Pattern and process of biotic homogenization in the New Pangaea

Human activities have reorganized the earth''s biota resulting in spatially disparate locales becoming more or less similar in species composition over time through the processes of biotic homogenization and biotic differentiation, respectively. Despite mounting evidence suggesting that this process may be widespread in both aquatic and terrestrial systems, past studies have predominantly focused on single taxonomic groups at a single spatial scale. Furthermore, change in pairwise similarity is itself dependent on two distinct processes, spatial turnover in species composition and changes in gradients of species richness. Most past research has failed to disentangle the effect of these two mechanisms on homogenization patterns. Here, we use recent statistical advances and collate a global database of homogenization studies (20 studies, 50 datasets) to provide the first global investigation of the homogenization process across major faunal and floral groups and elucidate the relative role of changes in species richness and turnover. We found evidence of homogenization (change in similarity ranging from −0.02 to 0.09) across nearly all taxonomic groups, spatial extent and grain sizes. Partitioning of change in pairwise similarity shows that overall change in community similarity is driven by changes in species richness. Our results show that biotic homogenization is truly a global phenomenon and put into question many of the ecological mechanisms invoked in previous studies to explain patterns of homogenization.     

Rise of the generalists: evidence for climate driven homogenization in avian communities

Aims Biogeographical evidence suggests a strong link between climate and patterns of species diversity, and climate change is known to cause range shifts. However, there is little understanding of how shifts affect community composition and we lack empirical evidence of recent impacts of climate change on the diversity of vertebrates. Using a long‐term comprehensive dataset on bird abundance, we explore recent patterns of change in different components of species diversity and avian communities, and postulate a process to explain the observed changes in diversity and specialization. Location Britain. Methods We used Breeding Bird Survey data for Britain from 1994 to 2006 to calculate site‐specific diversity and community specialization indices. We modelled these indices using generalized additive models to examine the relationship between local climate and spatial and temporal trends in community metrics and the relationship between changes in diversity and specialization. Results Local temperature was positively associated with alpha diversity, which increased over the study period, supporting empirical and theoretical predictions of the effect of climate warming. Diversity increased in all habitats, but the rate of increase was greatest in upland areas. However, temperature was negatively associated with community specialization indices, which declined over the same period. Our modelling revealed a nonlinear relationship between community specialization and species diversity. Main conclusions Our models of diversity and specialization provide stark empirical evidence for a link between warming climate and community homogenization. Over a 13‐year period of warming temperatures, diversity indices increased while average community specialization decreased. We suggest that the observed diversity increases were most likely driven by range expansion of generalist species and that future warming is likely to increase homogenization of community structure. When assessed in combination, diversity and specialization measures provide a powerful index for monitoring the impacts of climate change.     

Forest fragmentation drives Atlantic forest of northeastern Brazil to biotic homogenization

Aim To examine whether the tree flora of the Atlantic forest of northeastern Brazil has experienced detectable taxonomic homogenization via the proliferation of native pioneer species in response to habitat loss and fragmentation. Location Biotic homogenization (BH) was examined across the Atlantic forest of northeast Brazil, i.e. a 56,000 km² piece of tropical forest and a distinct centre of species endemism in South America. Methods We assessed a dataset consisting of 5122 tree records and compared the similarity of tree floras from 12 semi‐natural sub‐regions of the Atlantic forest between two time periods: pre‐1980 (plant records between 1902 and 1980), and post‐1980 (between 1981 and 2006). To understand the mechanisms leading to BH (1) tree floras were ordered (via non‐metric multidimensional scaling – NMDS) by date (pre/post 1980) based on species occurrence and frequency, (2) NMDS axes were regressed against the proportion of those species that increased their occurrence post‐1980 (i.e. the winner species), and (3) patterns of geographic distribution and frequency of particular life‐history traits were examined across winner species and a control group. Results Tree floras across the Atlantic forest became c. 20–40% more similar to each other post‐1980, but patterns of species similarity were also influenced by between‐plot geographical distance. NMDS ordination clearly segregated pre‐ and post‐1980 floras with a clear signal of floristic convergence. Furthermore, winner tree species were largely composed of short‐lived and small‐seeded pioneer species that exhibit wide geographic distributions. Main conclusions Our results suggest that tropical forest biotas are susceptible to taxonomic homogenization (i.e. increasing levels of similarity) in the context of severe human‐disturbance via the proliferation of particular groups of native species comprised mainly by ecologically‐plastic, generalist species. We are thus extending the concept of homogenization to address and highlight a pervasive biological shift in the structure of tropical forest communities currently taking place across hyper‐fragmented landscapes.     

