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Metacommunity models describe species occupancy frequency distribution (hereinafter ‘SOFD’). Our goal is to present how the differences in eight macroinvertebrate orders dispersal ability affect SOFD patterns. A total of 293 species from eight macroinvertebrate orders were observed in 14 eutrophic lakes in southern Finland. Species occupancy ranged from 1 to 14. About 30% (89 out of 293) of the species were found in only one lake, yielding a surprisingly high number of rare species. So, there were few widely distributed common species and numerous rare species with a restricted distribution. Combined data from eight macroinvertebrate orders supported the bimodal truncated SOFD pattern. Similarly, the low dispersal ability orders, watermites and mayflies, fitted the bimodal truncated SOFD pattern. However, bimodal symmetric SOFD pattern also fitted relatively well to the dragonflies and damselflies with high dispersal ability. It seems that differences in dispersal ability among different macroinvertebrate orders may partly explain observed differences. Moreover, our results supported slightly more a niche-based model rather than a metapopulation dynamics model in eutrophic lakes littoral macroinvertebrate metacommunities. Our results highlight that the dispersal ability is important trait for species conservation in patchily distributed habitat.

A useful method for characterizing biological numerous assemblages at regional scales is the species occupancy frequency distribution (SOFD). An SOFD shows the number or proportion of study sites each species occurred. Species that occur at only a few sites are termed satellite species, while species that occur at many sites are termed core species. This study is the first to document and assess SOFD patterns in aquatic macrophytes. It characterizes SOFD patterns of freshwater macrophyte assemblages in Finland at two spatial and two temporal scales. For this, I analyzed three published datasets on freshwater macrophyte distributions: two from studies conducted at a local scale and the third from large national surveys. One local study and the national study also included data on temporal variation in species occupancy frequencies. In the national study, the number of core and satellite species varied slightly between the older and the newer survey, respectively. Among the 113 waterbodies surveyed as part of the national study, the SOFD followed a unimodal satellite pattern. However, for the older dataset (from the 1930s), a bimodal symmetric pattern also fit the SOFD data well. At the local scale, I observed geographical variation in SOFD patterns. The dataset from southern Finland followed a unimodal satellite SOFD pattern; data from central Finland instead displayed a bimodal symmetric SOFD pattern, although they also fit equally well with a bimodal truncated pattern. Moreover, temporal patterns in central Finland seemed to demonstrate a shift from a bimodal symmetric to a bimodal asymmetric SOFD probably. Geographical variation in the SOFD pattern may be due to variation in the regional species pool. The temporal changes in SOFD pattern may be due to lake eutrophication and anthropogenic disturbance around waterbodies, which may increase number of macrophyte species., A useful method for characterizing biological numerous assemblages at regional scales is the species occupancy frequency distribution (SOFD). This study is the first to document and assess SOFD patterns in aquatic macrophytes. Geographical variation in the SOFD pattern may be due to variation in the regional species pool and the temporal changes in SOFD pattern may be due to lake eutrophication and anthropogenic disturbance around waterbodies, which may increase number of macrophyte species.

Species prevalence across the landscape is related to their local abundance, which is a result of deterministic and stochastic processes that select organisms capable of recolonizing sites where they were once extinct, a process known as the rescue effect. The occupancy-frequency distribution (OFD) describes these patterns and has been extensively used to understand organism's distribution but has been poorly tested on microorganisms. In order to test OFD on freshwater bacteria, we collected data from 60 shallow lakes distributed across a wide area in southeastern Brazil, to determine the bacterial operational taxonomic units (OTUs) that were present in all sites (core) and at only one site (satellite). Then, we analyzed the spatial abundance distributions of individual OTUs to understand the influence of local abundances on regional occupancy patterns. Finally, we tested the environmental factors that influenced occupancy and abundance. We found a significant bimodal OFD for freshwater bacteria using both OTUs (97% clustering) and amplicon sequence variants (ASVs, unique sequences), with 13 core OTUs and 1169 satellite OTUs, but only three core ASVs. Core organisms had a bimodal or gamma abundance distribution. The main driver of the core community was pH, while nutrients were key when the core community was excluded and the rest of the community (mild and satellite taxa) was considered. This study demonstrates the close relationship between local environmental conditions and the abundance and dispersion of microorganisms, which shapes their distribution across the landscape.

