Fine-scale Environmental Heterogeneity Research Articles

Ectoparasite infection levels differ between fish from upwelling-exposed and sheltered rocky reefs areas in Brazil

Ubiquitous and abundant within marine ecosystems, parasites play essential ecological roles such as shaping host population dynamics, altering competition between species, and influencing energy flows through communities. Their diversity and population dynamics are demonstrably shaped by both seasonal and geographic variations. These variations have been often explored at broad spatial scale. However, parasite communities can exhibit significant disparities even at small spatial scales, driven by factors such as wave exposure, temperature fluctuations, and benthic habitat composition. We investigated how crustacean parasites of fish – caligids and gnathiids - differed between two distinct habitats, which are separated solely by a few kilometres, at Arraial do Cabo, Brazil. These two habitats are characterised by a sheltered embayment (hereinafter referred to as “inside”) or an exposed upwelling habitat (hereinafter referred to as “outside”). Individual fish from four species were examined in both habitats. We found that the infestation rate of caligids varied among fish species and, gnathiids varied between the two sampling sites. Gnathiids were absent from fish outside, while they were present on fish inside the embayment. This disparity suggests a critical role of local environmental factors in shaping gnathiid distribution. Potential drivers include temperature fluctuations, substrate composition, and wave exposure, which differed markedly between the two sites. Conversely, caligid parasites infected fish in both locations. While environmental factors may also influence caligid abundance, they appear to exhibit greater tolerance compared to gnathiids. These findings emphasize the importance of incorporating fine-scale environmental heterogeneity when investigating parasite distribution patterns.

Alpine Extremophytes in Evolutionary Turmoil: Complex Diversification Patterns and Demographic Responses of a Halophilic Grass in a Central Asian Biodiversity Hotspot.

Diversification and demographic responses are key processes shaping species evolutionary history. Yet we still lack a full understanding of ecological mechanisms that shape genetic diversity at different spatial scales upon rapid environmental changes. In this study, we examined genetic differentiation in an extremophilic grass Puccinellia pamirica and factors affecting its population dynamics among the occupied hypersaline alpine wetlands on the arid Pamir Plateau in Central Asia. Using genomic data, we found evidence of fine-scale population structure and gene flow among the localities established across the high-elevation plateau as well as fingerprints of historical demographic expansion. We showed that an increase in the effective population size could coincide with the Last Glacial Period, which was followed by the species demographic decline during the Holocene. Geographic distance plays a vital role in shaping the spatial genetic structure of P. pamirica alongside with isolation-by-environment and habitat fragmentation. Our results highlight a complex history of divergence and gene flow in this species-poor alpine region during the Late Quaternary. We demonstrate that regional climate specificity and a shortage of nonclimate data largely impede predictions of future range changes of the alpine extremophile using ecological niche modeling. This study emphasizes the importance of fine-scale environmental heterogeneity for population dynamics and species distribution shifts.

Thermal plasticity over a marine-estuarine ecocline can buffer a tropical fish from warming

Intraspecific variation in thermal tolerance can favor species persistence in a warmer ocean, but is often overlooked in fine-scale studies. Nonetheless, local drivers (e.g. salinity) interact with temperature to shape species' thermal response. Here, we acclimated juveniles of Brazilian silversides Atherinella brasiliensis captured at the limits of a marine-estuarine ecocline under reciprocal-cross conditions, to test for phenotypic plasticity in heat tolerance. We also tested whether silversides acclimated to temperatures predicted for 2100 (+3–4.5 °C). Fish in warm-brackish waters showed higher CTMax (Critical Thermal Maximum) than those in cold-marine conditions, regardless of their origin. Silversides’ CTMax reached up to 40.6 °C, but it did not increase after exposure to temperatures predicted for 2100. Lack of acclimation response suggests that silversides heat tolerance has reached a “ceiling”, despite thermal plasticity. Our findings show that fine-scale environmental heterogeneity can promote phenotypic plasticity for tropical species, reducing the risk of short-term extirpation.

