Abstract Aim Species on islands are at high risk of extinction due to environmental changes, including global warming, land‐use alterations and invasions. At local scales, extinctions can be offset by strategies promoting in situ persistence. We explored how persistence‐related traits of plants—that is, linked to belowground resource conservation, growth, size and longevity—on edaphic islands respond to variation in insularity and the environment (soil and microclimate), including intraspecific variability, which is rarely considered in functional island biogeography. We hypothesised that plants facing strong insularity and harsh soil conditions are characterised by enhanced persistence abilities. Location Shallow‐soil temperate dry grasslands on granite outcrops, Central Europe. Methods We focussed on edaphic island specialist species belonging to different life histories, namely clonal and non‐clonal perennial plants. We used linear and linear mixed‐effect models to examine intra‐ and interspecific trait patterns versus variation in insularity, soil and microclimate. Results Insularity tended to promote smaller plants (non‐clonal species) and belowground resource‐conservative strategies (both clonal and non‐clonal species), increasing the likelihood of local persistence. Soil also contributed largely to explaining persistence‐related trait patterns: plants growing in harsh soil conditions tended to be resource conservative. Clonal species are distinguished by highly consistent responses to variation in insularity and soil conditions, whereas non‐clonal plants showed distinct species‐specific responses. Main conclusions Our findings have important implications for the conservation biogeography of edaphic island plant specialists. Clonal species may be susceptible to local extinction should insularity or soil conditions vary, for example, due to abrupt changes in the geographical setting (e.g. habitat loss) or local environmental factors (e.g. N‐deposition). Non‐clonal species may instead face environmental changes differently; some will go extinct, whereas others will survive, depending on the prevailing abiotic pressures. This seems to challenge previous views that predicted clonal species to be the winners and non‐clonal species the losers against local extinction.
Abstract Functional island biogeography can provide eco–evolutionary insights into which main drivers contribute to shaping the distribution of organisms' forms and functions on islands. It does so by examining trait patterns. As a result, traits are increasingly studied on islands, either along insularity gradients or by comparing patterns of island versus mainland biota. So far, functional island biogeography has investigated trends of trait values (i.e., average, functional diversity), whereas coordination between pairs of traits remains unexplored along insularity gradients. Yet, trait coordination analyses constitute the foundational tool to detect main functional spectra and strategies of organisms. In this perspective, we set out to offer a conceptual and analytical framework that should facilitate the inclusion of trait coordination (i.e., the co–variation of traits both at the intra– and interspecific level) in functional island biogeography. We illustrate, with a case study focused on persistence traits of edaphic island plant specialists, what type of insights can be gained by examining the response of trait coordination to variation in insularity. We asked two questions, namely whether, with increasing insularity, the strength of the relationship ( R 2 ) increases (Q1), and the direction of the relationship (slope) decreases (Q2). We positively answered our research questions, with lines of evidence suggesting a selective “forcing” towards tighter and more strongly coordinated strategies (Q1), and functional trade‐offs (Q2). We infer which ecological and biogeographic drivers could be behind the observed patterns, while acknowledging possible drawbacks. We conclude by identifying three main take–home messages and related future directions for integrating trait coordination in functional island biogeography to further advance the field.
Abstract Aim Positive relationships between island size and species richness have been found for oceanic and non‐oceanic islands. However, the effect of isolation, also predicted by island‐biogeographical theory, seems challenging to test. Species‐specific mechanisms drive the perception of distance and space for animals, and therefore isolation patterns are masked when described at the community level. We used epiphytic plants (canopy islands) and their spider communities to study how functional group and species‐specific use of space modifies the effect of island‐level predictors on species richness. Location A coffee plantation near Xalapa, Mexico. Taxon Spiders (Araneae), vascular epiphytes (Bromeliaceae, Piperaceae, Orchidaceae, Araceae, Pteridophyta). Methods We collected canopy islands from three trees and recorded their three‐dimensional position. Afterwards, we collected all epiphyte‐dwelling spiders. We used Generalized Linear Models (GLMs) to analyse the effect of island size and isolation on the species richness of spider communities and community subgroups (guilds, mobility groups and families). Isolation was calculated using two metrics (aerial and walking distances) associated with different ways of mobility in the canopy (bridging and wandering respectively). We evaluated species‐level associations with island features and tested for sample‐size effects on GLMs using null‐model analysis of ecological networks. Results Although unimportant at the community level, isolation was more important than island size for some spider subgroups, while having negligible effects for others. Species‐specific responses to isolation and island size were responsible for the variability in species richness patterns observed for broader subgroups (families, guilds and mobility groups). Specific responses of mobility groups to isolation metrics were in line with two ways of moving between islands; one using tree branches as pathways (walking isolation) and the other using silk threads to connect points directly across the air (aerial isolation). Main conclusions Spatial patterns of epiphyte‐dwelling spiders were explained better when functional traits were considered, and the effect of isolation became apparent only when analysed per functional group, whereas it was masked at the community level. This supports the hypothesis that hunting behaviour and mobility affect the specific perception of space and distance even at the small spatial scale of canopy islands.