Mass coral bleaching causes biotic homogenization of reef fish assemblages

Global climate change is altering community composition across many ecosystems due to nonrandom species turnover, typically characterized by the loss of specialist species and increasing similarity of biological communities across spatial scales. As anthropogenic disturbances continue to alter species composition globally, there is a growing need to identify how species responses influence the establishment of distinct assemblages, such that management actions may be appropriately assigned. Here, we use trait‐based analyses to compare temporal changes in five complementary indices of reef fish assemblage structure among six taxonomically distinct coral reef habitats exposed to a system‐wide thermal stress event. Our results revealed increased taxonomic and functional similarity of previously distinct reef fish assemblages following mass coral bleaching, with changes characterized by subtle, but significant, shifts toward predominance of small‐bodied, algal‐farming habitat generalists. Furthermore, while the taxonomic or functional richness of fish assemblages did not change across all habitats, an increase in functional originality indicated an overall loss of functional redundancy. We also found that prebleaching coral composition better predicted changes in fish assemblage structure than the magnitude of coral loss. These results emphasize how measures of alpha diversity can mask important changes in the structure and functioning of ecosystems as assemblages reorganize. Our findings also highlight the role of coral species composition in structuring communities and influencing the diversity of responses of reef fishes to disturbance. As new coral species configurations emerge, their desirability will hinge upon the composition of associated species and their capacity to maintain key ecological processes in spite of ongoing disturbances.     

Biotic homogenization of three insect groups due to urbanization

Cities are growing rapidly, thereby expected to cause a large‐scale global biotic homogenization. Evidence for the homogenization hypothesis is mostly derived from plants and birds, whereas arthropods have so far been neglected. Here, I tested the homogenization hypothesis with three insect indicator groups, namely true bugs, leafhoppers, and beetles. In particular, I was interested whether insect species community composition differs between urban and rural areas, whether they are more similar between cities than between rural areas, and whether the found pattern is explained by true species turnover, species diversity gradients and geographic distance, by non‐native or specialist species, respectively. I analyzed insect species communities sampled on birch trees in a total of six Swiss cities and six rural areas nearby. In all indicator groups, urban and rural community composition was significantly dissimilar due to native species turnover. Further, for bug and leafhopper communities, I found evidence for large‐scale homogenization due to urbanization, which was driven by reduced species turnover of specialist species in cities. Species turnover of beetle communities was similar between cities and rural areas. Interestingly, when specialist species of beetles were excluded from the analyses, cities were more dissimilar than rural areas, suggesting biotic differentiation of beetle communities in cities. Non‐native species did not affect species turnover of the insect groups. However, given non‐native arthropod species are increasing rapidly, their homogenizing effect might be detected more often in future. Overall, the results show that urbanization has a negative large‐scale impact on the diversity specialist species of the investigated insect groups. Specific measures in cities targeted at increasing the persistence of specialist species typical for the respective biogeographic region could help to stop the loss of biodiversity.     

On‐farm habitat restoration counters biotic homogenization in intensively managed agriculture

To slow the rate of global species loss, it is imperative to understand how to restore and maintain native biodiversity in agricultural landscapes. Currently, agriculture is associated with lower spatial heterogeneity and turnover in community composition (β‐diversity). While some techniques are known to enhance α‐diversity, it is unclear whether habitat restoration can re‐establish β‐diversity. Using a long‐term pollinator dataset, comprising ～9,800 specimens collected from the intensively managed agricultural landscape of the Central Valley of California, we show that on‐farm habitat restoration in the form of native plant ‘hedgerows’, when replicated across a landscape, can boost β‐diversity by approximately 14% relative to unrestored field margins, to levels similar to some natural communities. Hedgerows restore β‐diversity by promoting the assembly of phenotypically diverse communities. Intensively managed agriculture imposes a strong ecological filter that negatively affects several important dimensions of community trait diversity, distribution, and uniqueness. However, by helping to restore phenotypically diverse pollinator communities, small‐scale restorations such as hedgerows provide a valuable tool for conserving biodiversity and promoting ecosystem services.     

