Effects of Global Warming on Ancient Mammalian Communities and Their Environments

Background

Current global warming affects the composition and dynamics of mammalian communities and can increase extinction risk; however, long-term effects of warming on mammals are less understood. Dietary reconstructions inferred from stable isotopes of fossil herbivorous mammalian tooth enamel document environmental and climatic changes in ancient ecosystems, including C₃/C₄ transitions and relative seasonality.

Methodology/Principal Findings

Here, we use stable carbon and oxygen isotopes preserved in fossil teeth to document the magnitude of mammalian dietary shifts and ancient floral change during geologically documented glacial and interglacial periods during the Pliocene (∼1.9 million years ago) and Pleistocene (∼1.3 million years ago) in Florida. Stable isotope data demonstrate increased aridity, increased C₄ grass consumption, inter-faunal dietary partitioning, increased isotopic niche breadth of mixed feeders, niche partitioning of phylogenetically similar taxa, and differences in relative seasonality with warming.

Conclusion/Significance

Our data show that global warming resulted in dramatic vegetation and dietary changes even at lower latitudes (∼28°N). Our results also question the use of models that predict the long term decline and extinction of species based on the assumption that niches are conserved over time. These findings have immediate relevance to clarifying possible biotic responses to current global warming in modern ecosystems.     

Niches in the Anthropocene: passerine assemblages show niche expansion from natural to urban habitats

Human‐mediated habitat transformation is increasingly evident around the world. Yet, how this transformation influences species’ niche width and overlap remains unclear. On the one hand, human‐mediated habitat transformation promotes increased species similarity through trait‐based filtering, and an increased prevalence of generalist species with broad niches, resulting in functional homogenization. On the other hand, species that colonize transformed habitats could use empty niches, resulting in decreased species similarity and an expansion of assemblage‐level niche space. Here we explore these two alternatives in eight highly diverse passerine assembles in natural, rural and urban habitats in south and southwest China, a rapidly developing region of the world. Based on stable isotopes, we found that species’ niche width increased from natural to human‐made habitats, but there were no differences in niche overlap among habitats. Therefore, we found evidence for niche expansion, with generalists appearing to use empty niches created by human habitat modification, and with assemblages being comprised of complementary species. Further research is needed to determine whether increased between‐ or within‐individual niche variation is the main driver of niche expansion in transformed habitats.     

On the evolutionary stability of sink populations

The evolution of adaptive behaviours can influence population dynamics. Conversely, population dynamics can affect both the rate and direction of adaptive evolution. This paper examines reasons why sink populations – populations maintained by immigration, preventing local extinction – might persist in the habitat repertoire of a species over evolutionary time-scales. Two such reasons correspond to standard explanations for deviations from an ideal free habitat distribution: organisms may not be free to settle in whichever habitat has the highest potential fitness, and may be constrained by costs, perceptual limitations, or mode of dispersal in the acuity of their habitat selectivity. Here, I argue that a third general reason for persistent sink populations is provided by unstable population dynamics in source habitats. I present a simple model illustrating how use of a sink habitat may be selectively advantageous, when a source population has unstable dynamics (which necessarily reflects temporal variation in local fitnesses). Species with unstable local dynamics in high-quality habitats should be selected to utilize a broader range of habitats than species with stable local dynamics, and in particular in some circumstances should utilize sink habitats. This observation has implications for the direction of niche evolution, and the likelihood of niche conservatism.     

Impact of Ocean Warming and Ocean Acidification on Larval Development and Calcification in the Sea Urchin Tripneustes gratilla

Background

As the oceans simultaneously warm, acidify and increase in P _CO2, prospects for marine biota are of concern. Calcifying species may find it difficult to produce their skeleton because ocean acidification decreases calcium carbonate saturation and accompanying hypercapnia suppresses metabolism. However, this may be buffered by enhanced growth and metabolism due to warming.