This paper investigates species richness and species occupancy frequency distributions (SOFD) as well as patterns of abundance–occupancy relationship (SAOR) in Odonata (dragonflies and damselflies) in a subtropical area. A total of 82 species and 1983 individuals were noted from 73 permanent and temporal water bodies (lakes and ponds) in the Pampa biome in southern Brazil. Odonate species occupancy ranged from 1 to 54. There were few widely distributed generalist species and several specialist species with a restricted distribution. About 70% of the species occurred in, In this paper, we explore species richness, species occupancy frequency distributions (SOFD), and abundance–occupancy relationship (SAOR) patterns of dragonflies (Odonata) in a neotropical subtropical grassland area, especially addressing differences between permanent and temporal water bodies. We were able to show that both types of water bodies had odonate assemblages which fitted best with the unimodal satellite SOFD pattern while the SAOR pattern was positive and did not differ between permanent and temporal water bodies. Our results are consistent with a niche‐based model rather than a metapopulation dynamic model. This is to our knowledge, the first time SOFD and SAOR patterns are tested for aquatic invertebrates in a subtropical area.

More and more of the globe is becoming urbanized. Thus, characterizing the distribution and abundance of species occupying different towns is critically important. The primary aim of this study was to examine the effect of urbanization and latitude on the patterns of species occupancy frequency distribution (SOFD) in urban core zones of European towns (38 towns) along a 3850-km latitudinal gradient. We determined which of the three most common distributional models (unimodal-satellite dominant, bimodal symmetrical, and bimodal asymmetrical) provides the best fit for urban bird communities using the AICc-model selection procedure. Our pooled data exhibited a unimodal-satellite SOFD pattern. This result is inconsistent with the results from previous studies that have been conducted in more natural habitats, where data have mostly exhibited a bimodal SOFD pattern. Large-sized towns exhibited a bimodal symmetric pattern, whereas smaller-sized towns followed a unimodal- -satellite dominated SOFD pattern. The difference in environmental diversity is the most plausible explanation for this observation because habitat diversity of the study plots decreased as urbanization increased. Southern towns exhibited unimodal satellite SOFD patterns, central European towns exhibited bimodal symmetric, and northern towns exhibited bimodal asymmetric SOFD patterns. One explanation for this observation is that urbanization is a more recent phenomenon in the north than in the south. Therefore, more satellite species are found in northern towns than in southern towns. We found that core species in European towns are widely distributed, and their regional population sizes are large. Our results indicated that earlier urbanized species are more common in towns than the species that have urbanized later. We concluded that both the traits of bird species and characteristics of towns modified the SOFD patterns of urban-breeding birds. In the future, it would be interesting to study how the urban history impacts SOFD patterns and if the SOFD patterns of wintering and breeding assemblages are the same.

Aim Although species-occupancy distributions (SODs) and species-area relationships (SARs) arise from the two marginal sums of the same presence/absence matrices, the two biodiversity patterns are usually explored independently. Here, we aim to unify the two patterns for isolate-based data by constraining the SAR to conserve information from the SOD. Location Widespread. Methods Focusing on the power-model SAR, we first developed a constrained form that conserved the total number of occupancies from the SOD. Next, we developed an additive-constrained SAR that conserves the entire shape of the SOD within the power-model SAR function, using a single parameter (the slope of the endemics-area relationship). We then relate this additive-constrained SAR to multiple-sites similarity measures, based on a probabilistic view of Sorensen similarity. We extend the constrained and additive-constrained SAR framework to 23 published SAR functions. We compare the fit of the original and constrained forms of 12 SAR functions using 154 published data sets, covering various spatial scales, taxa and systems. Main conclusions In all 23 SAR functions, the constrained form had one parameter less than the original form. In all 154 data sets the model with the highest weight based on the corrected Akaike's information criteria (wAICc) had a constrained form. The constrained form received higher wAICc than the original form in 98.79% of valid pairwise cases, approaching the wAICc expected under identical log-likelihood. Our work suggests, both theoretically and empirically, that all SAR functions may have one unnecessary parameter, which can be excluded from the function without reduction in goodness-of-fit. The more parsimonious constrained forms are also easier to interpret as they reflect the probability of a randomly chosen occupancy to be found in an isolate. The additive-constrained SARs accounts for two complimentary turn-over components of occupancies: turnover between species and turnover between sites.

