Determinants of fine‐scale homogenization and differentiation of native freshwater fish faunas in a Mediterranean Basin: implications for conservation

Aim Increasing threats to freshwater biodiversity are rapidly changing the distinctiveness of regional species pools and local assemblages. Biotic homogenization/differentiation processes are threatening the integrity and persistence of native biodiversity patterns at a range of spatial scales and pose a challenge for effective conservation planning. Here, we evaluate the extent and determinants of fine‐scale alteration in native freshwater fish assemblages among stream reaches throughout a large river basin and consider the implications of these changes for the long‐term conservation of native fishes. Location Guadiana River basin (South‐Western Iberian Peninsula). Methods We quantified the magnitude of change in compositional similarity between observed and reference assemblages and its potential effect on natural patterns of compositional distinctiveness. Reference assemblages were defined as the native species expected to occur naturally (in absence of anthropogenic alterations) and were reconstructed using a multivariate adaptive regression splines predictive model. We also evaluated the role of habitat degradation and introduced species as determinants of biotic homogenization/differentiation. Results We found a significant trend towards homogenization for native fish assemblages. Changes in native fish distributions led to the loss of distinctiveness patterns along natural environmental gradients. Introduced species were the most important factor explaining the homogenization process. Homogenization of native assemblages was stronger in areas close to reservoirs and in lowland reaches where introduced species were more abundant. Main conclusions The implementation of efficient conservation for the maintenance of native fish diversity is seriously threatened by the homogenization processes. The identification of priority areas for conservation is hindered by the fact that the most diverse communities are vanishing, which would require the selection of broader areas to adequately protect all the species. Given the principal role that introduced species play in the homogenization process and their relation with reservoirs, special attention must be paid to mitigating or preventing these threats.     

Anthropogenic disturbance homogenizes seagrass fish communities

Anthropogenic activities have led to the biotic homogenization of many ecological communities, yet in coastal systems this phenomenon remains understudied. In particular, activities that locally affect marine habitat‐forming foundation species may perturb habitat and promote species with generalist, opportunistic traits, in turn affecting spatial patterns of biodiversity. Here, we quantified fish diversity in seagrass communities across 89 sites spanning 6° latitude along the Pacific coast of Canada, to test the hypothesis that anthropogenic disturbances homogenize (i.e., lower beta‐diversity) assemblages within coastal ecosystems. We test for patterns of biotic homogenization at sites within different anthropogenic disturbance categories (low, medium, and high) at two spatial scales (within and across regions) using both abundance‐ and incidence‐based beta‐diversity metrics. Our models provide clear evidence that fish communities in high anthropogenic disturbance seagrass areas are homogenized relative to those in low disturbance areas. These results were consistent across within‐region comparisons using abundance‐ and incidence‐based measures of beta‐diversity, and in across‐region comparisons using incidence‐based measures. Physical and biotic characteristics of seagrass meadows also influenced fish beta‐diversity. Biotic habitat characteristics including seagrass biomass and shoot density were more differentiated among high disturbance sites, potentially indicative of a perturbed environment. Indicator species and trait analyses revealed fishes associated with low disturbance sites had characteristics including stenotopy, lower swimming ability, and egg guarding behavior. Our study is the first to show biotic homogenization of fishes across seagrass meadows within areas of relatively high human impact. These results support the importance of targeting conservation efforts in low anthropogenic disturbance areas across land‐ and seascapes, as well as managing anthropogenic impacts in high activity areas.     

Fish assemblage changes over half a century in the Yellow River,China

Riverine environments have been threatened by anthropogenic perturbations worldwide, whereby their fish assemblages have been modified by habitat changes and nonendemic species invasions. We assessed changes in fish assemblages by comparing the species presence in historical and contemporary fish data in the Yellow River from 1965 to 2015. The temporal change in species assemblages was found with increased nonendemic species and fewer natives. Fish species richness of the river declined 35.4% over the past fifty years. Moreover, the decreased mean Bray–Curtis dissimilarity among reaches suggested that the fish assemblages of different reaches in the Yellow River were becoming more similar over time. However, temporal patterns of fish assemblages varied among reaches. In the upper Yellow River, higher species richness and more invasive species were found than those in the historical record, while the lower reaches experienced significant species loss. Dam constructions, exotic fish invasions, and flow reductions played the vital role in structuring the temporal fish assemblages in the Yellow River. It is suggested that river basins which experienced different types and levels of stressors by anthropogenic perturbations can produce varied effects on their temporal trends of species assemblages.     