Evaluation of fine-scale environmental heterogeneity and its effect on terrestrial mammal diversity in a grassland in the Chihuahuan Desert

Environmental heterogeneity is expected to increase the richness and diversity of species. The aim of this study is to identify environmental elements that influence the richness and diversity of small, medium and large species of mammals. For the sampling of mammals, camera and Sherman traps were used at 18 sampling stations. For vegetation sampling, eight large-scale explanatory variables were selected in a 1000 m2 plot at each station. The diversity (H’) of habitats (heterogeneity) and species were measured using the Shannon–Wiener diversity index. To relate the heterogeneity and environmental variables with species richness for small, medium and large mammals, a generalised linear model (GLM) was used while considering the Poisson probability distribution. A multiple linear regression model was used for data with normal distributions. The results suggested that environmental elements could favour rodent richness but may have a negative effect on community evenness and some environmental variables that are related to the diversity of rodent species and the species richness of medium and large mammals. It was concluded that the relationship between environmental heterogeneity and mammal richness is complex and that the response to the biotic elements of the landscape is diverse and different for each species.

Consistent trait-environment relationships within and across tundra plant communities.

A fundamental assumption in trait-based ecology is that relationships between traits and environmental conditions are globally consistent. We use field-quantified microclimate and soil data to explore if trait-environment relationships are generalizable across plant communities and spatial scales. We collected data from 6,720 plots and 217 species across four distinct tundra regions from both hemispheres. We combined these data with over 76,000 database trait records to relate local plant community trait composition to broad gradients of key environmental drivers: soil moisture, soil temperature, soil pH and potential solar radiation. Results revealed strong, consistent trait-environment relationships across Arctic and Antarctic regions. This indicates that the detected relationships are transferable between tundra plant communities also when fine-scale environmental heterogeneity is accounted for, and that variation in local conditions heavily influences both structural and leaf economic traits. Our results strengthen the biological and mechanistic basis for climate change impact predictions of vulnerable high-latitude ecosystems.

Fine-scale environmental heterogeneity and spatial niche partitioning among spring-flowering forest herbs.

Environmental heterogeneity influences plant distributions and diversity at several spatial scales. In temperate forests, fine-scale environmental variation may promote local coexistence among herbaceous species by allowing plants to spatially partition microsites within forest stands. Here we argue that shallow soils, low soil water-holding capacity and fertility, and reduced light near tree boles should favor short, shallow-rooted, evergreen species like Anemone acutiloba with low moisture, nutrient, and light requirements. Farther from trees, richer, deeper soils should favor taller, deeper-rooted herbs with greater moisture and nutrient demands, such as Sanguinaria canadensis and Trillium flexipes. We tested these hypotheses by mapping the fine-scale distributions of Anemone, Sanguinaria, and Trillium individuals within a 50 × 50 m plot, comparing local species' distributions with respect to soil depth and proximity to neighboring trees, and characterizing intraspecific and interspecific spatial associations. Local plant distributions were consistent with our predictions based on leaf height, physiology, and phenology. Anemone was found in microsites on shallower soils and closer to trees than either Sanguinaria or Trillium. In all three species, individual plants were spatially aggregated within 2 m, but spatially segregated from individuals of the other species beyond 2 m. Differential plant responses to fine-scale environmental heterogeneity and observed spatial associations suggest that local species-environment associations could facilitate coexistence. These findings illustrate how fine-scale environmental heterogeneity coupled with phenological and physiological differences likely contribute to spatial niche partitioning among spring-flowering forest herbs and maintain high local plant diversity within temperate forests.

Incorporating fine-scale environmental heterogeneity into broad-extent models.

