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.     

Inferring long-distance dispersal and topographic barriers during post-glacial colonization from the genetic structure of red maple (Acer rubrum L.) in New England

Aim  This study aims to assess the role of long-distance seed dispersal and topographic barriers in the post-glacial colonization of red maple ( Acer rubrum L.) using chloroplast DNA (cpDNA) variation, and to understand whether this explains the relatively higher northern diversity found in eastern North American tree species compared with that in Europe.
Location  North-eastern United States.
Methods  The distribution of intraspecific cpDNA variation in temperate tree populations has been used to identify aspects of post-glacial population spread, including topographic barriers to population expansion and spread by long-distance seed dispersal. We sequenced c.  370 cpDNA base pairs from 221 individuals in 100 populations throughout the north-eastern United States, and analysed spatial patterns of diversity and differentiation.
Results  Red maple has high genetic diversity near its northern range limit, but this diversity is not partitioned by topographic barriers, suggesting that the northern Appalachian Mountains were not a barrier to the colonization of red maple. We also found no evidence of the patchy genetic structure that has been associated with spread by rare long-distance seed dispersal in previous studies.
Main conclusions  Constraints on post-glacial colonization in eastern North America seem to have been less stringent than those in northern Europe, where bottlenecks arising from long-distance colonization and topographic barriers appear to have strongly reduced genetic diversity. In eastern North America, high northern genetic diversity may have been maintained by a combination of frequent long-distance dispersal, minor topographic obstacles and diffuse northern refugia near the ice sheet.     

Broad-scale patterns of body size in squamate reptiles of Europe and North America

Aim  To document geographical interspecific patterns of body size of European and North American squamate reptile assemblages and explore the relationship between body size patterns and environmental gradients.
Location  North America and western Europe.
Methods  We processed distribution maps for native species of squamate reptiles to document interspecific spatial variation of body size at a grain size of 110 × 110 km. We also examined seven environmental variables linked to four hypotheses possibly influencing body size gradients. We used simple and multiple regression, evaluated using information theory, to identify the set of models best supported by the data.
Results  Europe is characterized by clear latitudinal trends in body size, whereas geographical variation in body size in North America is complex. There is a consistent association of mean body size with measures of ambient energy in both regions, although lizards increase in size northwards whereas snakes show the opposite pattern. Our best models accounted for almost 60% of the variation in body size of lizards and snakes within Europe, but the proportions of variance explained in North America were less than 20%.
Main conclusions  Although body size influences the energy balance of thermoregulating ectotherms, inconsistent biogeographical patterns and contrasting associations with energy in lizards and snakes suggest that no single mechanism can explain variation of reptile body size in the northern temperate zone.     

Track analysis of the marine palaeofauna from the Turonian (Late Cretaceous)

Aim  To analyse the worldwide distribution patterns of Turonian marine biotas using a panbiogeographical approach.
Location  Turonian localities of southern and north-eastern Brazil, Mexico, Canada, central Europe, England and Morocco.
Method  Panbiogeographical track analysis.
Results  Nine generalized tracks and six nodes were found. The generalized tracks comprise two vicariant track patterns (one northern and one mid-southern) across the Atlantic.
Main conclusions  The generalized tracks show clearly two separate marine biotas, which were associated with the proto-South Atlantic and the proto-North Atlantic oceans. These generalized tracks, as well as the two vicariant track clusters between the north and south Atlantic, are identified by vicariant relationships shared by most of the taxa analysed, and illustrate the final break up of the Gondwana and Laurasia supercontinents and the consequences of vicariant events for the biogeography of the Atlantic Ocean.     

Zooplankton biomass and community structure in a Danube River floodplain system: effects of hydrology

1. Zooplankton density and biomass was examined in a Danube River floodplain section with highly variable hydrological dynamics. Temporal patterns were analysed to assess the effects of hydrological conditions on zooplankton community structure and the differential response of the two major zooplankton taxa, rotifers and crustaceans.
2. Calculated floodplain water age was used as an integrated parameter describing hydrological conditions and connectivity.
3. Total zooplankton biomass, crustacean biomass and crustacean species number were significantly positively related to water age. Rotifer biomass followed a hump-shaped relationship with water age, and rotifer species number decreased with increasing water age.
4. Rotifers dominated the community in periods of low to medium water ages. In periods of higher water ages the community was dominated by crustaceans.
5. We propose that the hydrological regime of floodplains is crucial for zooplankton biomass patterns and succession, through the alternation of washing-out effects, taxon-specific potential of reproduction and biological interactions. Flood events and high water levels reset the community to an early successional phase.     