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.     

Consistent effects of consumer species loss across different habitats

Our knowledge of the effects of consumer species loss on ecosystem functioning is limited by a paucity of manipulative field studies, particularly those that incorporate inter‐trophic effects. Further, given the ongoing transformation of natural habitats by anthropogenic activities, studies should assess the relative importance of biodiversity for ecosystem processes across different environmental contexts by including multiple habitat types. We tested the context‐dependency of the effects of consumer species loss by conducting a 15‐month field experiment in two habitats (mussel beds and rock pools) on a temperate rocky shore, focussing on the responses of algal assemblages following the single and combined removals of key gastropod grazers (Patella vulgata, P. ulyssiponensis, Littorina littorea and Gibbula umbilicalis). In both habitats, the removal of limpets resulted in a larger increase in macroalgal richness than that of either L. littorea or G. umbilicalis. Further, by the end of the study, macroalgal cover and richness were greater following the removal of multiple grazer species compared to single species removals. Despite substantial differences in physical properties and the structure of benthic assemblages between mussel beds and rock pools, the effects of grazer loss on macroalgal cover, richness, evenness and assemblage structure were remarkably consistent across both habitats. There was, however, a transient habitat‐dependent effect of grazer removal on macroalgal assemblage structure that emerged after three months, which was replaced by non‐interactive effects of grazer removal and habitat after 15 months. This study shows that the effects of the loss of key consumers may transcend large abiotic and biotic differences between habitats in rocky intertidal systems. While it is clear that consumer diversity is a primary driver of ecosystem functioning, determining its relative importance across multiple contexts is necessary to understand the consequences of consumer species loss against a background of environmental change. Synthesis The roles of species may vary with environmental context, making it difficult to predict how biodiversity loss affects ecosystem functioning across multiple habitats. We tested how natural algal assemblages in two distinct intertidal habitats responded to the removal of different combinations of key consumer species. Despite an initial habitat‐dependent effect of consumer loss, habitat type did not modify the longer‐term responses of algal assemblages to either the identity or number of consumer species removed. Our findings show that, in certain systems, consumer diversity remains a primary driver of ecosystem functioning across widely different environmental contexts.     

The varying role of population abundance in structuring indices of biotic homogenization

Aim An important component of human‐induced global change is the decrease or increase in community distinctiveness (taxonomic homogenization or differentiation, respectively) that follows the loss of native species and gain of non‐native species. We use simulation approaches to assess the extent to which conclusions about the outcome of the homogenization process depend on whether or not abundance data are incorporated. Location Data were produced through computer simulation. Methods The frequency with which occurrence‐based similarity indices and abundance‐based similarity indices give different views of changes in community similarity, and the conditions under which such differences occurred were assessed using both deterministic and stochastic modelling approaches to simulate species assemblage states. Results Occurrence‐based and abundance‐based indices were positively correlated across the set of simulations for both the deterministic and stochastic models. However, in both situations approximately one quarter (25%) of models resulted in contrasting outcomes for the two approaches of calculating changes in compositional similarity; that is, one data type showed a positive value (homogenization), whereas the other showed a negative value (differentiation). Main conclusions In the majority of cases, species abundances will not change drastically enough after perturbation to produce large differences between homogenization scores measured using occurrence versus abundance information. However, in cases where these changes are large, it is important to recognize that the choice of metric to analyse homogenization trends will influence the qualitative and quantitative conclusions drawn. Studies of real assemblages are therefore necessary to evaluate the role of species abundance in defining the magnitude and direction of changes in community composition across space, and the implications of these changes for native biodiversity.     