Community patterns in species-by-site matrices provide valuable clues for inferring ecological processes at work. One such pattern is the occupancy frequency distribution (OFD) depicting the frequency distribution of row sums (i.e., occupancy) with a quarter OFDs of bimodal forms. Another pattern that also reflects the structure of row sums is the ranked species occupancy curve (RSOC), and has been shown to imply a 50% of bimodality in OFDs. The use of RSOCs has been advocated in literature over the OFD based on two conclusions from a 6-model inference using only 24 matrices: (i) RSOCs have two general forms, with half representing bimodal OFDs; (ii) there are no effects of spatial and study scales on RSOCs of different forms. Using a much more representative dataset of 289 matrices, I cast doubt on these two conclusions. A missing but dominant RSOC model (the truncated power law) is added. The number of species and the nestedness of the community differ significantly among matrices of different RSOC forms; however, the number of sites and the taxa in the studies do not differ among RSOC or OFD forms. The quarter OFDs of bimodal forms is reassured, with the least frequent occupancy consistent with Raunkiaer’s law of frequency. Importantly, a RSOC is mathematically transferrable to an OFD, with the derivative of the occupancy ranking curve being equal to the negative reciprocal of the occupancy frequency. Based on the type of the community (null versus interactive) and site environment (homogenous versus heterogeneous), four scenarios are needed to identify pre-inferring assemblage mechanisms. The results highlight the need for shifting research from the emphasis of marginal sums to the analysis of matrix structure for an in-depth understanding of the community assemblage patterns and mechanisms.

Patterns of species turnover may reflect the processes driving community dynamics across scales. While the majority of studies on species turnover have examined pairwise comparison metrics (e.g., the average Jaccard dissimilarity), it has been proposed that the species-area relationship (SAR) also offers insight into patterns of species turnover because these two patterns may be analytically linked. However, these previous links only apply in a special case where turnover is scale invariant, and we demonstrate across three different plant communities that over 90% of the pairwise turnover values are larger than expected based on scale-invariant predictions from the SAR. Furthermore, the degree of scale dependence in turnover was negatively related to the degree of variance in the occupancy frequency distribution (OFD). These findings suggest that species turnover diverges from scale invariance, and as such pairwise turnover and the slope of the SAR are not redundant. Furthermore, models developed to explain the OFD should be linked with those developed to explain species turnover to achieve a more unified understanding of community structure.

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.

Species distributions are commonly measured as the number of sites, or geographic grid cells occupied. These data may then be used to model species distributions and to examine patterns in both intraspecific and interspecific distributions. Harte et al. (1999) used a model based on a bisection rule and assuming self-similarity in species distributions to do so. However, this approach has also been criticized for several reasons. Here we show that the self-similarity in species distributions breaks down according to a power relationship with spatial scales, and we therefore adopt a power-scaling assumption for modeling species occupancy distributions. The outcomes of models based on these two assumptions (self-similar and power-scaling) have not previously been compared. Based on Harte's bisection method and an occupancy probability transition model under these two assumptions (self-similar and power-scaling), we compared the scaling pattern of occupancy (also known as the area-of-occupancy) and the spatial distribution of species. The two assumptions of species distribution lead to a relatively similar interspecific occupancy frequency distribution pattern, although the spatial distribution of individual species and the scaling pattern of occupancy differ significantly. The bimodality in occupancy frequency distributions that is common in species communities, is confirmed to a result for certain mathematical and statistical properties of the probability distribution of occupancy. The results thus demonstrate that the use of the bisection method in combination with a power-scaling assumption is more appropriate for modeling species distributions than the use of a self-similarity assumption, particularly at fine scales.

