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.     

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.     

Taxonomic homogenization in ungulates: patterns and mechanisms at local and global scales

Aim The aim of this paper is to examine taxonomic homogenization in ungulates globally and at the local scale in South Africa. Specifically, we aim to examine the roles of distance, scale, time, extinctions vs. introductions, and extralimital vs. extraregional introductions in the homogenization of ungulate biotas, and to determine pathways of introduction of ungulate species globally and the proximate explanatory variables of ungulate introductions in South Africa. Location Forty‐one countries globally and three spatial resolutions in South Africa. Methods Indigenous, extirpated and established introduced ungulate species data were obtained for countries globally, and at a quarter‐degree grid‐cell resolution in South Africa. Homogenization was calculated using Jaccard’s index of similarity (JI) for countries globally and for three spatial resolutions in South Africa. Zoo holdings and transfer data from the International Species Information System database were used to investigate the relationship between non‐indigenous ungulate species introductions and the number of non‐indigenous ungulate species in zoos. Relationships between JI and species richness, and between numbers of introductions and several environmental and social factors were examined using generalized linear models. Results Homogenization in ungulates was 2% for countries globally and 8% at the coarsest resolution in South Africa. Homogenization increased with increasing resolution and with time, but it decreased with increasing percentage change in species richness. Globally, introductions contributed more to homogenization than did extinctions. Within South Africa, extralimital introductions contributed more to the homogenization of ungulate assemblages than did extraregional ones, and ungulates were typically introduced to high‐income areas with high human population and livestock densities. The same was not true in the past, when ungulates were introduced to ungulate species‐poor areas. The number of non‐indigenous ungulate species established in a country is significantly related to the number of non‐indigenous ungulate species in zoos in the country, possibly owing to sales of surplus animals from zoos. Main conclusions Ungulate faunas are homogenized at both the global scale and in South Africa, with extralimital introductions being of considerable significance regionally. In consequence, increasing attention will have to be given to the conservation consequences of ungulate translocations, both within particular geopolitical regions and across the globe.     

Spread of common species results in local‐scale floristic homogenization in grassland of Switzerland

Aim We assess changes in plant species richness and changes in species dissimilarity at local scale in Swiss grassland between the time periods 2001–2004 and 2006–2009. Further, we provide an ecological interpretation of the observed taxonomic homogenization of vascular plants. Location Switzerland. Methods Changes in species richness and changes in Simpson dissimilarity index of vascular plants in grassland (meadows and pastures) were examined. The analyses were based on species lists recorded on 339 10‐m² sample plots from a systematic sample covering the entire Switzerland. Each sample plot had been surveyed once in 2001–2004 and once in 2006–2009 with 5 years between the first and the second survey. Changes in species dissimilarity were interpreted by comparing the relative contribution of several indicator species groups. Results Mean species richness of vascular plants in grassland increased during the study period. In contrast, species dissimilarity of plants decreased, suggesting local‐scale floristic homogenization of grassland in Switzerland. It was mostly because of the spread of common species, namely the species that are tolerant to high nutrient levels, the species of low conservation value and the species adapted to moderate temperature levels that led to taxonomic homogenization. Target species for conservation did only marginally affect taxonomic homogenization. In contrast to the predictions from studies of taxonomic homogenization on larger scales, the taxonomic homogenization of grassland at local scale was not explained by the spread of neophytic species. Main conclusions The biotic diversity of grassland in Switzerland changed considerably between 2001–2004 and 2006–2009. The observed taxonomic homogenization was merely because of the spread of common species. Local‐scale changes in land use regimes implemented by agri‐environmental schemes and other conservation efforts on parts of the entire grassland area were, apparently, not enough to prevent the total grassland from recent taxonomic homogenization.     

