The positive correlation between avian species richness and human population density in Britain is not attributable to sampling bias

Aim  To assess whether spatial variation in sampling effort drives positive correlations between human population density and species richness.
Location  British 10 × 10 km squares.
Methods  We calculated three measures of species richness from atlas data of breeding birds in Britain: total species richness, species richness standardised for sampling effort, and the number of species only recorded in supplementary casual records in a manner not standardised for survey effort. We then assessed the form of the relationship between these richness estimates and human population density, both with and without taking spatial autocorrelation into account.
Results  Both total and standardised species richness exhibit similar species richness–human population density relationships; species richness generally increases with human population density, but decreases at the very highest densities. Supplementary species richness is very weakly correlated with human population density.
Main conclusions  In this example, sampling effort only slightly influences the form of species richness–human population density relationships. The positive correlation between species richness and human population density and any resultant conservation conflicts are thus not artefactual patterns generated by confounding human density and sampling effort.     

The effect of sample size and species characteristics on performance of different species distribution modeling methods

Species distribution models should provide conservation practioners with estimates of the spatial distributions of species requiring attention. These species are often rare and have limited known occurrences, posing challenges for creating accurate species distribution models. We tested four modeling methods (Bioclim, Domain, GARP, and Maxent) across 18 species with different levels of ecological specialization using six different sample size treatments and three different evaluation measures. Our assessment revealed that Maxent was the most capable of the four modeling methods in producing useful results with sample sizes as small as 5, 10 and 25 occurrences. The other methods compensated reasonably well (Domain and GARP) to poorly (Bioclim) when presented with datasets of small sample sizes. We show that multiple evaluation measures are necessary to determine accuracy of models produced with presence-only data. Further, we found that accuracy of models is greater for species with small geographic ranges and limited environmental tolerance, ecological characteristics of many rare species. Our results indicate that reasonable models can be made for some rare species, a result that should encourage conservationists to add distribution modeling to their toolbox.     

Sampling the rarest: threatened beetles in boreal forest biodiversity inventories

Presence or absence of threatened species in samples is information that is widely used in designing and implementing conservation actions. We explored the effectiveness of beetle (Coleoptera) inventories and contribution of different sampling methods in revealing occurrences of threatened and near threatened species in boreal forests. The number of species caught using traps in a particular area proved to be a useful indicator of the representativeness of data, the relationship between total number of species and the number of threatened and near threatened species being almost exponential. Samples containing less than 200 trapped species (or 2000 individuals) are almost useless in surveying threatened and near threatened species. The probability of finding such species increases considerably when the number of trapped species exceeds 400. Window traps attached directly on the trunks of dead trees proved to be the most efficient sampling method in trapping threatened beetles, whereas many other standard methods gave relatively poor results. We suggest that the best alternative in surveying threatened species in boreal forests is a combination of intensive direct searching and trunk window traps. Finding threatened beetles with rigorous probability requires very large sample sizes, even if the most effective sampling methods are used. For example, ranking 10 boreal forest areas to be protected according to the occurrence of threatened species with some reliability may require trapping of over 100000 beetle individuals. Collecting and identifying these large samples routinely in conservation actions is not feasible, which means that shortcuts (indicators etc.) are necessary. However, a lot of good-quality inventories with appropriate sampling efforts are needed before these shortcuts can be identified and elaborated. Such inventories are also crucial for the improvement of the classification of threatened species and full assessment on how past forest management has eventually affected the biota.     

Numerical analyses of distributions of Iberian and Balearic endemic monocotyledons

Chorological information concerning 182 taxa of monocotyledons endemic to the Iberian Peninsula and Balearic Islands was compiled and related to the 100×100 km, 50×50 km and 10×10 km UTM grids. Distributions were analysed using multivariate methods (two-way indicator species analysis and detrended correspondence analysis) for each scale. Comparison of results allows recognition of several floristic elements and sectors (i.e. Balearic, Murcian-Almerian, south western) common to all three scales, whereas other regions are assigned to different sectors depending on the grid size considered. As a consequence of the increase in detail, characteristics such as number of sectors, the outline of boundaries and continuity or fragmentation of the areas also change. These factors are discussed.     