Interannual variation in reproductive phenology in a riverine fish assemblage: implications for predicting the effects of climate change and altered flow regimes

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Ecological filtering or random extinction? Beta‐diversity patterns and the importance of niche‐based and neutral processes following habitat loss

Although both niche‐based and neutral processes are involved in community assembly, most models on the effects of habitat loss are stochastic, assuming neutral communities mainly affected by ecological drift and random extinction. Given that habitat loss is considered the most important driver of the current biodiversity crisis, unraveling the processes underlying the effects of habitat loss is critical from both a theoretical and an applied perspective. Here we unveil the importance of niche‐based and neutral processes to species extinction and community assembly across a gradient of habitat loss, challenging the predictions of neutral models. We draw on a large dataset containing the distribution of 3653 individuals of 42 species, representing 35% of the small mammal species of the Atlantic Forest hotspot, obtained in 68 sites across three continuously‐forested landscapes and three adjacent 10 000‐ha fragmented landscapes differing in the amount of remaining forest (50%, 30% and 10%). By applying a null‐model approach, we investigated β‐diversity patterns by detecting deviations of observed community similarity from the similarity between randomly assembled communities. Species extinction following habitat loss was decidedly non‐random, in contrast to the notion that fragmented communities are mainly driven by ecological drift. Instead, habitat loss led to a strong biotic homogenization. Moreover, species composition changed abruptly at the same level of landscape‐scale habitat loss that has already been associated with a drastic decline in species richness. Habitat loss, as other anthropogenic disturbances, can thus be seen as a strong ecological filter that increases (rather than decreases) the importance of deterministic processes in community assembly. As such, critical advances for the development of conservation science lie on the incorporation of the relevant niche traits associated with extinction proneness into models of habitat loss. The results also underscore the fundamental importance of pro‐active measures to prevent human‐modified landscapes surpassing critical ecological thresholds.     

The relationship between functional and taxonomic homogenization

Aim Species introductions and extinctions have reorganized the earth's biota, often leaving formerly spatially distinct assemblages more similar in species composition, a process termed biotic homogenization. The study of biotic homogenization has been almost entirely focused on the change in taxonomic similarity between assemblages through time. Here, we provide a trait‐based method for calculating functional similarity through time and compare these trends in functional attributes with those trends generated from a taxonomic perspective. Location Data were produced through computer simulation and gathered from North American Breeding Bird Survey (BBS) data and published accounts of North American birds for 10 locations across the east and west coast of the United States. Methods We simulated change in assemblages with different trait types (binary and continuous), levels of trait overlap, number of traits and species richness to determine the relationship between change in taxonomic similarity (ΔTS) and change in functional similarity (ΔFS). We also assess the relationship between ΔTS and ΔFS for bird assemblages across 10 locales in the USA between 1968 and 2008. We used simple linear regression to determine the slope and correlation between ΔTS and ΔFS and used multiple regression to assess the influence of trait overlap, number of traits, species richness and the ratio of traits to species on the relationship between ΔTS and ΔFS. Results Simulations reveal that trait redundancy governs the relationship between ΔTS and ΔFS. A decrease in trait overlap increases the slope of the regression between ΔTS and ΔFS and an increase in the ratio of traits to species in the regional pool increases the correlation. The relationship between ΔTS and ΔFS for breeding birds is comparable to simulations with low trait redundancy. Main conclusions We show that often losing or gaining species from an assemblage tells us very little about the loss or gain of function, and that this scenario most often occurs when the two assemblages have high trait redundancy. It remains to be seen how prevalent this scenario is within empirical examples; however, the implications for the continued delivery of ecosystem functions in the face of species introductions and extinctions are large.     

Anthropogenic stressors and riverine fish extinctions

Human activities are often implicated in the contemporary extinction of contemporary species. Concerning riverine fishes, the major biotic and abiotic threats widely cited include introduction of non-native species, habitat fragmentation and homogenization in stream flow dynamics due to the damming of rivers, dumping of organic loadings, degradation of the riverine habitat by agricultural practices and water abstraction for human and agricultural consumption. However, few studies have evaluated the role of each of these threats on fish extinction at large spatial scales. Focusing on Western Europe and the USA, two of the most heavily impacted regions on Earth, we quantify fish species loss per river basin and evaluate for the first time to what extent, if any, these threats have been promoting fish extinctions. We show that mean fish extinction rates during the last 110 years in both continents is ∼112 times higher than calculated natural extinction rates. However, we identified only weak effects of our selected anthropogenic stressors on fish extinctions. Only river fragmentation by dams and percentage of non-native species seem to be significant, although weak, drivers of fish species extinction. In our opinion, the most probable explanation for the weak effects found here comes from limitations of both biological and threats datasets currently available. Obtaining realistic estimates on both extinctions and anthropogenic threats in individual river basins is thus urgently needed.     

