Relationships between local population persistence, local abundance and regional occupancy of species: distribution patterns of diatoms in boreal streams

Aims We have two aims: (1) to examine the relationship between local population persistence, local abundance and regional occupancy of stream diatoms and (2) to characterize the form of the species–occupancy frequency distribution of stream diatoms. Location Boreal streams in Finland. There were three spatial extents: (1) across ecoregions in Finland, (2) within ecoregions in Finland, and (3) within a single drainage system in southern Finland. Methods Diatoms were sampled from stones (epilithon), sediment (epipelon) and aquatic plants (epiphyton) in streams using standardized sampling methods. To assess population persistence, diatom sampling was conducted monthly at four stream sites from June to October. The relationships between local population persistence, local abundance and regional occupancy were examined using correlation analyses. Results There was a significant positive relationship between local persistence and abundance of diatoms in epilithon, epipelon and epiphyton. Furthermore, local abundance and regional occupancy showed a significant positive relationship at multiple spatial extents; that is, across ecoregions, within ecoregions and within a drainage system. The relationships between occupancy and abundance did not differ appreciably among impacted and near pristine‐reference sites. The occupancy–frequency distribution was characterized by a large number of satellite species which occurred at only a few sites, whereas core species that occurred at most sites were virtually absent. Main conclusions The positive relationship between local population persistence and abundance suggested that a high local abundance may prevent local extinction or that high persistence is facilitated by a high local cell density. High local persistence and local abundance may also positively affect the degree of regional occupancy in stream diatoms. The results further showed that anthropogenic effects were probably too weak to bias the relationship between occupancy and abundance, or that the effects have already modified the distribution patterns of stream diatoms. The small number of core species in the species–occupancy frequency distribution suggested that the regional distribution patterns of stream diatoms, or perhaps unicellular microbial organisms in general, may not be fundamentally different from those described previously for multicellular organisms, mainly in terrestrial environments, although average global range sizes may differ sharply between these two broad groups of organisms.     

Untangling the relationships among regional occupancy,species traits,and niche characteristics in stream invertebrates

The regional occupancy and local abundance of species are affected by various species traits, but their relative effects are poorly understood. We studied the relationships between species traits and occupancy (i.e., proportion of sites occupied) or abundance (i.e., mean local abundance at occupied sites) of stream invertebrates using small‐grained data (i.e., local stream sites) across a large spatial extent (i.e., three drainage basins). We found a significant, yet rather weak, linear relationship between occupancy and abundance. However, occupancy was strongly related to niche position (NP), but it showed a weaker relationship with niche breadth (NB). Abundance was at best weakly related to these explanatory niche‐based variables. Biological traits, including feeding modes, habit traits, dispersal modes and body size classes, were generally less important in accounting for variation in occupancy and abundance. Our findings showed that the regional occupancy of stream invertebrate species is mostly related to niche characteristics, in particular, NP. However, the effects of NB on occupancy were affected by the measure itself. We conclude that niche characteristics determine the regional occupancy of species at relatively large spatial extents, suggesting that species distributions are determined by environmental variation among sites.     

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.     

Positive relationship between regional distribution and local abundance in stream insects: a consequence of niche breadth or niche position?

A positive relationship between regional distribution and local abundance of species is almost ubiquitous macroecological pattern, yet the mechanisms behind this pattern remain poorly understood. I tested for the relationship between regional distribution and local abundance of stream insect species in a boreal drainage system, with a specific aim to examine if this relationship follows the mechanistic basis of either the niche-based (niche breadth and niche position) or metapopulation models. There was a positive relationship between regional distribution and local abundance of stream insects, and there also were significant relationships between distribution/abundance and niche breadth or niche position. These results thus suggest that widely distributed species tend to be, on average, locally more abundant, have wider niches and lower marginality of niche position with regard to environmental factors than species that have more restricted distributions. However, although significant, there was much unexplained variability around these relationships, suggesting that other mechanisms (e.g. metapopulation dynamics) besides differences in species' niches are likely to affect the distribution and abundance of stream insects, at least within a drainage system. The results thus showed that 1) although niche position was more consistently related to the positive distribution-abundance relationship, ecologists should not abandon niche breadth as a potential mechanism behind this relationship, and 2) that several mechanisms are likely to act in concert in determining the relationship between distribution and abundance of species.     

