Latitudinal variation of diversity in European freshwater animals is not concordant across habitat types

Aim   We analysed the variation of species richness in the European freshwater fauna across latitude. In particular, we compared latitudinal patterns in species richness and β-diversity among species adapted to different habitat types.
Location   Europe.
Methods   We compiled data on occurrence for 14,020 animal species across 25 pre-defined biogeographical regions of European freshwaters from the Limnofauna Europaea . Furthermore, we extracted information on the habitat preferences of species. We assigned species to three habitat types: species adapted to groundwater, lotic (running water) and lentic (standing water) habitats. We analysed latitudinal patterns of species richness, the proportion of lentic species and β-diversity.
Results   Only lentic species showed a significant species–area relationship. We found a monotonic decline of species richness with latitude for groundwater and lotic habitats, but a hump-shaped relationship for lentic habitats. The proportion of lentic species increased from southern to northern latitudes. β-Diversity declined from groundwater to lentic habitats and from southern to northern latitudes.
Main conclusions   The differences in the latitudinal variation of species richness among species adapted to different habitat types are in part due to differences in the propensity for dispersal. Since lentic habitats are less persistent than lotic or groundwater habitats, lentic species evolved more efficient strategies for dispersal. The dispersal propensity of lentic species facilitated the recolonization of central Europe after the last glaciation. Overall, we stress the importance of considering the history of regions and lineages as well as the ecological traits of species for understanding patterns of biodiversity.     

Evolution of distribution and habitat patterns in endemic millipedes of the genus Dolichoiulus (Diplopoda: Julidae) on the Canary Islands, with notes on distribution patterns of other Ganarian species swarms

Forty-six species of the genus Dolichoiulus , all endemic, occur on the Canary Islands The highest number of species occur on the largest, highest island (Tenerife); fewest occur on Lanzarote, Fuerteventura (low, xeric), El Hierro and La Palma (small, remote). Most of the Dolichoiulus species live on one island only, as in other endemic Canarian species swarms. The scarcity of pluri-insular Dolichoiulus species, in connection with information on phylogeny, suggests that speciation has mainly taken place within individual islands. Distribution patterns are partly governed by habitat differences between species, but vicariance patterns between species living in the same kind of habitat are evident on La Gomera and Tenerife. Dolichoiulus species occur in all kinds of natural habitats. Laurisilva and cave species are generally paler than other species. In the laurisilva of eastern Tenerife, microhabitat differentiation between species is pronounced. In some, but far from all, cases, species coexisting in the same microhabitat are of different sizes. The ancestral colonizing species of Dolichoiulus is/are hypothesized to have been small and to have lived in coastal habitats. Colonization of higher altitudes was usually accompanied by an increase in body size. Invasion of the laurisilva was usually accompanied by a habitat shift from the ground layer to logs.     

Patterns in species richness and endemism of European freshwater fish

Aim  To analyse the patterns in species richness and endemism of the native European riverine fish fauna, in the light of the Messinian salinity crisis and the Last Glacial Maximum (LGM).
Location  European continent.
Methods  After gathering native fish faunistic lists of 406 hydrographical networks, we defined large biogeographical regions with homogenous fish fauna, based on a hierarchical cluster analysis. Then we analysed and compared the patterns in species richness and endemism among these regions, as well as species–area relationships.
Results  Among the 233 native species present in the data set, the Cyprinidae family was strongly dominant (> 50% of the total number of species). Seven biogeographical regions were defined: Western Peri-Mediterranea, Central Peri-Mediterranea, Eastern Peri-Mediterranea, Ponto-Caspian Europe, Northern Europe, Central Europe and Western Europe. The highest regional species richness was observed for Central Peri-Mediterranea and Ponto-Caspian Europe. The highest endemic richness was found in Central Peri-Mediterranea. Species–area relationships were characterized by high slope values for Peri-Mediterranean Europe and low values for Central and Western Europe.
Main conclusions  The results were in agreement with the 'Lago Mare' hypothesis explaining the specificity of Peri-Mediterranean fish fauna, as well as with the history of recolonization of Central and Western Europe from Ponto-Caspian Europe following the LGM. The results also agreed with the mechanisms of speciation and extinction influencing fish diversity in hydrographical networks. We advise the use of the seven biogeographical regions for further studies, and suggest considering Peri-Mediterranean Europe and Ponto-Caspian Europe as 'biodiversity hotspots' for European riverine fish.     