Compositional changes over space and time along an occurrence–abundance continuum: anthropogenic homogenization of the North American avifauna

Aim Changes in community attributes due to the influence of anthropogenic activities have been examined primarily using occurrence data with little consideration of associated changes in abundance. To determine how this influences our perception of biotic homogenization, we examined compositional patterns for avian assemblages over space and time along an occurrence–abundance continuum. Location The contiguous United States and southern Canada. Methods We examined avian assemblages at 951 Breeding Bird Survey (BBS) routes from 1970 to 2005 that contained a total of 443 species. We used five dissimilarity indices to estimate compositional patterns along an occurrence–abundance continuum of assemblage structure (from species occurrence to transformed abundance to raw abundance) for 396,925 unique combinations of BBS route pairs. We examined annual plots of dissimilarity by distance between BBS routes pairs to estimate spatial and temporal patterns for each index. Results Dissimilarity declined with increasing distance between route pairs for occurrence and transformed abundance, reaching an asymptote at approximately 2500 km. For raw abundance, dissimilarity peaked at intermediate distances (1000–2500 km) with no evidence of an asymptote. Avian assemblages became more similar over time at all points along the continuum. Occurrence and transformed abundance presented the weakest temporal trends, which were uniform or poorly delineated as a function of distance between routes. Raw abundance presented the strongest temporal trends, which declined in strength with increasing distance between routes. Main conclusions With the addition of abundance, there was a substantial and consistent pattern of degradation of β‐diversity for North American avifauna that differed considerably from that observed from occurrence data alone. The geographical expansion of a few species, which recently benefited from the direct and indirect consequences of anthropogenic activities, probably played a prominent role in these patterns. When broad‐scale expansions in occupancy are evident, minor gains in similarity based on species occurrence can mask more substantial gains in similarity based on local abundance. When abundance information is unavailable, its role can be estimated by how occupancy has responded geographically to anthropogenic activities and the expectations of the abundance–occupancy relationship. Our findings support previous work indicating that widespread and locally abundant species will tend to benefit more from anthropogenic activities, creating a possible synergism that enhances biotic homogenization.     

Upstream effects of a reservoir on fish assemblages 45 years following impoundment

Fish assemblage structure, rarefied species richness, species diversity and evenness of assemblages upstream of a reservoir in Oklahoma, U.S.A., were compared pre and post‐impoundment as well as in contemporary collections from streams above and below the reservoir. There were significant shifts in assemblage structure between historical and contemporary collections above the reservoir but not between contemporary assemblages above and below the impoundment. Indicator species analysis revealed that the sand shiner Notropis stramineus and fathead minnow Pimephales promelas have declined, whereas largemouth bass Micropterus salmoides and western mosquitofish Gambusia affinis have increased in relative abundance in assemblages upstream of the impoundment. Species richness was lower in contemporary assemblages compared with historical assemblages. Furthermore, contemporary assemblages below the dam had lower species richness, diversity and evenness compared with contemporary collections above the dam. These results highlight the spatial and temporal extent of reservoirs altering fish assemblages upstream of impoundments.     

Fish assemblage–environment relationships suggest differential trophic responses to heavy metal contamination

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Species loss of stoneflies (Plecoptera) in the Czech Republic during the 20th century