Reinvestigation on species richness and environmental correlates of bryophytes at a regional scale in China

Aims The aim of this article is 4-fold: (i) to update species richness of bryophytes for each of the Chinese provinces based on the most current knowledge on distributions of bryophytes in China, (ii) to provide a set of analyses based on the updated species richness data and the environmental variables used in a recent article on species richness of bryophytes in China, (iii) to expand the analysis presented in the recent article by relating species richness of bryophytes to over 15 additional climatic variables and (iv) to determine the degree to which the relationships between bryophyte species richness and environmental variables that were reported in the recent article might have been biased.Methods Over 180 literatures with national, provincial and local species lists of bryophytes in China were used in this study. Taxonomy and nomenclature of bryophytes in China were standardized according to The Plant List. Correlation and regression analyses were used to examine the relationships between species richness or species density of bryophytes in Chinese provinces and environmental variables.Important findings On average, each Chinese province possesses 700.6 species of bryophytes, which is 112.1 species more than previously reported. With the updated species richness data reported in this study, stronger relationships between species richness of bryophytes and environmental variables have been found, compared with those found in a recently published study for China. When single environmental variables were considered, precipitation-related variables were, on average, more strongly correlated with species richness and species density than were temperature-related variables. Environmental variables were on average correlated more strongly with species density than with species richness of bryophytes at the regional scale in China. Our study showed that measures quantifying the average and variation of environmental conditions within each Chinese province explained 82.7% and 71.1% of the variation in species richness of liverworts and mosses, respectively, and explained 86.5% and 70.7% of the variation in species density of liverworts and mosses, respectively.     

Stream community richness predicts apex predator occupancy dynamics in riparian systems

Streams and adjacent riparian habitats represent linked terrestrial and aquatic ecosystems that exchange materials and energy. Recognized relationships among apex predators and ecosystem biodiversity led us to hypothesize that these predators in riparian‐stream systems were more likely to be found in sites with high stream quality, defined as increased ecosystem function and integrity. In our freshwater study system, river otter Lontra canadensis and mink Neovison vison play critical roles as apex predators. We used multi‐season occupancy modelling across three sampling years (2012–2014) to compare aspects of the stream communities that explain occupancy dynamics of river otter and mink, including their interactions with other semi‐aquatic mammals. We surveyed for semi‐aquatic mammals at 77 sites in 12 major watersheds in southern Illinois, USA (44 526 km²). Naïve occupancy differed among years but generally increased for river otter, and remained high (≥93.5%) for mink. Increasing substrate availability increased detectability of river otter, whereas mink detection varied by survey period. Occupancy of river otter during the initial survey period was higher in sites closer to reintroduction points. Probability of colonization of river otter was positively associated with macroinvertebrate index of biotic integrity, fish species richness, and beaver presence. Sites with high species richness of fish families preferred by river otter also had increased river otter persistence. Mink occupied sites with increased fish richness, muskrat presence and mussel community index. Taken together, our results show occupancy of both mink and river otter were predicted by aspects of prey diversity and presence, indicating the importance of community composition in occupancy dynamics of riparian predators. Ultimately, these relationships suggest that habitat heterogeneity and system stability are important to apex predator site use. However, the relative role of bottom–up and top–down forcing in stream systems remains to be resolved.     

Predictability of stream insect distributions is dependent on niche position,but not on biological traits or taxonomic relatedness of species

Species distributions can be analysed under two perspectives: the niche‐based approach, which focuses on species–environment relationships; and the dispersal‐based approach, which focuses on metapopulation dynamics. The degree to which each of these two components affect species distributions may depend on habitat fragmentation, species traits and phylogenetic constraints. We analysed the distributions of 36 stream insect species across 60 stream sites in three drainage basins at high latitudes in Finland. We used binomial generalised linear models (GLMs) in which the predictor variables were environmental factors (E models), within‐basin spatial variables as defined by Moran's eigenvector maps (M models), among‐basin variability (B models), or a combination of the three (E + M + B models) sets of variables. Based on a comparative analysis, model performance was evaluated across all the species using Gaussian GLMs whereby the deviance accounted for by binomial GLMs was fitted on selected explanatory variables: niche position, niche breadth, site occupancy, biological traits and taxonomic relatedness. For each type of model, a reduced Gaussian GLM was eventually obtained after variable selection (Bayesian information criterion). We found that niche position was the only variable selected in all reduced models, implying that marginal species were better predicted than non‐marginal species. The influence of niche position was strongest in models based on environmental variables (E models) or a combination of all types of variables (E + M + B models), and weakest in spatial autocorrelation models (M models). This suggests that species–environment relationships prevail over dispersal processes in determining stream insect distributions at a regional scale. Our findings have clear implications for biodiversity conservation strategies, and they also emphasise the benefits of considering both the niche‐based and dispersal‐based approaches in species distribution modelling studies.     

