a large shift in the latitudinal richness peak in Europe, from 60° N to 45° N latitude, when only preserved specimens are selected and duplicates based on date of collection and coordinates are excluded. Our results stress the importance of justifying the criteria used for filtering biodiversity data retrieved from biodiversity databases to avoid detecting misleading patterns. Curating records under particular criteria compromises the information in some areas displaying different spatial information of mosses. This problem can be ameliorated if data filtering is combined with identifying well-surveyed cells, render relatively constant results under different combinations of filtering even for less well-known groups such as mosses.

Forest conversion into pasture selects dung beetle traits at different biological scales depending on species pool composition.

The conversion of forests into open areas has large effects on the diversity and structure of native communities. The intensity of these effects may vary between regions, depending on the existence of native species adapted to open habitats in the regional pool or the time since habitat change.We assess the differences in species richness and functional diversity of dung beetle communities (Coleoptera: Scarabaeinae) between native forests and novel pasturelands of the Atlantic Forest and the Cerrado, two biomes with contrasting histories of human occupation in Brazil. We conducted standardized surveys in seven forest fragments and adjacent pastures in each region and measured 14 traits in individuals collected in each type of habitat at each particular site. We calculated functional richness, functional evenness, functional divergence, and community-weighted mean of traits for each area, and analyzed individual variation through nested variance decomposition and Trait Statistics.Communities were richer and more numerous at the Cerrado. We did not find any consistent relationship between functional diversity and forest conversion beyond the changes in species diversity. Although landscape changes were more recent at the Cerrado, the colonization of the new habitat by native species already adapted to open habitats lessens the functional loss in this biome. This indicates that habitat change's effects on trait diversity depend on the regional species pool rather than on time since land conversion.Forest conversion effects were primarily due to internal filtering. The effects of external filtering only appear at the intraspecific variance level, with contrasting differences between the Cerrado, where traits related to relocation behavior and size are selected, and the Atlantic Forest, where selection operates for traits related to relocation behavior and flight. These results evidence the importance of considering individual variance to address the responses of dung beetle communities to forest conversion.

A conversão de florestas em áreas abertas tem grandes efeitos sobre a diversidade e estrutura das comunidades nativas. A intensidade desses efeitos pode variar entre regiões, dependendo da existência de espécies nativas adaptadas a habitats abertos no pool regional ou do tempo de mudança de habitat.Avaliamos as diferenças na riqueza de espécies e diversidade funcional de comunidades de rola­bosta (Coleoptera: Scarabaeinae) entre florestas nativas e pastagens da Mata Atlântica e do Cerrado, dois biomas com histórias contrastantes de ocupação humana no Brasil. As comunidades foram amostradas em sete fragmentos florestais e pastagens adjacentes de cada região. Medimos quatorze caracteres de indivíduos coletados em cada ponto amostral de cada habitat e região. Posteriormente, calculamos a riqueza funcional, a equidade funcional, a divergência funcional e a média dos caracteres ponderada pela comunidade (CWM) para cada ponto amostral e analisamos a variação individual por meio de decomposição de variância aninhada e estatística T.As comunidades do Cerrado foram mais ricas e numerosas. Não encontramos nenhuma relação consistente entre diversidade funcional e conversão florestal além das mudanças na diversidade de espécies. Surpreendentemente, embora as mudanças na paisagem tenham sido mais recentes no Cerrado, a colonização do novo habitat por espécies nativas já adaptadas a habitats abertos diminuiu a perda funcional nos novos habitats deste bioma em comparação com a Mata Atlântica, onde as pastagens foram colonizadas apenas por espécies florestais generalistas e invasores exóticos. Isso indica que os efeitos da mudança de habitat na diversidade de caracteres dependem do conjunto regional de espécies e não do tempo desde a conversão da paisagem.Além disso, os efeitos da conversão florestal foram principalmente devidos à efeitos de filtros internos. Os efeitos de filtros externos apareceram apenas no nível de variância intraespecífica, com diferenças contrastantes entre o Cerrado, onde são selecionados caracteres relacionados ao comportamento de realocação e tamanho (Comprimento da Metatíbia, Comprimento, Volume e Largura do Pronoto) e a Mata Atlântica, onde a seleção opera em caracteres relacionadas ao comportamento de realocação e voo (Metatibia Length e Wing Load). Esses resultados evidenciam a importância de se considerar a variância individual para abordar as respostas das comunidades de rola­bostas à conversão florestal.

