Abundance, spatial variance and occupancy: arthropod species distribution in the Azores

1. The positive abundance-occupancy and abundance-variance relationships are two of the most widely documented patterns in population and community ecology. 2. Recently, a general model has been proposed linking the mean abundance, the spatial variance in abundance, and the occupancy of species. A striking feature of this model is that it consists explicitly of the three variables abundance, variance and occupancy, and no extra parameters are involved. However, little is known about how well the model performs. 3. Here, we show that the abundance-variance-occupancy model fits extremely well to data on the abundance, variance and occupancy of a large number of arthropod species in natural forest patches in the Azores, at three spatial extents, and distinguishing between species of different colonization status. Indeed, virtually all variation about the bivariate abundance-occupancy and abundance-variance relationships is effectively explained by the third missing variable (variance in abundance in the case of the abundance-occupancy relationship, and occupancy in the case of the abundance-variance relationship). 4. Introduced species tend to exhibit lower densities, less spatial variance in these densities, and occupy fewer sites than native and endemic species. None the less, they all lie on the same bivariate abundance-occupancy and abundance-variance, and trivariate abundance-variance-occupancy, relationships. 5. Density, spatial variance in density, and occupancy appear to be all the things one needs to know to describe much of the spatial distribution of species.     

Temporally stable abundance–occupancy relationships and occupancy frequency patterns in stream insects

The interspecific relationship between local abundance and regional distribution, as well as the occupancy frequency distribution, are widely studied topics in macroecology. A positive abundance-occupancy relationship has been found in a majority of studies, and satellite species modes are typically dominant in occupancy frequency distributions. However, there are a number of exceptions to these "general" findings, and only a few studies have examined these patterns and their temporal variability in stream organisms. I examined both abundance-occupancy relationships and occupancy frequency distributions in stream insects in a boreal drainage system over six consecutive years. I found that the positive interspecific abundance-occupancy relationship was highly stable temporally, with coefficients of determination ranging from 0.25 to 0.47 over the years. There were no strong differences in the strength and slope of the abundance-occupancy relationship between non-predatory and predatory insect species in each year. Temporally stable abundance-occupancy relationships were paralleled by among-year patterns in both abundance and occupancy, with locally abundant and widely distributed species remaining locally abundant and widely distributed over the years, while locally uncommon and regionally rare species showed the opposite. Occupancy frequency distributions were strongly right-skewed, mirroring the dominance of the left-most satellite mode of regionally rare species. That the abundance-occupancy relationship, species' abundances and distributions, as well as the dominance of satellite species in occupancy frequency distribution were temporally stable suggest that niche-based models are strong candidates for explaining these patterns in stream insects. By contrast, metapopulation-based models that predict clear temporal variability in species' abundance and occupancy, as well as bimodal occupancy frequency distributions, are less plausible candidates for explaining the observed patterns. The present findings are the opposite to those in some terrestrial studies, but they are in agreement with other terrestrial studies and with a few previous studies on stream organisms.     

The consistency and stability of abundance-occupancy relationships in large-scale population dynamics

1. Abundance-occupancy relationships comprise some of the most general and well-explored patterns in macro-ecology. The theory governing these relationships predicts that species will exhibit a positive interspecific and intraspecific relationship between regional occupancy and local abundance. Abundance-occupancy relationships have important implications in using distributional surveys, such as atlases, to understand and document large-scale population dynamics and the consequences of environmental change. A basic need for interpreting such data bases is a better understanding of whether changes in regional occupancy reflect changes in local abundance across species of varying life-history characteristics. 2. Our objective was to test the predictions of the abundance-occupancy rule using two independent data sets, the New York State Breeding Bird Atlas and the North American Breeding Bird Survey. The New York State Breeding Bird Atlas consists of 5332 25-km(2) survey blocks and is one of the first atlases in the USA to be completed for two time periods (1980-85 and 2000-05). The North American Breeding Survey is a large-scale annual survey intended to document the relative abundance and population change of songbirds throughout the USA. 3. We found that regional occupancy was positively correlated with relative abundance across 98 (beta = 0.60 +/- 0.11 SE, P < 0.001, R(2) = 0.60) and 85 species (beta = 0.67 +/- 0.06 SE, P < 0.001, R(2) = 0.57) in two separate time periods. This relationship proved stable over time and was notably consistent between breeding habitat groups and migratory guilds. 4. Between 1980 and 2005, changes in regional occupancy were highly correlated with long-term abundance trend estimates for 75 species (beta = 5.73 +/- 0.24 SE, P < 0.001, R(2) = 0.88). Over a 20-year period, woodland and resident birds showed an increase in occupancy while grassland species showed the greatest decline; these patterns were mirrored by changes in local abundance. 5. Although exceptions existed, we found most changes in occupancy parallel changes in local abundance. These findings support the basic predictions of the abundance-occupancy rule and demonstrate its consistency and stability in species and groups of varying life-history characteristics.     