Patterns of post-glacial spread and the extent of glacial refugia of European beech (Fagus sylvatica)

Aim  Establishing possible relationships between the magnitudes of the glacial distribution of the European beech, Fagus sylvatica L., and its post-glacial spread.
Location  Europe.
Methods  A database of over 400 pollen records has been used to locate Fagus populations at the end of the last glacial and during the post-glacial in Europe and to assess the areal extent of their past distribution.
Results  The rate of late-glacial and post-glacial increase in the number of pollen sites where Fagus was locally present conforms well to a logistic model of population growth. This suggests that the area occupied by beech populations expanded exponentially from the glacial refugia for a duration of over 10,000 years, until about 3500 yr bp . In the past three millennia beech populations increased at a slower rate, tending towards an equilibrium value.
Main conclusions  The conformity of the increase in beech distribution to the classical logistic model of population growth indicates that: (1) a multiplicative biological process was the main factor shaping the pattern of the post-glacial expansion of F. sylvatica in Europe, (2) climate conditions, human activity and competition may have influenced its rate of spread, and (3) beech populations did not expand with a moving closed front, but with a diffuse spread from scattered nuclei. The distribution of Fagus in Europe at the end of the last glacial appears to have been of two orders of magnitude less extensive than at present. Pleistocene refugia were likely to have been a mosaic of sparse stands of small populations scattered in multiple regions. Fagus populations appear to have increased very slowly and to a moderate extent in southern Europe, where they are now declining slightly. The central European populations increased quickly and extensively, reaching northern Europe, and are now approaching their carrying capacity.     

Glacial refugia for summer-green trees in Europe and south-west Asia as proposed by ECHAM3 time-slice atmospheric model simulations

Aim  To generate maps of potential refugia for summer-green trees during the Last Glacial Maximum (LGM).
Locations  Southern Europe and south-western Asia.
Methods  Time-slice simulations of the atmospheric climate with the ECHAM3 model are used for the LGM. Limiting factors beyond which cool and warm groups of deciduous trees cannot grow (such as temperature in growing degree days, minimum monthly temperature and precipitation in summer) are chosen. A limited validation by fossil pollen and charcoal records from LGM sites was done.
Results  Two sets of maps extending from Europe to the Caspian region for cool and warm summer-green trees are presented. Three criteria are combined using contour lines to indicate confidence levels. Small areas within the three southern peninsulas of Europe (Spain, Italy and Greece) are highlighted as possible refugia for summer-green trees. Further, areas that have remained poorly known are now proposed as refugia, including the Sakarya–Kerempe region in northern Turkey, the east coast of the Black Sea and the area south of the Caspian Sea.
Main conclusions  The maps produced in this study could be used to facilitate better long-term management for the protection of European and south-western Asian biodiversity.     

Patterns of parasitism by insect parasitoids in patchy environments

Abstract. 1. This paper shows how the different spatial patterns of per cent parasitism in patches of different host density can be explained within a single model framework that takes into account the parasitoid's aggregative response, and the factors limiting the degree of host exploitation within patches.
2. Two contrasting laboratory examples are presented in which the distribution of searching parasitoids and the resulting levels of parasitism in different patches are both known for a range of parasitoid densities.
3. A model is described predicting the number of hosts parasitized per patch, in which the number of parasitoids searching is determined from a simple expression allowing different degrees of aggregation.
4. The model generates patterns of parasitism encompassing the two laboratory examples and a wide range of examples from the field.
5. The importance of density dependent spatial distributions of parasitism to population stability is briefly discussed.     

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.     

Butterfly extinctions in European states: do socioeconomic conditions matter more than physical geography?