Geographical shifts in the successional dynamics of inland dune shrub communities.

Species' environmental requirements and large-scale spatial and evolutionary processes determine the structure and composition of local communities. However, ecological interactions also have major effects on community assembly at landscape and local scales. We evaluate whether two xerophytic shrub communities occurring in SW Portugal follow constrained ecological assembly dynamics throughout large geographical extents, or their composition is rather driven by species' individualistic responses to environmental and macroecological constraints. Inland dune xerophytic shrub communities were characterized in 95 plots. Then, we described the main gradients of vegetation composition and assessed the relevance of biotic interactions. We also characterized the habitat suitability of the dominant species, Stauracanthus genistoides, and Ulex australis, to map the potential distribution of the xerophytic shrub communities. Finally, we examined the relationships between the vegetation gradients and a broad set of explanatory variables to identify the relative importance of each factor driving changes in community composition. We found that xerophytic shrubs follow uniform successional patterns throughout the whole geographical area studied, but each community responds differently to the main environmental gradients in each region. Soil organic matter is the main determinant of community variations in the northern region, Setúbal Peninsula, whereas aridity is so in the South/South-Western region. In contrast, in the central region, Comporta, the variation between S. genistoides and U. australis communities is explained mainly by aridity and temperature seasonality, followed by the individualistic responses of the dominant species and soil organic matter. Overall, these results indicate that, the relative importance of the main factors causing community-level responses varies according to regional processes and the suitability of the environmental conditions for the dominant species in these communities. These responses are also determined by intrinsic community mechanisms that result in a high degree of similarity in the gradient-driven community stages in different regions.

A trait-based framework for dung beetle functional ecology.

Traits are key for understanding the environmental responses and ecological roles of organisms. Trait approaches to functional ecology are well established for plants, whereas consistent frameworks for animal groups are less developed. Here we suggest a framework for the study of the functional ecology of animals from a trait-based response-effect approach, using dung beetles as model system. Dung beetles are a key group of decomposers that are important for many ecosystem processes. The lack of a trait-based framework tailored to this group has limited the use of traits in dung beetle functional ecology. We review which dung beetle traits respond to the environment and affect ecosystem processes, covering the wide range of spatial, temporal and biological scales at which they are involved. Dung beetles show trait-based responses to variation in temperature, water, soil properties, trophic resources, light, vegetation structure, competition, predation and parasitism. Dung beetles' influence on ecosystem processes includes trait-mediated effects on nutrient cycling, bioturbation, plant growth, seed dispersal, other dung-based organisms and parasite transmission, as well as some cases of pollination and predation. We identify 66 dung beetle traits that are either response or effect traits, or both, pertaining to six main categories: morphology, feeding, reproduction, physiology, activity and movement. Several traits pertain to more than one category, in particular dung relocation behaviour during nesting or feeding. We also identify 136 trait-response and 77 trait-effect relationships in dung beetles. No response to environmental stressors nor effect over ecological processes were related with traits of a single category. This highlights the interrelationship between the traits shaping body-plans, the multi-functionality of traits, and their role linking responses to the environment and effects on the ecosystem. Despite current developments in dung beetle functional ecology, many knowledge gaps remain, and there are biases towards certain traits, functions, taxonomic groups and regions. Our framework provides the foundations for the thorough development of trait-based dung beetle ecology. It also serves as an example framework for other taxa.

Firefly (Coleoptera, Lampyridae) species from the Atlantic Forest hotspot, Brazil.