The influence of spatial resolution on macroecological patterns of range size variation: a case study using parrots (Aves: Psittaciformes) of the world

Aim To assess the extent to which the resolution at which geographical range sizes are measured influences macroecological patterns in this variable. Location Global. Methods Data on the geographical ranges of parrot species were digitized, and a Geographic Information System used to produce nine range size estimates for each species using different degrees of spatial resolution. The inter‐correlation of these estimates was then compared, together with their patterns of covariation with population size, body mass and migratory behaviour (across species and controlling for phylogeny), their pattern of phylogenetic correlation, and the frequency distributions of the different measures. Results Strong correlations exist among all nine range size measures across species, albeit that measures of similar spatial resolution are more strongly correlated. All measures show similar patterns of covariation with population size, body mass and migratory behaviour, and similar patterns of phylogenetic correlation. The skewness of frequency distributions increases towards zero as the resolution of the range size measure declines. Main conclusions The results of macroecological analyses are little affected by the resolution with which geographical range sizes are calculated, at least for the parrots of the world. Previously published studies based on crude measures of range size would be unlikely to have produced markedly different conclusions had they used more refined range size metrics.     

Strengthening the Natural and National Park system of Iberia to conserve vascular plants

The diversity of the Iberian vascular flora has been investigated using WORLDMAP versions 3.08 and 3.18. Two data sets scoring plant distributions as presences within the Iberian Peninsula were compiled; one for 2133 species at 50 × 50 km grid and the other for 801 species at 10 × 10 km map grids. Patterns of biodiversity were determined using the diversity measures of species richness, range-size rarity and character richness diversity. Using the diversity measures, combined with an area selection method, maps of priority areas were calculated using iterative procedures. Near minimum sets (NMSs) for both scales were calculated. Comparison of the NMS for the 10 × 10 km grid with the near minimum set for existing reserves (NMSER) showed that at least 2% more of the land surface would be required above and beyond the existing protected area network, currently comprising 6% of the area, to ensure representation of all species at least once as listed within the present data-base. It is demonstrated that reserve systems selected on a variety of different criteria are suboptimal when compared to particular groups of target organisms with a definite goal of representation for conservation. Calculating efficiency of existing reserve systems and accounting for all taxa identifies precisely the extra required areas for the protected area system to satisfy particular goals of representation.     

Using measurement error models to account for georeferencing error in species distribution models

Georeferencing error is prevalent in datasets used to model species distributions, inducing uncertainty in covariate values associated with species occurrences that result in biased probability of occurrence estimates. Traditionally, this error has been dealt with at the data‐level by using only records with an acceptable level of error (filtering) or by summarizing covariates at sampling units by using measures of central tendency (averaging). Here we compare those previous approaches to a novel implementation of a Bayesian logistic regression with measurement error (ME), a seldom used method in species distribution modeling. We show that the ME model outperforms data‐level approaches on 1) specialist species and 2) when either sample sizes are small, the georeferencing error is large or when all georeferenced occurrences have a fixed level of error. Thus, for certain types of species and datasets the ME model is an effective method to reduce biases in probability of occurrence estimates and account for the uncertainty generated by georeferencing error. Our approach may be expanded for its use with presence‐only data as well as to include other sources of uncertainty in species distribution models.     