Spatial aspects of trait homogenization within the German flora

Aim Biotic homogenization, the replacement of local biota by non‐indigenous and locally expanding species, is among the major effects of species invasions. Almost all studies related to this topic are focused on the species level, on taxonomic homogenization. Homogenization effects on trait diversity (functional homogenization) may be very important for ecosystem functioning, but they are rarely analysed, especially not at different spatial scales within a single study. This paper aims to examine how the presence of alien species (species introduced since 1500 ad ) affects the distribution of ploidy levels in the German flora at different spatial scales. Location Germany. Methods We used a resampling method and the Morisita–Horn dissimilarity index to calculate dissimilarities of ploidy level within and between alien and indigenous plants at three different spatial scales in Germany: (1) the entire country as one location, (2) between grid cells, and (3) within grid cells of a lattice covering Germany. Results We found a significant differentiation effect within grid cells of c. 130 km², with alien plants increasing the variability of ploidy levels. The differentiation effect varies according to the scale used. At the coarsest scale (entire Germany), ploidy levels of alien plants tend to be more homogeneous in composition compared with those of native plants. At the intermediate scale (between grid cells), and even more pronounced at a small scale (within grid cells), ploidy levels are more heterogeneously distributed across Germany than those of native plant species. Main conclusions Homogenization of ploidy levels at a large scale (entire Germany) and differentiation at a small scale (within grid cells) is comparable with the patterns found in taxonomic homogenization analyses. This analysis is a first step towards understanding the impacts of plant invasions on a trait level. Differentiation and homogenization of ploidy levels might mirror the distribution of related ecological traits and therefore might have an impact on ecosystem functioning.     

Elk parturition site selection at local and landscape scales

Selection of habitat components by ungulates associated with parturition sites varies among and within species depending upon vulnerability to predators, variation in local topography and climate regimes, and the length of time that the maternal–neonatal unit spends at or near the parturition location. We marked 169 parturition locations of elk (Cervus elaphus nelsoni) in western Wyoming using vaginal implant transmitters and evaluated parturition-specific habitat selection at macro- and microhabitat scales using a resource selection function modeling approach. Elk calved in a variety of habitats, yet demonstrated selection at both spatial scales. We found the strongest support for models that incorporated multiple habitat features and focused on topographical and vegetative cover types that provide physical and thermal cover at the macrohabitat scale and for visual cover models at the microhabitat scale. Models based solely on forage availability or quality were least supported at both scales, which may be indicative of a brief occupation of the parturition location or low heterogeneity in the availability of forage resources on parturition ranges. Results of early elk natural history studies may have represented a bias introduced by variable sightability and accessibility of females with calves and a lack of differentiation between calving and neonatal periods. More clearly defining calving site selection and removing biases toward more open habitats where sightability of neonates is greater may be used by wildlife or land managers to improve or protect calving habitats, which is often a stated objective of management actions. The results of this study suggest that microhabitat is more important to elk and that temporal closures over broad areas versus closures focused on specific macrohabitats may be more effective in protecting calving animals. © 2011 The Wildlife Society.     

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.     

From plots to islands: species diversity at different scales

Aim To investigate how plant diversity of whole islands (‘gamma’) is related to alpha and beta diversity patterns among sampling plots within each island, thus exploring aspects of diversity patterns across scales. Location Nineteen islands of the Aegean Sea, Greece. Methods Plant species were recorded at both the whole‐island scale and in small 100 m² plots on each island. Mean plot species richness was considered as a measure of alpha diversity, and six indices of the ‘variation’‐type beta diversity were also applied. In addition, we partitioned beta diversity into a ‘nestedness’ and a ‘replacement’ component, using the total species richness recorded in all plots of each island as a measure of ‘gamma’ diversity. We also applied 10 species–area models to predict the total observed richness of each island from accumulated plot species richness. Results Mean alpha diversity was not significantly correlated with the overall island species richness or island area. The range of plot species richness for each island was significantly correlated with both overall species richness and area. Alpha diversity was not correlated with most indices of beta diversity. The majority of beta diversity indices were correlated with whole‐island species richness, and this was also true for the ‘replacement’ component of beta diversity. The rational function model provided the best prediction of observed island species richness, with Monod’s and the exponential models following closely. Inaccuracy of predictions was positively correlated with the number of plots and with most indices of beta diversity. Main conclusions Diversity at the broader scale (whole islands) is shaped mainly by variation among small local samples (beta diversity), while local alpha diversity is not a good predictor of species diversity at broader scales. In this system, all results support the crucial role of habitat diversity in determining the species–area relationship.     