Background: We compiled a database of firefly species records from the Atlantic Forest hotspot in Brazil and made it available at GBIF. Data were gathered from literature and from several key entomological collections, including: Coleção entomológica Prof. José Alfredo Pinheiro Dutra (DZRJ/UFRJ) and Coleção do Laboratório de Ecologia de Insetos from Universidade Federal do Rio de Janeiro (CLEI/UFRJ); Coleção Entomológica do Instituto Oswaldo Cruz (CEIOC); Museu de Zoologia da Universidade de São Paulo (MZSP); Coleção Entomológica Pe. Jesus Santiago Moure from Universidade Federal do Paraná (DZUP/UFPR); and Coleção Entomológica from Universidade Federal Rural de Pernambuco (UFRPE). This database represents the largest contribution to a public repository of recorded occurrences from Neotropical fireflies. New information: This dataset shows the occurrence and abundance of firefly species in the Atlantic Forest hotspot. Firefly species endemic to this biome are also present and considered in the study. These data can assist scientific and societal needs, by supporting future research projects and conservation decision-making.

Moss establishment success is determined by the interaction between propagule size and species identity.

Colonization of new habitat patches is a key aspect of metacommunity dynamics, particularly for sessile organisms. Mosses can establish in new patches through fragmentation, with different vegetative structures acting as propagules. Despite the importance of these propagules for successful colonization the specific aspects that favour moss colonization by vegetative propagules remain poorly understood, including the effect of propagule size. We examine the intra- and interspecific variation of establishment and colonization success in culture of propagules of different sizes in six widespread soil moss species of contrasting growth form (Dicranum scoparium, Homalothecium aureum, Hypnum cupressiforme, Ptychostomum capillare, Syntrichia ruralis and Tortella squarrosa). We obtained three different size classes of propagules from artificially fragmented vegetative material, and assessed their establishment under controlled light and temperature conditions. We characterize the size, shape, apparent viability, morphological type and size changes due to hydration states of the propagules, all of them traits with potentially significant influence in their dispersal pattern and establishment. Then we assess the effect of these traits on moss establishment, using indicators of surface establishment (number of established shoots and colonized surface) and biomass production (viable biomass) as proxies of colonization success. The establishment indicators related to colonization surface and biomass production differ among species and propagule sizes. The magnitude of the interspecific differences of all indicators of establishment success was larger at the smaller propagule size class. T. squarrosa was the most successful species, and D. scoparium showed the lowest performance. We also found interspecific differences in the hydration dynamics of the propagules. The process of establishment by vegetative fragments operates differently among moss species. Besides, differences between hydration states in propagules of some species could be part of syndromes for both dispersal and establishment. This study unveils several functional traits relevant for moss colonization, such as wet versus dry area and length of fragments, which may improve our understanding of their spatial dynamics.

Too simple models may predict the island rule for the wrong reasons.

Biddick & Burns (2021) proposed a null/neutral model that reproduces the island rule as a product of random drift. We agree that it is unnecessary to assume adaptive processes driving island dwarfing or gigantism, but several flaws make their approach unrealistic and thus unsuitable as a stochastic model for evolutionary size changes.

Disturbed habitats locally reduce the signal of deep evolutionary history in functional traits of plants.

The functioning of present ecosystems reflects deep evolutionary history of locally cooccurring species if their functional traits show high phylogenetic signal (PS). However, we do not understand what drives local PS. We hypothesize that local PS is high in undisturbed and stressful habitats, either due to ongoing local assembly of species that maintained ancestral traits, or to past evolutionary maintenance of ancestral traits within habitat species-pools, or to both. We quantified PS and diversity of 10 traits within 6704 local plant communities across 38 Dutch habitat types differing in disturbance or stress. Mean local PS varied 50-fold among habitat types, often independently of phylogenetic or trait diversity. Mean local PS decreased with disturbance but showed no consistent relationship to stress. Mean local PS exceeded species-pool PS, reflecting nonrandom subsampling from the pool. Disturbance or stress related more strongly to mean local than to species-pool PS. Disturbed habitats harbour species with evolutionary divergent trait values, probably driven by ongoing, local assembly of species: environmental fluctuations might maintain different trait values within lineages through an evolutionary storage effect. If functional traits do not reflect phylogeny, ecosystem functioning might not be contingent on the presence of particular lineages, and lineages might establish evolutionarily novel interactions.

Knowledge gaps hamper understanding the relationship between fragmentation and biodiversity loss: the case of Atlantic Forest fruit-feeding butterflies.