Broad-scale patterns in geographical and habitat distribution of migrant and resident passerines in Britain and Ireland

We analysed patterns of breeding habitat use in migrant and resident passerines on a broad geographical scale in Britain. Migrants were less widely distributed than residents and used fewer habitats. Although resident genera that were more widespread used more nesting habitats, this did not hold for migrant genera though it did at the species level. Although migrancy status in Europe has been associated with the use of 'open' structural habitat types (e.g. scrubland), evidence from Britain was contradictory. Within lowland woodlands the ratio of migrants to residents and the density of migrants was highest in habitats with large areas of young growth. The highest migrant ratios of all, however, were recorded in mature upland woods, with little or no undergrowth. These upland woods carried the lowest densities of residents. Regression models accounting for variation in the migrant ratios and densities in woodland habitats varied according to the geographical scale on which they were constructed. The extent of young growth was a strong predictor of migrant ratios and densities in south-east England but not for woods in the north and west. Resident densities in northern and western woods were inversely related to altitude.     

How does habitat diversity affect the species–area relationship?

Aim   To examine the way in which 'area' and 'habitat diversity' interact in shaping species richness and to find a simple and valid way to express this interaction.
Location   The Natura 2000 network of terrestrial protected areas in Greece, covering approximately 16% of the national territory.
Methods   We used the Natura 2000 framework, which provides a classification scheme for natural habitat types, to quantify habitat heterogeneity. We analysed data for the plant species composition in 16,143 quadrats in which 5044 species and subspecies of higher plants were recorded. We built a simple mathematical model that incorporates the effect of habitat diversity on the species–area relationship (SAR).
Results   Our analysis showed that habitat diversity was correlated with area. However, keeping habitat diversity constant, species richness was related to area; while keeping area constant, species richness was related to habitat diversity. Comparing the SAR of the 237 sites we found that the slope of the species–area curve was related to habitat diversity.
Main conclusions   Discussion of the causes of the SAR has often focused on the primacy of area per se versus habitat heterogeneity, even though the two mechanisms are not mutually exclusive and should be considered jointly. We find that increasing habitat diversity affects the SAR in different ways, but the dominant effect is to increase the slope of the SAR. While a full model fit typically includes a variety of terms involving both area and habitat richness, we find that the effect of habitat diversity can be reduced to a linear perturbation of the slope of the species accumulation curve.     

Niche properties of Central European spiders: shading, moisture and the evolution of the habitat niche

Aim  Niche theory emphasizes the importance of environmental conditions for the distribution and abundance of species. Using a macroecological approach our study aimed at identifying the important environmental gradients for spiders. We generated numerical values of niche position and niche width. We also investigated relationships between these niche properties as well as the degree of phylogenetic conservatism in order to draw conclusions about the evolution of the habitat niche.
Location  Central Europe: lowlands of Austria, Belgium, Germany, Luxembourg, the Netherlands and Switzerland.
Methods  We analysed 244 published spider communities from 70 habitat types by correspondence analysis. The resulting community scores were used to test for correlations with habitat characteristics. Species scores were used to derive niche position (mean scores) and niche width (standard deviation of scores). To test for niche conservatism we estimated variance components across the taxonomic hierarchy.
Results  The first two axes of the correspondence analysis were correlated with shading and moisture, respectively. Niche width had a hump-shaped relationship to both environmental gradients. β-diversity was strikingly higher in open habitats than in forests. Habitat niche conservatism was lower than phylogenetic conservatism in body size.
Main conclusions  Environmental factors are important drivers for the β-diversity of spiders, especially across open habitats. This underlines the importance of preserving the whole range of moisture conditions in open habitats. Narrow niches of species occurring at the ends of both environmental gradients indicate that adaptations to extreme habitats lead to constraints in ecological flexibility. Nevertheless, the habitat niche of species seems to evolve much faster than morphological or physiological traits.     

