Search publications of the members of the Royal Belgian institute of natural Sciences
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Parallel habitat specialization within the wolf spider genus Hogna from the Galapagos
- Within most island archipelagos, such as the Galapagos, similar ecological gradients are found on geographically isolated islands. Species radiations in response to these ecological gradients may follow different scenarios being (i) a single habitat specialization event followed by secondary colonization of each ecotype on the different islands or (ii) repeated and parallel habitat specialization on each island separately. This latter scenario has been considered less likely as gene flow might hamper such ecotypic differentiation. At least for the Galapagos, the extent to which this process is involved in species radiations remains yet poorly understood. Within the wolf spider genus Hogna, seven species are described that can be divided into three different ecotypes based on general morphology and habitat preference i.e. species that inhabit the pampa vegetation in the highlands, species that occur in coastal dry habitats and one generalist species. Comparison of the species phylogeny based on one mitochondrial (COI) and one nuclear (28S) gene fragment convincingly demonstrates that 'pampa' and 'coastal dry' species evolved in parallel on the islands Santa Cruz and San Cristobal. Despite the observation that allozymes analysis indicated that each species forms a distinct genetic cluster, phylogenetic divergence within these species complexes was very low and paraphyletic and most likely due to hybridization rather than incomplete lineage sorting, as demonstrated for the Santa Cruz species complex. This suggests that within-island speciation occurred under low levels of gene flow. Species phylogeny in general did not follow the progression of island emergence as a molecular clock analysis suggested that island endemic species may have diverged after as well as before the emergence of the islands. This represents the first clear example of parallel and within-island speciation because of habitat specialization on the Galapagos and that such divergence most likely occurred under historic gene flow.
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Patterns of habitat occupancy, genetic variation and predicted movement of a flightless bush cricket, Pholidoptera griseoaptera, in an agricultural mosaic landscape
- Habitat fragmentation has been generally regarded detrimental to the persistence of many species, especially those with limited dispersal abilities. Yet, when exactly habitat elements become functionally disconnected very much depends on the dispersal ability of a species in combination with the landscape's composition in which it occurs. Surprisingly, for many small and ground-walking generalists knowledge at what spatial scale and to what extent landscape structure affects dispersal is very scarce. Because it is flightless, the bush cricket Pholidoptera griseoaptera may be regarded susceptible to fragmentation. We applied habitat occupancy surveys, population genetic analyses and movement modelling to investigate the performance of P. griseoaptera in an agricultural mosaic landscape with suitable habitat patches of varying size and isolation. Despite its presumed dispersal limitation we could show that P. griseoaptera occupied the majority of suitable habitats, including small and isolated patches, showed a very low and non-significant genetic differentiation (F (ST) = 0.0072) and, in the model, managed to colonize around 73\% of all suitable habitat patches within one generation under weak and strong landscape-effect scenarios. We conclude that P. griseoaptera possesses the behavioural attributes (frequent inter-patch dispersal) necessary to persist in this landscape characterized by a patchy distribution of habitat elements. Yet, sound recommendations to landscape planning and conservation require more research to determine whether this represents a general behaviour of the species or a behavioural adaptation to this particular landscape.
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Pervasive effects of dispersal limitation on within- and among-community species richness in agricultural landscapes
- Aim To determine whether the effect of habitat fragmentation and habitat heterogeneity on species richness at different spatial scales depends on the dispersal ability of the species assemblages and if this results in nested species assemblages. Location Agricultural landscapes distributed over seven temperate Europe countries covering a range from France to Estonia. Methods We sampled 16 local communities in each of 24 agricultural landscapes (16 km(2)) that differ in the amount and heterogeneity of semi-natural habitat patches. Carabid beetles were used as model organisms as dispersal ability can easily be assessed on morphological traits. The proximity and heterogeneity of semi-natural patches within the landscape were related to average local (alpha), between local (beta) and landscape (gamma) species richness and compared among four guilds that differ in dispersal ability. Results For species assemblages with low dispersal ability, local diversity increased as the proximity of semi-natural habitat increased, while mobile species showed an opposite trend. Beta diversity decreased equally for all dispersal classes in relation to proximity, suggesting a homogenizing effect of increased patch isolation. In contrast, habitat diversity of the semi-natural patches affected beta diversity positively only for less mobile species, probably due to the low dispersal ability of specialist species. Species with low mobility that persisted in highly fragmented landscapes were consistently present in less fragmented ones, resulting in nested assemblages for this mobility class only. Main conclusions The incorporation of dispersal ability reveals that only local species assemblages with low dispersal ability show a decrease of richness as a result of fragmentation. This local species loss is compensated at least in part by an increase in species with high dispersal ability, which obscures the effect of fragmentation when investigated across dispersal groups. Conversely, fragmentation homogenizes the landscape fauna for all dispersal groups, which indicates the invasion of non-crop habitats by similar good dispersers across the whole landscape. Given that recolonization of low dispersers is unlikely, depletion of these species in modern agricultural landscapes appears temporally pervasive.
