Limits to the estimation of species richness: The use of relative abundance distributions

The present study demonstrates the possibility of estimating species numbers of animal or plant communities from samples using relative abundance distributions. We use log‐abundance–species‐rank order plots and derive two new estimators that are based on log‐series and lognormal distributions. At small to moderate sample sizes these estimators appear to be more precise than previous parametric and nonparametric estimators. We test our estimators using samples from 171 published medium‐sized to large animal and plant communities taken from the literature. By this we show that our new estimators define also limits of precision.     

Confronting different models of community structure to species-abundance data: a Bayesian model comparison

Species abundances are undoubtedly the most widely available macroecological data, but can we use them to distinguish among several models of community structure? Here we present a Bayesian analysis of species‐abundance data that yields a full joint probability distribution of each model's parameters plus a relatively parameter‐independent criterion, the posterior Bayes factor, to compare these models. We illustrate our approach by comparing three classical distributions: the zero‐sum multinomial (ZSM) distribution, based on Hubbell's neutral model, the multivariate Poisson lognormal distribution (MPLN), based on niche arguments, and the discrete broken stick (DBS) distribution, based on MacArthur's broken stick model. We give explicit formulas for the probability of observing a particular species‐abundance data set in each model, and argue that conditioning on both sample size and species count is needed to allow comparisons between the two distributions. We apply our approach to two neotropical communities (trees, fish). We find that DBS is largely inferior to ZSM and MPLN for both communities. The tree data do not allow discrimination between ZSM and MPLN, but for the fish data ZSM (neutral model) overwhelmingly outperforms MPLN (niche model), suggesting that dispersal plays a previously underestimated role in structuring tropical freshwater fish communities. We advocate this approach for identifying the relative importance of dispersal and niche‐partitioning in determining diversity of different ecological groups of species under different environmental conditions.     

Scale‐dependent frequency distributions of plant species in dune slacks: Dispersal and niche limitation

Question: How do distribution patterns change with increasing scale level, and can this shift be attributed to dispersal and/or niche limitation? Location: Dune slacks at the Belgian and North French coast. Method: Frequency distribution patterns of species were tested over different scale levels (ranging from 0.008 to 45 km²). Analyses were executed for the total species pool and for subsets of species with high and low dispersal rates and habitat generalist and specialist species. Results: Species distributions for the total species pool on scale levels larger than 1.5 km² were unimodal, with an overrepresentation of rare species. With decreasing scale level, the proportion of common species increased, leading to weak bimodality. Distributions of subsets of slowly dispersing and habitat specialist species are on all scale levels characterised by a strongly unimodal pattern. The subset of species with high dispersal rates and habitat generalist species has a higher proportion of common species, leading to a significant core peak. Conclusions: On all scale levels both dispersal and niche limitation are hampering the wider distribution of rare species. However, since isolation and habitat heterogeneity are limited on the smaller scale levels, a larger proportion of well dispersing and habitat generalist species is able to occupy a high number of patches, resulting in a significant core peak for the total species pool.     

Impact of the Timing of Male Emergence on Mating Capacity of Males in Trichogramma evanescens Westwood

In species with highly structured population, such as Trichogramma spp., mating occurs mostly at emergence on the patch and early emerging males have an advantage, as they are present when the first females emerge. However, early emergence of male could also enable males to mature sexually before the emergence of females or enhance their capacity to induce higher receptivity in females. We measured time of emergence of male and female Trichogramma evanescens Westwood (Hymenoptera: Trichogrammatidae) from hosts that were parasitized within a 30 min period. We also measured the effect of male age on their capacity to mate, and the ability of inseminated females to produce daughters. To verify if early emerging males induced higher receptivity in females, we observed the females mate choice between males of different ages. Our results indicated that the mean time of emergence between males and females was 29 min for individuals that develop in 9 days and 10 min for individuals that develop in 10 days. The protandry observed in this species could be the result of either a faster development of males or a male first strategy by the ovipositing female. In T. evanescens, protandry does not permit males to mature sexually nor to induce higher receptivity in females. The main advantage of protandry for males thus appears to be early access to females as they emerge.     