Phyloecology of urban alien floras

1.  Understanding the mechanisms that affect invasion success of alien species is a major issue in current ecological research. Although many studies have searched for either functional or habitat attributes that drive invasion mechanisms, few researchers have addressed the role of phylogenetic diversity of alien species.
2.  Here, using data from 21 urban floras located in Europe and eight in the USA, we show that the phylogenetic diversity of alien species is significantly lower than that of native species, both at the continental scale and at the scale of single cities.
3.  Second, we show that if archaeophytes and neophytes (non-native species introduced into Europe before and after AD 1500, respectively) are analysed separately, archaeophytes show lower phylogenetic diversity than neophytes, while the phylogenetic structure of neophytes is indistinguishable from a random sample of species from the entire species pool.
4.  Our results suggest that urban aliens are subject to environmental filters that constrain their phylogenetic diversity, although these filters act more strongly upon archaeophytes than neophytes.
5.   Synthesis. Despite the huge taxonomic diversity of plants imported into European and American cities, the strong environmental filters imposed by cities constrain the functional diversity of urban floras, which is reflected in their generally low phylogenetic diversity. Urban alien floras are mainly composed of phylogenetically related species that are well adapted to anthropogenic habitats, although these filters are stronger for species groups with longer residence times.     

Multi-scale habitat selection and foraging ecology of the eurasian hoopoe (<Emphasis Type="Italic">Upupa epops</Emphasis>) in pine plantations

Bird conservation can be challenging in landscapes with high habitat turnover such as planted forests, especially for species that require large home ranges and juxtaposition of different habitats to complete their life cycle. The eurasian hoopoe (Upupa epops) has declined severely in western Europe but is still abundant in south-western France. We studied habitat selection of hoopoes in pine plantation forests using a multi-scale survey, including point-counts at the landscape level and radio-tracking at the home-range scale. We quantified habitat use by systematically observing bird behaviour and characterized foraging sites according to micro-habitat variables and abundance of the main prey in the study area, the pine processionary moth (Thaumetopoea pityocampa). At the landscape scale, hoopoes selected habitat mosaics of high diversity, including deciduous woods and hedgerows as main nesting sites. At the home-range scale, hoopoes showed strong selection for short grassland vegetation along sand tracks as main foraging habitats. Vegetation was significantly shorter and sparser at foraging sites than random, and foraging intensity appeared to be significantly correlated with moth winter nest abundance. Hoopoe nesting success decreased during the three study years in line with processionary moth abundance. Thus, we suggest that hoopoes need complementation between foraging and breeding habitats to establish successfully in pine plantations. Hoopoe conservation requires the maintenance of adjacent breeding (deciduous woods) and foraging habitats (short swards adjacent to plantation edges), and consequently depends on the maintenance of habitat diversity at the landscape scale.     

Can forest management have season-dependent effects on bird diversity?

Forest management modified the original structure of most European forests, and in the most extreme cases, genuinely natural and semi-natural forests were turned into plantations through clear felling and replanting, often using non-native species. We compared the bird community structure of native oak woods of northern Italy with that of their anthropogenic counter-parts: black locust and sweet chestnut woods. The three stand types were compared in terms of vegetation structure, bird species richness, diversity and abundance of foraging guilds. We analysed both the overwintering and the breeding community, to assess whether management had specific seasonal effects on bird diversity. Forestry-imposed disturbances affected bird diversity more consistently in winter than in breeding time: bird species richness and diversity were significantly greater in oak and chestnut stands, which were the preferred habitat for bark foragers and foliage gleaners. In the breeding period, bird diversity of black locust woodlands increased, and inter-stand differences were not significant. At this time of year, understorey gleaners were more abundant in black locust stands (where shrubs were denser). In winter, species richness, diversity and the abundance of several guilds were positively correlated with stand age, whereas in the breeding period canopy gleaners preferred younger woodlots. Despite the lack of inter-stand differences in breeding bird diversity, young-managed woods benefited generalist birds that need no particular conservation efforts. Conversely, priority species for forest conservation such as specialised bark foragers positively selected native and mature stands throughout the year. We suggest that detailed year-round studies on diversity and community composition could sharpen the precision with which it is possible to prescribe conservation measures in forested areas.     

