Darwin's Naturalization Conundrum in the Azores: Scale‐Dependent Patterns of Arthropod Invasion

ABSTRACT Aim To test the mechanisms governing the assembly of exotic arthropods within endemic‐rich island communities by evaluating Darwin's naturalization conundrum, which predicts that invasion success occurs through either functional dissimilarity from native species (Naturalization Hypothesis) or functional similarity with them (Pre‐adaptation Hypothesis). Location Azores archipelago, Portugal, North Atlantic Ocean. Methods We analysed long‐term spatial and temporal data from two standardized monitoring programmes in Azorean native forests: the BALA project (spatial dataset) and the SLAM project (temporal dataset). The dataset included more than 100 species of spiders and beetles sampled across multiple islands over two decades. Using functional trait approaches, we quantified exotic–endemic community structure across spatial and temporal scales and compared observed patterns with null expectations to assess functional similarity or dissimilarity. Results Across both datasets, most Standard Effect Sizes (SES) values clustered around zero, indicating that exotic assemblages were largely indistinguishable from null expectations relative to endemic communities. This neutral pattern was consistent for exotic spiders across all spatial and temporal scales, providing no support for either Darwinian hypothesis. Exotic beetles, however, showed weak but detectable functional overdispersion at broader spatial scales and through time, suggesting limited and context‐dependent support for the Naturalization Hypothesis. Overall, the pattern found is consistent with stochastic processes that appear to dominate exotic–endemic functional relationships, although this cannot be demonstrated directly from SES values alone. Main Conclusions Our results suggest that deterministic competition plays a minor role in structuring exotic arthropod assemblages in Azorean native forests. Instead, landscape‐level processes, particularly the pressure from surrounding anthropogenic habitats and environmental filtering linked to forest fragmentation, likely shape invasion dynamics. These findings highlight the importance of human‐modified landscapes in influencing community assembly and provide insights into how invasion processes operate in insular ecosystems.

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Journal
Diversity and Distributions
Published
2026-09-30
DOI
https://doi.org/10.1111/ddi.70265
Primary Topic
Ecology and Vegetation Dynamics Studies
Type
article
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article

Darwin's Naturalization Conundrum in the Azores: Scale‐Dependent Patterns of Arthropod Invasion

Ana M. C. Santos, Jagoba Malumbres‐Olarte, Ricardo Costa, Isabel Rosário Amorim et al.
Diversity and Distributions
Ecology and Vegetation Dynamics Studies
article

Darwin's Naturalization Conundrum in the Azores: Scale‐Dependent Patterns of Arthropod Invasion

Ana M. C. Santos, Jagoba Malumbres‐Olarte, Ricardo Costa, Isabel Rosário Amorim, Guilherme Oyarzabal, Mário Boieiro, Sébastien Lhoumeau, Gábor Pozsgai, Paulo A. V. Borges, Pedro Cardoso, Rosalina Maria de Almeida Gabriel, François Rigal
article en

Abstract

ABSTRACT Aim To test the mechanisms governing the assembly of exotic arthropods within endemic‐rich island communities by evaluating Darwin's naturalization conundrum, which predicts that invasion success occurs through either functional dissimilarity from native species (Naturalization Hypothesis) or functional similarity with them (Pre‐adaptation Hypothesis). Location Azores archipelago, Portugal, North Atlantic Ocean. Methods We analysed long‐term spatial and temporal data from two standardized monitoring programmes in Azorean native forests: the BALA project (spatial dataset) and the SLAM project (temporal dataset). The dataset included more than 100 species of spiders and beetles sampled across multiple islands over two decades. Using functional trait approaches, we quantified exotic–endemic community structure across spatial and temporal scales and compared observed patterns with null expectations to assess functional similarity or dissimilarity. Results Across both datasets, most Standard Effect Sizes (SES) values clustered around zero, indicating that exotic assemblages were largely indistinguishable from null expectations relative to endemic communities. This neutral pattern was consistent for exotic spiders across all spatial and temporal scales, providing no support for either Darwinian hypothesis. Exotic beetles, however, showed weak but detectable functional overdispersion at broader spatial scales and through time, suggesting limited and context‐dependent support for the Naturalization Hypothesis. Overall, the pattern found is consistent with stochastic processes that appear to dominate exotic–endemic functional relationships, although this cannot be demonstrated directly from SES values alone. Main Conclusions Our results suggest that deterministic competition plays a minor role in structuring exotic arthropod assemblages in Azorean native forests. Instead, landscape‐level processes, particularly the pressure from surrounding anthropogenic habitats and environmental filtering linked to forest fragmentation, likely shape invasion dynamics. These findings highlight the importance of human‐modified landscapes in influencing community assembly and provide insights into how invasion processes operate in insular ecosystems.

Diversity and DistributionsVol. 32(10)
Universidade dos Açores (PT), University of Helsinki (FI), Institut des Sciences Analytiques et de Physico-Chimie pour l'Environnement et les Matériaux (FR), Sustainability Institute (ZA), Finnish Museum of Natural History (FI), Hospital de Santo Espirito de Angra do Heroísmo (PT), Universidad Autónoma de Madrid (ES)
Life below water
Openalex Percentile: Top 8%
Ecology and Vegetation Dynamics Studies
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