Assessment of species diversity from species abundance distributions at different localities

We show how the spatial structure of species diversity can be analyzed using the correlation between the log abundances of the species in the communities, assuming that two communities at different localities can be described by a bivariate lognormal species abundance distribution. A useful property of this approach is that the log abundances of the species at two localities can be considered as samples from a bivariate normal distribution defined by only five parameters. The variances and the correlation can be estimated by maximum likelihood methods even if there is no information about the sampling intensity and the number of unobserved species. This method also enables estimation of over-dispersion in the sampling relative to a Poisson distribution that allows sampling adjustment of the estimate of β-diversity. Furthermore, we also obtain a partitioning of species diversity into additive components of α-, β- and γ-diversity. For instance, if the correlation between the log abundances of the species is close to one, the same species will be common and rare in the two communities and the β-diversity will be low. We illustrate this approach by analysing similarities of communities of rare and endangered species of oak-living beetles in south-eastern Norway. The number of recorded species was estimated to be only 48.1% of the total number of species actually present in these communities. The correlations among communities dropped rather quickly with distance with a scaling of order 200 km. This illustrates large spatial heterogeneity in species composition, which should be accounted for in the design of schemes of such devices for assessing species diversity in these habitat-types.     

Effects of sample size on estimates of population growth rates calculated with matrix models

Background

Matrix models are widely used to study the dynamics and demography of populations. An important but overlooked issue is how the number of individuals sampled influences estimates of the population growth rate (λ) calculated with matrix models. Even unbiased estimates of vital rates do not ensure unbiased estimates of λ–Jensen''s Inequality implies that even when the estimates of the vital rates are accurate, small sample sizes lead to biased estimates of λ due to increased sampling variance. We investigated if sampling variability and the distribution of sampling effort among size classes lead to biases in estimates of λ.

Methodology/Principal Findings

Using data from a long-term field study of plant demography, we simulated the effects of sampling variance by drawing vital rates and calculating λ for increasingly larger populations drawn from a total population of 3842 plants. We then compared these estimates of λ with those based on the entire population and calculated the resulting bias. Finally, we conducted a review of the literature to determine the sample sizes typically used when parameterizing matrix models used to study plant demography.

Conclusions/Significance

We found significant bias at small sample sizes when survival was low (survival = 0.5), and that sampling with a more-realistic inverse J-shaped population structure exacerbated this bias. However our simulations also demonstrate that these biases rapidly become negligible with increasing sample sizes or as survival increases. For many of the sample sizes used in demographic studies, matrix models are probably robust to the biases resulting from sampling variance of vital rates. However, this conclusion may depend on the structure of populations or the distribution of sampling effort in ways that are unexplored. We suggest more intensive sampling of populations when individual survival is low and greater sampling of stages with high elasticities.     

Territory tenure in song sparrows is related to song sharing with neighbours, but not to repertoire size

Song repertoires may be a product of sexual selection and several studies have reported correlations of repertoire size and reproductive success in male songbirds. This hypothesis and the reported correlations, however, are not sufficient to explain the observation that most species have small song repertoire sizes (usually fewer than 10, often fewer than five song types). We examined a second important aspect of a male's song repertoire, the extent to which he shares songs with his neighbours. Song sharing has not been measured in previous studies and it may be partially confounded with repertoire size. We hypothesized that in song sparrows, Melospiza melodia, song sharing rather than repertoire size per se is crucial for male territorial success. Our longitudinal study of 45 song sparrows followed from their first year on territory showed that the number of songs a bird shares with his neighbourhood group is a better predictor of lifetime territory tenure than is his repertoire size. We also found that song sharing increases with repertoire size up to but not beyond eight to nine song types, which are the most common repertoire sizes in the population (range in our sample 5-13). This partial confound of song sharing and repertoire size may account for some earlier findings of territory tenure-repertoire size correlations in this species and other species having small- or medium-sized repertoires. Copyright 2000 The Association for the Study of Animal Behaviour.     