Bioclimatic analyses of distributions of a parasitoid Peristenus digoneutis and its host species Lygus spp. in Europe and North America

• 1 Peristenus digoneutis Loan is a parasitoid of Lygus plant bugs, which was successfully introduced from Europe into North America in the 1980s for controlling native Lygus populations. Surveys confirmed that P. digoneutis populations have become established throughout eastern North America and that the spread of the parasitoid continues. For unknown reasons, previous releases of P. digoneutis in Western Canada were not successful.
• 2 A bioclimate (climex ®; Hearne Scientific Software Pty Ltd, Australia) model for P. digoneutis in North America was developed, based on climate and ecological parameters, and then validated with actual distribution records. The current distribution of P. digoneutis in eastern North America was consistent with the predicted distribution. The model suggests that P. digoneutis will probably continue its spread westwards throughout the U.S.A. along the Great Lakes.
• 3 The southern distribution of P. digoneutis is expected to be limited by hot summer temperatures, whereas its northern range is limited by the number of Lygus host generations rather than cold stress.
• 4 Peristenus digoneutis has the potential to occur in the southern parts of the prairie ecozone of western Canada; however, Ecoclimatic Index values in the prairies indicate mainly marginal or unfavourable conditions, which may explain why earlier releases of P. digoneutis in Western Canada failed.

     

Some genetic consequences of skewed fecundity distributions in plants

Summary Most plant populations show a skewedrd distribution of fecundity amongst their members, in contrast to the poisson distribution assumed by most population genetical theory. We examine by simulation the consequences of skewed fecundity for plant evolution when combined with sieve selection. In comparison with poisson-based theory, plant populations are likely to show a faster response to selection, especially when the favoured allele is at a low frequency. Selection against a deleterious immigrant allele will also be more effective, reducing its equilibrium frequency in a population. In the special case of heterozygote disadvantage traits will evolve that could not under poisson theory. However, random variation is also higher, giving a 10-plant population an effective population size of about 6.4 under poisson theory. The conclusions are not qualitatively changed by different assumptions on the exact shape of the fecundity distribution, or on heritability, or on reproduction by the smallest plants of the population.     

An index to compare geographical distributions of species

We present an index for the dissimilarity/distance between geographical distributions based on reporting rates recorded on a regular lattice. Reporting rate data are common, for example, in bird atlas projects where observers fill in check lists of encountered species in a particular area. Our index is a variation of the Euclidean distance, with the contribution of each grid cell weighted by the number of checklists collected for the grid cell, and a scaling factor to ensure that the dissimilarity ranges between zero and one. Reporting rates were transformed to ordered percentile classes to make species with different mean reporting rates comparable. The index was developed for the comparison of distributions of The Atlas of Southern African Birds . We illustrate the dissimilarity index comparing distributions of whydahs and indigobirds (widowfinches), which are specialized brood parasites, to the distributions of their hosts: waxbills and other finches.     

Break zones in the distributions of alleles and species in alpine plants

Aim We test for the congruence between allele‐based range boundaries (break zones) in silicicolous alpine plants and species‐based break zones in the silicicolous flora of the European Alps. We also ask whether such break zones coincide with areas of large elevational variation. Location The European Alps. Methods On a regular grid laid across the entire Alps, we determined areas of allele‐ and species‐based break zones using respective clustering algorithms, identifying discontinuities in cluster distributions (breaks), and quantifying integrated break densities (break zones). Discontinuities were identified based on the intra‐specific genetic variation of 12 species and on the floristic distribution data from 239 species, respectively. Coincidence between the two types of break zones was tested using Spearman’s correlation. Break zone densities were also regressed on topographical complexity to test for the effect of elevational variation. Results We found that two main break zones in the distribution of alleles and species were significantly correlated. Furthermore, we show that these break zones are in topographically complex regions, characterized by massive elevational ranges owing to high mountains and deep glacial valleys. We detected a third break zone in the distribution of species in the eastern Alps, which is not correlated with topographic complexity, and which is also not evident from allelic distribution patterns. Species with the potential for long‐distance dispersal tended to show larger distribution ranges than short‐distance dispersers. Main conclusions We suggest that the history of Pleistocene glaciations is the main driver of the congruence between allele‐based and species‐based distribution patterns, because occurrences of both species and alleles were subject to the same processes (such as extinction, migration and drift) that shaped the distributions of species and genetic lineages. Large elevational ranges have had a profound effect as a dispersal barrier for alleles during post‐glacial immigration. Because plant species, unlike alleles, cannot spread via pollen but only via seed, and thus disperse less effectively, we conclude that species break zones are maintained over longer time spans and reflect more ancient patterns than allele break zones.     