Aim  To distinguish the effects of physical geography and socioeconomic conditions on the extinction of butterflies in European states, and to compare patterns influencing extinctions with patterns influencing species richness.
Location  Europe.
Method  Per-state species richness and extinctions were taken from the Red Data Book of European Butterflies , and their relationships with physical geography and socioeconomic predictors were analysed using regression analysis. Two hypothesis were explored: (1) extinctions are related primarily to identical physical geography factors that influence species richness; and (2) extinctions are influenced primarily by human pressure on natural biotopes and follow correlates of modern land use.
Results  Extinctions and richness are not correlated. Richness increased towards low latitudes and with biotope and topographic heterogeneity, and decreased in states affected by Quaternary glaciation and on islands. The only socioeconomic correlate was human density, exhibiting a weak negative effect. Extinctions were negatively correlated with area and with biotope and topographic heterogeneity. They peaked in regions with mild climate in central latitudes. The strongest socioeconomic correlate was high density of railways, interpreted as a proxy of early industrialization. Further correlates were human density and urban employment.
Main conclusion  Topographic and biotope heterogeneity predicts both high species richness and low extinction rates. Losses of butterflies result from a complex interplay of geography and relatively recent economic history, as low topographic heterogeneity and flat relief favoured the early advent of industrialization and intensive land use.     

Range-wide phylogeography of the European temperate-montane herbaceous plant Meum athamanticum Jacq.: evidence for periglacial persistence

Aim  The aim of this study is to analyse the genetic population structure of Meum athamanticum Jacq. in order to explore the alternative hypotheses (1) that the central and northern highland populations are the result of post-glacial recolonization from southern refugia, and the disjunct distribution of M. athamanticum can be explained by modern ecological conditions, or (2) that extant populations north of the Alps and Pyrenees persisted in situ during glacial periods.
Location  Europe.
Methods  Variation of amplified fragment length polymorphisms (AFLPs) was analysed for 23 populations from the entire range of the species. We used band-based approaches and methods based on allele frequencies to measure genetic diversity and to identify population structure.
Results  Our analyses reveal a north–south differentiation within M. athamanticum . High levels of genetic diversity, as well as private fragments, are found in populations both north and south of the Alps. Differentiation among populations is lower in the northern than in the southern population group, and significant isolation-by-distance is found only in the latter group.
Main conclusions  Our results indicate that M. athamanticum survived the last ice age in multiple refugia throughout its contemporary range and did not expand into areas north of the Alps from southern refugia. We found evidence that regional-scale migration in northern, formerly periglacial, parts of the species range has resulted in the intermingling of populations. In contrast, southern populations are characterized by long-term isolation. The south-west Alps represent an area where immigration and mixing of populations from northern and southern refugia appears to have taken place.     

Patterns of molecular diversity in wild stocks of the redclaw crayfish (Cherax quadricarinatus) from northern Australia and Papua New Guinea: impacts of Plio-Pleistocene landscape evolution

1. Analysis of mitochondrial and nuclear DNA (microsatellites) in 379 individuals, collected from 15 localities in northern Australia and Papua New Guinea (PNG), demonstrated that wild redclaw crayfish ( Cherax quadricarinatus ) populations consist of two highly divergent Australian lineages and two PNG lineages.
2. The disjunction between the two Australian lineages occurs over a distance of approximately 200 km in the south-western corner of the Gulf of Carpenteria. These data conflict with an earlier study that detected no significant differentiation in 23 variable allozyme loci in redclaw sampled from northern Australia, but concur broadly with the previous recognition of two morphologically distinct species ( C. quadricarinatus and C. bicarinatus ) across northern Australia, and a third species in PNG ( C. albertsii ).
3. The inferred timing and patterns of divergence evident in the molecular data presented here closely align with a similar pattern reported in a co-distributed freshwater decapod crustacean, and broadly reflect patterns in some vertebrate taxa with similar distributions across northern Australia and PNG.
4. These congruent patterns most probably reflect periodic Plio-Pleistocene land and freshwater connections between Australia and New Guinea.     

Inverse density dependent parasitism in a patchy environment: a laboratory system

Abstract. 1. Two species of parasitoids (Anisopteromalus calandrae (Howard) and Heterospilus prosopidis Vier) attacking the bruchid beetle, Callosobruchus chinensis (L.), show marked inverse density dependent relationships between per cent parasitism and host density per patch.
2. These patterns are well described quantitatively using data on the spatial distribution of searching time by the parasitoids and their attack rates on patches of different host density.
3. A model of optimal foraging predicts just the opposite (i.e. density dependent) patterns of parasitism.
4. Both density dependent and inversely density dependent spatial patterns of parasitism can be explained mechanistically in terms of (a) the allocation of searching time in patches of different host density and (b) the maximum attack rate per parasitoid that constrains the extent of host exploitation within a patch.     