Spatial overlap enhances geographic range size conservatism

Phylogenetic conservatism or heritability of the geographic range sizes of species has been predicted to occur because of the phylogenetic conservatism of niche traits. However, evidence for range size conservatism is mixed, and even when statistically significant is often rather weak and of questionable biological importance. Here, we test the prediction that such conservatism will be more strongly expressed when the amount of spatial overlap between sister species increases. We used the global distributions of 1136 avian species (>10% of extant members of this Class), and tested the conservatism of geographic range sizes using the coefficients of correlation between values for pairs of sister species. We used a null model to test whether the range sizes of sister species were more similar to one another than expected by chance. We found that sister species showed a significant positive relationship between their geographic range sizes whatever the degree of spatial overlap. However, as predicted, the level of conservatism increases with the level of range overlap between sister species. More precisely, the strong increase in the coefficient of correlation between sister species' range sizes when we add species with some range overlap to the pool of pairs without any such overlap indicates an important threshold effect. These results suggest that niche conservatism is more likely to lead to marked heritability of the range sizes of species when similar niche traits are expressed under more similar environmental conditions. These results have significant implications because they suggest 1) that previous analyses of conservatism of range sizes have been confounded by the level of spatial overlap, and 2) that closely related species experiencing similar conditions may tend to expand or restrict their geographic ranges in parallel when faced with climate change.     

Sensitivity of predictive species distribution models to change in grain size

Predictive species distribution modelling (SDM) has become an essential tool in biodiversity conservation and management. The choice of grain size (resolution) of environmental layers used in modelling is one important factor that may affect predictions. We applied 10 distinct modelling techniques to presence-only data for 50 species in five different regions, to test whether: (1) a 10-fold coarsening of resolution affects predictive performance of SDMs, and (2) any observed effects are dependent on the type of region, modelling technique, or species considered. Results show that a 10 times change in grain size does not severely affect predictions from species distribution models. The overall trend is towards degradation of model performance, but improvement can also be observed. Changing grain size does not equally affect models across regions, techniques, and species types. The strongest effect is on regions and species types, with tree species in the data sets (regions) with highest locational accuracy being most affected. Changing grain size had little influence on the ranking of techniques: boosted regression trees remain best at both resolutions. The number of occurrences used for model training had an important effect, with larger sample sizes resulting in better models, which tended to be more sensitive to grain. Effect of grain change was only noticeable for models reaching sufficient performance and/or with initial data that have an intrinsic error smaller than the coarser grain size.     

Sample size for collecting seeds in germplasm conservation: the case of the Lima bean (Phaseolus lunatus L.)

 The design of optimum sampling strategies integrating criteria of efficiency relevant to multilocus models and many target populations has been investigated with respect to the number of plants and the number of seeds per plant to be sampled for a Lima bean (Phaseolus lunatus L.) gene pool. This study, using five populations and six polymorphic enzyme loci, shows that the number of plants rather than the number of seeds collected per plant primarily determines the success of seed sampling, suggesting that plant number plays an essential part in maintaining the allelic multiplicity of predominantly selfing species like Lima bean. According to the results, it appears that among Lima bean populations an efficient sampling procedure is achieved by collecting 1–4 seeds from 200 to 300 plants. These sample sizes will retain 8–10 alleles, regardless of their frequencies. When we consider polymorphism at the 5% level, it is expected that sampling 10–80 plants will collect combinations of 4–8 alleles. Based on data from genetic and demographic studies, we suggest an efficient sampling scheme for Lima bean germplasm at both population and geographical levels. Received: 10 March 1998 / Accepted: 1 April 1998     

插网网目尺寸及采样时间对潮沟游泳动物采样的影响

在长江河口九段沙盐沼湿地的一条潮沟中,我们比较了两种不同网目尺寸（4mm和8mm）的插网（fyke net）及日夜潮对游泳动物群落采样效率的影响。共捕获鱼类16种5476尾、虾2种4982只。采用方差分析、柯尔莫诺夫-斯米尔诺夫非参数检验法和无度量多维标定分析对两种网目插网的捕获效率,样品的体长分布,群落结构及日夜潮间的差异进行了分析。结果表明,网目尺寸对样品中游泳动物的物种数和鱼类的多度没有影响,但是小网目的插网对虾类有较高的捕获率。两种网目尺寸的插网均在夜潮中捕获较多的鱼类个体,但虾类在日潮捕获较多。在日潮中,网目尺寸对捕获物中五种优势游泳动物的体长分布无显著影响。在夜间,小网目的插网对斑尾复缎虎和脊尾白虾的较大个体具有较高的捕获率,但对较大个体鲅的捕获率不如大网目的插网。样品中游泳动物的群落组成在日夜潮间存在明显差异,大网目捕获物的样本间差异比小网目小。     