A key aim of ecology is to understand the drivers of ecological patterns, so that we can accurately predict the effects of global environmental change. However, in many cases, predictors are measured at a finer resolution than the ecological response. We therefore require data aggregation methods that avoid loss of information on fine‐grain heterogeneity.We present a data aggregation method that, unlike current approaches, reduces the loss of information on fine‐grain spatial structure in environmental heterogeneity for use with coarse‐grain ecological datasets. Our method contains three steps: (a) define analysis scales (predictor grain, response grain, scale‐of‐effect); (b) use a moving window to calculate a measure of variability in environment (predictor grain) at the process‐relevant scale (scale‐of‐effect); and (c) aggregate the moving window calculations to the coarsest resolution (response grain). We show the theoretical basis for our method using simulated landscapes and the practical utility with a case study. Our method is available as the grainchanger r package.The simulations show that information about spatial structure is captured that would have been lost using a direct aggregation approach, and that our method is particularly useful in landscapes with spatial autocorrelation in the environmental predictor variable (e.g. fragmented landscapes) and when the scale‐of‐effect is small relative to the response grain. We use our data aggregation method to find the appropriate scale‐of‐effect of land cover diversity on Eurasian jay Garrulus glandarius abundance in the UK. We then model the interactive effect of land cover heterogeneity and temperature on G. glandarius abundance. Our method enables us quantify this interaction despite the different scales at which these factors influence G. glandarius abundance.Our data aggregation method allows us to integrate variables that act at varying scales into one model with limited loss of information, which has wide applicability for spatial analyses beyond the specific ecological context considered here. Key ecological applications include being able to estimate the interactive effect of drivers that vary at different scales (such as climate and land cover), and to systematically examine the scale dependence of the effects of environmental heterogeneity in combination with the effects of climate change on biodiversity.

Environmental heterogeneity mediates scale-dependent declines in kelp diversity on intertidal rocky shores.

Biodiversity loss is driven by interacting factors operating at different spatial scales. Yet, there remains uncertainty as to how fine-scale environmental conditions mediate biological responses to broad-scale stressors. We surveyed intertidal rocky shore kelp beds situated across a local gradient of wave action and evaluated changes in kelp diversity and abundance after more than two decades of broad scale stressors, most notably the 2013–2016 heat wave. Across all sites, species were less abundant on average in 2017 and 2018 than during 1993–1995 but changes in kelp diversity were dependent on wave exposure, with wave exposed habitats remaining stable and wave sheltered habitats experiencing near complete losses of kelp diversity. In this way, wave exposed sites have acted as refugia, maintaining regional kelp diversity despite widespread local declines. Fucoids, seagrasses and two stress-tolerant kelp species (Saccharina sessilis, Egregia menziesii) did not decline as observed in other kelps, and the invasive species Sargassum muticum increased significantly at wave sheltered sites. Long-term monitoring data from a centrally-located moderate site suggest that kelp communities were negatively impacted by the recent heatwave which may have driven observed losses throughout the region. Wave-sheltered shores, which saw the largest declines, are a very common habitat type in the Northeast Pacific and may be especially sensitive to losses in kelp diversity and abundance, with potential consequences for coastal productivity. Our findings highlight the importance of fine-scale environmental heterogeneity in mediating biological responses and demonstrate how incorporating differences between habitat patches can be essential to capturing scale-dependent biodiversity loss across the landscape.

Cultural monuments and nature conservation: a review of the role of kurgans in the conservation and restoration of steppe vegetation

Steppe is among the most endangered biomes of the world, especially in Eastern Europe, where more than 90 % of original steppes have been destroyed due to conversion into croplands, afforestation and other human activities. Currently, steppe vegetation is often restricted to places unsuitable for ploughing, such as ancient burial mounds called kurgans. The aim of our study was to collect and synthesise existing knowledge on kurgans by a review of research papers and grey literature. The proportion of kurgans covered by steppe vegetation increases from west to east and from lowlands to uplands. Despite their small size, kurgans act as biodiversity hotspots and harbour many red-listed species. High overall species richness and a high proportion of grassland specialists are maintained by a pronounced fine-scale environmental heterogeneity. The main factors threatening the biodiversity of kurgans are intensified agriculture and construction works. We conclude that kurgans can play a crucial role in preserving steppe vegetation, especially in intensively used agricultural landscapes in the western part of the steppe zone. Despite the vital role of kurgans in sustaining steppe vegetation, we identified serious knowledge gaps on their distribution, vegetation, flora and fauna and their potential role in steppe restoration.