More is less: net gain in species richness,but biotic homogenization over 140 years

While biodiversity loss continues globally, assessments of regional and local change over time have been equivocal. Here, we assess changes in plant species richness and beta diversity over 140 years at the level of regions within a country. Using 19th‐century flora censuses for 14 Danish regions as a baseline, we overcome previous criticisms concerning short time series and neglect of completely altered habitats. We find that species composition has changed dramatically and directionally across all regions. Substantial species losses were more than offset by large gains, resulting in a net increase in species richness in all regions. The occupancy of initially widespread species increased, while initially rare species lost terrain. These changes were accompanied by strong biotic homogenization; i.e. regions are more similar now than they were 140 years ago. Species declining in Denmark were found to be in similar decline all over Northern Europe.     

Taxonomic and functional homogenization of an endemic desert fish fauna

Aim The highly endemic fishes of the arid Southwest USA have been heavily impacted by human activities resulting in one of the most threatened fish faunas in the world. The aim of this study was to examine the patterns and drivers of taxonomic and functional beta diversity of freshwater fish in the Lower Colorado River Basin across the 20th century. Location Lower Colorado River Basin (LCRB). Methods The taxonomic and functional similarities of watersheds were quantified to identify patterns of biotic homogenization or differentiation over the period 1900–1999. Path analysis was used to identify the relative influence of dam density, urban land use, precipitation regimes and non‐native species richness on observed changes in fish faunal composition. Results The fish fauna of the LCRB has become increasingly homogenized, both taxonomically (1.1% based on β_sim index) and functionally (6.2% based on Bray–Curtis index), over the 20th century. The rate of homogenization varied substantially; range declines of native species initially caused taxonomic differentiation (?7.9% in the 1960s), followed by marginal homogenization (observed in the 1990s) in response to an influx of non‐native species introductions. By contrast, functional homogenization of the basin was evident considerably earlier (in the 1950s) because of the widespread introduction of non‐native species sharing similar suites of biological traits. Path analysis revealed that both taxonomic and functional homogenization were positively related to the direct and indirect (facilitation by dams and urbanization) effects of non‐native species richness. Main conclusions Our study simultaneously examines rates of change in multiple dimensions of the homogenization process. For the endemic fish fauna of the LCRB, we found that the processes of taxonomic and functional homogenization are highly dynamic over time, varying both in terms of the magnitude and rate of change over the 20th century.     

Invasiveness and homogenization: synergism of wide dispersal and high local abundance

Aim To determine whether invasive and locally abundant non‐native species have a more homogenizing effect on plant communities than non‐invasive and less abundant non‐native species. Location California and Florida counties, conservation areas in the USA, and eight US cities. Methods Species lists among counties, conservation areas and cities were compared to see whether invasive and abundant non‐native species increased the Jaccard index of similarity between localities beyond any increases caused by non‐invasive and less abundant non‐native species. Results For all comparisons, we found that invasive non‐native species have a significantly greater homogenizing effect than non‐invasive non‐native species. For the US conservation areas, we found that locally abundant invasive species tend to be more widespread and more widely shared than less abundant invasive species. There is also a positive relationship between homogenization by invasive species and the magnitude of human disturbance. Main conclusions Invasive non‐native species tend to be disproportionately shared among communities relative to non‐invasive non‐native species. This effect is enhanced by human disturbance, as measured by the ratio of non‐native to native species. There is a synergism between abundance and geographical range which enhances the homogenizing effects of abundant species. Invasive species, with wide ecological niches, are more widely shared among communities and more locally abundant. Abundant invasive species are thus more spatially homogenizing, and more ecologically dominant (functionally homogenizing). Also, ‘perceived homogenization’ is probably greater than homogenization measured by the increase in shared species. The abundant species typically seen by the casual observer in a biological community are probably more commonly shared between communities than less common species. Studies that lack abundance data and measure homogenization only on the basis of shared species, which includes most homogenization studies to date, probably underestimate the homogenizing impacts of non‐native species as perceived by people.     