Numerous hypotheses have been proposed to explain the shape of occupancy frequency distributions (distributions of the numbers of species occupying different numbers of areas). Artefactual effects include sampling characteristics, whereas biological mechanisms include organismal, niche-based and meta-population models. To date, there has been little testing of these models. In addition, although empirically derived occupancy distributions encompass an array of taxa and spatial scales, comparisons between them are often not possible because of differences in sampling protocol and method of construction. In this paper, the effects of sampling protocol (grain, sample number, extent, sampling coverage and intensity) on the shape of occupancy distributions are examined, and approaches for minimising artefactual effects recommended. Evidence for proposed biological determinants of the shape of occupancy distributions is then examined. Good support exists for some mechanisms (habitat and environmental heterogeneity), little for others (dispersal ability), while some hypotheses remain untested (landscape productivity, position in geographic range, range size frequency distributions), or are unlikely to be useful explanations for the shape of occupancy distributions 'species specificity and adaptation to habitat, extinction-colonization dynamics). The presence of a core (class containing species with the highest occupancy) mode in occupancy distributions is most likely to be associated with larger sample units, and small homogenous sampling areas positioned well within and towards the range centers of a sufficient proportion of the species in the assemblage. Satellite (class with species with the lowest occupancy) modes are associated with sampling large, heterogeneous areas that incorporate a large proportion of the assemblage range. However, satellite modes commonly also occur in the presence of a core mode, and rare species effects are likely to contribute to the presence of a satellite mode at most sampling scales. In most proposed hypotheses, spatial scale is an important determinant of the shape of the observed occupancy distribution. Because the attributes of the mechanisms associated with these hypotheses change with spatial scale, their predictions for the shape of occupancy distributions also change. To understand occupancy distributions and the mechanisms underlying them, a synthesis of pattern documentation and model testing across scales is thus needed. The development of null models, comparisons of occupancy distributions across spatial scales and taxa, documentation of the movement of individual species between occupancy classes with changes in spatial scale, as well as further testing of biological mechanisms are all necessary for an improved understanding of the distribution of species and assemblages within their geographic ranges.

Much ecological research relies on existing multispecies distribution datasets. Such datasets, however, can vary considerably in quality, extent, resolution or taxonomic coverage. We provide a framework for a spatially-explicit evaluation of geographical representation within large-scale species distribution datasets, using the comparison of an occurrence atlas with a range atlas dataset as a working example. Specifically, we compared occurrence maps for 3773 taxa from the widely-used Atlas Florae Europaeae (AFE) with digitised range maps for 2049 taxa of the lesser-known Atlas of North European Vascular Plants. We calculated the level of agreement at a 50-km spatial resolution using average latitudinal and longitudinal species range, and area of occupancy. Agreement in species distribution was calculated and mapped using Jaccard similarity index and a reduced major axis (RMA) regression analysis of species richness between the entire atlases (5221 taxa in total) and between co-occurring species (601 taxa). We found no difference in distribution ranges or in the area of occupancy frequency distribution, indicating that atlases were sufficiently overlapping for a valid comparison. The similarity index map showed high levels of agreement for central, western, and northern Europe. The RMA regression confirmed that geographical representation of AFE was low in areas with a sparse data recording history (e.g., Russia, Belarus and the Ukraine). For co-occurring species in south-eastern Europe, however, the Atlas of North European Vascular Plants showed remarkably higher richness estimations. Geographical representation of atlas data can be much more heterogeneous than often assumed. Level of agreement between datasets can be used to evaluate geographical representation within datasets. Merging atlases into a single dataset is worthwhile in spite of methodological differences, and helps to fill gaps in our knowledge of species distribution ranges. Species distribution dataset mergers, such as the one exemplified here, can serve as a baseline towards comprehensive species distribution datasets.