1. Rapid expansion and intensification of anthropogenic activities in the 20th century has caused profound changes in freshwater assemblages. Unfortunately, knowledge of the extent and causes of species loss (SL) is limited due to the lack of reliable historical data. An unusual data set allows us to compare changes in the most sensitive of aquatic insect orders, the Plecoptera, at some 170 locations in the Czech Republic between two time periods, 1955–1960 and 2006–2010. Historical data (1890–1911) on assemblages of six lowland rivers allow us to infer even earlier changes. 2. Regional stonefly diversity decreased in the first half of the 20th century. Streams at lower altitudes lost a substantial number of species, which were never recovered. In the second half of the century, large‐scale anthropogenic pressure caused SL in all habitats, leading to a dissimilarity of contemporary and previous assemblages. The greatest changes were found at sites affected by organic pollution and a mixture of organic pollution and channelisation or impoundment. Colonisation of new habitats was observed in only three of the 80 species evaluated. 3. Species of moderate habitat specialisation and tolerance to organic pollution were most likely to be lost. Those with narrow specialisations in protected habitats were present in both historical and contemporary collections. 4. Contemporary assemblages are the consequence of more than a 100 years of anthropogenic impacts. In particular, streams at lower altitude and draining intensively exploited landscapes host a mere fragment of the original species complement. Most stonefly species are less frequently present than before, although their assemblages remain almost intact in near‐natural mountain streams. Our analyses demonstrate dramatic restriction of species ranges and, in some cases, apparent changes in altitudinal preference throughout the area.     

Low‐head dams induce biotic homogenization/differentiation of fish assemblages in subtropical streams

Extensive distribution of widespread species and the loss of native species driven by anthropogenic disturbances modify community similarity, resulting in a decrease or increase in community distinctiveness. Data from four basins in the Wannan Mountains, China, were used to evaluate the effects of low‐head dams on patterns of fish faunal homogenization and differentiation based on abundance data. We aimed to examine the spatial changes in taxonomic and functional similarities of fish assemblages driven by low‐head dams and to examine whether the changes in the similarity of fish assemblages differed between taxonomic and functional components. We found that low‐head dams significantly decreased the mean taxonomic similarity but increased the mean functional similarity of fish assemblages in impoundments using abundance‐based approaches, suggesting that taxonomic differentiation accompanied functional homogenization in stream fish assemblages. These results show the importance of population abundance in structuring fish faunal homogenization and differentiation at small scales, especially when the major differences among assemblages are in species abundance ranks rather than species identities. Additionally, we also found only a weak positive correlation between changes in mean taxonomic and functional similarities, and partial pairs exhibited considerable variation in patterns of fish faunal homogenization and differentiation for taxonomic and functional components. In conclusion, this study highlighted that the observed taxonomic differentiation of current fish assemblages (short‐term phenomenon) is probably an early warning sign of further homogenization in regions where native species are completely predominated and that changes in taxonomic similarity cannot be used to predict changes in functional similarity.     

Distribution of skates and sharks in the North Sea: 112 years of change

How have North Sea skate and shark assemblages changed since the early 20th century when bottom trawling became widespread, whilst their environment became increasingly impacted by fishing, climate change, habitat degradation and other anthropogenic pressures? This article examines long‐term changes in the distribution and occurrence of the elasmobranch assemblage of the southern North Sea, based on extensive historical time series (1902–2013) of fishery‐independent survey data. In general, larger species (thornback ray, tope, spurdog) exhibited long‐term declines, and the largest (common skate complex) became locally extirpated (as did angelshark). Smaller species increased (spotted and starry ray, lesser‐spotted dogfish) as did smooth‐hound, likely benefiting from greater resilience to fishing and/or climate change. This indicates a fundamental shift from historical dominance of larger, commercially valuable species to current prevalence of smaller, more productive species often of low commercial value. In recent years, however, some trends have reversed, with the (cold‐water associated) starry ray now declining and thornback ray increasing. This shift may be attributed to (i) fishing, including mechanised beam trawling introduced in the 1960s–1970s, and historical target fisheries for elasmobranchs; (ii) climate change, currently favouring warm‐water above cold‐water species; and (iii) habitat loss, including potential degradation of coastal and outer estuarine nursery habitats. The same anthropogenic pressures, here documented to have impacted North Sea elasmobranchs over the past century, are likewise impacting shelf seas worldwide and may increase in the future; therefore, parallel changes in elasmobranch communities in other regions are to be expected.     