Taxonomic homogenization and differentiation across Southern Ocean Islands differ among insects and vascular plants

Aim To investigate taxonomic homogenization and/or differentiation of insect and vascular plant assemblages across the Southern Ocean Islands (SOI), and how they differ with changing spatial extent and taxonomic resolution. Location Twenty‐two islands located across the Southern Ocean, further subdivided into five island biogeographical provinces. These islands are used because comprehensive data on both indigenous and non‐indigenous insect and plant species are available. Methods An existing database was updated, using newly published species records, identifying the indigenous and non‐indigenous insect and vascular plant species recorded for each island. Homogenization and differentiation were measured using Jaccard’s index (JI) of similarity for assemblages across all islands on a pairwise basis, and for island pairs within each of the biogeographical provinces. The effects of taxonomic resolution (species, genus, family) and distance on levels of homogenization or differentiation were examined. To explore further the patterns of similarity among islands for each of the taxa and groupings (indigenous and non‐indigenous), islands were clustered based on JI similarity matrices and using group averaging. Results Across the SOI, insect assemblages have become homogenized (0.7% increase in similarity at species level) while plant assemblages have become differentiated at genus and species levels. Homogenization was recorded only when pairwise distances among islands exceeded 3000 km for insect assemblages, but distances had to exceed 10,000 km for plant assemblages. Widely distributed non‐indigenous plant species tend to have wider distributions across the SOI than do their insect counterparts, and this is also true of the indigenous species. Main conclusions Insect assemblages across the SOI have become homogenized as a consequence of the establishment of non‐indigenous species, while plant assemblages have become more differentiated. The likely reason is that indigenous plant assemblages are more similar across the SOI than are insect assemblages, which show greater regionalization. Thus, although a suite of widespread, typically European, weedy, non‐indigenous plant species has established on many islands, the outcome has largely been differentiation. Because further introductions of insects and vascular plants are probable as climates warm across the region, the patterns documented here are likely to change through time.     

The complexity of forest borders determines the understorey vegetation

Questions

What are the most important drivers of plant species richness (gamma‐diversity) and species turnover (beta‐diversity) in the field layer of a forest edge? Does the tree and shrub species richness structure and complexity affect the richness of forest and grassland specialist species?

Location

Southeast Sweden.

Methods

We sampled 50 forest edges with different levels of structural complexity in agricultural landscapes. In each border we recorded trees, shrubs and herb layer species in a 50‐m transect parallel with the forest. We investigated species composition and species turnover in relation to the proportions of gaps in the border and the diversity of trees and shrubs.

Results

Total plant species richness in the field layer was mainly explained by the proportion of gaps to areas with full canopy cover and tree diversity. Increasing number of gaps promoted higher diversity of grassland specialist species within the field layer, resulting in open forest borders with the highest overall species richness. Gaps did however have a negative impact on forest species richness. Conversely, increasing forest species richness was positively related to tree diversity, but the number of grassland specialist species was negatively affected by tree diversity.

Conclusions

Managing forest borders, and therefore increasing the area of semi‐open habitats in fragmented agricultural landscapes, provides future opportunities to create a network of suitable habitats for both grassland and deciduous forest specialist species. Such measures therefore have the potential to increase functional connectivity and support dispersal of species in homogeneous forest/agricultural landscapes.     

On defining and quantifying biotic homogenization

Ongoing species invasions and extinctions are changing biological diversity in different ways at different spatial scales. Biotic homogenization (or BH) refers to the process by which the genetic, taxonomic or functional similarities of regional biotas increase over time. It is a multifaceted process that encompasses species invasions, extinctions and environmental alterations, focusing on how the identities of species (or their genetic or functional attributes) change over space and time. Despite the increasing use of the term BH in conservation biology, it is often used erroneously as a synonym for patterns of species invasions, loss of native species or changes in species richness through time. This reflects the absence of an agreed-upon, cogent definition of BH. Here, we offer an operational definition for BH and review the various methodologies used to study this process. We identify the strengths and weaknesses of these approaches, and make explicit recommendations for future studies. We conclude by citing the need for researchers to: (1) consider carefully the definition of BH by recognizing the genetic, taxonomic and functional realms of this process; (2) recognize that documenting taxonomic homogenization requires tracking the identity of species (not species richness) comprising biotas through space and time; and (3) employ more rigorous methods for quantifying BH.