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Prediction uncertainty of environmental change effects on temperate European biodiversity
- Observed patterns of species richness at landscape scale (gamma diversity) cannot always be attributed to a specific set of explanatory variables, but rather different alternative explanatory statistical models of similar quality may exist. Therefore predictions of the effects of environmental change (such as in climate or land cover) on biodiversity may differ considerably, depending on the chosen set of explanatory variables. Here we use multimodel prediction to evaluate effects of climate, land-use intensity and landscape structure on species richness in each of seven groups of organisms (plants, birds, spiders, wild bees, ground beetles, true bugs and hoverflies) in temperate Europe. We contrast this approach with traditional best-model predictions, which we show, using cross-validation, to have inferior prediction accuracy. Multimodel inference changed the importance of some environmental variables in comparison with the best model, and accordingly gave deviating predictions for environmental change effects. Overall, prediction uncertainty for the multimodel approach was only slightly higher than that of the best model, and absolute changes in predicted species richness were also comparable. Richness predictions varied generally more for the impact of climate change than for land-use change at the coarse scale of our study. Overall, our study indicates that the uncertainty introduced to environmental change predictions through uncertainty in model selection both qualitatively and quantitatively affects species richness projections.
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Present conservation status of red wood ants in north-western Belgium: Worse than previously, but not a lost cause
- The number of colonies of red wood ants (Formica rufa group) in the north western part of Belgium has declined by more than half over the past 20 years. In the study area of 20 x 50 km there are currently only 36 colonies of two species of red wood ants (Formica rufa and F. polyctena). Both species form both monodomous and polydomous colonies. Monodomous F. rufa colonies occur in large areas of open woodland dominated by Pinus sylvestris. The other three types of red wood ant (F. rufa polydomous, F. polyctena mono-and polydomous) occur in clearings or along the margins of small areas of deciduous or mixed woodland. The extinction of red wood ants cannot be attributed to a loss of their main habitat. Their decline in abundance, however, appears to be due to the deterioration in their habitat caused by several factors. Shading by overgrowing shrubs and the closure of the tree canopy, nearby intensive agriculture, urbanisation and use of the forest for recreation are the more important factors threatening and ultimately leading to the extinction of colonies. At poor quality sites there are no or very few nests of Serviformica ants, which initially serve as host ants and are necessary for recolonization by red wood ants. Thus, it is unlikely that these sites will be recolonized. To conserve red wood ants and maintain populations there is an urgent need to undertake certain measures, especially in areas outside nature reserves. For this there is a need to create clearings in the woodland, reduce shading due to overgrowing shrubs and closure of the tree canopy, and mitigate the effects of intensive agriculture, urbanisation and recreation. It is important that the effects of these measures should be monitored.
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Quantifying the impact of environmental factors on arthropod communities in agricultural landscapes across organizational levels and spatial scales
- 1. In landscapes influenced by anthropogenic activities, such as intensive agriculture, knowledge of the relative importance and interaction of environmental factors on the composition and function of local communities across a range of spatial scales is important for maintaining biodiversity. 2. We analysed five arthropod taxa covering a broad range of functional aspects (wild bees, true bugs, carabid beetles, hoverflies and spiders) in 24 landscapes (4 x 4 km) across seven European countries along gradients of both land-use intensity and landscape structure. Species-environment relationships were examined in a hierarchical design of four main sets of environmental factors (country, land-use intensity, landscape structure, local habitat properties) that covered three spatial scales (region, landscape, local) by means of hierarchical variability partitioning using partial canonical correspondence analyses. 3. Local community composition and the distribution of body size classes and trophic guilds were most affected by regional processes, which highly confounded landscape and local factors. After correcting for regional effects, factors at the landscape scale dominated over local habitat factors. Land-use intensity explained most of the variability in species data, whereas landscape characteristics (especially connectivity) accounted for most of the variability in body size and trophic guilds. 4. Synthesis and applications. Our results suggest that management effort should be focused on land-use intensity and habitat connectivity in order to enhance diversity in agricultural landscapes. Since these factors are largely independent, specific conservation programmes may be developed with regards to socio-economic and agri-environmental requirements. Changes in either of these factors will enhance diversity but will also result in specific effects on local communities related to dispersal ability and the resource use of species.