Abundance-occupancy dynamics in a human dominated environment: linking interspecific and intraspecific trends in British farmland and woodland birds

1. Range size, population size and body size, the key macroecological variables, vary temporally both within and across species in response to anthropogenic and natural environmental change. However, resulting temporal trends in the relationships between these variables (i.e. macroecological patterns) have received little attention. 2. Positive relationships between the local abundance and regional occupancy of species (abundance-occupancy relationships) are among the most pervasive of all macroecological patterns. In the absence of formal predictions of how abundance-occupancy relationships may vary temporally, we outline several scenarios of how changes in abundance within species might affect interspecific patterns. 3. We use data on the distribution and abundance of 73 farmland and 55 woodland bird species in Britain over a 32-year period encompassing substantial habitat modification to assess the likelihood of these scenarios. 4. In both farmland and woodland habitats, the interspecific abundance-occupancy relationship changed markedly over the period 1968-99, with a significant decline in the strength of the relationship. 5. Consideration of intraspecific dynamics shows that this has been due to a decoupling of abundance and occupancy particularly in rare and declining species. Insights into the intraspecific processes responsible for the interspecific trend are obtained by analysis of temporal trends in the distribution of individuals between sites, which show patterns consistent with habitat quality declines. 6. This study shows that a profitable approach to ascertaining the nature of human impacts is to link intra- and interspecific processes. In the case of British farmland and woodland birds, changes to the environment lead to species-specific responses in large-scale distributions. These species-specific changes are the driver of the observed changes in the form and strength of the interspecific relationship.     

Abundance-variance and abundance-occupancy relationships in a marine host-parasite system: the importance of taxonomy and ecology of transmission

Abundance-occupancy and abundance-variance relationships are two of the most general macroecological patterns capturing essential fundamentals of the structuring of species distributions and are widely documented for free-living animal and plant species populations at different spatial scales. However, empirical data for parasites have been gathered using appropriate sampling designs only recently. We performed analyses across species of the variation in infection parameters and patterns of aggregation of the most widespread parasites in the marine sparid fish Boops boops across seven localities of two marine biogeographical regions, the North East Atlantic and the Mediterranean. We used a large dataset of multiple population samples replicated over time for 20 parasite species and carried out assessments both intraspecifically and interspecifically, across taxonomic and ecological groupings. This taxonomically diverse complex of species representing five major metazoan higher taxa with differing transmission ecologies allowed us to assess the effect of taxonomic and ecological determinants on the abundance-occupancy and abundance-variance relationships in the model marine host-parasite system. The results revealed that: (i) a power function, relating spatial variance to mean abundance, represents a suitable model for the spatial distribution of the species; (ii) prevalence, abundance and the degree of spatial heterogeneity are true species characteristics and differ consistently between higher level taxonomic groupings; (iii) infection parameters and abundance-variance relationship are dependent on host specificity and regional distribution patterns of the parasites; and (iv) the observed infection parameters agree well with predictions from the epidemiological negative binomial abundance-occupancy model built on parameters of Taylor's power law both within and across species.     

Do developmental mode and dispersal shape abundance-occupancy relationships in marine macroinvertebrates?