Homogenization of dispersal ability across bird species in response to landscape change

Human land use is known to homogenize biotic communities, increasing similarity in their genetic, taxonomic and functional diversity. Explanations have focused almost exclusively on human‐mediated extinction and range expansion. However, homogenization could also be produced by land use driving selection for similar traits across species. We propose a novel hypothesis to explain how human land use homogenizes dispersal ability across species. With habitat loss and increasing human land use intensities there should be larger increases in the costs of dispersal for dispersive than sedentary species, because dispersive species interact with non‐habitat more frequently. In contrast, the benefits of dispersal should increase more for sedentary than dispersive species, because sedentary species are at greater risk from inbreeding depression, predation and competition associated with habitat loss. Therefore we predict that sedentary species become more dispersive in a human‐altered landscape, and dispersive species more sedentary. We tested this prediction using wing pointedness to estimate the initial dispersal ability and change in dispersal ability for 21 North American passerines over the 20th century. More pointed wings are associated with stronger dispersal ability. Thus our prediction would be supported by a negative cross‐species relationship between these two measurements. We found a strong, negative relationship, as predicted. This resulted in declines in the variability in wing pointedness among species over time. Although other things can cause wing shape to change, including changes in habitat availability, none of these explained the observed relationship. Our result provides the first evidence that human landscape alteration is homogenizing bird communities, driving selection for intermediate dispersal ability across species. It also implies that more dispersive species are more at‐risk from human landscape use because, when rates of landscape alteration are faster than a species’ ability to adapt to that change, the costs of dispersal increase more for dispersive than sedentary species.     

Bird species richness is associated with phylogenetic relatedness,plant species richness,and altitudinal range in Inner Mongolia

Bird species richness is mediated by local, regional, and historical factors, for example, competition, environmental heterogeneity, contemporary, and historical climate. Here, we related bird species richness with phylogenetic relatedness of bird assemblages, plant species richness, topography, contemporary climate, and glacial‐interglacial climate change to investigate the relative importance of these factors. This study was conducted in Inner Mongolia, an arid and semiarid region with diverse vegetation types and strong species richness gradients. The following associated variables were included as follows: phylogenetic relatedness of bird assemblages (Net Relatedness Index, NRI), plant species richness, altitudinal range, contemporary climate (mean annual temperature and precipitation, MAT and MAP), and contemporary‐Last Glacial Maximum (LGM) change in climate (change in MAT and change in MAP). Ordinary least squares linear, simultaneous autoregressive linear, and Random Forest models were used to assess the associations between these variables and bird species richness across this region. We found that bird species richness was correlated negatively with NRI and positively with plant species richness and altitudinal range, with no significant correlations with contemporary climate and glacial–interglacial climate change. The six best combinations of variables ranked by Random Forest models consistently included NRI, plant species richness, and contemporary‐LGM change in MAT. Our results suggest important roles of local ecological factors in shaping the distribution of bird species richness across this semiarid region. Our findings highlight the potential importance of these local ecological factors, for example, environmental heterogeneity, habitat filtering, and biotic interactions, in biodiversity maintenance.     

The spatial scale of homogenisation and differentiation in Chihuahuan Desert fish assemblages

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Toward a mechanistic understanding and prediction of biotic homogenization

The widespread replacement of native species with cosmopolitan, nonnative species is homogenizing the global fauna and flora. While the empirical study of biotic homogenization is substantial and growing, theoretical aspects have yet to be explored. Consequently, the breadth of possible ecological mechanisms that can shape current and future patterns and rates of homogenization remain largely unknown. Here, we develop a conceptual model that describes 14 potential scenarios by which species invasions and/or extinctions can lead to various trajectories of biotic homogenization (increased community similarity) or differentiation (decreased community similarity); we then use a simulation approach to explore the model's predictions. We found changes in community similarity to vary with the type and number of nonnative and native species, the historical degree of similarity among the communities, and, to a lesser degree, the richness of the recipient communities. Homogenization is greatest when similar species invade communities, causing either no extinction or differential extinction of native species. The model predictions are consistent with current empirical data for fish, bird, and plant communities and therefore may represent the dominant mechanisms of contemporary homogenization. We present a unifying model illustrating how the balance between invading and extinct species dictates the outcome of biotic homogenization. We conclude by discussing a number of critical but largely unrecognized issues that bear on the empirical study of biotic homogenization, including the importance of spatial scale, temporal scale, and data resolution. We argue that the study of biotic homogenization needs to be placed in a more mechanistic and predictive framework in order for studies to provide adequate guidance in conservation efforts to maintain regional distinctness of the global biota.