Effect of landscape structure on the spatial distribution of Mediterranean dung beetle diversity

Aim  Landscape structure influences the distribution of animals, altering their movements and their ability to reach habitat patches. We analysed the spatial patterns of dung beetle species diversity in three differently structured natural landscapes in a Mediterranean protected area in the centre of the Iberian Peninsula.
Location  Cabañeros National Park, Central Spain.
Methods  Diversity components within (α) and among (β) the three main vegetation types in the reserve were compared by using a hierarchical nested design. These were forests, scrublands and grasslands embedded in three different landscapes, where each was the most dominant and structurally connected habitat.
Results  Species richness of grassland habitat did not vary across landscapes, but forest habitat showed lower species richness in the grassland-dominated landscape. Scrubland was the least species-rich habitat, but here again there was no significant variation across landscapes. However, in all cases, there was a significant influence of habitat context (configuration of habitat patches within landscape matrix) on similarity of species composition. These tended to be more similar to the dominant landscape matrix where they were embedded, rather than to the same habitat type in other landscapes. Additive partitioning of diversity showed higher than expected values of β in all landscapes, which indicated a structured response. Highest values of β in the grassland-dominated landscape suggest that this was the least connected landscape for dung beetles.
Main conclusions  Our results suggest that in homogeneous conditions of climate and trophic resources, landscape structure may well be more important than habitat type as a determinant of dung beetle distribution in the Mediterranean.     

Ecological correlates of seed persistence in soil in the north-west European flora

1 Using the data in a recently published seed bank database for north-west Europe, we describe how a species' seed bank behaviour can be characterized by a single 'longevity index', and investigate how representative the information in the database is of the north-west European flora. We also test the hypotheses that seeds of short-lived species are more persistent than those of long-lived species, and long-lived seeds are characteristic of species living in disturbed habitats.
2 The data are not representative of the north-west European flora as a whole; they are a fair reflection of a research effort that has been largely directed towards grassland and arable weeds. Data for rare species, non-agricultural habitats in general and wetlands, rocky habitats and native woodland in particular, are scarce or absent.
3 Annuals and biennials almost always have more persistent seeds than related perennials, and this difference is most striking when, as in Anagallis arvensis and Aphanes arvensis , the short-lived species have moved well away from the 'core' habitat of the family. Confamilial monocarps and polycarps do not differ consistently in seed mass.
4 Gradients of habitat disturbance are accompanied by predictable changes in seed persistence, which are themselves often (but not always) accompanied by parallel shifts in seed size. These results suggest that increasing habitat disturbance (i.e. increasing density-independent mortality) always selects for increased seed persistence, confirming both theory and previous analyses. However, increased seed persistence is not always associated with reduced seed size. This is because persistence depends not only on seed size, but on other traits, many of them physiological. In many habitats the probability of seed burial is strongly linked to seed size and shape, but in arable habitats cultural practices have broken this link.     

Habitat availability does not explain the species richness patterns of European lentic and lotic freshwater animals

Aim In Europe, the relationships between species richness and latitude differ for lentic (standing water) and lotic (running water) species. Freshwater animals are highly dependent on suitable habitat, and thus the distribution of available habitat should strongly influence large‐scale patterns of species richness. We tested whether habitat availability can account for the differences in species richness patterns between European lentic and lotic freshwater animals. Location Europe. Methods We compiled occurrence data of 1959 lentic and 2445 lotic species as well as data on the amount of lentic and lotic habitats across 25 pre‐defined biogeographical regions of European freshwaters. We used the range of elevation of each region as a proxy for habitat diversity. We investigated the relationships between species richness, habitat availability and habitat diversity with univariate and multiple regression analyses. Results Species richness increased with habitat availability for lentic species but not for lotic species. Species richness increased with elevational range for lotic species but decreased for lentic species. For both groups, neither habitat availability nor diversity could account for previously reported latitudinal patterns in species richness. For lotic species, richness declined with latitude, whereas there was no relationship between habitat availability and latitude. For lentic species, richness showed a hump‐shaped relationship with latitude, whereas available habitat increased with latitude. Main conclusions Habitat availability and diversity are poor predictors of species richness of the European freshwater fauna across large scales. Our results indicate that the distributions of European freshwater animals are probably not in equilibrium and may still be influenced by history, namely the recurrent European glaciations and possible differences in post‐glacial recolonization. The distributions of lentic species appear to be closer to equilibrium than those of lotic species. This lends further support to the hypothesis that lentic species have a higher propensity for dispersal than lotic species.     