Spatial and temporal variability of aquatic habitat use by amphibians in a hydrologically modified landscape

1. Habitat loss is a major driver of biodiversity decline worldwide. Temporary waterbodies are especially vulnerable because they are sensitive both to human impact and to climatic variations. Pond‐breeding amphibians are often dependent on temporary waterbodies for their reproduction, and hence are sensitive to loss of temporary ponds. 2. Here we present the results of a 5‐year study regarding the use of temporary aquatic habitats by amphibians in a hydrologically modified area of Eastern Europe (Romania). The annual number of aquatic habitats varied between 30 and ～120. Each aquatic habitat was characterised by a number of variables such as: ‘type’ (pond, drainage ditch and archaeological ditch), ‘hydroperiod’ (number of weeks the ponds were filled in a given year), ‘depth’ (cm), ‘area’ (m²) and the density of predatory insects (‘predation’). The turnover rate for each amphibian species for each wetland was calculated based on the pond occupancy. 3. Eight amphibian species were recorded from the aquatic habitats. Hydroperiod was the most important variable, positively influencing wetland use by amphibians and their reproductive success. Most species preferred drainage ditches for reproduction, and the reproductive success was highest in this habitat type every year. For most of the species, the local extinction rate was higher than the colonisation rate in the first 4 years, but the situation reversed in the last year of the study when wetland use by amphibians sharply increased because of high rainfall. 4. This study confirms the importance for amphibians of maintaining and managing aquatic habitat diversity at small spatial scales. Man‐made aquatic habitats such as drainage ditches may be important habitats for amphibians, and this should be considered in restoration activities.     

Control of stream insect assemblages: roles of spatial configuration and local environmental factors

Abstract 1. Current views in ecology emphasise that community structure is the sum of multiple processes, with imprints of both regional and local drivers. However, the degree to which stream insect assemblages are structured by spatial configuration (complying with the dispersal‐based neutral hypothesis) and local environmental features (complying with the niche‐based species sorting hypothesis) has not been rigorously examined based on surveys in multiple years. 2. Stream sites in a boreal drainage system were surveyed during three consecutive years and the relative contribution of spatial configuration and local environmental variables to aquatic insect assemblage structure (characterised by both abundance and presence–absence data) was assessed. Separate analyses were conducted for mayflies (Ephemeroptera), stoneflies (Plecoptera), caddisflies (Trichoptera), and non‐biting midges (Diptera: Chironomidae) in each year. 3. There were no relationships between the spatial location and local environmental features of streams in Mantel tests, facilitating exploration of their independent effects on assemblage structure. The study found virtually no effects of spatial location on stream insect assemblages across the study drainage system, as evidenced by Mantel tests and canonical correspondence analyses (CCA). The environmental variables were also rather weakly associated with assemblage structure, with the total amount of explained variation ranging from 9.8% to 31.7% in the CCAs. There were no appreciable differences in the amount of environment‐related explained variation in assemblage structure between mayflies, stoneflies, caddisflies, and midges, but some between‐year differences were noticeable in most insect groups. The environmental variables that were significantly related to assemblage structure exhibited some between‐group and between‐year variability. In general, patterns shown by abundance and presence–absence data were highly similar. 4. It appears that stream insect assemblages comply with the niche‐based species sorting hypothesis in the context of metacommunity ecology. In contrast, the absence of spatial structuring suggests that stream insect assemblages do not comply with the neutral hypothesis, being not strongly dispersal limited at the within‐drainage basin scale.     

Odonate species occupancy frequency distribution and abundance–occupancy relationship patterns in temporal and permanent water bodies in a subtropical area

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 <10% of the water bodies, yielding a surprisingly high number of rare species, often making up the majority of the communities. No difference in species richness was found between temporal and permanent water bodies. Both temporal and permanent water bodies had odonate assemblages that fitted best with the unimodal satellite SOFD pattern. It seems that unimodal satellite SOFD pattern frequently occurred in the aquatic habitats. 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.     