BACKGROUND: A key challenge for conservation biology in the Neotropics is to understand how deforestation affects biodiversity at various levels of landscape fragmentation. Addressing this challenge requires expanding the coverage of known biodiversity data, which remain to date restricted to a few well-surveyed regions. Here, we assess the sampling coverage and biases in biodiversity data on fruit-feeding butterflies at the Brazilian Atlantic Forest, discussing their effect on our understanding of the relationship between forest fragmentation and biodiversity at a large-scale. We hypothesize that sampling effort is biased towards large and connected fragments, which occur jointly in space at the Atlantic forest. METHODS: We used a comprehensive dataset of Atlantic Forest fruit-feeding butterfly communities to test for sampling biases towards specific geographical areas, climate conditions and landscape configurations. RESULTS: We found a pattern of geographical aggregation of sampling sites, independently of scale, and a strong sampling bias towards large and connected forest fragments, located near cities and roads. Sampling gaps are particularly acute in small and disconnected forest fragments and rare climate conditions. In contrast, currently available data can provide a fair picture of fruit-feeding butterfly communities in large and connected Atlantic Forest remnants. DISCUSSION: Biased data hamper the inference of the functional relationship between deforestation and biodiversity at a large-scale, since they are geographically clustered and have sampling gaps in small and disconnected fragments. These data are useful to inform decision-makers regarding conservation efforts to curb biodiversity loss in the Atlantic Forest. Thus, we suggest to expand sampling effort to small and disconnected forest fragments, which would allow more accurate evaluations of the effects of landscape modification.

Short- and long-term temporal changes in the assemblage structure of Amazonian dung beetles.

Species diversity varies in space and time. Temporal changes in the structure and dynamics of communities can occur at different scales. We investigated the temporal changes of dung beetle assemblages in the Amazonian region along seasons, years, and successional stages. We evaluated if assemblage structure changes between temporal scales and whether such changes affect the functional structure of communities. To achieve these goals, we sampled dung beetles using linear transects of baited pitfall traps during the dry and rainy seasons at two natural reserves in the Amazon region, each representing different time scales: one covering successional variations (80, 30, 5, and 1 years of recovery from logging) and the other one encompassing three consecutive years at two successional stages (20 and 10 years from logging). We used Generalized Linear Models to analyze interannual and successional changes in diversity, described assemblage structure with a NMDS, and examined compositional variation by partitioning beta diversity into its nestedness and turnover components. Abundance and richness decrease from the rainy to the dry season and towards earlier successional stages but do not differ between years. Assemblage diversity changes differently in interannual and successional scales. During succession, dung beetle assemblages change drastically, following a nested structure due to the appearance of species and functional groups in later successional stages. In contrast, functional group composition does not show consistent changes between years, displaying a turnover structure. This pattern supports non-deterministic changes in dung beetle assemblage structure along forest succession.

COVID-19 effective reproduction number dropped during Spain's nationwide dropdown, then spiked at lower-incidence regions.

COVID-19 pandemic has rapidly spread worldwide. Spain has suffered one of the largest nationwide bursts, particularly in the highly populated areas of Madrid and Barcelona (two of the five largest conurbations in Europe). We used segmented regression analyses to identify shifts in the evolution of the effective reproduction number (Rt) reported for 16 Spanish administrative regions. We associate these breaking points with a timeline of key containment measures taken by national and regional governments, applying time lags for the time from contagion to case detection, with their associated errors. Results show an early decrease of Rt that preceded the nationwide lockdown; a generalized, sharp decrease in Rt associated with such lockdown; a low impact of the strengthened lockdown, with a flattening of Rt evolution in high-incidence regions, and even increases in Rt at low-incidence regions; and an increase in Rt associated to the relaxation of the lockdown measures in ten regions. These results evidence the importance of generalized lockdown measures to contain COVID-19 spread, and the limited effect of the subsequent application of a stricter lockdown (restrictions to all non-essential economic activities). Most importantly, they highlight the importance of maintaining strong social distancing measures and strengthening public health control during lockdown de-escalation.