1. Dispersal is a crucial process in maintaining population structures in many organisms, and is hypothesized as a process underlying the interspecific relationship between abundance and distribution. Here we examined whether there was a link between the dispersal and developmental modes of marine macroinvertebrates and the slopes and elevations of interspecific abundance-occupancy relationships. We predicted that if within-site retention of larvae ranks in the order brooders > lecithotrophs > planktotrophs, for any given level of mean abundance, occupancy should increase in the order brooders < lecithotrophs < planktotrophs. We also predicted that propensity to form metapopulations should be greater for planktonic dispersers (i.e. lecithotrophs and planktotrophs combined) than for non-planktonic (i.e. brooders), resulting in steeper abundance-occupancy relationships for the former. 2. Predictions were tested using a data set for 362 subtidal marine macroinvertebrates occurring across 446 1-km(2) grid squares around the British Isles; analyses were performed on the data set as a whole and for separate phyla. 3. The total data set had a Z-transformed effect size of 0.79, within the confidence intervals described by Blackburn et al. (2006; Journal of Animal Ecology, 75, 1426-1439), and was consistently present with relatively homogeneous effect size in separate analyses of polychaetes, crustaceans, molluscs and echinoderms. 4. In all cases, planktonic dispersing organisms showed an abundance-occupancy relationship with greater elevation than that for non-planktonic organisms; in polychaetes the elevation of slopes was in the rank order planktotrophs > lecithotrophs > brooders. No differences between the slopes of the abundance-occupancy relationship were apparent for different dispersal modes either within, or across phyla. 5. We conclude that dispersal capacity may play an important part in determining the elevation of the abundance-occupancy relationship, the corollary of low dispersal in the marine realm being greater local retention of larvae and greater local population abundance at low extents of geographical distribution.     

Interspecific abundance-occupancy relationships and the effects of disturbance: a test using microcosms

The effects of disturbance on interspecific relationships between abundance and occupancy are tested using the results of a factorial experiment carried out with microcosm communities of protists. A positive relationship was documented whether marked disturbance was present or not; the pattern was neither a product of disturbance, nor was it destroyed by disturbance. Coefficients of determination, and the slopes and intercepts of abundance-occupancy relationships did not appear to change systematically with treatment. This robustness of the relationship may reflect the tendency for species to maintain approximately the same relative abundances and levels of occupancy in a broad range of circumstances. Received: 4 October 1996 / Accepted: 29 April 1997     

Range-size,local abundance and germination niche-breadth in Mediterranean plants of two life-forms

The relationships between rarity (i.e., range-size, local abundance) and niche-breadth can be important to assess the risks the species face under global change, namely those resulting from fire regime change. In fire areas, germination is critical for establishing after fire for many species. We examined the relationships between rarity and germination niche-breadth for 53 plant species of two life-forms (chamaephytes and hemicryptophytes) growing in Eastern Spain. Rarity was measured as geographic range-size and as local abundance. Local abundance was evaluated at two sites differing in their post-fire successional status (a recently burned area and a long-unburned one). Germination niche-breadth was measured as the mean germination evenness index from four germination experiments that subjected the seeds to various germinating conditions characteristics of fire environments. Correlations between rarity (range-size, local abundance) and niche-breadth were calculated in cross-species mode or by way of phylogenetically independent contrasts, and either for the 53 species (all-species set) or for each of two life-form groups (chamaephytes, hemicryptophytes). In general, no significant correlations were found between the rarity measures and germination niche-breadth for the all-species set. However, significant correlations emerged when the analyses were done separating species by life-forms. Germination niche-breadth was positively correlated with range-size for chamaephytes, and negatively for hemicryptophytes. In addition, germination niche-breadth was uncorrelated, or negatively so, with local abundance for chamaephytes and hemicryptophytes, respectively. While no correlation between range-size and local abundance was found for chamaephytes, a negative one was obtained for hemicryptophytes. We conclude that rarity/germination-niche relationships varied by life-form. This pattern of relationships was obscured when all species were joined in a single group. Based on the contrasting patterns of correlations obtained for each life-form we argue that the changes in the germinating environment caused by alterations in fire regime are likely to differentially affect these two groups of species.     

Abundance–occupancy patterns in a riverine fish assemblage

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Distributions of habitat suitability and the abundance-occupancy relationship

Positive abundance-occupancy relationships (a relationship between the number of sites a species occupies and the average density of individuals in occupied sites) are widespread through a range of taxa. The simplest model for this is the "vital rates" model, which proposes that habitat suitability varies spatially; increasing average habitat quality thus leads to simultaneous increases in average densities within occupied areas, as well as the total area that is habitable. This model has not been tested. We develop a general analytical version of this model and show that it predicts that the skewness of population size or aggregation of individuals within sites should vary systematically with density and occupancy, depending on the distribution of habitat suitability, and that the variance in occupancy should be highest at low densities. We compare these predictions with data from the British Trust for Ornithology's Common Birds Census, and we find systematic changes in both variance and skewness of density, both intra- and interspecifically.     