Forecasting the future suitable growth areas and constructing ecological corridors for the vulnerable species Ephedra sinica in China

Ephedra sinica is a rare and vulnerable species in China, and the habitat of Ephedra sinica is seriously threatened (by climate change and human activities). Predicting the suitable growth areas and constructing ecological corridors for Ephedra sinica in China will help to protect it scientifically. Based on 306 valid distribution records and 13 selected environmental factors, the maximum entropy (MaxEnt) model was used to simulate the potential current habitat zones and future (2050 and 2070) habitat zones of Ephedra sinica under four climate change scenarios. The minimum cumulative resistance (MCR) model was applied to extract important ecological corridors of Ephedra sinica. The results indicate that: (1) Under the current environment, the total area of the suitable habitat for Ephedra sinica in China is 42.24 × 10⁵ km², mainly distributed in Northwest China and North China. (2) Suitable area increases as the RCP rises. The center of mass of the habitat zone moved northward from Shaanxi Province to Ordos City in Inner Mongolia Autonomous Region. (3) Of the 13 environmental factors selected, the primary factor was elevation (20.8 %), followed by wettest month precipitation (18.2 %) and temperature seasonality (15.2 %). (4) Built 19 ecological corridors, with a total corridor length of 430.2 km, including seven long-distance passages and 12 short-distance corridors. All corridors are far from the artificial surface, mostly near high-altitude areas. The 19 ecological corridors constructed using the MCR model will also provide considerable importance for the survival of Ephedra sinica on a longer time scale in the future.     

Integration of plant and soil characteristics and the ecological success of two shape Prosopis species

The integration of mineral element composition was compared in two tree species of contrasting ecology. The native Prosopis cineraria is slow growing and non-invasive, while the introduced P. juliflora is fast growing, highly aggressive and invasive, and causes substratum degradation in the semi-arid and arid areas of north and north-west India. The two species were sampled from 18 sites in this region. Their leaf samples were analysed for leaflet weight, area and leaflet specific weight, and ash, Ca, Mg, Na, K, Zn, Cu and P concentrations. The soil samples were analysed for pH, electrical conductivity, exchangeable Ca, Mg, Na, K, Zn, PO$_4$, HCO$_3$, and organic carbon contents. Analysis of the data showed that while P. cineraria exhibited numerous correlations amongst plant and soil characteristics, it was not so in P. juliflora. This lack of integration amongst plant and soil characteristics and the ability to meet its nutrient requirements in all situations could be the basis of the phenomenal spread of P. juliflora across varying environmental conditions, in contrast to P. cineraria. This study illustrates the use of such information in the assessment of the ecological success of plant species, especially its evaluation in alien environments.     

Dynamics of core and occasional species in the marine plankton: tintinnid ciliates in the north-west Mediterranean Sea

Aim To assess short‐term variability in the community composition and community structure of tintinnid ciliates, herbivores of the microzooplankton. Location North‐west Mediterranean Sea. Methods We sampled on 18 dates over a 4‐week period in 2004 at an open‐water site. Species were classified as ‘core species’, found on every date, or ‘occasional species’, absent on one or more dates. Species abundance distributions of the entire community, and separately the core and occasional species, were compared with geometric, log‐series and log‐normal distributions. Core and occasional species were compared in terms of the shell or lorica oral diameter (LOD), analogous to gape size. Results We found 11 core and 49 occasional species. Diversity metrics were stable compared with shifts in abundances. Core species accounted for the majority of individuals in all samples. On each date, 9–22 occasional species, representing 10–15% of the population, were found. Species richness of the occasionals was positively related to population size. The identities of the occasional species found were unrelated to the time between sampling. The species abundance distribution of the occasional population was best fit by a log‐series distribution, while that of the core species was best fit by a log‐normal distribution. The species abundance distribution of the entire community was best fit by a log‐series distribution. Most of the occasional species had LODs distinct from that of a core species and occupied size classes left empty by the core population. However, the most abundant and frequent of the occasional species had a LOD similar to that of a core species. Main conclusions Among tintinnids, which are planktonic protists, occasional species have a species abundance distribution pattern distinct from that of core species. Occasional species appeared to be composed of two groups, one of relatively abundant species and similar to core species, and a second group of ephemeral species with morphologies distinct from core species. The existence of two categories of occasional or rare species may be common: (1) those similar to, and thus perhaps able to replace, dominant species in the absence of a change in the environment; and (2) those distinct from dominant species and requiring different conditions to prosper.     

Small-scale environmental heterogeneity and the analysis of species distributions along gradients

Abstract. The observed distribution of a species along an environmental gradient is strongly affected by environmental variability within a quadrat. Because a quadrat does not represent a point along an environmental gradient, but rather a range of conditions, it is likely to contain species not typically associated with the mean conditions in the quadrat. Systematic relationships exist between a species' true distribution, the observed distribution as a function of mean quadrat environment, and the frequency distribution of the environment within that quadrat. The observed species habitat breadth increases and the observed maximum abundance decreases as within-quadrat environmental heterogeneity increases. If species distributions or beta diversities are to be compared among species or coenoclines, they should be correctedforintra-quadratheterogeneity.Wederive simple corrections for environmental heterogeneity. The distributions of hardwood forest understory species along a soil acidity gradient in the North Carolina piedmont are presented as an example.     