Deconstructing occupancy frequency distributions in stream insects: effects of body size and niche characteristics in different geographical regions

1. Occupancy frequency distributions (OFDs) are one means to study species distribution patterns, allowing the delineation of rare and common species. Very few studies have deconstructed entire assemblages by ecological or biological characteristics and subsequently examined OFDs in subgroups of species. 2. The effect of deconstruction of entire assemblages by niche breadth, niche position or body size classes on OFDs in stream insects in three drainage basins was examined. It was hypothesized that OFDs should not vary between different drainage basins, but they should be affected by deconstruction into different niche breadth, niche position or body size classes. 3. The OFDs were typically strongly right‐skewed in all drainage basins. The set of small‐sized species was strongly dominated by rare species, whereas the importance of rare species decreased with increasing body size. Further, while the OFDs of sets of species with marginal niche position or small niche breadth were strongly dominated by rare species, those of species with non‐marginal niche position or large niche breadth showed highly variable degrees of occupancy. The OFDs of non‐marginal species were even uniform in the entire data and one drainage basin, providing partial support to the a priori hypothesis. 4. Niche‐based explanations are likely to account for occupancies of marginal and small‐niched species, whereas the distributions of non‐marginal and broad‐niched species may be not only affected by niche‐based mechanisms but also by spatial dynamics. Deconstruction of OFDs by ecological and biological traits thus showed that the patterns may vary between different subgroups of species.     

Persistence at distributional edges: Columbia spotted frog habitat in the arid Great Basin,USA

A common challenge in the conservation of broadly distributed, yet imperiled species is understanding which factors facilitate persistence at distributional edges, locations where populations are often vulnerable to extirpation due to changes in climate, land use, or distributions of other species. For Columbia spotted frogs (Rana luteiventris) in the Great Basin (USA), a genetically distinct population segment of conservation concern, we approached this problem by examining (1) landscape‐scale habitat availability and distribution, (2) water body‐scale habitat associations, and (3) resource management‐identified threats to persistence. We found that areas with perennial aquatic habitat and suitable climate are extremely limited in the southern portion of the species’ range. Within these suitable areas, native and non‐native predators (trout and American bullfrogs [Lithobates catesbeianus]) are widespread and may further limit habitat availability in upper‐ and lower‐elevation areas, respectively. At the water body scale, spotted frog occupancy was associated with deeper sites containing abundant emergent vegetation and nontrout fish species. Streams with American beaver (Castor canadensis) frequently had these structural characteristics and were significantly more likely to be occupied than ponds, lakes, streams without beaver, or streams with inactive beaver ponds, highlighting the importance of active manipulation of stream environments by beaver. Native and non‐native trout reduced the likelihood of spotted frog occupancy, especially where emergent vegetation cover was sparse. Intensive livestock grazing, low aquatic connectivity, and ephemeral hydroperiods were also negatively associated with spotted frog occupancy. We conclude that persistence of this species at the arid end of its range has been largely facilitated by habitat stability (i.e., permanent hydroperiod), connectivity, predator‐free refugia, and a commensalistic interaction with an ecosystem engineer. Beaver‐induced changes to habitat quality, stability, and connectivity may increase spotted frog population resistance and resilience to seasonal drought, grazing, non‐native predators, and climate change, factors which threaten local or regional persistence.     

Modeling stream fish distributions using interval‐censored detection times

Controlling for imperfect detection is important for developing species distribution models (SDMs). Occupancy‐detection models based on the time needed to detect a species can be used to address this problem, but this is hindered when times to detection are not known precisely. Here, we extend the time‐to‐detection model to deal with detections recorded in time intervals and illustrate the method using a case study on stream fish distribution modeling. We collected electrofishing samples of six fish species across a Mediterranean watershed in Northeast Portugal. Based on a Bayesian hierarchical framework, we modeled the probability of water presence in stream channels, and the probability of species occupancy conditional on water presence, in relation to environmental and spatial variables. We also modeled time‐to‐first detection conditional on occupancy in relation to local factors, using modified interval‐censored exponential survival models. Posterior distributions of occupancy probabilities derived from the models were used to produce species distribution maps. Simulations indicated that the modified time‐to‐detection model provided unbiased parameter estimates despite interval‐censoring. There was a tendency for spatial variation in detection rates to be primarily influenced by depth and, to a lesser extent, stream width. Species occupancies were consistently affected by stream order, elevation, and annual precipitation. Bayesian P‐values and AUCs indicated that all models had adequate fit and high discrimination ability, respectively. Mapping of predicted occupancy probabilities showed widespread distribution by most species, but uncertainty was generally higher in tributaries and upper reaches. The interval‐censored time‐to‐detection model provides a practical solution to model occupancy‐detection when detections are recorded in time intervals. This modeling framework is useful for developing SDMs while controlling for variation in detection rates, as it uses simple data that can be readily collected by field ecologists.     