Positive interspecific relationship between temporal occurrence and abundance in insects

One of the most studied macroecological patterns is the interspecific abundance-occupancy relationship, which relates species distribution and abundance across space. Interspecific relationships between temporal distribution and abundance, however, remain largely unexplored. Using data for a natural assemblage of tabanid flies measured daily during spring and summer in Nova Scotia, we found that temporal occurrence (proportion of sampling dates in which a species occurred in an experimental trap) was positively related to temporal mean abundance (number of individuals collected for a species during the study period divided by the total number of sampling dates). Moreover, two models that often describe spatial abundance-occupancy relationships well, the He-Gaston and negative binomial models, explained a high amount of the variation in our temporal data. As for the spatial abundance-occupancy relationship, the (temporal) aggregation parameter, k, emerged as an important component of the hereby named interspecific temporal abundance-occurrence relationship. This may be another case in which a macroecological pattern shows similarities across space and time, and it deserves further research because it may improve our ability to forecast colonization dynamics and biological impacts.     

Invasive bivalves increase benthic communities complexity in neotropical reservoirs

Invasive bivalves often act as ecosystem engineers, generally causing physical alterations in the ecosystems in which they establish themselves. However, the effects of these physical alterations over benthic macroinvertebrate communities’ structure are less clear. The objective of this study was to characterize the ecological effects of the invasive bivalves Corbicula fluminea and Limnoperna fortunei on the structure of benthic macroinvertebrate communities in neo-tropical reservoirs. Three hypotheses were tested: (1) invasive bivalves act as facilitator species to other benthic macroinvertebrates, resulting in communities with higher number of species, abundance and diversity; (2) invasive bivalves change the taxonomic composition of benthic macroinvertebrate communities; (3) invasive bivalves increase the complexity of benthic macroinvertebrate communities. For that it was used data from 160 sampling sites from four reservoirs. We sampled sites once in each area, during the dry season from 2009 to 2012. The first hypothesis was rejected, as the presence of invasive bivalves significantly decreased the host benthic communities’ number of species and abundance. The second hypothesis was corroborated, as the composition of other benthic macroinvertebrates was shown to be significantly different between sites with and without invasive bivalves. We observed a shift from communities dominated by common soft substrate taxa, such as Chironomidae and Oligochaeta, to communities dominated by the invasive Gastropoda Melanoides tuberculata. The biomass data corroborated that, showing significantly higher biomass of M. tuberculata in sites with invasive bivalves, but significantly lower biomass of native species. Benthic macroinvertebrate communities presenting invasive bivalves showed significantly higher eco-exergy and specific eco-exergy, which corroborate the third hypothesis. These results suggest that while the presence of invasive bivalves limits the abundance of soft bottom taxa such as Chironomidae and Oligochaeta, it enhances benthic communities’ complexity and provide new energetic pathways to benthic communities in reservoirs. This study also suggests a scenario of invasion meltdown, as M. tuberculata was facilitated by the invasive bivalves.     

Macroecology of local insect communities

The inter-relationships between animal body weight, range size, species richness and abundance are currently the basis of macroecology. Using 41 099 insects sampled from 31 Acacia tree canopies in north-east Tanzania, we first documented the basic macroecological patterns. The relationship between body weight and both species richness and abundance was polygonal with the highest insect species richness and abundance occurring at intermediate body weights. Across individual tree communities, the most statistically robust relationships were found between range size, abundance and species richness and they were all linear. In a second part, we focused on the positive abundance-range size relationship and we could test predictions of six of the eight proposed hypotheses to explain this widely documented pattern of community structure. The relationship is most likely explained by the metapopulation hypothesis stating that with more patches being occupied, local abundance in a given patch increases due to a higher rate of immigration from nearby patches. In addition, we found high slopes for the species-area relationship, typical of island systems and thus it seems reasonable to characterise Acacia trees in the savannah as habitat islands for insects.     

Distribution and abundance of parasite nematodes: ecological specialisation,phylogenetic constraint or simply epidemiology?

