Local Colonization Rates Research Articles

Nutrient addition drives declines in grassland species richness primarily via enhanced species loss

Abstract Declines in grassland diversity in response to nutrient addition are a general consequence of global change. This decline in species richness may be driven by multiple underlying processes operating at different time‐scales. Nutrient addition can reduce diversity by enhancing the rate of local extinction via competitive exclusion, or by reducing the rate of colonization by constraining the pool of species able to colonize under new conditions. Partitioning net change into extinction and colonization rates will better delineate the long‐term effect of global change in grasslands. We synthesized changes in richness in response to experimental fertilization with nitrogen, phosphorus and potassium with micronutrients across 30 grasslands. We quantified changes in local richness, colonization, and extinction over 8–10 years of nutrient addition, and compared these rates against control conditions to isolate the effect of nutrient addition from background dynamics. Total richness at steady state in the control plots was the sum of equal, relatively high rates of local colonization and extinction. On aggregate, 30%–35% of initial species were lost and the same proportion of new species were gained at least once over a decade. Absolute turnover increased with site‐level richness but was proportionately greater at lower‐richness sites relative to starting richness. Loss of total richness with nutrient addition, especially N in combination with P or K, was driven by enhanced rates of extinction with a smaller contribution from reduced colonization. Enhanced extinction and reduced colonization were disproportionately among native species, perennials, and forbs. Reduced colonization plateaued after the first few (<5) years after nutrient addition, while enhanced extinction continued throughout the first decade. Synthesis. Our results indicate a high rate of colonizations and extinctions underlying the richness of ambient communities and that nutrient enhancement drives overall declines in diversity primarily by exclusion of previously established species. Moreover, enhanced extinction continues over long time‐scales, suggesting continuous, long‐term community responses and a need for long‐term study to fully realize the extinction impact of increased nutrients on grassland composition.

Species distribution models and a 60‐year‐old transplant experiment reveal inhibited forest plant range shifts under climate change

AbstractAimClimate change causes species to shift their distributions. Individual species, however, greatly vary in their capacity to track the macroclimatic temperature increase due to differences in demography and dispersal. To better predict range shifts to climate change we need a complementary integration of long‐term empirical data and predictive modelling.LocationBelgium and North‐West Europe.TaxonHyacinthoides non‐scripta¸ forest understorey plants.MethodsComplementing species distribution models with demographic data from an exceptional 60‐year‐old over‐the‐range‐edge transplant experiment measured not less than 45 and 60 years after installation, we evaluated the long‐term consequences of climate change on one of the most emblematic but also among the slowest colonizing plant species of European forests, bluebellHyacinthoides non‐scripta.ResultsWe found bluebell able to establish viable populations beyond its natural range. These results were confirmed by the SDM, showing that bluebell’s potential range is considerably larger than its current range. Colonization rates of only 2 m century−1were observed in the transplanted populations. Beyond bluebell’s current range, we observed decreasing trends in population growth rates over the past 15 years. By the end of the 21st century, substantial decreases in the southern parts of bluebell’s range were predicted.Main conclusionsBased on empirical and modelling results, we expect serious population declines in large parts of its current natural distribution of bluebell. Although the species is able to establish viable populations beyond the natural range edge, slow demography and local colonization rates four orders of magnitude lower than the velocity of climate change make fast enough range shifts virtually impossible in this species.

Phantom species: adjusting estimates of colonization and extinction for pseudo‐turnover

Ecologists rely on field surveys to monitor long‐term ecological change but finite sampling and the prevalence of rare species mean that surveys inevitably miss some species present at a given location. These ‘phantom species’ produce pseudo‐turnover by inflating observed rates of local colonization and extinction in resurvey studies, especially among rare species. In this paper, we quantify the probability that pseudo‐turnover occurs due to imprecise plot relocation and/or shifts in where individuals are located. Using sampling models derived from the binomial distribution, we estimate probabilities of missing species as a function of local abundance and sampling intensity. Spatially explicit simulations confirm that our binomial model is robust to non‐random sampling schemes and clumped species distributions. False absences decline predictably as species abundance and sampling intensity increase allowing us to statistically adjust naïve estimates of colonization and extinction for expected rates of pseudo‐turnover. To illustrate the model's real‐world utility, we analyze apparent colonizations and extinctions for 331 species distributed over 83 sites in southern Wisconsin forests surveyed in both the 1950s and 2000s. Limited sampling in the 1950s means expected rates of pseudo‐colonization are appreciable. Accounting for pseudo‐turnover thus reduces estimated community‐wide colonization rates by 51% compared to naïve (observed) rates. More complete sampling in the 2000s limited overestimates of local extinction to 14%. We distinguish three zones of inference based on sampling intensity and species abundance where: 1) naïve estimates approximate true values of turnover, 2) adjustments are important, and 3) species are too rare relative to sampling to reliably infer turnover. Accounting for phantom species substantially improves our ability to accurately estimate local colonization and extinction rates, enhancing our ability to infer community dynamics and monitor long‐term ecological change.