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Relationship between fluctuating asymmetry and fitness within and between stressed and unstressed populations of the wolf spider Pirata piraticus
- Although developmental instability, measured as fluctuating asymmetry (FA), is expected to be positively related to stress and negatively to fitness, empirical evidence is often lacking or contradictory when patterns are compared at the population level. We demonstrate that two important properties of stressed populations may mask such relationships: (i) a stronger relationship between FA and fitness, resulting in stronger selection against low quality (i.e. developmental unstable) individuals and (ii) the evolution of adaptive responses to environmental stress. In an earlier study, we found female wolf spiders Pirata piraticus from metal exposed populations to be characterized by both reduced clutch masses and increased egg sizes, the latter indicating an adaptive response to stress. By studying the relationship between these two fitness related traits and levels of FA at individual level, we here show a significant negative correlation between FA and clutch mass in metal stressed populations but not in unstressed reference populations. As a result, levels of population FA may be biased downward under stressful conditions because of the selective removal of developmentally unstable (low quality) individuals. We further show that females that produced larger eggs in stressed populations exhibited lower individual FA levels. Such interaction between individual FA and fitness with stress may confound the effect of metal stress on FA, resulting in an absence of relationships between FA, fitness and stress at the population level.
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Response of snails and slugs to fragmentation of lowland forests in NW Germany
- Habitat fragmentation is a major cause for species loss, but its effect on invertebrates with low active dispersal power, like terrestrial gastropods, has rarely been studied. Such species can not cross a hostile habitat matrix, for which the predictions of island theory, such as positive relations between species richness and patch size, should apply. In order to test this prediction, we studied gastropod species diversity by assessing gastropod assemblage characteristics from 35 sites in 19 fragments of deciduous old-growth forests in the Lower Rhine Embayment, Germany. Assemblages differed between larger (a parts per thousand yen700 ha) and smaller forests ( 400 ha), those of large forests held a higher percentage of forest species. Although alpha-diversity was similar between the two forest size classes, small forests often comprised matrix species, resulting in a higher beta-diversity. Edge effects on the species richness of matrix species were noticeable up to 250 m into the forest. Hierarchical partitioning revealed that distance to disturbances (external edge, internal edges like roads) explained most assemblage variables, whereas forest size and woodland cover within a 1 km radius from the sites explained only a few assemblage variables. Densities of two forest-associated species, Discus rotundatus and Arion fuscus, decreased with forest size. Yet, forest size was positively correlated with richness of typical forest species and densities of Limax cinereoniger. The latter species seems to need forests of 1,000 ha, i.e., well above the size of most fragments. In conclusion, the prediction is valid only for forest species. The response to fragmentation is species specific and seems to depend on habitat specialization and macroclimatic conditions.
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Site Fidelity of Formica rufa: Micro-Scaled and Persistent Despite Disturbance (Hymenoptera: Formicidae)
- Long-term site fidelity or Ortstreue is an individual foraging strategy typical of ants exploiting stable and predictable food sources in space and time. Red wood ant workers (Formica s. str.) are central-place foragers feeding on honeydew secreted by stable aphid populations. In this field study, temporal site fidelity of Formica rufa honeydew tenders was investigated on a micro-scaled level for a period of time. Additionally, the effect of exchanging honeydew workers between two trees on site fidelity was examined. Site fidelity of the honeydew tenders was very high and they returned repeatedly to particular micro-sites (branches, set of leaves) on a tree. The transfer to a new feeding site apparently did not influence site fidelity as honeydew tenders mainly return to their initial tree. Small-scale site fidelity decreased with time and was not so rigid as larger-scale fidelity. We discuss how site fidelity on one hand and flexibility in foraging and recruiting on the other hand promote foraging efficiency of the colony.