We investigate the patterns of abundance‐spatial occupancy relationships of adult parasite nematodes in mammal host populations (828 populations of nematodes from 66 different species of terrestrial mammals). A positive relationship between mean parasite abundance and host occupancy, i.e. prevalence, is found which suggests that local abundance is linked to spatial distribution across species. Moreover, the frequency distribution of the parasite prevalence is bimodal, which is consistent with a core‐satellite species distribution. In addition, a strong positive relationship between the abundance (log‐transformed) and its variance (log‐transformed) is observed, the distribution of worm abundance being lognormally distributed when abundance values have been corrected for host body size.
Hanski et al. proposed three distinct hypotheses, which might account for the positive relationship between abundance and prevalence in free and associated organisms: 1) ecological specialisation, 2) sampling artefact, and 3) metapopulation dynamics. In addition, Gaston and co‐workers listed five additional hypotheses. Four solutions were not applicable to our parasitological data due to the lack of relevant information in most host‐parasite studies. The fifth hypothesis, i.e. the confounded effects exerted by common history on observed patterns of parasite distributions, was considered using a phylogeny‐based comparison method. Testing the four possible hypotheses, we obtained the following results: 1) the variation of parasite distribution across host species is not due to phylogenetic confounding effects; 2) the positive relationship between mean abundance and prevalence of nematodes may not result from an ecological specialisation, i.e. host specificity, of these parasites; 3) both a positive abundance‐prevalence relationship and a negative coefficient of variation of abundance‐prevalence relationship are likely to occur which corroborates the sampling model developed by Hanski et al. We argue that demographic explanations may be of particular importance to explain the patterns of bimodality of prevalence when testing Monte‐Carlo simulations using epidemiological modelling frameworks, and when considering empirical findings. We conclude that both the bimodal distribution of parasite prevalence and the mean‐variance power function simply result from demographic and stochastic patterns (highlighted by the sampling model), which present compelling evidence that nematode parasite species might adjust their spatial distribution and burden in mammal hosts for simple epidemiological reasons.     

Interspecific abundance-range size relationships: range position and phylogeny

A number of mechanisms have been proposed to explain the widely observed positive interspecific relationship between local abundance and extent of geographic distribution in animals Here, we use data on British birds to assess two of these hypotheses that the relationship results from the relative position of a study area with respect to the geographic ranges of the species which occur there, and that the relationship results from a simple difference between taxonomic groups, rather than any general tendency for more abundant species to have larger range sizes We find support for neither hypothesis Phylogenetically controlled comparative analyses reveal that the positive abundance-range size relationship is consistently found within taxa, even when abundance and range size are calculated at a variety of spatial and temporal scales Analyses both across species and within taxa show that bird species for which Britain is near to the centre of their distribution in Europe tend to have larger British range sizes and higher abundances than do species where Britain is close to the edge of their range in Europe However, these relationships do not cause that between abundance and range size, because this latter relationship persists within different range position categories Whether a species is near the centre or edge of its geographic range in Britain may affect its position on the abundance-range size relationship, but does not produce the relationship Range position in Britain does, however, seem to be related to the magnitude of temporal changes in the range sizes of British birds There is some evidence to suggest that species for which Britain is nearer to their European range centre have shown smaller changes in distribution over the period 1970–1990 than have species for which Britain is close to their European range edge     

Species–area relationship within benthic habitat patches of a tropical floodplain river: An experimental test

The curvilinear relationship between species richness and habitat area (species–area relationship (SAR)) is a fundamental ecological pattern. The relationship is often viewed from a long‐term perspective across relatively large spatial scales, reflecting a balance between immigration and extinction dynamics. We explored whether predictions of SAR also manifest over short time periods (days) in benthic habitat patches of a dynamic floodplain river where littoral faunal assemblages are continuously assembled and disassembled with changing water levels. We examined the relationship of patch size with faunal abundance (i.e. fish and aquatic invertebrates), taxonomic richness, trophic group richness and overall assemblage composition. Strong taxa–area relationships emerged despite the relatively short experimental time period (21 days); larger patches had more taxa and trophic groups. For the smallest patches, taxonomic richness was especially sensitive to abundance of individuals; abundance of individuals was a less important predictor of taxonomic and trophic group richness for the largest patches. Despite the relatively short time frame for study within this temporally dynamic ecosystem, our findings indicate a strong SAR for fishes and macroinvertebrates inhabiting patchy habitats in the littoral zone of this tropical river.     