Beyond the classical nurse species effect: diversity assembly in a Mediterranean semi-arid dwarf shrubland

Questions Shrublands often exhibit very high levels of species richness. We ask which mechanisms underlie plant species co-existence in a hyper-diverse Mediterranean shrubland, and the scales at which individual species exert their effects during organization of diversity in these communities. Location Brea de Tajo, Madrid, Spain. Methods We computed the individual species–area relationships (ISARs) in a completely mapped Mediterranean dwarf shrubland. Using appropriate species-specific null models, we evaluated how diversity was organized around individual species, covering a wide array of spatial scales. Results We mapped 7858 individuals from 48 perennial species. Our analyses showed that most species (80%) acted as accumulators and exhibited more diversity than expected, whereas 20% were neutral at fine scales. At larger spatial scales, most species exhibited no significant effects in terms of diversity accumulation. Conclusions In contrast to other species-rich ecosystems such as tropical forests, our results suggest that positive interactions and niche complementarities are the main drivers of community assembly in these communities. The most important finding was the existence of non-hierarchical multi-specific clumping that was independent of the fine-scale environmental heterogeneity, which is a pattern that has not been described previously, to the best of our knowledge. The abundance of accumulator species and the similarity among the ISAR curves suggest that nurse-like species were not present, thereby indicating that a different ecological mechanism led to the emergence of this pattern.

Ground layer plant species turnover and beta diversity in southern-European old-growth forests.

Different assembly processes may simultaneously affect local-scale variation of species composition in temperate old-growth forests. Ground layer species diversity reflects chance colonization and persistence of low-dispersal species, as well as fine-scale environmental heterogeneity. The latter depends on both purely abiotic factors, such as soil properties and topography, and factors primarily determined by overstorey structure, such as light availability. Understanding the degree to which plant diversity in old-growth forests is associated with structural heterogeneity and/or to dispersal limitation will help assessing the effectiveness of silvicultural practices that recreate old-growth patterns and structures for the conservation or restoration of plant diversity. We used a nested sampling design to assess fine-scale species turnover, i.e. the proportion of species composition that changes among sampling units, across 11 beech-dominated old-growth forests in Southern Europe. For each stand, we also measured a wide range of environmental and structural variables that might explain ground layer species turnover. Our aim was to quantify the relative importance of dispersal limitation in comparison to that of stand structural heterogeneity while controlling for other sources of environmental heterogeneity. For this purpose, we used multiple regression on distance matrices at the within-stand extent, and mixed effect models at the extent of the whole dataset. Species turnover was best predicted by structural and environmental heterogeneity, especially by differences in light availability and in topsoil nutrient concentration and texture. Spatial distances were significant only in four out of eleven stands with a relatively low explanatory power. This suggests that structural heterogeneity is a more important driver of local-scale ground layer species turnover than dispersal limitation in southern European old-growth beech forests.

The geography and ecology of plant speciation: range overlap and niche divergence in sister species