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Spiders do not escape reproductive manipulations by Wolbachia
- Background: Maternally inherited bacteria that reside obligatorily or facultatively in arthropods can increase their prevalence in the population by altering their hosts' reproduction. Such reproductive manipulations have been reported from the major arthropod groups such as insects (in particular hymenopterans, butterflies, dipterans and beetles), crustaceans (isopods) and mites. Despite the observation that endosymbiont bacteria are frequently encountered in spiders and that the sex ratio of particular spider species is strongly female biased, a direct relationship between bacterial infection and sex ratio variation has not yet been demonstrated for this arthropod order. Results: Females of the dwarf spider Oedothorax gibbosus exhibit considerable variation in the sex ratio of their clutches and were infected with at least three different endosymbiont bacteria capable of altering host reproduction i.e. Wolbachia, Rickettsia and Cardinium. Breeding experiments show that sex ratio variation in this species is primarily maternally inherited and that removal of the bacteria by antibiotics restores an unbiased sex ratio. Moreover, clutches of females infected with Wolbachia were significantly female biased while uninfected females showed an even sex ratio. As female biased clutches were of significantly smaller size compared to non-distorted clutches, killing of male embryos appears to be the most likely manipulative effect. Conclusions: This represents to our knowledge the first direct evidence that endosymbiont bacteria, and in particular Wolbachia, might induce sex ratio variation in spiders. These findings are pivotal to further understand the diversity of reproductive phenotypes observed in this arthropod order.
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Spinnen en zware metalen: 1. Een kort overzicht van de literatuur
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Storage mediums affect metal concentration in woodlice (Isopoda)
- Terrestrial invertebrates are becoming widely established as tools to assess heavy metal pollution at contaminated sites. A practical and time saving method to sample terrestrial invertebrates consist of pitfall traps, often filled with a 4\% formaldehyde solution and some detergent. The reliability of metal concentrations based on organisms captured and stored in this solution might however be questioned and we therefore tested the effect of formaldehyde on Zn, Cu, Cd and Pb concentration experimentally in three isopod species. Our results showed that in many cases, significant decreases in Cu concentrations compared to animals stored in a freezer were observed that could be as high as 40\%, while Zn, Cd and Ph concentrations increased. A regression analysis of individual dry weight on individual size revealed that formaldehyde decreases the dry weight substantially and in that way causes increased measurements of Zn, Cd and Pb concentrations. We conclude that pitfall traps with formaldehyde should better not be used to collect animals in which concentrations of heavy metals or other toxic substances will be determined. (C) 2002 Elsevier Science Ltd. All rights reserved.
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Survival differences of the two male morphs in the dwarf spider Oedothorax gibbosus Blackwall, 1841 (Erigoninae, Linyphiidae, Araneae)
- Oedothorax gibbosus is a rare dwarf spider species in Flanders bound to oligo- and mesotrophic alder carrs. This dwarf spider is characterised by the appearance of a male dimorphism; there is a gibbosus male morph with a hunch on the last third of the carapace, anterior to which is a hairy groove, and a tuberosus morph without these features. Earlier studies have already indicated some differences between both male morphs. According to VANACKER et al. (2001) the juvenile phase (the period between the emergence of the spiders and the last moult) of the gibbosus morph is significantly longer than the juvenile phase of the tuberosus morph, and this is presumably necessary for the production of the hunch and the hairy groove. This hairy groove probably secretes a fluid that is important for the gustatoric courtship behaviour and gibbosus would therefore have a reproductive advantage (HEINEMANN & UHL, 2000). In this paper we report another interesting difference between both morphs. To examine if the male morphs also differ in survival, some survival experiments, in normal food and moisture conditions and in two extreme conditions, namely in foodless and low moisture environments, were set up. The gibbosus male is significantly more susceptible to foodless conditions than the tuberosus male, and also in normal conditions tuberosus lives significantly longer than gibbosus. This is demonstrated by means of survival curves. In low moisture conditions, however, we could not prove that survival rate differs between both male morphs. The higher survival rate of tuberoses is a possible explanation for the morph ratio in favour of tuberosus in the studied field population.