Methodology/Principal Findings

We examined the interactive effects of near-future ocean warming and increased acidification/P _CO2 on larval development in the tropical sea urchin Tripneustes gratilla. Larvae were reared in multifactorial experiments in flow-through conditions in all combinations of three temperature and three pH/P _CO2 treatments. Experiments were placed in the setting of projected near future conditions for SE Australia, a global change hot spot. Increased acidity/P _CO2 and decreased carbonate mineral saturation significantly reduced larval growth resulting in decreased skeletal length. Increased temperature (+3°C) stimulated growth, producing significantly bigger larvae across all pH/P _CO2 treatments up to a thermal threshold (+6°C). Increased acidity (-0.3-0.5 pH units) and hypercapnia significantly reduced larval calcification. A +3°C warming diminished the negative effects of acidification and hypercapnia on larval growth.

Conclusions and Significance

This study of the effects of ocean warming and CO₂ driven acidification on development and calcification of marine invertebrate larvae reared in experimental conditions from the outset of development (fertilization) shows the positive and negative effects of these stressors. In simultaneous exposure to stressors the dwarfing effects of acidification were dominant. Reduction in size of sea urchin larvae in a high P _CO2 ocean would likely impair their performance with negative consequent effects for benthic adult populations.     

Relationships of climate,residence time,and biogeographical origin with the range sizes and species richness patterns of exotic plants in Great Britain

Based on atlas data with a 10-km cell resolution for 1,406 exotic plant species inhabiting Great Britain, we investigate the extent to which arrival time (residence time) and biogeographical origin (climate suitability) are associated with range sizes of exotic plants and how exotic plant richness is related to current climate and the human footprint. We grouped species according to four arrival periods (archaeophytes and three classes of neophytes), and three broad biogeographical origins, each reflecting a different macroclimate similarity with the study region (northern Holarctic > Mediterranean > and tropical–subtropical). While we found that mean range sizes increased with residence time, no strong effect of the region of origin on range size was detected. Also, across all groups, species richness was primarily and positively associated with temperature, whereas relationships with human footprint were much weaker, albeit also positive in all cases. The proportion of variance explained by environmental models of richness increased from groups comprising recently arrived species to those that arrived earlier, and from tropical–subtropical species to exotics coming from the Holarctic. Our data also illustrate how these trends translate into richness patterns and their association with climate, which become more similar to native richness patterns as residence time and macroclimatic matching increase. In contrast, broad-scale human alteration of ecosystems appeared to be less important for variation in exotic richness than climate, although we did not evaluate anthropogenic effects at finer scales.     

Topographic Variation in Aboveground Biomass in a Subtropical Evergreen Broad-Leaved Forest in China

The subtropical forest biome occupies about 25% of China, with species diversity only next to tropical forests. Despite the recognized importance of subtropical forest in regional carbon storage and cycling, uncertainties remain regarding the carbon storage of subtropical forests, and few studies have quantified within-site variation of biomass, making it difficult to evaluate the role of these forests in the global and regional carbon cycles. Using data for a 24-ha census plot in east China, we quantify aboveground biomass, characterize its spatial variation among different habitats, and analyse species relative contribution to the total aboveground biomass of different habitats. The average aboveground biomass was 223.0 Mg ha⁻¹ (bootstrapped 95% confidence intervals [217.6, 228.5]) and varied substantially among four topographically defined habitats, from 180.6 Mg ha⁻¹ (bootstrapped 95% CI [167.1, 195.0]) in the upper ridge to 245.9 Mg ha⁻¹ (bootstrapped 95% CI [238.3, 253.8]) in the lower ridge, with upper and lower valley intermediate. In consistent with our expectation, individual species contributed differently to the total aboveground biomass of different habitats, reflecting significant species habitat associations. Different species show differently in habitat preference in terms of biomass contribution. These patterns may be the consequences of ecological strategies difference among different species. Results from this study enhance our ability to evaluate the role of subtropical forests in the regional carbon cycle and provide valuable information to guide the protection and management of subtropical broad-leaved forest for carbon sequestration and carbon storage.     