A goal of evolutionary biology is to understand the roles of geography and ecology in speciation. The recent shared ancestry of sister species can leave a major imprint on their geographical and ecological attributes, possibly revealing processes involved in speciation. We examined how ecological similarity, range overlap and range asymmetry are related to time since divergence of 71 sister species pairs in the California Floristic Province (CFP). We found that plants exhibit strikingly different age-range correlation patterns from those found for animals; the latter broadly support allopatric speciation as the primary mode of speciation. By contrast, plant sisters in the CFP were sympatric in 80% of cases and range sizes of sisters differed by a mean of 10-fold. Range overlap and range asymmetry were greatest in younger sisters. These results suggest that speciation mechanisms broadly grouped under 'budding' speciation, in which a larger ranged progenitor gives rise to a smaller ranged derivative species, are probably common. The ecological and reproductive similarity of sisters was significantly greater than that of sister-non-sister congeners for every trait assessed. However, shifts in at least one trait were present in 93% of the sister pairs; habitat and soil shifts were especially common. Ecological divergence did not increase with range overlap contrary to expectations under character displacement in sympatry. Our results suggest that vicariant speciation is more ubiquitous in animals than plants, perhaps owing to the sensitivity of plants to fine-scale environmental heterogeneity. Despite high levels of range overlap, ecological shifts in the process of budding speciation may result in low rates of fine-scale spatial co-occurrence. These results have implications for ecological studies of trait evolution and community assembly; despite high levels of sympatry, sister taxa and potentially other close relatives, may be missing from local communities.

Effects of fine-scale environmental heterogeneity on local genetic structure in Macoma balthica from the Gulf of Gdañsk (southern Baltic Sea)

It is increasingly recognized that populations of marine organisms with potential for large-scale dispersal may exhibit fine-scale genetic structure. The Gulf of Gdansk (Poland) is an interesting setting to study fine-grained population structure in marine organisms as it is characterized by chronic anthropogenic pollution and strong salinity gradients. We investigated, at two nested spatial scales (35 and 7 km), genetic structure among populations of the infaunal tellinid bivalve Macoma balthica. The COI mitochondrial marker revealed a likely mix of evolutionary lineages in the Gulf, with no detectable spatial structure. Seven microsatellite markers detected weak population structure, separating deep and shallow populations within short distances (7 km) and assignment tests suggested asymmetric gene flow among these populations, with no shallow recruits being detected in deep waters. Given the specific environmental conditions encountered at deeper depths in the Gulf (increased salinity, lower temperatures, oxygen depletion, hydrogen sulfide pollution), we suggest that deeper populations may be subjected to local adaptation.

Positive diversity-invasibility relationship in species-rich semi-natural grassland at the neighbourhood scale

Attempts to answer the old question of whether high diversity causes high invasion resistance have resulted in an invasion paradox: while large-scale studies often find a positive relationship between diversity and invasibility, small-scale experimental studies often find a negative relationship. Many of the small-scale studies are conducted in artificial communities of even-aged plants. Species in natural communities, however, do not represent one simultaneous cohort and occur at various levels of spatial aggregation at different scales. This study used natural patterns of diversity to assess the relationship between diversity and invasibility within a uniformly managed, semi-natural community. In species-rich grassland, one seed of each of ten species was added to each of 50 contiguous 16 cm(2) quadrats within seven plots (8 × 100 cm). The emergence of these species was recorded in seven control plots, and establishment success was measured in relation to the species diversity of the resident vegetation at two spatial scales, quadrat (64 cm(2)) within plots (800 cm(2)) and between plots within the site (approx. 400 m(2)) over 46 months. Invader success was positively related to resident species diversity and richness over a range of 28-37 species per plot. This relationship emerged 7 months after seed addition and remained over time despite continuous mortality of invaders. Biotic resistance to plant invasion may play only a sub-ordinate role in species-rich, semi-natural grassland. As possible alternative explanations for the positive diversity-invasibility relationship are not clear, it is recommended that future studies elaborate fine-scale environmental heterogeneity in resource supplies or potential resource flows from resident species to seedlings by means of soil biological networks established by arbuscular mycorrhizal fungi.