The effects of gerbil foraging on the natural seedbank and consequences on the annual plant community

We explicitly tested the following predictions using Gerbillus allenbyi and G. pyramidum foraging on artificial arrays of natural seedbank on a semi-stabilized sand dune: 1. that the gerbils will forage using a quitting harvest rate rule, 2. that larger seeds are preferred due to higher encounter rates, and 3. that there are community level consequences for the annual plants as a direct result of foraging by the desert rodents. Natural seedbank, separated into two size classes, was placed in seed trays in the field at four different densities (1/16×, 1/4×, 1×, and 2×normal). Following exposure to granivory, the remaining seeds were weighed and germinated to test for community level effects. Only the 1× and 2×normal density plots were heavily foraged, and at both seed sizes, which suggests that the gerbils employed a quitting harvest rate rule. In support of our second prediction, the two species of gerbils tended to consume more of the larger seeds, particularly at higher densities. The mean number and total number of plant species that germinated in plots exposed to granivores was not significantly different from unexposed samples. At the community level however, there was no net association of germinated plant species in the four treatment groups exposed to granivores, but a significant net positive association in the unexposed control seedbank. Gerbil foraging on annual plant seedbank may thus subtly change the entire community structure (from positive to neutral), although not necessarily shift the species distributions significantly. Our results corroborate other studies involving artificial food types such as millet and suggest that foraging decisions may affect the plant community.     

Worker size and seed size selection by harvester ants in a neotropical forest

The rules ants follow when selecting seeds are important both to theories of colony organization and to the shaping of their impact as harvesters. Two Costa Rican forests yielded the most diverse harvester ant assemblages yet studied (41 and 44±4 species). I assayed seed size preferences within and between species using milled barley. Seed size selection varied at a number of levels. First, species differed in seed size use: small species mainly carried off small seeds; large species retrieved a broad range of seed sizes. Within species, one-third of those tested yielded inter-colony differences in preferred seed sizes. Finally, workers of polymorphic species tested showed significant worker/seed size correlations. Species, colony, and worker level differences are common among harvester ants. Their significance to community organization and colony ergonomics however is hotly debated and requires a better understanding of the dynamics of food limitation. From the plant's perspective, small seeds will be harvested by a wider variety of ants than large seeds.     

Effects of patch size on colonisation in estuaries: revisiting the species-area relationship

The effects of patch size on the colonisation and succession of intertidal invertebrates and algae were investigated in an estuary near Sydney, New South Wales, Australia. The specific aim was to test explicitly for the presence of a species-area relationship, and examine whether this could be explained by the random placement hypothesis (i.e. that the number of species per unit area was the same on patches of different sizes). In addition, I tested the extent to which differences in numbers of species reflected differences in the composition of assemblages. Wooden panels of three different sizes (10 × 10 cm, 20 × 20 cm and 40 × 40 cm) were placed in the field on intertidal oyster leases in each of two different experimental trials: spring (October 1994) and summer (January 1995). Independent replicate measures of the number of colonising species on panels were obtained after different periods of time, up to 25 months. I also obtained measures of abundance of individual species and composition of assemblages on panels of different sizes. This allowed specific tests of the hypothesis that the size of the patch being colonised is important in structuring these assemblages. The strength of the species-area relationship increased through time on panels submersed in October, but the trend was reversed for panels submersed in January. There was a significant interaction between factors of patch size and time of submersion for multivariate measures of differences in composition among replicates. The random placement hypothesis was supported in certain situations, but not in others. When rejected, it was for different reasons on panels submersed in the two different trials. Panels initiated in October tended to have proportionally greater numbers of species per unit area on larger panels, while the panels initiated in January tended to have more species per unit area on smaller panels. There was an identifiable relationship between differences in numbers of species and differences in species composition for panels submersed in October. This was not true, however, for panels submersed in January, where the species-area relationship did not hold after longer periods. The succession of organisms through time was, overall, more important in structuring the assemblages than was the size of the patch being colonised. The species-area relationship should not necessarily be regarded as a truism – it did not always hold in this system. The initial timing of experiments with respect to recruitment and succession influenced the results. Received: 16 March 1998 / Accepted: 5 October 1998     