Can amphibians take the heat? Vulnerability to climate warming in subtropical and temperate larval amphibian communities

Predicting the biodiversity impacts of global warming implies that we know where and with what magnitude these impacts will be encountered. Amphibians are currently the most threatened vertebrates, mainly due to habitat loss and to emerging infectious diseases. Global warming may further exacerbate their decline in the near future, although the impact might vary geographically. We predicted that subtropical amphibians should be relatively susceptible to warming‐induced extinctions because their upper critical thermal limits (CT_max) might be only slightly higher than maximum pond temperatures (T_max). We tested this prediction by measuring CT_max and T_max for 47 larval amphibian species from two thermally distinct subtropical communities (the warm community of the Gran Chaco and the cool community of Atlantic Forest, northern Argentina), as well as from one European temperate community. Upper thermal tolerances of tadpoles were positively correlated (controlling for phylogeny) with maximum pond temperatures, although the slope was steeper in subtropical than in temperate species. CT_max values were lowest in temperate species and highest in the subtropical warm community, which paradoxically, had very low warming tolerance (CT_max–T_max) and therefore may be prone to future local extinction from acute thermal stress if rising pond T_max soon exceeds their CT_max. Canopy‐protected subtropical cool species have larger warming tolerance and thus should be less impacted by peak temperatures. Temperate species are relatively secure to warming impacts, except for late breeders with low thermal tolerance, which may be exposed to physiological thermal stress in the coming years.     

Altitudinal distribution of evergreen broad-leaved trees and their leaf-size pattern on a humid subtropical mountain,Mt. Emei,Sichuan, China

Altitudinal distribution of evergreen broad-leaved trees and changes in their leaf sizes were studied on a humid subtropical mountain, Mt. Emei (3099 m a.s.l., 29°34.5 N, 103°21.5 E), Sichuan, China. Among the total woody flora of ca. 540 species, evergreen broad-leaved trees account for 88 species in 39 genera and 23 families, corresponding to the northern limit of subtropical evergreen broad-leaved trees. The number of evergreen broad-leaved tree species greatly decreased from the low-altitudinal, evergreen broad-leaved forest zone (600–1500 m) to the mid-altitudinal, mixed forest zone (1500–2500 m), and to the high-altitudinal, coniferous forest zone (2500–3099 m). The overall trend of reduced leaf size toward upper zones was analyzed and documented in detail. The 88 species were assigned to three leaf-size classes: notophylls (48%), microphylls (36%), and mesophylls (16%). The leaf size was relatively small and the specific leaf weight (SLW, mg cm^–2) was much larger in high altitude as compared to low altitude. No overall correlation was found between leaf size and SLW, but leaf size decreased as SLW increased toward high altitude for certain species having relatively wide altitudinal ranges. Moreover, leaf size varied with forest stratification: canopy trees were predominantly notophyllous species, while subcanopy and understorey trees were mainly microphyllous species. The tendency is compatible with the trend found in other mountains of East Asia.     

Demographic Amplification of Climate Change Experienced by the Contiguous United States Population during the 20th Century

Better understanding of the changing relationship between human populations and climate is a global research priority. The 20^th century in the contiguous United States offers a particularly well-documented example of human demographic expansion during a period of radical socioeconomic and environmental change. One would expect that as human society has been transformed by technology, we would become increasingly decoupled from climate and more dependent on social infrastructure. Here we use spatially-explicit models to evaluate climatic, socio-economic and biophysical correlates of demographic change in the contiguous United States between 1900 and 2000. Climate-correlated variation in population growth has caused the U.S. population to shift its realized climate niche from cool, seasonal climates to warm, aseasonal climates. As a result, the average annual temperature experienced by U.S. citizens between 1920 and 2000 has increased by more than 1.5°C and the temperature seasonality has decreased by 1.1°C during a century when climate change accounted for only a 0.24°C increase in average annual temperature and a 0.15°C decrease in temperature seasonality. Thus, despite advancing technology, climate-correlated demographics continue to be a major feature of contemporary U.S. society. Unfortunately, these demographic patterns are contributing to a substantial warming of the climate niche during a period of rapid environmental warming, making an already bad situation worse.     