The Chemical Pathway of Carotenoids: From Plants to Birds

Carotenoids are interesting in avian ecology due their dietary origin and the resulting environmental dependence of certain functions and traits such as plumage pigmentation and the immune system. Carotenoids are frequently assumed to be limited for birds, but data on availability is rarely collected. Thus, to understand the role of the carotenoids as mediators of trade-offs, additional information on the whole chemical pathway of carotenoids is needed. In conjunction with previous studies on carotenoid ecology and physiology of the Great Tit Parus major living in Swedish urban vs. rural environments, I here present data on 1) carotenoid concentration in two primary carotenoid producers, birch Betula verrucosa and oak Quercus robur and 2) vegetation cover on Great Tit territories. Previous studies found lower carotenoid concentrations in caterpillars and plumage pigmentation in urban compared to rural areas. Correspondingly, urban trees had significantly lower total carotenoid concentration in comparison to rural trees, but there was no environmental difference in the relative amount of lutein and zeaxanthin (Lut:Zx ratio) in sun-exposed leaves. Second, urban Great Tits had significantly more shade in their territories than rural birds, which may affect the leaves Lut:Zx composition. The results, taken together, show that the environmental difference in carotenoid concentration of caterpillars and Great Tit yellow breast coloration, may occur because urban trees synthesise less or have used up more carotenoids in response to environmental stress compared to rural trees. Thus, tracking carotenoid content in different steps of the food chain raises questions about how fine-scale environmental heterogeneity can affect optimal foraging behaviour and individual variation in carotenoid-dependent traits.

CHANGING THE GAP DYNAMICS PARADIGM: VEGETATIVE REGENERATION CONTROL ON FOREST RESPONSE TO DISTURBANCE

Understanding the manner in which changes in disturbance regimes will affect forest biodiversity is an important goal of global change research. Prevailing theories of recruitment after disturbance center on the role of pioneer species; predictions of forest biodiversity focus almost exclusively on dispersal and shade tolerance while vegetative reproduction is virtually omitted from models and serious discussions of the topic. However, the persistence of live damaged trees increases understory shade, generates fine-scale environmental heterogeneity, and moderates ecosystem responses to damage, while the sprouting of damaged trees offers a shortcut to reestablishment of the canopy. While a number of studies document snapshots of post-disturbance vegetative reproduction, we lack an understanding of the underlying demographic processes needed in order to both comprehend and predict observed patterns. In this study we quantify the abundance, competitive ability, and interspecific variability of vegetative reproduction in 18 replicated experimental gaps in the southern Appalachians and Carolina Piedmont, USA, in order to assess the potential role of sprouting in driving gap dynamics. Annual rates of damaged adult survival, sprout initiation, growth, and mortality were monitored over four years and compared to the performance of gap-regenerating saplings. Recruitment from sprouts was found to constitute 26–87% of early gap regeneration and forms the dominant pathway of regeneration for some species. Sprouts from recently damaged trees also grow significantly faster than the saplings with which they compete. For all measured demographic rates (damaged tree survival, sprout initiation, number, growth, and survival) differences among species are large and consistent across sites, suggesting that vegetative reproduction is an important and non-neutral process in shaping community composition. Sprouting ability does not correlate strongly with other life-history trade-offs, thus sprouting potentially provides an alternate trait axis in promoting diversity.

Analysis of spatial genetic structure in an expanding Pinus halepensis population reveals development of fine‐scale genetic clustering over time

We analysed the change of spatial genetic structure (SGS) of reproductive individuals over time in an expanding Pinus halepensis population. To our knowledge, this is the first empirical study to analyse the temporal component of SGS by following the dynamics of successive cohorts of the same population over time, rather than analysing different age cohorts at a single time. SGS is influenced by various factors including restricted gene dispersal, microenvironmental selection, mating patterns and the spatial pattern of reproductive individuals. Several factors that affect SGS are expected to vary over time and as adult density increases. Using air photo analysis, tree-ring dating and molecular marker analysis we reconstructed the spread of reproductive individuals over 30 years beginning from five initial individuals. In the early stages, genotypes were distributed randomly in space. Over time and with increasing density, fine-scale (< 20 m) SGS developed and the magnitude of genetic clustering increased. The SGS was strongly affected by the initial spatial distribution and genetic variation of the founding individuals. The development of SGS may be explained by fine-scale environmental heterogeneity and possibly microenvironmental selection. Inbreeding and variation in reproductive success may have enhanced SGS magnitude over time.