Chasing the ghost of infection past: identifying thresholds of change during the COVID-19 infection in Spain.

One of the largest nationwide bursts of the first COVID-19 outbreak occurred in Spain, where infection expanded in densely populated areas through March 2020. We analyse the cumulative growth curves of reported cases and deaths in all Spain and two highly populated regions, Madrid and Catalonia, identifying changes and sudden shifts in their exponential growth rate through segmented Poisson regressions. We associate these breakpoints with a timeline of key events and containment measures, and data on policy stringency and citizen mobility. Results were largely consistent for infections and deaths in all territories, showing four major shifts involving 19-71% reductions in growth rates originating from infections before 3 March and on 5-8, 10-12 and 14-18 March, but no identifiable effect of the strengthened lockdown of 29-30 March. Changes in stringency and mobility were only associated to the latter two shifts, evidencing an early deceleration in COVID-19 spread associated to personal hygiene and social distancing recommendations, followed by a stronger decrease when lockdown was enforced, leading to the contention of the outbreak by mid-April. This highlights the importance of combining public health communication strategies and hard confinement measures to contain epidemics.

Assessing spatial and temporal biases and gaps in the publicly available distributional information of Iberian mosses.

One of the most valuable initiatives on massive availability of biodiversity data is the Global Biodiversity Information Facility, which is creating new opportunities to develop and test macroecological knowledge. However, the potential uses of these data are limited by the gaps and biases associated to large-scale distributional databases (the so-called Wallacean shortfall). Describing and quantifying these limitations are essential to improve knowledge on biodiversity, especially in poorly-studied groups, such as mosses. Here we assess the coverage of the publicly-available distributional information on Iberian mosses, defining its eventual biases and gaps. For this purpose, we compiled IberBryo v1.0, a database that comprises 82,582 records after processing and checking the geospatial and taxonomical information. Our results show the limitations of data and metadata of the publicly-available information. Particularly, ca. 42% of the records lacked collecting date information, which limits data usefulness for time coverage analyses and enlarges the existing knowledge gaps. Then we evaluated the overall coverage of several aspects of the spatial, temporal and environmental variability of the Iberian Peninsula. Through this assessment, we demonstrate that the publicly-available information on Iberian mosses presents significant biases. Inventory completeness is strongly conditioned by the recorders' survey bias, particularly in northern Portugal and eastern Spain and the spatial pattern of surveys is also biased towards mountains. Besides, the temporal pattern of survey effort intensifies from 1970 onwards, encompassing a progressive increase in the geographic coverage of the Iberian Peninsula. Although we just found 5% of well-surveyed cells of 30' of resolution over the 1970-2018 period, they cover about a fifth of the main climatic gradients of the Iberian Peninsula, which provides a fair - though limited - coverage. Yet, the well-surveyed cells are biased towards anthropised areas and some of them are located in areas under intense land-use changes, mainly due to the wood-fires of the last decade. Despite the overall increase, we found a noticeable gap of information in the south-west of Iberia, the Ebro river basin and the inner plateaus. All these gaps and biases call for a careful use of the available distributional data of Iberian mosses for biogeographical and ecological modelling analysis. Further, our results highlight the necessity of incorporating several good practices to increase the coverage of high-quality information. These good practices include digitalisation of specimens and metadata information, improvement on the protocols to get accurate data and metadata or revisions of the vouchers and recorders' field notebooks. These procedures are essential to improve the quality and coverage of the data. Finally, we also encourage Iberian bryologists to establish a series of re-surveys of classical localities that would allow updating the information on the group, as well as to design their future surveys considering the most important information gaps on IberBryo.

Assessing the functional relationship between dung beetle traits and dung removal, burial, and seedling emergence.