Modelization of the Current and Future Habitat Suitability of Rhododendron ferrugineum Using Potential Snow Accumulation

Mountain areas are particularly sensitive to climate change. Species distribution models predict important extinctions in these areas whose magnitude will depend on a number of different factors. Here we examine the possible impact of climate change on the Rhododendron ferrugineum (alpenrose) niche in Andorra (Pyrenees). This species currently occupies 14.6 km² of this country and relies on the protection afforded by snow cover in winter. We used high-resolution climatic data, potential snow accumulation and a combined forecasting method to obtain the realized niche model of this species. Subsequently, we used data from the high-resolution Scampei project climate change projection for the A2, A1B and B1 scenarios to model its future realized niche model. The modelization performed well when predicting the species’s distribution, which improved when we considered the potential snow accumulation, the most important variable influencing its distribution. We thus obtained a potential extent of about 70.7 km² or 15.1% of the country. We observed an elevation lag distribution between the current and potential distribution of the species, probably due to its slow colonization rate and the small-scale survey of seedlings. Under the three climatic scenarios, the realized niche model of the species will be reduced by 37.9–70.1 km² by the end of the century and it will become confined to what are today screes and rocky hillside habitats. The particular effects of climate change on seedling establishment, as well as on the species’ plasticity and sensitivity in the event of a reduction of the snow cover, could worsen these predictions.     

Damage and recovery of coral reefs affected by El Niño related seawater warming in the Thousand Islands,Indonesia

Extensive coral bleaching occurred during sea-water warming (as a result of the 1982/3 El Niño Southern Oscillation event) in 1983 on the shallow reefs in the Java Sea. Mean seawater temperatures rose by 2–3° C over a six month period with values greater than 33° C being recorded between 1200–1500 h. As many as 80–90% of corals died on the reef flats at the study sites, with the major casualties being branching species in the genera Acropora and Pocillopora. Five years after the event the community structure of the study sites has recovered significantly, though coral cover is still 50% of its former level. Contrasting patterns of recovery at two selected sites, in close proximity to each other, are discussed.     

European Bison as a Refugee Species? Evidence from Isotopic Data on Early Holocene Bison and Other Large Herbivores in Northern Europe

According to the refugee species concept, increasing replacement of open steppe by forest cover after the last glacial period and human pressure had together forced European bison (Bison bonasus)—the largest extant terrestrial mammal of Europe—into forests as a refuge habitat. The consequent decreased fitness and population density led to the gradual extinction of the species. Understanding the pre-refugee ecology of the species may help its conservation management and ensure its long time survival. In view of this, we investigated the abundance of stable isotopes (δ¹³C and δ¹⁵N) in radiocarbon dated skeletal remains of European bison and other large herbivores—aurochs (Bos primigenius), moose (Alces alces), and reindeer (Rangifer tarandus)—from the Early Holocene of northern Europe to reconstruct their dietary habits and pattern of habitat use in conditions of low human influence. Carbon and nitrogen isotopic compositions in collagen of the ungulate species in northern central Europe during the Early Holocene showed significant differences in the habitat use and the diet of these herbivores. The values of the δ¹³C and δ¹⁵N isotopes reflected the use of open habitats by bison, with their diet intermediate between that of aurochs (grazer) and of moose (browser). Our results show that, despite the partial overlap in carbon and nitrogen isotopic values of some species, Early Holocene large ungulates avoided competition by selection of different habitats or different food sources within similar environments. Although Early Holocene bison and Late Pleistocene steppe bison utilized open habitats, their diets were significantly different, as reflected by their δ¹⁵N values. Additional isotopic analyses show that modern populations of European bison utilize much more forested habitats than Early Holocene bison, which supports the refugee status of the species.     