以林相图为基础的不同取样方式对植被排序分析结果的影响

以6种不同方式对林相图中同一样带取样,采用CA、DCA和CCA 3种排序方法,研究了取样方式对排序轴的解释效果、物种与环境因子、环境因子之间、环境因子与坐标轴之间关系的影响.结果表明,样方大小和形状的变化在不同程度上改变了排序结果.大样方和长方形样方都增强了排序轴的解释效果,并对双序图中稀有种、独特种的位置有较大的影响;环境因子中土壤因子对样方的大小和形状都很敏感,坡度、经纬度只对样方大小敏感,坡位、海拔、温度和降水则对样方形状敏感;随着样方面积的增加,海拔、温度和降水的作用降低,而坡向的作用增加.     

How far do songbirds disperse?

Dispersal distances determine the scales over which many population processes occur. Knowledge of these distances may therefore be crucial in determining the appropriate spatial scales for research and management. However, dispersal distances are difficult to measure, especially for vagile organisms like songbirds. For these species, the use of traditional mark–recapture and radio‐telemetry methods is problematic. We used positive one‐year time‐lagged correlations in abundance to estimate natal dispersal distances. Using the North American Breeding Bird Survey database, we examined one‐year time‐lagged correlations between pairs of North American songbird samples separated by 10–100 km. We submit that consistent positive one‐year time‐lagged correlations reflect the exchange of individuals through dispersal. We found positive one‐year time‐lagged correlations between pairs of samples from 25 different songbird species. The median distances of these correlations ranged from 15 to 95 km, depending on the species. These distances were positively correlated with body size and wing length. Dispersal appears to be the most parsimonious explanation for the time‐lagged correlations we observed in these species. The putative dispersal distances we measured are generally an order of magnitude longer than those reported in the literature.     

Patterns of co‐occurrences in a killifish metacommunity are more related with body size than with species identity

Body size may be more important than species identity in determining species interactions and community structure. However, co‐occurrence of organisms has commonly been analysed from a taxonomic perspective and the body size is rarely taken into account. On six sampling occasions, we analysed patterns of killifish co‐occurrences in nestedness (tendency for less rich communities to be subsamples of the richest), checkerboard structure (tendency for species segregation), and modularity (tendency for groups to co‐occur more frequently than random expectation) in a pond metacommunity located in Uruguay. We contrasted co‐occurrence patterns among species and body size‐classes (individuals from different species were combined into size categories). The analysis was performed at two spatial scales: ponds (communities) and sample units within ponds. Observed nestedness was frequently smaller than the null expectation, with significantly greater deviations for body size‐classes than for species, and for sample units than for communities. At the sample unit level, individuals tended to segregate (i.e. clump into a checkerboard pattern) to a larger extent by body size rather than by taxonomy. Modularity was rarely detected, but nevertheless indicated a level of taxonomic organization not evident in nestedness or checkerboard indices. Identification of the spatial scale and organization at which ecological forces determine community structure is a basic requirement for advancement of robust theory. In our study system, these ecological forces probably structured the community by body sizes of interacting organisms rather than by species identities.