Null model of biotic homogenization: a test with the European freshwater fish fauna

In recent years, there has been growing concern about how species invasions and extinctions could change the distinctiveness of formerly disparate fauna and flora, a process called biotic homogenization. In the present study, a null model of biotic of homogenization was developed and applied to the European freshwater fish fauna. We found that non-native fish species led to the greatest homogenization in south-western Europe and greatest differentiation in north-eastern Europe. Comparing these observed patterns to those expected by our null model empirically demonstrated that biotic homogenization is a non-random ecological pattern, providing evidence for previous assumptions. The place of origin of non-native species was also considered by distinguishing between exotic (originating from outside Europe) and translocated species (originating from within Europe). We showed that exotic and translocated species generated distinct geographical patterns of biotic homogenization across Europe because of their contrasting effects on the changes in community similarity among river basins. Translocated species promoted homogenization among basins, whereas exotic species tended to decrease their compositional similarity. Quantifying the individual effect of exotic and translocated species is therefore an absolute prerequisite to accurately assess the spatial dynamics of biotic homogenization.     

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.     

Native communities determine the identity of exotic invaders even at scales at which communities are unsaturated

Aim To determine why some communities are more invasible than others and how this depends on spatial scale. Our previous work in serpentine ecosystems showed that native and exotic diversity are negatively correlated at small scales, but became positively correlated at larger scales. We hypothesized that this pattern was the result of classic niche partitioning at small scales where the environment is homogeneous, and a shift to the dominance of coexistence mechanisms that depend on spatial heterogeneity in the environment at large scales. Location Serpentine ecosystem, Northern California. Methods We test the above hypotheses using the phylogenetic relatedness of natives and exotics. We hypothesized that (1) at small scales, native and exotic species should be more distantly related than expected from a random assemblage model because with biotic resistance, successful invaders should have niches that are different from those of the natives present and (2) at large scales, native and exotic species should not be more distantly related than expected. Result We find strong support for the first hypothesis providing further evidence of biotic resistance at small scales. However, at large scales, native and exotic species were also more distantly related than expected. Importantly, however, natives and exotics were more distantly related at small scales than they were at large scales, suggesting that in the transition from small to large scales, biotic resistance is relaxed but still present. Communities at large scales were not saturated in the sense that more species could enter the community, increasing species richness. However, species did not invade indiscriminately. Exotic species closely related to species already established the community were excluded. Main conclusions Native communities determine the identity of exotic invaders even at large spatial scales where communities are unsaturated. These results hold promise for predicting which species will invade a community given the species present.     

Sensitivity and response of northern hemisphere altitudinal and polar treelines to environmental change at landscape and local scales

The sensitivity and response of northern hemisphere altitudinal and polar treelines to environmental change are increasingly discussed in terms of climate change, often forgetting that climate is only one aspect of environmental variation. As treeline heterogeneity increases from global to regional and smaller scales, assessment of treeline sensitivity at the landscape and local scales requires a more complex approach than at the global scale. The time scale (short‐, medium‐, long‐term) also plays an important role when considering treeline sensitivity. The sensitivity of the treeline to a changing environment varies among different types of treeline. Treelines controlled mainly by orographic influences are not very susceptible to the effects of warming climates. Greatest sensitivity can be expected in anthropogenic treelines after the cessation of human activity. However, tree invasion into former forested areas above the anthropogenic forest limit is controlled by site conditions, and in particular, by microclimates and soils. Apart from changes in tree physiognomy, the spontaneous advance of young growth of forest‐forming tree species into present treeless areas within the treeline ecotone and beyond the tree limit is considered to be the best indicator of treeline sensitivity to environmental change. The sensitivity of climatic treelines to climate warming varies both in the local and regional topographical conditions. Furthermore, treeline history and its after‐effects also play an important role. The sensitivity of treelines to changes in given factors (e.g. winter snow pack, soil moisture, temperature, evaporation, etc.) may vary among areas with differing climatic characteristics. In general, forest will not advance in a closed front but will follow sites that became more favourable to tree establishment under the changed climatic conditions.     