Conservation of arthropod diversity in montane wetlands: effect of altitude, habitat quality and habitat fragmentation on butterflies and grasshoppers

1. The patterns of arthropod diversity were investigated in 24 montane wetlands in Switzerland. These differed in altitude, management regime (cattle-grazing vs. mowing), vegetation structure (index combining vegetation height and density) and degree of habitat fragmentation.
2. The general arthropod diversity was determined by net sampling at 10 sampling points per site. The diversity of grasshoppers and butterflies was measured by counting species richness at the site and species density (species richness per unit area) on transects. The species richness of grasshoppers and butterflies was found to be more sensitive to the geographical attributes of the site whereas species density was more affected by the habitat quality.
3. Grasshopper diversity decreased within the observed altitudinal range (800–1400 m) and was higher at grazed sites, whereas butterfly diversity was higher at mown sites. Arthropod diversity but not abundance of arthropods was positively related to the vegetation structure.
4. The species richness of butterflies was negatively influenced by the degree of habitat fragmentation: both the size of habitat as well as the area of wetland habitats within 4 km were related positively to the number of specialist wetland butterflies.
5. Late mowing as well as low-density cattle-grazing are appropriate management actions to maintain arthropod diversity in montane wetlands. In order to establish site-specific management plans, the biology of the present target species as well as the historical context should be considered.
6. We suggest that the best protection for the species examined in this study would be a network of wetland sites managed using a variety of traditional, non-intensive methods. This can only be achieved by coordinated planning of conservation measures among sites.     

Using potential distributions to explore determinants of Western Palaearctic migratory songbird species richness in sub-Saharan Africa

Aim  Although the breeding ranges of most Western Palaearctic migratory passerines are well documented in Europe, their overwintering ranges and patterns of species richness in Africa remain poorly understood. To illustrate potential patterns of species richness despite severely limited data, we extrapolated species ranges from a new and unique data bank of locality records that documents overwintering locations of these birds in Africa.
Location  Sub-Saharan Africa.
Methods  We predicted potential geographical distributions of 60 species of passerine birds based on overwintering records using bioclimatic models. We then combined these predictions to estimate potential species richness and explored response shapes using spatial linear regression. We also evaluated the evidence for a mid-domain effect using a one-dimensional null model.
Results  Spatial linear regression analyses of the species richness pattern revealed non-linear relationships to seasonality in precipitation, minimum net primary productivity, minimum average temperature, habitat heterogeneity, percentage of tree cover, distance from the Sahara Desert and inter-annual variability in net primary productivity. The explanatory power of these variables decreased with geographic range size. The one-dimensional null model of species richness based on distance from the Sahara Desert did not show evidence of a mid-domain effect.
Main conclusions  Distributions of migrants seem generally strongly determined by distance from the Sahara Desert working in concert with climatic effects, but this cannot adequately explain richness patterns of species with small ranges in Africa, many of which are of substantial conservation concern.     

Human land use, agriculture, pesticides and losses of imperiled species

Aim  Anthropogenic habitat loss is usually cited as the most important cause of recent species' extinctions. We ask whether species losses are in fact more closely related to habitat loss than to any other aspect of human activity such as use of agricultural pesticides, or human population density (which reflects urbanization).
Location  Canada.
Methods  We statistically compared areas in Canada where imperiled species currently occur, versus areas where they have been lost. Using multiple regressions, we relate the numbers of species that had suffered range reductions in an ecoregion to variables that represent present habitat loss, pesticide use and human population density.
Results  We find high losses of imperiled species in regions with high proportions of agricultural land cover. However, losses of imperiled species are significantly more strongly related to the proportion of the region treated with agricultural pesticides. The relationship between species losses and area treated with pesticides remains significant after controlling for area in agriculture.
Main conclusions  Our results are consistent with the hypothesis that agricultural pesticide use, or something strongly collinear with it (perhaps intensive agriculture more generally), has contributed significantly to the decline of imperiled species in Canada. Habitat conversion per se may be a less important cause of species declines than how that converted habitat is used.     

