skip to main content
US FlagAn official website of the United States government
dot gov icon
Official websites use .gov
A .gov website belongs to an official government organization in the United States.
https lock icon
Secure .gov websites use HTTPS
A lock ( lock ) or https:// means you've safely connected to the .gov website. Share sensitive information only on official, secure websites.


This content will become publicly available on November 1, 2026

Title: Evidence for environmental filtering and limiting similarity depends on spatial scale and dissimilarity metrics
Abstract Darwin's theory of natural selection provides two seemingly contradictory hypotheses for explaining the success of biological invasions: (1) the pre‐adaptation hypothesis posits that introduced species that are closely related to native species will be more likely to succeed due to shared advantageous characteristics; (2) the limiting similarity hypothesis posits that invaders that are more similar to resident species will be less likely to succeed due to competitive exclusion. Previous studies assessing this conundrum show mixed results, possibly stemming from inconsistent study spatial scales and failure to integrate both functional and phylogenetic information. Here, we address these limitations using a 33‐year grassland successional survey at Cedar Creek Ecosystem Science Reserve (USA). We incorporate functional dissimilarities, phylogenetic distances, environmental covariates, and species origin data for 303 vascular plant taxa (256 native, 47 introduced), collected from 2700 plots. In contrast with other studies, we test both hypotheses at two fine spatial scales—neighborhood (0.5 m2) and site (~40 m2)—to better capture competition and environmental filtering, respectively. Findings related to Darwin's naturalization conundrum depended on spatial scale and dissimilarity metric. Our results agreed with the pre‐adaptation hypothesis at site scale (40 m2)—a much finer resolution than typically used to test the pre‐adaptation hypothesis—highlighting the role of environmental filtering. At the neighborhood scale (0.5 m2), support for the limiting similarity hypothesis emerged when using functional dissimilarity, while phylogenetic distance aligned with the pre‐adaptation hypothesis, demonstrating that different dissimilarity metrics can yield contrasting conclusions. In addition, native and introduced species showed different abundance patterns in relation to functional ranked dissimilarities, with introduced species reaching higher cover when they were taller than co‐occurring species, had higher leaf dry matter content (LDMC) and lower seed mass. Introduced species also reached high cover with higher soil N concentrations and a shorter time after colonization, relative to native species. Our results suggest that inconsistent findings related to Darwin's naturalization conundrum may arise from an overreliance on single dissimilarity metrics and the use of spatial scales failing to capture underlying ecological processes. This highlights the need for more nuanced methodologies when testing the pre‐adaptation and limiting similarity hypotheses.  more » « less
Award ID(s):
1831944 2425352
PAR ID:
10659509
Author(s) / Creator(s):
 ;  ;  ;  ;  
Publisher / Repository:
Wiley
Date Published:
Journal Name:
Ecology
Volume:
106
Issue:
11
ISSN:
0012-9658
Format(s):
Medium: X
Sponsoring Org:
National Science Foundation
More Like this
  1. Abstract AimAlternative hypotheses of Darwin's Naturalization Conundrum (DNC) predict that the non‐native species that successfully establish within a community are those either more closely or more distantly related to the resident native species. Despite the increasing number of studies using phylogenetic data to testDNCand evaluate community assembly, it remains unknown whether phylogenetic relationships alone can be used to predict invasion susceptibility across communities differing environmentally and in disturbance history. In this study, we evaluate whether phylogenetic structure of diverse native communities predicts the occurrence of non‐native species and offers insight into community assembly. LocationEastern United States of America. MethodsWe examine multiple communities across a north–south transect of the eastern United States to test whether non‐native species richness and abundance are associated with phylogenetic diversity measures of the native community. We also test whether non‐native species are consistently closely or distantly related to native species using two approaches differing in phylogenetic scale and whether this differs with ecologically successful species. ResultsOur analyses did not unambiguously resolveDNC. Non‐native species richness and abundance decreased with increasing native species phylogenetic diversity. Within some communities, non‐native species were significantly more closely related to native species than expected by chance, and tended to be more often closely related to a native species than that native species was to other native relatives. When considering species abundance, only one community showed that ecologically successful non‐native species were closely related to resident species. Main conclusionsPhylogenetic relationships can reveal important details about community assembly in diverse ecological settings. However, given the multifaceted nature of community assembly, phylogenetic metrics alone have limited utility as a general predictive tool for community invasion. Our study highlights a need to incorporate additional types of data to better understand why some communities are more susceptible to non‐native species establishment. 
    more » « less
  2. Understanding the mechanisms governing biological invasions has implications for population dynamics, biodiversity, and community assembly. The enemy escape hypothesis posits that escape from enemies such as herbivores and predators that were limiting in the native range helps explain rapid spread in the introduced range. While the enemy escape hypothesis has been widely tested aboveground, data limitations have prevented comparisons of belowground mechanisms for invasive and noninvasive introduced species, which limits our understanding of why only some introduced species become invasive. We assessed the role of soil biota in driving plant invasions in a phylogenetic meta−analysis, incorporating phylogeny in the error structure of the models, and comparing live and sterilized conditioned soils. We found 29 studies and 396 effect size estimates across 103 species that compared live and sterilized soils. We found general positive effects of soil biota for plants (0.099, 95% CI = 0.0266, 0.1714), consistent with a role of soil mutualists. The effect size of soil biota among invaders was 3.2× higher than for natives, the strength of effects was weaker for older conditioning species with a longer introduced history, and enemy escape was stronger for distant relatives. In addition, invasive species had a weaker allocation tradeoff than natives. By demonstrating that the net effect of soil biota is more positive for invasive than native and noninvasive introduced species, weakens over time since introduction, and strengthens as phylogenetic distance increasing, we provide mechanistic insights into the considerable role of soil biota in biological invasions, consistent with the predictions of the enemy escape hypothesis. 