Photosymbiotic ascidians in Singapore: turbid?waters?may reduce living space

The photosymbiotic ascidian fauna at Changi Beach, Pulau Semakau, Sentosa and St. John’s Island, Singapore were surveyed. A total of five species, Diplosoma simile, Lissoclinum bistratum, Lissoclinum punctatum, Lissoclinum timorense and Trididemnum cyclops, were recorded, with Lissoclinum timorense and Trididemnum cyclops being newly recorded in Singapore. However, no photosymbiotic species were found at Changi Beach probably due to the polluted waters in the region. Coastal development has caused Singapore waters to become turbid, leading to decrease in suitable habitats for photosymbiotic ascidians. Clean waters in Pulau Semakau probably provide a better environment for the growth of photosymbiotic ascidians and this area has a greater variety of these ascidians than the other areas in Singapore. Each of the five species has also been recorded in the Ryukyu Archipelago (Japan) and three species (Diplosoma simile, Lissoclinum bistratum and Trididemnum cyclops) have also been recorded in Taiwan.     

Potential Effects of Climate Warming on Fish Habitats in Temperate Zone Lakes with Special Reference to Lake 239 of the Experimental Lakes Area (ELA), North-Western Ontario

We used simple statistics (e.g. mean temperature, degree days, cumulative volume days) to describe present thermal habitats for cool water (yellow perch, Perca flavescens) and cold water (lake trout, Salvelinus namaycush) fish of a small boreal lake. We then modelled changes in the vertical and temporal extent of these habitats under various scenarios of climatic change that included increases in air temperature of 2°C, 4°C, and 9°C, and positive and negative deviations from present levels of 10% in solar radiation and relative humidity, and 20% in wind speed and the lake water extinction coefficient. Model simulations indicated pronounced changes in the temporal and vertical availability of fish thermal niche space. These changes were mainly driven by the large increases in mean mixed layer temperatures that corresponded to 85% of the increases in air temperature, but, in particular, changes in light attenuation also resulted in some non-linear, unexpected effects in the distribution and seasonal availability of thermal niche space. As expected, classical lake trout thermal habitat (5–15°C) was progressively reduced and almost disappeared in littoral areas in spring and early summer. Perch thermal niche space expanded for air temperature increases of up to 4°C, but largely disappeared for the 9°C increase. We discuss changes in thermal habitat with regard to the life history of lake trout and yellow perch, and include other determinants of fish habitat to evaluate the potential of these species for long-term ecological success under climatic warming.     

Subtropical Front fluctuations south of Australia (45°09′S, 146°17′E) for the last 130 ka years based on calcareous nannoplankton

Calcareous nannoplankton assemblages from a Late Quaternary deep-sea core (GC07; 46°09′S, 146°17′E) south of Australia provide information on regional palaeoceanography and palaeoclimate changes in the Southern Ocean, in particular the movement of the Subtropical Front for the past 130 ka years. Marine Isotope Stages 1–5 are identified through changes in calcareous nannoplankton assemblages, supported by ¹⁴C dates, and oxygen isotope and %CaCO data.Two distinct assemblages are recognised: a warm water assemblage with higher abundances of Calcidiscus leptoporus, Emiliania huxleyi, Helicosphaera.carteri, Syracosphaera pulchra, Gephyrocapsa caribbeanica and Gephyrocapsa oceanica; and, a cold water assemblage with higher abundances of Gephyrocapsa muellerae and Coccolithus pelagicus. Alternation between these two assemblages downcore in GC07 reflect movement of the Subtropical Front across the location and can be correlated to Marine Isotope Stages (MIS) 1–5. Sediments with a cold water assemblage indicate the position of the Subtropical Front equatorward of the site when transitional to sub-antarctic waters were overlying the site. Conversely sediments with a warm water assemblage indicate the Subtropical Front was poleward of GC07 when warmer, subtropical waters were over the site. MIS 1 and 5 are interpreted as warmer than MIS 3 (based on species composition) with the Subtropical Front more poleward than for MIS 3. During MIS 3 the Subtropical Front is interpreted as adjacent to or immediately poleward of GC07. Some species including C. leptoporus and C. pelagicus show negative covariance and are considered to be reliable species in identifying glacial and interglacial intervals in this region.Comparison with established biostratigraphy based on calcareous nannoplankton showed the datum event for the reversal between E. huxleyi and G. muellerae of 73 ka in transitional waters is not applicable in this region. The reversal between these two species occurs between 48 and 30 cm downcore in GC07 with a ¹⁴C date of 11 020 year BP at 49–48 cm, i.e. the reversal event is younger than this date.     