An occupancy-based quantification of the highly imperiled status of desert fishes of the southwestern United States.

Desert fishes are some of the most imperiled vertebrates worldwide due to their low economic worth and because they compete with humans for water. An ecological complex of fishes, 2 suckers (Catostomus latipinnis, Catostomus discobolus) and a chub (Gila robusta) (collectively managed as the so-called three species) are endemic to the U.S. Colorado River Basin, are affected by multiple stressors, and have allegedly declined dramatically. We built a series of occupancy models to determine relationships between trends in occupancy, local extinction, and local colonization rates, identify potential limiting factors, and evaluate the suitability of managing the 3 species collectively. For a historical period (1889-2011), top performing models (AICc) included a positive time trend in local extinction probability and a negative trend in local colonization probability. As flood frequency decreased post-development local extinction probability increased. By the end of the time series, 47% (95% CI 34-61) and 15% (95% CI 6-33) of sites remained occupied by the suckers and the chub, respectively, and models with the 2 species of sucker as one group and the chub as the other performed best. For a contemporary period (2001-2011), top performing (based on AICc ) models included peak annual discharge. As peak discharge increased, local extinction probability decreased and local colonization probability increased. For the contemporary period, results of models that split all 3 species into separate groups were similar to results of models that combined the 2 suckers but not the chub. Collectively, these results confirmed that declines in these fishes were strongly associated with water development and that relative to their historic distribution all 3 species have declined dramatically. Further, the chub was distinct in that it declined the most dramatically and therefore may need to be managed separately. Our modeling approach may be useful in other situations in which targeted data are sparse and conservation status and best management approach for multiple species are uncertain.

To predict the niche, model colonization and extinction.

Ecologists frequently try to predict the future geographic distributions of species. Most studies assume that the current distribution of a species reflects its environmental requirements (i.e., the species' niche). However, the current distributions of many species are unlikely to be at equilibrium with the current distribution of environmental conditions, both because of ongoing invasions and because the distribution of suitable environmental conditions is always changing. This mismatch between the equilibrium assumptions inherent in many analyses and the disequilibrium conditions in the real world leads to inaccurate predictions of species' geographic distributions and suggests the need for theory and analytical tools that avoid equilibrium assumptions. Here, we develop a general theory of environmental associations during periods of transient dynamics. We show that time-invariant relationships between environmental conditions and rates of local colonization and extinction can produce substantial temporal variation in occupancy-environment relationships. We then estimate occupancy-environment relationships during three avian invasions. Changes in occupancy-environment relationships over time differ among species but are predicted by dynamic occupancy models. Since estimates of the occupancy-environment relationships themselves are frequently poor predictors of future occupancy patterns, research should increasingly focus on characterizing how rates of local colonization and extinction vary with environmental conditions.