Contribution of rare and common species to richness patterns at local scales

Several studies have recently reported that common species are more important for species richness patterns than rare species. However, most such studies have been based on broad‐scale atlas data. We studied the contribution of different species occupancy, i.e. number of plots occupied, to species richness patterns emerging from species data in 50 by 50 m plots within six 140–200 ha forests in Norway. The study included vascular plants, lichens, bryophytes, and polypore fungi. We addressed the following questions: 1) are common species more correlated with species richness than rare species? 2) How do occupancy classes combine at various levels of species richness? 3) Which occupancy class is best in identifying the overall most species‐rich sites (hotspots) by sampling? The results showed that rare species were better correlated with species richness than common species when the information content was accounted for, that high species richness was associated with a higher proportion of less frequent species, and that the best occupancy class for local hotspot identification was species present in 10–30% of the plots within a forest. We argue that the observed correlations between overall richness and sub‐assembly richness are primarily structured by the combination of the distributions of species richness and species occupancy. Although these distributions result from general ecological processes, they may also be strongly affected by idiosyncratic elements of the individual datasets caused by the specific environmental composition of a study area. Hence, different datasets collected in different areas may lead to different results regarding the relative importance of common versus rare species, and such effects should be expected on both broad and fine spatial scales. Despite these effects, we suggest that infrequent species will tend to be more strongly correlated to species richness at local scales than at broader scales as a result of more right‐skewed species‐occupancy distributions.     

The barriers to oceanic island radiation in bryophytes: insights from the phylogeography of the moss Grimmia montana

Aim In contrast to angiosperms, bryophytes do not appear to have radiated in Macaronesia and the western Mediterranean. We evaluate if: (1) the apparent lack of radiation in bryophytes reflects our failure to recognize cryptic endemic species; (2) bryophytes are characterized by extremely low evolutionary rates; or (3) bryophytes have a high dispersal ability, which prevents genetic isolation. Location Worldwide, with a special emphasis on Macaronesia and the western Mediterranean. Methods Three chloroplast regions were sequenced from samples of the moss Grimmia montana from its entire distribution range. Network analyses, Fst and Nst statistics were used to describe and interpret the phylogeographical signal in the data. Results Despite significant phylogeographical signal in the chloroplast genome, which demonstrates limits to gene flow at the continental scale, repeated sister group relationships observed among accessions from different geographical areas suggest recurrent colonization patterns. These observations are consistent with mounting evidence that intercontinental distributions exhibited by many bryophyte species result from long‐distance dispersal rather than continental drift. Madeiran and western Mediterranean island haplotypes are either shared by, or closely related to, European and North American ones. Fst values between Madeira, western Mediterranean islands, North America and Europe are not significantly different from zero, and suggest that Madeira and the south‐western Mediterranean are subject to strong transatlantic gene flow. By contrast, haplotypes found in the Canary Islands are shared or closely related to those of populations from south‐western Europe or southern Africa. Main conclusions Multiple origins and colonization events are not consistent with the hypothesis of a relictual origin of the Macaronesian moss flora. One possible reason for the failure of taxa that experienced multiple colonization events to radiate is niche pre‐emption. We suggest that strong gene flow, coupled with the occupancy of all suitable niches, either by earlier conspecific colonizers or by other species, could be the mechanism preventing island radiation in G. montana and other cryptogams with high long‐distance dispersal abilities.     

Regional occupancy in unicellular eukaryotes: a reflection of niche breadth,habitat availability or size‐related dispersal capacity?

1. The distribution patterns of unicellular and multicellular organisms have recently been shown to differ profoundly, with the former probably being mostly cosmopolitan, whereas the latter are mostly restricted to certain regions. However, the within‐region distribution patterns of these two organism groups may be rather similar. 2. We predicted that the degree of regional occupancy in unicellular eukaryotes would be related to niche characteristics, dispersal ability and size, as has been found previously for multicellular organisms. The niche characteristics we considered were niche position, that measures marginality in species habitat distribution, and niche breadth, that measures amplitude in species habitat distribution. Niche characteristics were determined using Outlying Mean Index (OMI) analysis. 3. We found that the regional occupancy in our model group of unicellular eukaryotes, stream diatoms, was primarily a reflection of the niche position of a species or, more generally, habitat availability. Thus, non‐marginal species (i.e. species that occupied common habitat conditions across the region) tended to be more widely distributed than marginal species (i.e. species that were restricted to a limited range of rare habitat conditions). This finding was further supported by the general linear model, with niche position, niche breadth, maximum size and attachment mode as explanatory variables: niche position was by far the most important variable accounting for variability in regional occupancy, with significant amounts of additional variation related to niche breadth and maximum size of diatoms. 4. Thus, the degree of regional occupancy among unicellular eukaryotes may be primarily governed by habitat availability, supporting former findings for multicellular organisms.     