An evaluation of methods for modelling species distributions

Aim Various statistical techniques have been used to model species probabilities of occurrence in response to environmental conditions. This paper provides a comprehensive assessment of methods and investigates whether errors in model predictions are associated to specific kinds of geographical and environmental distributions of species. Location Portugal, Western Europe. Methods Probabilities of occurrence for 44 species of amphibians and reptiles in Portugal were modelled using seven modelling techniques: Gower metric, Ecological Niche Factor Analysis, classification trees, neural networks, generalized linear models, generalized additive models and spatial interpolators. Generalized linear and additive models were constructed with and without a term accounting for spatial autocorrelation. Model performance was measured using two methods: sensitivity and Kappa index. Species were grouped according to their spatial (area of occupancy and extent of occurrence) and environmental (marginality and tolerance) distributions. Two‐way comparison tests were performed to detect significant interactions between models and species groups. Results Interaction between model and species groups was significant for both sensitivity and Kappa index. This indicates that model performance varied for species with different geographical and environmental distributions. Artificial neural networks performed generally better, immediately followed by generalized additive models including a covariate term for spatial autocorrelation. Non‐parametric methods were preferred to parametric approaches, especially when modelling distributions of species with a greater area of occupancy, a larger extent of occurrence, lower marginality and higher tolerance. Main conclusions This is a first attempt to relate performance of modelling techniques with species spatial and environmental distributions. Results indicate a strong relationship between model performance and the kinds of species distributions being modelled. Some methods performed generally better, but no method was superior in all circumstances. A suggestion is made that choice of the appropriate method should be contingent on the goals and kinds of distributions being modelled.     

Dispersal limitation and geographical distributions of mammal species

Aim To relate the dispersal limitation of endemic terrestrial mammals in Mexico to species life‐history traits and latitude. Location Mexico. Methods We modelled species ecological niches projected as potential distributions (P) using point occurrence data and 19 environmental variables for 89 endemic mammal species, and compared the areas covered by these ecological niche models with maps of species actual distributions (R) based on minimum convex polygons connecting marginal records based on museum specimens. We correlated body mass, food habits (herbivore, omnivore, insectivore, frugivore/granivore), volant and non‐volant (fossorial, arboreal, terrestrial) habits and mean latitude to the proportion of occupancy of species potential distributional areas (R/P). Results R and P were significantly positively correlated, with an overall average R/P ratio of 0.49. Less than half of the endemics (41 species) had a high occupancy (R/P values ranging from 0.50 to 0.90); a few (four species) showed full occupancy (> 0.90). Body mass and food habits were not correlated with R/P, but latitude showed significant correlations with R/P; volant mammals tended to show higher R/P values than non‐volant mammals. Main conclusions Few species filled most of the spatial extent of their ecological niches. Life‐history traits were generally poor predictors of proportional occupancy of species potential distributions. Endemics occurring at higher latitudes showed higher occupancy, suggesting that abiotic factors are likely to limit their distributions. Conversely, species at lower latitudes showed lower occupancy, suggesting that their distributions are limited by biotic factors and/or by geographical or historical barriers that prevent dispersal. The dispersal abilities of volant compared with non‐volant endemics can explain the higher occupancy in species potential distributions in the former group. These trends provide a baseline for exploring the importance of life‐history traits and abiotic versus biotic factors in limiting species distributions.     

Assessing transportation by the covariance of species with comments on contagious and random distributions

The paleoecologist must be able to distinguish a transported assemblage from an in situ one. A method is proposed to assess whether the post-mortem transport of individuals affected their observed spatial (horizontal) distribution. The spatial distribution of a species can be random or contagious. The spatial distribution of individuals of a species in the death assemblage produced by the cumulation of many temporally discrete inputs will be random if the individual inputs are random and contagious if the inputs are contagious. The spatial distribution patterns of several species should not covary in the absence of physical disturbance regardless of their own distributions, however. The degree of covariance between individuals of several species of similar hydrodynamic propensity is dependent on the amount and intensity of postmortem movement. The more species that covary, and the larger the size classes that covary, the more likely that transportation played an important role in the species' distribution patterns. Conversely, the absence of covariance suggests that, for at least some species, biological factors determined the species' spatial distributions. Similarly, covariance of vertical distribution patterns might suggest homogenization. by bioturbational or physical mixing.