Herbivores Enforce Sharp Boundaries Between Terrestrial and Aquatic Ecosystems

The transitions between ecosystems (ecotones) are often biodiversity hotspots, but we know little about the forces that shape them. Today, often sharp boundaries with low diversity are found between terrestrial and aquatic ecosystems. This has been attributed to environmental factors that hamper succession. However, ecosystem properties are often controlled by both bottom-up and top-down forces, but their relative importance in shaping riparian boundaries is not known. We hypothesize that (1) herbivores may enforce sharp transitions between terrestrial and aquatic ecosystems by inhibiting emergent vegetation expansion and reducing the width of the transition zone and (2) the vegetation expansion, diversity, and species turnover are related to abiotic factors in the absence of herbivores, but not in their presence. We tested these hypotheses in 50 paired grazed and ungrazed plots spread over ten wetlands, during two years. Excluding grazers increased vegetation expansion, cover, biomass, and species richness. In ungrazed plots, vegetation cover was negatively related to water depth, whereas plant species richness was negatively related to the vegetation N:P ratio. The presence of (mainly aquatic) herbivores overruled the effect of water depth on vegetation cover increase but did not interact with vegetation N:P ratio. Increased local extinction in the presence of herbivores explained the negative effect of herbivores on species richness, as local colonization rates were unaffected by grazing. We conclude that (aquatic) herbivores can strongly inhibit expansion of the riparian vegetation and reduce vegetation diversity over a range of environmental conditions. Consequently, herbivores enforce sharp boundaries between terrestrial and aquatic ecosystems.

Edge creation and tree dieback influence the patch-tracking metapopulation dynamics of a red-listed epiphytic bryophyte

1. Edges in landscapes have an effect on the abundance of many species but the underlying ecological mechanisms are poorly known for most taxonomic groups. One way to gain insight into the mechanisms is to examine how key demographic or metapopulation parameters are affected by proximity to edge. The main objective of this study was to investigate the effects of edge creation through clearcutting on the dynamics of forest species’ metapopulations. 2. We used the red-listed epiphytic moss Neckera pennata as a model species. Based on data from repeated surveys of a metapopulation and its host tree network in a hemiboreal forest, we tested the effect of edge creation on key metapopulation parameters: rates of local colonization and extinction, local abundance growth and patch destruction through the fall of host trees. We predicted the long-term consequences of the edge effects using simulations with Bayesian statistical models. We also explored the potential effects of the pathogen Chalara fraxinea causing ash dieback, a tree disease currently spreading in Europe. 3. The colonization probability on host trees unoccupied by the moss increased with increasing connectivity to occupied trees. The growth of local populations on occupied trees decreased with increasing proximity to edge, and with initial local abundance. Stochastic extinctions of the epiphyte from standing trees were very rare and only occurred near the edge; most of the observed extinctions were deterministic due to tree fall. Tree fall decreased with increasing distance from the edge into the forest, and with increasing tree diameter. 4. Under edge conditions, simulations predicted decreases in the total number of host trees, number of occupied host trees, and in the total abundance of the epiphyte over a 30-year period. We suggest that ash dieback increases the tree fall rate and thereby the local extinction rate, leading to increased metapopulation extinction risk. 5. Synthesis and applications. The results show that small protected forest areas such as woodland key habitats may not allow long-term persistence of red-listed epiphytes if they are influenced by edge creation through clearcutting. Delineating uncut buffers of 50–100 m around the protected areas may alleviate such effects.

Rates of local colonization and extinction reveal different plant community assembly mechanisms on road verges in central Spain

Question: Does community assembly on roadsides differ between two contrasting habitats along a stress–productivity gradient? Is establishment success determined by the regional pool, environmental filters or historical events? Location: Highway roadcuts and embankments in central Spain (401290N, 031340W). Methods: Species composition was recorded annually in 45 plots distributed on steep slopes with newly exposed surfaces (roadcuts) or newly built topsoiled substrates (embankments), for 4 years following hydroseeding with standard or native seed mixture. Frequency of appearance and local colonization and extinction rates of individual species were clustered in establishment success groups. We examined the correspondence between descriptive plant traits and species performance on both roadslope types. Results: Roadslope species richness showed a sustained increase over time, although at consistently lower levels and rates on the more productive embankments. Sixty per cent of the colonization events involved species from the surrounding vegetation. Hydroseeded species persisted through time, but did not modify community composition or dynamics. A higher establishment success rate was found in wind-pollinated species with large seeds and in exotic species. Those species growing on embankments showed an equal or higher establishment success rate on roadcuts and, conversely, species from roadcuts exhibited an equal or lower success rate on embankments. Conclusions: At the initial stages of community assemblage on unproductive newly created areas, species richness was shaped by the regional species pool. Communities on less harsh topsoiled embankments were subjected, however, to the filtering effect of competitive exclusion. In a reclamation context, efforts to increase site productivity may have detrimental consequences for in situ conservation of local diversity.