ABSTRACT Species extinction risk at local scales can be partially offset by strategies promoting in-situ persistence. We explored how persistence-related traits of clonal and non-clonal plants in temperate dry grasslands respond intra- and interspecifically to variation in environmental conditions (soil, climate) and insularity. We focused on edaphic island specialist species, hypothesizing that plants experiencing harsh soil environments and strong insularity are distinguished by traits supporting enhanced persistence, such as small stature, long lifespan and resource-conservative strategies. We used linear mixed-effect models and bivariate ordinary least squares linear models to explore the response of species triats to environmental and biogeographic predictors. We found general support for this hypothesis. Soil properties and insularity emerged as the most important drivers of trait patterns. However, clonal species showed more consistent responses to variation in environmental conditions and insularity than non-clonal plants, which were characterized by distinct species-specific responses. Soil properties and insularity confirmed their major role in shaping the persistence strategies of edaphic island plant species. These drivers may exert their effect on specific functions (e.g. belowground resource conservation captured by BDMC). Additionally, we unambiguously identified that clonal species had different persistence strategies than non-clonal ones.
The theory of island biogeography postulates that size and isolation are key drivers of biodiversity on islands. This theory has been applied not only to true (e.g. oceanic) islands but also to terrestrial island‐like systems (e.g. edaphic islands). Recently, a debate has opened as to whether terrestrial island‐like systems function like true islands. However, identifying the effect of insularity in terrestrial systems is conceptually and methodologically challenging because recognizing species source(s) and measuring isolation is not as straightforward as for true islands. We contribute to the debate by proposing an approach to contextualize the definition of insularity and to identify the role of isolation in terrestrial island‐like systems. To test this approach, we explored the relationship between insularity predictors and specialist species richness of edaphic islands in three systems in Europe (spring fens, mountaintops, and outcrops). We detected that insularity affected specialist richness of edaphic islands through island size and target effect (i.e. an emergent property of islands depending on their isolation and size). As predicted by the Theory of Island Biogeography, species richness decreased with increasing isularity. Given the comprehensiveness and ease of implementation of our approach, we encourage its extension to other island‐like systems.
Abstract Aim Epiphytic plants are isolated from each other by nonhabitat canopy elements and are thus expected to act as islands, the biodiversity of their inhabitants (e.g., spiders) conforming to island biogeographic predictions of species‐richness patterns. Although it has been shown that arthropod diversity decreases with decreasing epiphyte size, the effects of isolation have not yet been addressed. We studied the joint effect of isolation, spatial position, and size of epiphytic plants (canopy islands) on species richness, relative rareness, and similarity of spider communities. Location A shade‐coffee plantation in central Veracruz, Mexico. Taxon Spiders (Araneae), vascular epiphytes (Bromeliaceae, Piperaceae, Orchidaceae, Araceae, Pteridophyta). Methods We collected all canopy islands occurring on three trees and recorded their three‐dimensional spatial position. In the laboratory, we disassembled the plants and collected all spiders present. We analysed the effects of island size, isolation, and spatial position on the species richness and the relative rareness of spider communities using generalized linear models ( GLM ), on the distribution of different spider guilds using a CCA , and on community composition using a permutational multivariate analysis of variance (Permanova). Isolation and spatial position were addressed using five distance measures representing isolation from different potential species sources. We tested how the similarity in spider community composition changed with the distance between islands using Mantel correlograms. Results As predicted by island biogeography theory, spider species richness increased with canopy‐island size and decreased with isolation. In comparison to island size, the effect of isolation was weak, though significant. Relative rareness was hardly affected by island size but more by isolation. Compositional similarity was affected by island size and decreased with increasing spatial distance up to ca. 4 m. Guilds separated along the main CCA axes, this ordination being driven by epiphyte size and position. Main conclusions Epiphytic plants behaved like canopy islands in that their size and isolation influenced the diversity and composition of spider communities. However, the effect of isolation was only a fraction of that of island size, perhaps because spatial relationships are taxon‐specific. This may be due to differences in hunting behaviour and dispersal capacities, for example, between guilds of hunting and web‐building spiders, which is a dimension deserving more attention. For a better understanding of biogeographic principles driving the diversity of canopy island inhabitants, further research on this topic should include position and isolation, at scales matching the mobility of different functional groups, as part of their explanatory variables.