There appears to be a general relationship between abundance and distribution that has two parts. First, within species, population density tends to be greatest in the center of the range and to decline gradually toward the boundaries. This pattern holds over a range of spatial scales from steep environmental gradients within local regions to the entire geographic range. Exceptions include: (1) abrupt changes in abundance that usually correspond to sharp, discontinuous changes in single environmental variables; and (2) multimodal patterns of abundance that are caused by environmental patchiness. The second general relationship is that among closely related, ecologically similar species spatial distribution is positively correlated with average abundance. Again this pattern holds over a variety of spatial scales from local regions to entire geographic ranges. These empirical patterns have already been reported in the literature, but their generality is demonstrated by analysis of additional data for divers...

Odonate (damselfly and dragonfly) species richness and species occupancy frequency distributions (SOFDs) were analyzed in relation to geographical location in standing waters (lakes and ponds) in Fennoscandia, from southern Sweden to central Finland. In total, 46 dragonfly and damselfly species were recorded from 292 waterbodies. Species richness decreased to the north and increased with waterbody area in central Finland, but not in southern Finland or in Sweden. Species occupancy ranged from 1 up to 209 lakes and ponds. Over 50% of the species occurred in

Urbanization is negatively affecting biodiversity worldwide and general ecological patterns may also differ between urban and more natural areas. The main aim of this study was to examine if urbanization has effects on the wintering species occupancy frequency distribution (SOFD) and species abundance-occupancy relationship (SAOR), and if the observed patterns varied between winters, different sizes of towns and regionally in Finland. In this study temporal variation of the SOFD and SAOR patterns were studied in 29 town and village centers along a 950 km (60-68˚ N) latitudinal gradient during mid-winters in Finland. Wintering bird were counted during three winters (1991-1992, 1999-2000, and 2009-2010) from the same study sites and with the same survey methods. A total of 35 wintering bird species and 13285 individuals were detected. The bimodal symmetric SOFD pattern explained best the distribution of species in the pooled data, and the observed pattern was constant between the study winters, different sized of towns and towns located in southern and northern Finland. Three species (Parus major, Pica pica and Passer domesticus) were core species during all winters, irrespective of the size of town or latitude. There was slightly higher number of species belonging in satellite species group in the southern than in the northern towns. No changes of species from the core- to satellite-species were detected, and vice versa. However, occupancy rate of some species belonging to the intermediate species group either moved towards satellite-species or core-species across winters. The SAOR pattern was positive and stable over study winters and did not differ between different sizes of towns or town location. Our results indicated that urbanization leads a structure of winter bird community, where there is few widely distributed sedentary core species and many partially migratory or migratory satellite species with a restricted distribution. Our results also demonstrated that urbanization stabilize between-winter community structure, probably because of intensive winter feeding activities. Our results give support to the meta-population model, which predicts a bimodal SOFD pattern.

A new concept is introduced to analyse species' regional distributions and to relate the pattern of distributions to niche relations. Several sets of data indicate that average local abundance is positively correlated with regional distribution, i.e. the fraction of patchily distributed population sites occupied by the species. This observation is not consistent with the assumptions of a model of regional distribution introduced by Levins. A corrected model is now presented, in which the probability of local extinction is a decreasing function of distribution, and a stochastic version of the new model is analysed. If stochastic variation in the rates of local extinction and/or colonization is sufficiently large, species tend to fall into two distinct types, termed the "core" and the "satellite" species. The former are regionally common and locally abundant, and relatively well spaced-out in niche space, while opposite attributes characterize satellite species. This dichotomy, if it exists, provides null hypotheses to test theories about community structure, and it may help to construct better structured theories. Testing the core-satellite hypothesis and its connection to the r-K theory and to Raunkiaer's "law of frequency" are discussed.