Colonization and Extinction Patterns of a Metapopulation of Gold-spotted Pond Frogs, Rana plancyi chosenica

We investigated colonization and extinction patterns in a meta population of the gold-spotted pond frog (Rana planeyi ehoseniea) near the Korea National University of Education, Chungbuk, Korea, by surveying the frogs in the nine occupied habitat patches in the study area four times per breeding season for three years (20062008) and recording whether the patches were occupied by frogs as well as how many frogs were calling in the patches. We then developed five a priori year-specific models using the Akaike Information Criterion (AIC). The models predicted that: 1) probabilities of colonization and local extinction of the frogs were better explained by year-dependent models than by constant models, 2) there are high local extinction and low colonization probabilities, 3) approximately 31% number of patches will be occupied at equilibrium, and 4) that considerable variation in occupation rate should occur over a 30-year period, due to demographic stochasticity (in our model, the occupation rate ranged from 0.222 to 0.889). Our results suggest that colonization is important in this metapopulation system, which is governed by mainly stochastic components, and that more constructive conservation effects are needed to increase local colonization rates.

The Park Grass Experiment 1856–2006: its contribution to ecology

Summary The Park Grass Experiment, begun in 1856, is the oldest ecological experiment in existence. Its value to science has changed and grown since it was founded to answer agricultural questions. In recent times the experiment has shown inter alia how: plant species richness, biomass and pH are related; community composition responds to climatic perturbation and nutrient additions; soil is acidified and corrected by liming. It also provided one of the first demonstrations of the evolution of adaptation at a very local scale and contains a putative case of the evolution of reproductive isolation by reinforcement. The application of molecular genetic markers to archived plant material promises to reveal a whole new chapter of genetic detail about the long‐term dynamics of plant populations. Over the range of values observed at Park Grass, biomass (productivity) has a negative effect upon species richness. Any positive effect of species richness on productivity could only be weak by comparison. The experiment provides support for both the competitive exclusion and pool size hypotheses for determination of species density. Instantaneous comparisons of species richness between plots do not accurately reflect temporal rates of loss which may be multiplicative rather than additive. This suggests that comparisons among sites, nutrient inputs, especially N treatments, or soil acidity may in general underestimate the threat posed to plant species diversity by long‐term changes in plant nutrient availability, both enrichment and depletion. Differences between plots at the community level are maintained despite a flow of propagules between plots. There is no strong evidence for a spatial mass effect. Guild (grass/legume/other) compositions of plant communities have equilibrated, but the species composition within guilds is more dynamic and continues to change over time, suggesting that species and guild abundances are independently regulated. At least some members of all the major trophic levels, including predators (spiders), herbivores (leafhoppers) and detritivores (springtails) are treatment‐specific in their distributions. Plant populations on Park Grass are subdivided by treatments which, to some degree, have led to plots becoming genetically isolated from one another and decoupled demographically. This subdivision has created a metapopulation structure in each species, characterized by species‐specific rates of local colonization and extinction. Inverse clines in flowering time occur in the grass Anthoxanthum odoratum across some plot boundaries. These suggest that reproductive isolation between plots has been reinforced by natural selection. Drift as well as selection may have taken place in A. odoratum, especially on plots where effective population size is restricted by population bottlenecks caused by drought. Park Grass illustrates how long‐term experiments grow in value with time and how they may be used to investigate scientific questions that were inconceivable at their inception. This is as likely to be true of the future of Park Grass as it has proved to be of its past.

Vegetation dynamics following resource manipulations in herb-rich woodland

Changes in species richness, turnover, composition and above-ground biomass of herb-rich woodland were documented following fertilizer application and water addition over three growing seasons. Addition of fertilizer significantly reduced species richness relative to unmanipulated control and water addition plots after 3 years. This change coincided with significant increases in biomass, which were largely due to increased growth of exotic annual grasses. The reductions in richness observed in the fertilized plots were a consequence of both lower rates of local colonization and enhanced rates of local extinction of the resident species. Species loss was not random; native species were lost after nutrient addition whereas exotic species were not. Nutrient limitation was more important for species coexistence in these communities than was water availability.