    more » « less
  3. Understanding the mechanisms governing biological invasions has implications for population dynamics, biodiversity, and community assembly. The enemy escape hypothesis posits that escape from enemies such as herbivores and predators that were limiting in the native range helps explain rapid spread in the introduced range. While the enemy escape hypothesis has been widely tested aboveground, data limitations have prevented comparisons of below- ground mechanisms for invasive and noninvasive introduced species, which limits our understanding of why only some introduced species become invasive. We assessed the role of soil biota in driving plant invasions in a phylogenetic meta−analysis, incorpo- rating phylogeny in the error structure of the models, and comparing live and sterilized conditioned soils. We found 29 studies and 396 effect size estimates across 103 species that compared live and sterilized soils. We found general positive effects of soil biota for plants (0.099, 95% CI 0.0266, 0.1714), consistent with a role of soil mutualists. The effect size of soil biota among invaders was 3.2× higher than for natives, the strength of effects was weaker for older conditioning species with a longer introduced history, and enemy escape was stronger for distant relatives. In addition, invasive species had a weaker allocation tradeoff than natives. By demonstrating that the net effect of soil biota is more positive for invasive than native and noninvasive introduced species, weakens over time since introduction, and strengthens as phy- logenetic distance increasing, we provide mechanistic insights into the considerable role of soil biota in bio- logical invasions, consistent with the predictions of the enemy escape hypothesis. 
    more » « less
  4. Abstract The spread of nonindigenous species by shipping is a large and growing global problem that harms coastal ecosystems and economies and may blur coastal biogeographical patterns. This study coupled eukaryotic environmental DNA (eDNA) metabarcoding with dissimilarity regression to test the hypothesis that ship‐borne species spread homogenizes port communities. We first collected and metabarcoded water samples from ports in Europe, Asia, Australia and the Americas. We then calculated community dissimilarities between port pairs and tested for effects of environmental dissimilarity, biogeographical region and four alternative measures of ship‐borne species transport risk. We predicted that higher shipping between ports would decrease community dissimilarity, that the effect of shipping would be small compared to that of environment dissimilarity and shared biogeography, and that more complex shipping risk metrics (which account for ballast water and stepping‐stone spread) would perform better. Consistent with our hypotheses, community dissimilarities increased significantly with environmental dissimilarity and, to a lesser extent, decreased with ship‐borne species transport risks, particularly if the ports had similar environments and stepping‐stone risks were considered. Unexpectedly, we found no clear effect of shared biogeography, and that risk metrics incorporating estimates of ballast discharge did not offer more explanatory power than simpler traffic‐based risks. Overall, we found that shipping homogenizes eukaryotic communities between ports in predictable ways, which could inform improvements in invasive species policy and management. We demonstrated the usefulness of eDNA metabarcoding and dissimilarity regression for disentangling the drivers of large‐scale biodiversity patterns. We conclude by outlining logistical considerations and recommendations for future studies using this approach. 
    more » « less
  5. Abstract Cultivation and spread of non‐native plant species may result in either phylogenetic homogenization (increasing similarity) or differentiation (decreasing similarity) of urban floras. However, it is unknown how non‐native species influence homogenization of cultivated versus spontaneously occurring species in cities, and which traits are associated with species that promote homogenization versus differentiation. In this study, we compared homogenization effects of cultivated and spontaneous non‐native species in yard floras across and within seven widely distributed U.S. cities. Additionally, we explored which traits explained their particular contribution to homogenization. We recorded plant presence/absence in 178 private yards distributed among seven metropolitan statistical areas in the United States. We compared phylogenetic homogenization effects of non‐native species within both the cultivated and spontaneous species pools using phylogenetic dissimilarities and the homogenization index. Then, we expressed contributions of non‐native species to the homogenization of each pool as a function of two different sets of plant functional traits using phylogenetic generalized least square (PGLS) models across and within cities. Across cities, spontaneous non‐native species homogenized, and cultivated non‐native species differentiated, yard floras. Within the spontaneous pool, short, small‐seeded non‐native plants and non‐native grasses significantly homogenized yard floras. Within the cultivated pool, species contribution to homogenization was best predicted by plant height, presence of showy flowers, and growth form, with non‐native grasses significantly homogenizing cultivated yard floras. Within cities, non‐native species—whether they were cultivated or spontaneous—consistently homogenized yard floras of the three northern cities and differentiated yard floras of three of the four southern cities, suggesting that homogenization processes are context‐ and scale‐dependent. Likewise, traits explaining homogenization differed substantially among cities. The inconsistent patterns among cities in the plant traits that promoted homogenization of both cultivated and spontaneous species suggest that local environmental and anthropogenic conditions of individual cities imposed strong constraints on trait selection. Linking plant functional traits that promote homogenization with residents’ preferences for vegetation may further enhance understanding of how yard plant communities assemble at regional and local scales. 
    more » « less