Spider diversity in cereal fields: comparing factors at local, landscape and regional scales

Aim Factors acting at various scales may affect biodiversity, demanding analyses at multiple spatial scales in order to understand how community richness is determined. Here, we adopted a hierarchical approach to test the contribution of region, landscape heterogeneity, local management (organic vs. conventional) and location within field (edge vs. centre) to the species richness and abundance of spiders in cereals. Location Three regions of western and central Germany: Leine Bergland, Soester Boerde, and Lahn‐Dill Bergland. Methods Forty‐two paired organic and conventional winter wheat fields were compared. Field pairs were located in areas ranging from structurally simple to structurally complex landscapes. In May and June 2003, spiders were sampled using pitfall traps. Linear mixed models were used to determine the relationship of spider diversity and abundance with regional spatial factors and landscape heterogeneity within a 500‐m radius, as well as with local management and within‐field location. Results Within‐field location of the traps and landscape heterogeneity were the best predictors of species richness: more species were found in field edges and in heterogeneous landscapes. Region and local management had no effect on species richness. Activity density was higher in field edges and differed among regions. Main conclusions The diversity of farmland spiders was influenced by differences at two of the spatial scales (edge vs. centre, simple vs. complex landscapes), but not at the two others (field management, region), emphasizing the importance of analyses at a variety of spatial scales for an adequate explanation of patterns in biodiversity. Our study suggests that promoting heterogeneity in land use at landscape scales is one of the keys to promoting spider diversity in agroecosystems.     

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.     

Processes at multiple scales affect richness and similarity of non‐native plant species in mountains around the world

Aim To investigate how species richness and similarity of non‐native plants varies along gradients of elevation and human disturbance. Location Eight mountain regions on four continents and two oceanic islands. Methods We compared the distribution of non‐native plant species along roads in eight mountainous regions. Within each region, abundance of plant species was recorded at 41–84 sites along elevational gradients using 100‐m² plots located 0, 25 and 75 m from roadsides. We used mixed‐effects models to examine how local variation in species richness and similarity were affected by processes at three scales: among regions (global), along elevational gradients (regional) and with distance from the road (local). We used model selection and information criteria to choose best‐fit models of species richness along elevational gradients. We performed a hierarchical clustering of similarity to investigate human‐related factors and environmental filtering as potential drivers at the global scale. Results Species richness and similarity of non‐native plant species along elevational gradients were strongly influenced by factors operating at scales ranging from 100 m to 1000s of km. Non‐native species richness was highest in the New World regions, reflecting the effects of colonization from Europe. Similarity among regions was low and due mainly to certain Eurasian species, mostly native to temperate Europe, occurring in all New World regions. Elevation and distance from the road explained little of the variation in similarity. The elevational distribution of non‐native species richness varied, but was always greatest in the lower third of the range. In all regions, non‐native species richness declined away from roadsides. In three regions, this decline was steeper at higher elevations, and there was an interaction between distance and elevation. Main conclusions Because non‐native plant species are affected by processes operating at global, regional and local scales, a multi‐scale perspective is needed to understand their patterns of distribution. The processes involved include global dispersal, filtering along elevational gradients and differential establishment with distance from roadsides.     

Body shape differentiation at global and local geographic scales in the invasive cichlid Oreochromis mossambicus

The Mozambique tilapia Oreochromis mossambicus (Teleostei, Cichlidae) has been transplanted worldwide during the 20th century, and now belongs to the list of the most invasive species. Using a geometric morphometric approach, we describe body shape differentiation among 15 populations from native (Mozambique) and invaded (New Caledonia and Guadeloupe) ranges. A dominant phylogeographic signal is detected, despite the broad range of environmental conditions at the local scale. This result suggests that phylogeographic background rather than phenotypic plasticity responding to environmental variation constitutes the main factor correlated with shape divergence. This could result from successive founder events that occurred during the process of colonization of new geographic areas, and therefore strongly suggests heritable phenotypic differentiation. In addition, shape changes along a major axis of divergence hypothetically refer to different swimming abilities, possibly related to divergent functional requirements between the native and invaded ranges. Overall, patterns of contemporary shape diversification in O. mossambicus probably result from both phylogenetic constraints and adaptive divergence processes. We show that critically taking into account recent phylogenetic history of populations as a constraint on rapid phenotypic divergence is necessary for an improved view of contemporary evolution. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 369–381.