Effect of starvation time on the prey capture behaviour, functional response and population growth of Asplanchna sieboldi (Rotifera)

1. We examined the effect of different periods of prior starvation(from 30 min to 16 h) on the prey capture behaviour, and functional and numerical responses of the predatory rotifer Asplanchna sieboldi using the rotifer Brachionus calyciflorus as prey.
2. Feeding activity (i.e. encounter, attack, capture and ingestion) by Asplanchna increased significantly with increasing prey densities from 2 to 16 mL⁻⁻¹ and with increasing prior starvation periods from 0.5 to 8 h.
3.  Asplanchna sieboldi showed a type II functional response at all the prior starvation periods tested. The asymptotic prey density was highest after 8 h of starvation.
4. The instantaneous population growth rate of A. sieboldi ranged from 0.089 ± 0.044 (when starved for 8 h in every 24 h and at a prey density of 2 individuals mL⁻⁻¹ for the other 16 h) to 1.015 ± 0.142 in the control (no starvation and at a prey density of 16 individuals mL⁻⁻¹). The effect of starvation time on the numerical response was evident only at the higher prey density.     

An ecosystem model-based estimate of changes in water availability differs from water proxies that are commonly used in species distribution models

Aim  To assess whether the water availability measures commonly used in species distribution models might be misleading because they do not account for the hydrological effects of changes in vegetation structure and functioning.
Location  Europe.
Methods  We compared different methods for estimating water availability in species distribution models with the soil water content predicted by a process-based ecosystem model. The latter also accounted for the hydrological effects of dynamic changes in vegetation structure and functioning, including potential physiological effects of increasing CO₂.
Results  All proxies showed similar patterns of water availability across Europe for current climate, but when projected into the future, the changes in the simpler water availability measures showed no correlation with those projected by the more complex ecosystem model, even if CO₂ effects were switched off.
Main conclusions  Results from species distribution modelling studies concerning future changes in species ranges and biodiversity should be interpreted with caution, and more process-based representations of the water balance of terrestrial ecosystems should be considered within these models.     

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.     

Density-dependent melanism in sub-arctic populations of winter moth larvae (Operophtera brumata)

Abstract.  1. The aim of this 4-year observational study was to test for the presence of direct and delayed density-dependent larval melanism in the geometrid moth species Operophtera brumata (winter moth) in northern Norway.
2. Data from many populations with a wide range of population densities in time and space facilitated statistical analyses that could separate the effects of current and past density. The data also included different phases of the 10-year population cycle of this species so that eventual non-linear density effects due to population phase could be detected.
3. The results showed that the prevalence of melanism had a strong positive, linear relation to population density within years, whereas there was no evidence for a delayed effect from the year before or dependency on the phase of the population cycle.
4. In combination, these results limit the range of possible explanations of larval melanism in this outbreaking species. The possible reasons why winter moth larvae might benefit from crowding-induced melanism are discussed.     

Spoor counts as indices of large carnivore populations: the relationship between spoor frequency, sampling effort and true density

1. Studies of large carnivore populations and, particularly, reliable estimates of population density are necessary for effective conservation management. However, these animals are difficult to study, and direct methods of assessing population size and density are often expensive and time-consuming.
2. Indirect sampling, by counting spoor, could provide repeatable and inexpensive measures of some population parameters. The relationship between true population density and indirect sampling results has seldom been described in large carnivore studies.
3. In northern Namibia the population densities of leopards, lions and wild dogs were measured through recognition of individuals and groups. Spoor counts were then conducted independently, to assess the relationship between true density and the distribution of spoor.
4. Sampling effort, both in terms of the number of roads and total road distance in a sample zone, and the intensity of sampling, had a marked effect on the accuracy and precision of spoor frequency calculations.
5. In a homogeneous habitat, leopard spoor were evenly spread along different roads and spoor frequency was independent of road length. Taking into account very low sample sizes, the spoor density of leopards, lions and wild dogs showed a strong linear correlation with true density. The slope of the regression for leopards was different to that of lions and wild dogs.     

The 'edgeeffect' in butterfly oviposition: causality in Anthocharis cardarnines and related species

Abstract. 1. An 'edge-effect', where outlying host plants receive proportionately most attention from insect herbivores, is described from the literature, and from data on Anthochoris cardamines (L.), in northern England.
2. Six hypotheses are examined as explanations for the effect in A.curdamines . Four are shown to be refuted by studies on female behaviour and the survival of larvae.
3. It is argued that the 'edge-effect' comes about due to two or more aspects of the searching behaviour of females. The effect may be seen as an outcome of behaviour adapted to searching for host plants of low density.