Effect of Nitrogen and Water Stress on Photosynthesis and Nitrogen Content in Wheat

The relationships between photosynthesis, leaf nitrogen content and water stress were studied in ten genotypes of wheat differing in the presence of dwarfing genes. Net photosynthetic rate (P_N) was mostly higher at ear emergence stage than at anthesis stage. P_N decreased with water stress (leaf water potential from –2.0 to –2.5 MPa), and with reduced leaf N content in all genotypes studied. Among the various genotypes, single dwarf and wild types showed higher P_N rate and maintained higher leaf N content under different N doses and water supply as compared to the other types studied.     

Satellite data identify decadal trends in the quality of Pygoscelis penguin chick‐rearing habitat

Pygoscelis penguins are experiencing general population declines in their northernmost range whereas there are reported increases in their southernmost range. These changes are coincident with decadal‐scale trends in remote sensed observations of sea ice concentrations (SIC) and sea surface temperatures (SST) during the chick‐rearing season (austral summer). Using SIC, SST, and bathymetry, we identified separate chick‐rearing niche spaces for the three Pygoscelis penguin species and used a maximum entropy approach (MaxEnt) to spatially and temporally model suitable chick‐rearing habitats in the Southern Ocean. For all Pygoscelis penguin species, the MaxEnt models predict significant changes in the locations of suitable chick‐rearing habitats over the period of 1982–2010. In general, chick‐rearing habitat suitability at specific colony locations agreed with the corresponding increases or decreases in documented population trends over the same time period. These changes were the most pronounced along the West Antarctic Peninsula where there has been a rapid warming event during at least the last 50 years.     

Diversity-productivity dependent resistance of an alpine plant community to different climate change scenarios

Here we report from a experiment imposing different warming scenarios [control with ambient temperature, constant level of moderate warming for 3 years, stepwise increase in warming for 3 years, and one season of high level warming (pulse) simulating an extreme summer event] on an alpine ecosystem to study the impact on species diversity–biomass relationship, and community resistance in terms of biomass production. Multiple linear mixed models indicate that experimental years had stronger influence on biomass than warming scenarios and species diversity. Species diversity and biomass had almost humpback relationships under different warming scenarios over different experimental years. There was generally a negative diversity–biomass relationship, implying that a positive diversity–biomass relationship was not the case. The application of different warming scenarios did not change this tendency. The change in community resistance to all warming scenarios was generally negatively correlated with increasing species diversity, the strength of the correlation varying both between treatments and between years within treatments. The strong effect of experimental years was consistent with the notion that niche complementarity effects increase over time, and hence, higher biomass productivity over experimental years. The strongest negative relationship was found in the first year of the pulse treatment, indicating that the community had weak resistance to an extreme event of one season of abnormally warm climate. Biomass production started recovering during the two subsequent years. Contrasting biomass-related resistance emerged in the different treatments, indicating that micro sites within the same plant community may differ in their resistance to different warming scenarios.     