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Synecology of spiders of gravel banks and environmental constraints along a lowland river system, the Intermeuse (Belgium, the Netherlands) (Araneae)
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The effect of successional stage and salinity on the vertical distribution of seeds in salt marsh soils
- Seed bank density and similarity between above-ground vegetation and seed bank with depth were compared between two adjacent salt marshes that differ in age. In addition, the effect of salinity on the variation in seed bank density and similarity between above-ground vegetation and seed bank with depth was compared between euhaline against mesohaline conditions in three salt marshes. Ten plots of 2 m x 2 m were situated in a new salt marsh (existing since 2002) and 80 plots in three old salt marshes. Soil samples were collected at three different depths (0-5, 5-10, 10-15 cm) in spring 2006. Soil seed bank was investigated in germination experiments under greenhouse conditions. Germination experiments lasted 6 months and all seedlings were identified and removed after identification. Above-ground vegetation composition was determined during the growing season in all plots. Viable seed density was calculated for each plot and for the three different depths; the similarity between seed bank and floristic composition of the above-ground vegetation was calculated. A general linear model was used to investigate the effect of soil depth, salinity and age of the salt marsh on density and similarity between seed bank and above-ground vegetation. The results showed that seed density decreased with depth in all salt marshes, irrespective of their age and soil salinity. Seed density and similarity between seed bank and above-ground vegetation were higher in the new salt marsh than in the old one in the same study area. This is because in young as well as in old successional stages, the seed bank was mostly composed of new colonizers, while most perennial species were absent from the soil seed bank, although they were dominant in the standing vegetation of the old salt marsh. The characteristics of the seed bank of a mesohaline salt marsh were found not to be fundamentally different from that of both euhaline salt marshes. (C) 2010 Elsevier GmbH. All rights reserved.
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The effects of forest patch size and matrix type on changes in carabid beetle assemblages in an urbanized landscape
- In this study we compared ground beetles (Carabidae) from a range of different forest fragments along an urbanization gradient in Brussels, Belgium. We address the following questions: (i) How does the degree of urbanization in the surrounding habitat affect forest beetles, and does it interact with the effects of patch size and distance to forest edge? (ii) Do these factors have a different effect at the level of individual species, habitat affinity groups or total community? During 2002 we sampled 13 forest plots in 10 forest patches, ranging in size from 5.27 to 4383 ha. The beetles were captured using transects of pitfall traps from the edge to a distance of 100 m into each woodland and identified to species level. Effects of urbanization, forest size and forest edge were evaluated on total species number, abundance and habitat affinity groups and ten abundant, widespread model carabid species. Overall, the effects of urbanization, forest size and edge effects slightly influenced total species richness and abundance but appeared to have a major effect on ground beetle assemblages through species specific responses. More urbanized sites had significantly fewer forest specialists and more generalist species. Large forest fragments were favoured by forest specialist species while generalist species and species frequently associated with forest (forest generalists) dominated the smaller forests. Forest edges mainly harboured generalist species while forest specialist species were more frequent into the forests if the forest patches were large enough, otherwise they disappeared due to the destruction or impoverishment of their habitat. Our results show the importance of differentiating between habitat affinity, especially habitat generalists versus specialists, the latter having a higher value in nature conservation, and merely the quantity of species represented in human-dominated areas. (c) 2008 Elsevier Ltd. All rights reserved.
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The importance of biological factors affecting trace metal concentration as revealed from accumulation patterns in co-occurring terrestrial invertebrates
- As physicochemical properties of the soil highly influence the bioavailable fraction of a particular trace metal, measured metal body burdens in a particular species are often assumed to be more reliable estimators of the contamination of the biota. To test this we compared the Cd, Cu and Zn content of three spiders (generalist predators) and two amphipods (detritivores), co-occurring in seven tidal marshes along the river Schelde, between each other and with the total metal concentrations and the concentrations of four sequential extractions of the soils. Correlations were significant in only one case and significant site x species interactions for all metals demonstrate that factors affecting metal concentration were species and site specific and not solely determined by site specific characteristics. These results emphasize that site and species specific biological factors might be of the utmost importance in determining the contamination of the biota, at least for higher trophic levels. A hypothetical example clarifies these findings. (C) 2003 Elsevier Ltd. All rights reserved.
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The richness and paucity of the spider fauna of the Belgian coast.
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La Formation des schistes noires de Matagne et l'extinction du Dévonien Supérieur
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A single tooth replacement pattern generates diversity in the dentition in cichlids of the tribe Eretmodini, endemic to Lake Tanganyika.pdf