Invasion impacts local species turnover in a successional system

AbstractExotic plant invasions are often associated with declines in diversity within invaded communities. However, few studies have examined the local community dynamics underlying these impacts. Changes in species richness associated with plant invasions must occur through local changes in extinction and/or colonization rates within the community. We used long‐term, permanent plot data to evaluate the impacts of the exotic vine Lonicera japonica. Over time, species richness declined with increasing L. japonica cover. L. japonica reduced local colonization rates but had no effect on extinction rates. Furthermore, we detected significant reductions in the immigration of individual species as invasion severity increased, showing that some species are more susceptible to invasion than others. These findings suggest that declines in species richness associated with L. japonica invasion resulted from effects on local colonization rates only and not through the competitive displacement of established species.

Large‐scale dynamics in colonization and extinction for breeding birds in Britain

1 A number of generalizations have been made as to the effects of the area of occupancy, population size, dispersal ability and body size of species on their relative rates of local colonization and extinction. 2 Here, data on the breeding bird assemblage of Britain are used to test these generalizations. The complete geographical ranges of British birds have been censused twice, in the periods 1968–72 and 1988–91, allowing rates of colonization and extinction between these periods to be estimated. 3 The local colonization dynamics of species are influenced independently by their range sizes and the dispersal abilities of adult birds: species with smaller range sizes and larger dispersal distances were more likely to have colonized new areas between the two census periods. 4 The local extinction dynamics of species are influenced independently by their population sizes and body masses: species with smaller population sizes and body sizes were more likely to have gone extinct from areas inhabited in the first census period. 5 These results remain when controlling for the effects of phylogenetic relatedness. 6 These analyses uphold many commonly held generalizations about the correlates of local colonization and extinction, and suggest that the long-term evolutionary history of these bird species has influenced their potential to respond to current ecological conditions.

Finite metapopulation models with density–dependent migration and stochastic local dynamics

The effects of small density-dependent migration on the dynamics of a metapopulation are studied in a model with stochastic local dynamics. We use a diffusion approximation to study how changes in the migration rate and habitat occupancy affect the rates of local colonization and extinction. If the emigration rate increases or if the immigration rate decreases with local population size, a positive expected rate of change in habitat occupancy is found for a greater range of habitat occupancies than when the migration is density-independent. In contrast, the reverse patterns of density dependence in respective emigration and immigration reduce the range of habitat occupancies where the metapopulation will be viable. This occurs because density-dependent migration strongly influences both the establishment and rescue effects in the local dynamics of metapopulations.

Association between Climate and Enterobacter Colonization in Swedish Neonatal Units

To study risk factors for the highly variable local colonization rates with unrelated Enterobacter species strains previously found in 22 Swedish neonatal units (0% to 32.4% of the infants).The fecal Enterobacter species carriage rates among 953 infants in the 22 special-care neonatal units were correlated with variables related to the ward (size, crowding, staffing, work load, antibiotic usage, level of care, hygienic precautions), and the hospital (temperature of water supplied, geographical location).The average Enterobacter species carriage rate was highest at seven days of age (17% of the infants) and then declined to 3%. Only location of the hospital in an area with warmer climate according to horticultural zone showed an association with Enterobacter species carriage in multivariate analysis (P = 0.005).Although Enterobacter species mainly cause nosocomially acquired infections, the occurrence of the organism in special-care neonatal units seemed to be determined more by extrahospital than by intrahospital factors.

Association between climate and Enterobacter colonization in Swedish neonatal units.

To study risk factors for the highly variable local colonization rates with unrelated Enterobacter species strains previously found in 22 Swedish neonatal units (0% to 32.4% of the infants). The fecal Enterobacter species carriage rates among 953 infants in the 22 special-care neonatal units were correlated with variables related to the ward (size, crowding, staffing, work load, antibiotic usage, level of care, hygienic precautions), and the hospital (temperature of water supplied, geographical location). The average Enterobacter species carriage rate was highest at seven days of age (17% of the infants) and then declined to 3%. Only location of the hospital in an area with warmer climate according to horticultural zone showed an association with Enterobacter species carriage in multivariate analysis (P = 0.005). Although Enterobacter species mainly cause nosocomially acquired infections, the occurrence of the organism in special-care neonatal units seemed to be determined more by extrahospital than by intrahospital factors.