Symbiont bleaching in fossil planktonic foraminifera

Size restricted carbon isotopes (δ¹³C) are used to track changes in the ontogenetic life strategies of two species of extinct planktonic foraminifera and demonstrate that the species Morozovelloides crassatus lost their photosymbiotic association prior to their extinction in the latest middle Eocene. M. crassatus exhibit a strong positive correlation between test size and δ¹³C between 39.5 Ma and 38.7 Ma and a Δδ¹³C shift of 1.0‰/100 μm, this is analogous with modern species that possess an association with algal photosymbionts. Turborotalia cerroazulensis is interpreted as an asymbiotic, thermocline dweller and consistently shows no size related δ¹³C trends and greater δ¹⁸O values in comparison to Morozovelloides. We show a long-term (1.5 million year) deterioration of Morozovelloides ecology that culminated in their extinction at 38.021 Ma. The Δδ¹³C /100 μm in M. crassatus is dramatically reduced from 1.0‰ at 39.53 to only 0.2‰ at 38.026 Ma, 5 kyr before their extinction. The decline in ontogenetic δ¹³C suggests diminished photosymbiotic activity (bleaching) and disruption of foraminiferal ecology in the interval preceding their extinction. We conclude that the demise of Morozovelloides was directly related to the deterioration of photosymbiotic partnerships with algae.     

Spiny invaders – Patterns and determinants of cacti invasion in Europe

In this paper, we analyse the patterns and determinants of cacti invasion in 22 European countries. We compiled a checklist for each country. Cacti were classified for each country according to their invasion status as casuals, locally established (1–5 localities of small population size) and widely established (>5 localities of considerable population size).We used generalised linear models (GLM) from the Poisson family with a log-link function and a set of seven country-specific explanatory variables to account for geographical, climatic, habitat-related and economic determinants to test which features of the recipient area determine invasion success and if distribution patterns of species at different invasion stages are governed by the same interplay of explanatory variables. Separate models were fitted with the same predictor variables for casual, locally established, widely established and all cacti. Further, we analysed the temporal invasion trend, and tested if niche breadth (expressed as the number of habitat types colonized) is influenced by the range size (measured as the number of countries invaded). Finally, we reviewed the consequences of cacti invasion for nature conservation.In total, 26 cacti species have been recorded in Europe. Ten species are more widespread and occur in at least three countries, Opuntia humifusa (six countries) being the most widespread species. The country with most cacti is Spain (21 species), whereas in 13 countries no cactus species have been recorded. By far the most important genus is Opuntia with 20 alien species. The temporal invasion pattern shows an exponential increase of the cumulative number of invasion events, increasing from three (1801–50) to nine (1951–2000) invasion events over a 50-year period.Regardless of the invasion stage, the factor explaining most of the variance in the models is the presence of the Mediterranean biogeographic region, and a significant positive effect of the country size on species numbers was identified.Considering the invasion stage, some interesting deviations in the models can be observed. Invasion of casual cacti is only influenced by the presence of the Mediterranean biogeographic region. For locally established cacti, precipitation is negatively correlated with the invasion rate, and the presence of the Alpine biogeographic region is positively correlated; the latter is due the local occurrence of few hardy cacti (Opuntia phaeacantha, O. humifusa) in low-lying valleys of the Alps. As all widely established cactus species are restricted to the Mediterranean region, only this factor was included in the model.All cacti are confined to dry, open habitats on acid siliceous bedrock. Thus, the predominant habitats invaded are rock vegetation, dry grassland, open Mediterranean scrub and dry ruderal habitats. The niche breadth of cacti increases with the numbers of countries colonized. Further, the niche breadth of cacti exhibits a geographic gradient towards the Mediterranean region.Until the 19th century, the dominant pathway of invasion was agriculture, as some cactus species had been introduced for the production of forage and fruits. However, in the last decades horticulture and deliberate planting in the wild have become the dominant pathways.The invasion of cacti in natural and semi-natural habitats in the Mediterranean region changes habitat structure and species composition. However, dense and extensive stands of cacti are restricted to few species (e.g. Opuntia ficus-indica).