The relationship between biodiversity and ecosystem functioning is often assessed through trait diversity. However, the relationship between traits and functions is typically assumed but seldom tested. We analyze the relationship between dung beetle traits and three ecological functions: dung removal, dung burial, and seedling emergence. We set up a laboratory experiment using nine Scarabaeidae species (three endocoprids, four paracoprids, and two telecoprids). We placed a sexual pair of beetles in each experimental unit, together with a mixture of dung and seeds, and measured the amount of dung removed and buried, burial depth, and the number of emerged seedlings. Sixteen morphological traits related to dung removal and burial were measured in each individual. Results indicate that these traits were related to dung beetle performance in dung removal and burial. Most traits were positively related to dung removal, indicating the existence of a general trait syndrome associated with dung manipulation and digging capability. Dung exploitation strategies did not provide further explanatory power. Seedling emergence showed a negative but weak relationship with dung burial amount and depth and species identity. This implies that specific differences in dung-soil interface activity may be important in secondary seed dispersal by dung beetles.

A niche perspective on the range expansion of symbionts.

Range expansion results from complex eco-evolutionary processes where range dynamics and niche shifts interact in a novel physical space and/or environment, with scale playing a major role. Obligate symbionts (i.e. organisms permanently living on hosts) differ from free-living organisms in that they depend on strong biotic interactions with their hosts which alter their niche and spatial dynamics. A symbiotic lifestyle modifies organism-environment relationships across levels of organisation, from individuals to geographical ranges. These changes influence how symbionts experience colonisation and, by extension, range expansion. Here, we investigate the potential implications of a symbiotic lifestyle on range expansion capacity. We present a unified conceptual overview on range expansion of symbionts that integrates concepts grounded in niche and metapopulation theories. Overall, we explain how niche-driven and dispersal-driven processes govern symbiont range dynamics through their interaction across scales, from host switching to geographical range shifts. First, we describe a background framework for range dynamics based on metapopulation concepts applied to symbiont organisation levels. Then, we integrate metapopulation processes operating in the physical space with niche dynamics grounded in the environmental arena. For this purpose, we provide a definition of the biotope (i.e. living place) specific to symbionts as a hinge concept to link the physical and environmental spaces, wherein the biotope unit is a metapopulation patch (either a host individual or a land fragment). Further, we highlight the dual nature of the symbionts' niche, which is characterised by both host traits and the external environment, and define proper conceptual variants to provide a meaningful unification of niche, biotope and symbiont organisation levels. We also explore variation across systems in the relative relevance of both external environment and host traits to the symbiont's niche and their potential implications on range expansion. We describe in detail the potential mechanisms by which hosts, through their function as biotopes, could influence how some symbionts expand their range - depending on the life history and traits of both associates. From the spatial point of view, hosts can extend symbiont dispersal range via host-mediated dispersal, although the requirement for among-host dispersal can challenge symbiont range expansion. From the niche point of view, homeostatic properties of host bodies may allow symbiont populations to become insensitive to off-host environmental gradients during host-mediated dispersal. These two potential benefits of the symbiont-host interaction can enhance symbiont range expansion capacity. On the other hand, the central role of hosts governing the symbiont niche makes symbionts strongly dependent on the availability of suitable hosts. Thus, environmental, dispersal and biotic barriers faced by suitable hosts apply also to the symbiont, unless eventual opportunities for host switching allow the symbiont to expand its repertoire of suitable hosts (thus expanding its fundamental niche). Finally, symbionts can also improve their range expansion capacity through their impacts on hosts, via protecting their affiliated hosts from environmental harshness through biotic facilitation.

Quantitative genetics of body size evolution on islands: an individual-based simulation approach.

According to the island rule, small-bodied vertebrates will tend to evolve larger body size on islands, whereas the opposite happens to large-bodied species. This controversial pattern has been studied at the macroecological and biogeographical scales, but new developments in quantitative evolutionary genetics now allow studying the island rule from a mechanistic perspective. Here, we develop a simulation approach based on an individual-based model to model body size change on islands as a progressive adaptation to a moving optimum, determined by density-dependent population dynamics. We applied the model to evaluate body size differentiation in the pigmy extinct hominin Homo floresiensis, showing that dwarfing may have occurred in only about 360 generations (95% CI ranging from 150 to 675 generations). This result agrees with reports suggesting rapid dwarfing of large mammals on islands, as well as with the recent discovery that small-sized hominins lived in Flores as early as 700 kyr ago. Our simulations illustrate the power of analysing ecological and evolutionary patterns from an explicit quantitative genetics perspective.