Global and regional patterns in lotic meiofauna

• 1 Parsimony analysis of endemicity (PAE) was used to assess patterns in the distribution of harpacticoid copepods (all freshwater species and stream species only) at global and regional scales. These analyses provided a focus for reviewing large scale patterns and processes in freshwater meiofauna.
• 2 On a global scale, PAE suggested that large‐scale biogeographical events have been most important in shaping present‐day distributions in the Canthocamptidae. A small proportion (4%) of canthocamptid species were widespread (i.e. occurred in more than one biogegraphical region), suggesting that dispersal may also play a role in determining distribution at the species level. Global distribution patterns for other meiofauna suggest varying roles for dispersal and vicariant events. No consistent latitudinal trends in species diversity were evident, although a lack of distributional data for many regions, and uncertainty over the status of many cosmopolitan species, precludes more robust analyses. Molecular techniques should prove useful in identifying truly cosmopolitan taxa.
• 3 On a regional scale, a PAE within Western Europe demonstrated a clear link between the distribution of canthocamptid species and the extent of the Last (Wiechselian) glaciation. Northern and southern areas of Europe contain distinctive harpacticoid faunas and the recolonisation of northern Europe appears to have been from the Balkans rather than other Mediterranean peninsulae. The high harpacticoid diversity in southern Europe, may reflect a lack of glacial disruption of groundwater habitats.
• 4 A PAE of lotic data for harpacticoid copepods within the Holarctic reflected the global PAE for freshwater harpacticoids as a whole, but not the regional PAE. A high proportion of stream‐dwelling harpacticoids are widespread species, but only one (Bryocamptus zschokkei) was found in streams across the Holarctic. Other cosmopolites were restricted to streams in Europe or North America, suggesting that species‘ niche requirements might differ among regions. There appeared to be some convergence in the composition of lotic copepod communities in terms of the number of species within genera.
• 5 We conclude that large‐scale processes inevitably have a major influence on the local composition of lotic meiofaunal communities, but that the relative importance of small scale vs. large scale processes is unclear at present, largely due to a paucity of suitable data.

     

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.     

Temporal variability of nestedness and idiosyncratic species in stream insect assemblages

Aims The nested subset pattern has been widely studied in the last 20 years, and recent syntheses have challenged the prevalence of this pattern in nature. We examined the degree of nestedness, its temporal variability and its environmental correlates in stream insects of a boreal drainage system. We also examined differences between nested and idiosyncratic species in site occupancy, niche position and niche breadth. Location Koutajoki drainage basin in northern Finland. Methods We used (i) nestedness analyses with three null models for testing the significance of nestedness; (ii) Spearman rank correlation to examine the correlates of nestedness; (iii) outlying mean index analysis to analyse the niche characteristics of species; (iv) and t‐test to examine differences in niche breadth, niche position and site occupancy of idiosyncratic and other nested species. Results Stream insect assemblages were significantly nested in each of the three study years. The maximally packed matrices were significantly nested according to the nestedness calculator based on null models I (species frequencies and site richness equiprobable) and II (species frequencies fixed and site richness equiprobable), but non‐significant based on a conservative null model III (species frequencies and site richness fixed to those of the observed matrix). The most important correlate of nestedness was stream size, whereas isolation, productivity (total phosphorus) and habitat heterogeneity exhibited non‐significant relationship with nestedness. Idiosyncratic species occurred, on average, at more sites than nested species, mirroring the restricted distributions of several nested species that were inclined towards species‐rich sites. Idiosyncratic and nested species also differed in niche position and niche breadth, with idiosyncratic species having, on average, less marginal niche positions and wider niches than nested species. Main conclusions Stream size correlated with nestedness, possibly because small streams were inhabited only by species able to persist under, or colonize shortly after, disturbances, while most species could occur at larger sites where disturbances are less severe. From the conservation perspective, our findings suggest that stream size really matters, given that sites with high species richness and many rare species are more likely to occur in larger streams. However, also the requirements of idiosyncratic species should be accommodated in conservation planning.