Distribution of alien vs. native plant species in roadside communities along an altitudinal gradient in Tenerife and Gran Canaria (Canary Islands)

Roadside plant communities were studied along two roads following an altitudinal gradient in Gran Canaria and Tenerife (Canary Islands). Our aim was to investigate variation in plant species richness, particularly of the alien flora, along a gradient from coastal shrubland to summit vegetation (1950 m a.s.l. in Gran Canaria, 2300 m in Tenerife) in relation to variation in habitat factors (altitude, habitat structure, roadside disturbance, distance to urban nuclei). We compared different species groups that were classified in terms of their biogeographical status, origin and life form. Altitude was the most important factor determining species richness and composition along both roadside transects. Alien plants showed a unimodal distribution pattern along the altitudinal gradient, with less species and lower abundance at low and high altitudes, and highest abundance at intermediate altitude. Alien plant species were also relatively more frequent near urban centres. The number of native and alien species was significantly positively correlated along the altitudinal gradient. Both alien and native, non-endemic species showed differences in their distribution along the altitudinal gradient according to their biogeographical affinities and climatic tolerances. Despite considerable differences in species pools these patterns were consistent among the two islands. Environmental (abiotic) stress is proposed as a primary, altitude-related factor acting as a filter against most alien plants at coastal and high-mountain altitudes. A higher frequency or intensity of disturbance at intermediate altitudes may be a further causal factor promoting alien plants in this zone. Future management efforts to control alien plants along roads should, therefore, concentrate on intermediate altitudinal zones of the higher Canary Islands.     

Biosystematics of the Muellerianella complex (Homoptera, Delphacidae): host-plants, habitats and phenology

Abstract. 1. Ecological differences between closely related species and forms of the planthopper genus MueIlerianella (Delphacidae) were studied.
2. In the field M.brevipennis and M.faimairei are restricted to Deschampsia cespitosa and Holcus lanahis/H.mollis . respectively. M.extrusa is mainly found on Molinia caerulea , but has also been recorded from other grass-species. Triploid all-female forms which reproduce by gynogenesis usually live on Holcus together with M.fairmairei . Other gynogenetic populations were found on Bromus ramosus, Deschampsia flexuosa and Calamagrostis canescens .
3. Breeding experiments showed that the diploid species cannot be maintained on each other's host-plants, but accept other grass-species.
4. Due to overlapping habitat requirements of the host-plants, ecological isolation between the Muellerianella species is incomplete.
5. M.extrusa lives in bogs, moors and forests on poor acid soil. M.brevipennis is found in forests, carr, fens and in extensively managed grasslands. M.faimairei and associated triploid forms occur mainly in grasslands and other man-made habitats. The habitats of all three species are usually wet.
6. In most parts of Europe M.brevipennis and M.extrusa are monovoltine and M.fairmairei is mostly bi-or polyvoltine.     

Microsatellite analysis of relatedness structure in young of the year of the endangered Zingel asper (Percidae) and implications for conservation

1. Zingel asper is one of the most endangered freshwater fish of Western Europe. Because of critical habitat loss and fragmentation, stocking or reintroduction protocols are sometimes considered even though few data are available about the behaviour and biology of the species.
2. The present study is the first attempt to explore the population genetic structure of juveniles (young of the year, YOY) of this endangered species using genetic markers (microsatellite loci). In the Beaume River (ardeche, France), Z. asper exhibits very low densities (10–80 fish per hectare), and it was expected that annual recruitment would comprise only few clutches (full- or half-sibs groups).
3. However, sibship reconstruction in one YOY cohort showed that more than 60% of the adult population reproduced. Dispersal rate in early YOY was high: relatedness estimates showed that only few months after hatching, YOY sibs were mixed among study sites. A large number of breeders together with high dispersal rates explain the high genetic variability observed in small populations of Z. asper .
4. Conservation strategies for the species are discussed based on these results, emphasizing the importance of conserving population genetic variability and preserving habitat connectivity for juveniles.