Effects of land use and fine-scale environmental heterogeneity on net ecosystem production over a temperate coniferous forest landscape

Effects of land use and fine-scale environmental heterogeneity on net ecosystem production over a temperate coniferous forest landscape

Effects of land use and fine-scale environmental heterogeneity on net ecosystem production over a temperate coniferous forest landscape

In temperate coniferous forests, spatial variation in net ecosystem production (NEP) is often associated with variation in stand age and heterogeneity in environmental factors such as soil depth. However, coarse spatial resolution analyses used to evaluate the terrestrial contribution to global NEP do not generally incorporate these effects. In this study, a fine-scale (25 m grid) analysis of NEP over a 164-km2 area of productive coniferous forests in the Pacific Northwest region of the United States was made to evaluate the effects of including fine scale information in landscape-scale NEP assessments. The Enhanced Thematic Mapper (ETM×) sensor resolved five cover classes in the study area and further differentiated between young, mature and old-growth conifer stands. ETM× was also used to map current leaf area index (LAI) based on an empirical relationship of observed LAI to spectral vegetation indices. A daily time step climatology, based on 18 years of meteorological observations, was distributed (1 km resolution) over the mountainous terrain of the study area using the DAYMET model. Estimates of carbon pools and flux associated with soil, litter, coarse woody debris and live trees were then generated by running a carbon cycle model (Biome-BGC) to a state that reflected the current successional status and LAI of each grid cell, as indicated by the remote sensing observations. Estimated annual NEP for 1997 over the complete study area averaged 230 g C m−2, with most of the area acting as a carbon sink. The area-wide NEP is strongly positive because of reduced harvesting in the last decade and the recovery of areas harvested between 1940 and 1990. The average value was greater than would be indicated if the entire area was assumed to be a mature conifer stand, as in a coarse-scale analysis. The mean NEP varied interannually by over a factor of two. This variation was 38% less than the interannual variation for a single point. The integration of process models with ground surface information provided by remote sensing provides a framework for investigating mechanisms regulating NEP and evaluating coarse resolution globally applied NEP scaling efforts.

Extending the quasi-neutral concept

In caricature, the equilibrium paradigm of community ecology states that plant communities are stable entities consisting of competing species - and that such species coexist because each has its "niche". This paradigm, in its extreme, has been dead for some time. Nevertheless, it has yet to be replaced by a credible "non" - equilibrium paradigm. The quasi-neutral concept of plant communities, proposed by Kristjan Zobel, Folia Geobot. 36: 3-8, 2001, possesses some of the key ingredients of a nonequilibrium theory of diversity. It recognizes that there are inescapable relationships between diversity at different scales, that similarity can influence the rate of competitive exclusion, that successional change is typically associated with changes in life form, and that rarefaction (i.e. "sampling artifacts") can have strong effects on fine-scale diversity. However, the current formulation of the quasi-neutral concept is incomplete in that it relies on an unrealistic definition of community, it assumes that random sampling means that species richness at one scale will be linearly related to richness at finer scales, it ignores the possibility of fine-scale processes producing broader-scale patterns, and it avoids the subject of fine-scale environmental heterogeneity. But the most serious limitation of the quasi-neutral concept is that similarity of species alone is not sufficient to allow indefinite coexistence. I present the results of a simple simulation to demonstrate that: (1) identical species will eventually be lost from communities due to stochastic "drift", (2) slight variations in reproductive rates accelerate this loss, but (3) adding a miniscule "cost of commonness" to the model allows the indefinite coexistence of species. I conclude that the quasi-neutral model cannot work without some kind of trade-off.