黄河三角洲植物生态位和生态幅对物种分布-多度关系的解释

物种分布与多度间的正相关格局非常普遍,但该格局的生态机制却一直不太明确。研究者提出了很多假说来解释这种分布-多度关系,其中物种的生态幅和生态位（资源可利用性）机制的研究较多。为了验证物种的生态幅和生态位是否能解释物种的多度-分布格局,本文研究了黄河三角洲地区湿地植物分布、多度、生态位和生态幅间的关系,结果表明：该区物种分布与多度呈显著正相关,且均与生态幅显著正相关,物种分布与生态位显著负相关,但物种多度与生态位相关性不显著。这说明物种的分布越广,其多度越高,环境容忍度越大;而可利用资源更多的物种分布更广,环境容忍度越大的物种多度越多,资源可利用性对该区物种多度影响不大。本研究说明物种生态幅能解释物种分布-多度正相关格局,而生态位假说不能很好的解释这一格局;应该还有其他因素一起解释这一物种分布-多度正相关格局。     

Predator diversity and abundance provide little support for the enemies hypothesis in forests of high tree diversity

Predatory arthropods can exert strong top-down control on ecosystem functions. However, despite extensive theory and experimental manipulations of predator diversity, our knowledge about relationships between plant and predator diversity--and thus information on the relevance of experimental findings--for species-rich, natural ecosystems is limited. We studied activity abundance and species richness of epigeic spiders in a highly diverse forest ecosystem in subtropical China across 27 forest stands which formed a gradient in tree diversity of 25-69 species per plot. The enemies hypothesis predicts higher predator abundance and diversity, and concomitantly more effective top-down control of food webs, with increasing plant diversity. However, in our study, activity abundance and observed species richness of spiders decreased with increasing tree species richness. There was only a weak, non-significant relationship with tree richness when spider richness was rarefied, i.e. corrected for different total abundances of spiders. Only foraging guild richness (i.e. the diversity of hunting modes) of spiders was positively related to tree species richness. Plant species richness in the herb layer had no significant effects on spiders. Our results thus provide little support for the enemies hypothesis--derived from studies in less diverse ecosystems--of a positive relationship between predator and plant diversity. Our findings for an important group of generalist predators question whether stronger top-down control of food webs can be expected in the more plant diverse stands of our forest ecosystem. Biotic interactions could play important roles in mediating the observed relationships between spider and plant diversity, but further testing is required for a more detailed mechanistic understanding. Our findings have implications for evaluating the way in which theoretical predictions and experimental findings of functional predator effects apply to species-rich forest ecosystems, in which trophic interactions are often considered to be of crucial importance for the maintenance of high plant diversity.     

Intraspecific abundance–occupancy relationships: case studies of six bird species in Britain

Although acknowledged to be common, intraspecific relationships between local abundance and site occupancy have been examined in detail for few species. Here we report such analyses for six widespread species of breeding birds in Britain, using data from the Common Birds Census. These exhibit a range of temporal trends, including different combinations of increase and decrease in abundance and occupancy. Overall, two species have a statistically significant positive abundance–occupancy relationship on farmland but no relationship in woodland (collared dove, tree sparrow), one a significant positive relationship on farmland and in woodland (magpie), two a significant positive relationship on farmland and a negative one in woodland (redstart, song thrush), and one a significant negative abundance–occupancy relationship on farmland but no relationship in woodland (sparrowhawk). The population dynamics associated with these patterns are used to discern their underlying mechanisms.     

Interspecific differences in intraspecific abundance-range size relationships of British breeding birds

A general positive interspecific relationship between local abundance and geographic range size in animals has prompted speculation that a similar relationship might exist intraspecifically, such that a species is widespread at times when it is locally abundant, and more restricted in distribution when it is locally rare. Current evidence suggests that intraspecific relationships often are positive, but that there is considerable variation in the pattern exhibited by species. Here, we use data on British birds to test the hypotheses that species showing a high mean or wide spread of local densities or range sizes will be more likely to show strong intraspecific relationships between abundance and geographic range size. These data show only inconsistent support for an effect of the range of densities or of occupancies on intraspecific abundance-range size relationships. However, the strength of an intraspecific relationship does seem to be related to the mean occupancy of species, and whether or not a species exhibits temporal trends in density, with the strongest relationships found in species with simultaneous trends in both density and occupancy. We suggest that these results are explained by time lags in the loss or gain of species at occupied sites in response to reductions or increases in density.