Refuge from cannibalism in complex-structured habitats: implications for the accumulation of invertebrate predators

Abstract.  1. Like many invertebrate predators, the wolf spider Pardosa littoralis Banks (Araneae: Lycosidae) accumulates in complex-structured habitats replete with leaf litter (thatch). Here we test the hypothesis that P. littoralis accumulates in complex habitats to gain refuge from cannibalism.
2. A laboratory experiment examined the effects of habitat complexity (thatch present or absent) and size-class pairing of conspecific spiders (large vs. small, small vs. small, and large vs. large) on the incidence of cannibalism. Spider survival was significantly higher (22%) in complex-structured habitats with thatch than in simple-structured habitats lacking thatch. Furthermore, cannibalism occurred more frequently in P. littoralis when the size of conspecifics was asymmetric (large vs. small spiders) than when spiders were of equal size. There was no interactive effect of habitat complexity and size-class pairing on spider survival.
3. A field experiment examined the effects of habitat complexity, conspecific density, and access to alternative prey on the prevalence of cannibalism in P. littoralis . Access to alternative prey significantly increased the number of spiders recovered from field enclosures, as did the presence of leaf litter thatch. That fewer spiders were recovered when thatch and alternative prey were absent suggests that cannibalism was most prevalent under these conditions.
4. Overall, results suggest that habitat complexity reduces agonistic interactions and cannibalism among wolf spiders, providing encouragement to pest managers that the structure of agricultural habitats can be managed to maximise densities of generalist predators for enhanced pest suppression.     

Spatial and temporal variation in a suite of life-history traits in two species of wolf spider

Abstract.  1. Life-history traits and density were assayed in seven populations of two sympatric species of wolf spider for three consecutive years. The goal of the study was to quantify population dynamics and its relation to spatial and temporal life-history variation.
2. Adult female body size and fecundity varied significantly, among field sites and among years, in both species. Female spiders of both species differed in mean relative reproductive effort among sites, but not among years. The size of offspring was invariable, with no significant differences due to site or year.
3. All populations of both species tended to either decrease or increase in density during a given year and this was tightly correlated with changes in prey consumption rates.
4. Since life-history patterns are determined primarily by selection, it is concluded that size at sexual maturity for females is phenotypically plastic and responds to changes in prey availability. Offspring size however is not plastic and it is likely that other selection forces have determined offspring size. Temporal fluctuations in population size are correlated over a large area relative to dispersal capabilities for these species and conservation efforts for invertebrates must take this into consideration.     

Development, growth, and egg production of Ageneotettix deorum (Orthoptera: Acrididae) in response to spider predation risk and elevated resource quality

Abstract.  1. Predation risk to insects is often size- or stage-selective and usually decreases as prey grow. Any factor, such as food quality, that accelerates developmental and growth rates is likely to reduce the period over which prey are susceptible to size-dependent predation.
2. Using field experiments, several hypotheses that assess growth, development, and egg production rates of the rangeland grasshopper Ageneotettix deorum (Scudder) were tested in response to combinations of food quality and predation risk from wolf spiders to investigate performance variation manifested through a behaviourally mediated path affecting food ingestion rates.
3. Grasshoppers with nutritionally superior food completed development ≈ 8–18% faster and grew 15–45% larger in the absence of spiders, in comparison with those subjected to low quality food exposed to spider predators. Growth and development did not differ for grasshoppers feeding on high quality food when predators were present in comparison with lower quality food unimpeded by predators. Responses indicated a compensatory relationship between resource quality and predation risk.
4. Surviving grasshoppers produced fewer eggs compared with individuals not exposed to spiders. Because no differences were found in daily egg production rate regardless of predation treatment, lower egg production was attributed to delayed age of first reproduction. Results compare favourably with responses observed in natural populations.
5. Risk of predation from spiders greatly reduced growth, development, and ultimately egg production. Increased food quality counteracts the impact of predation risk on grasshoppers through compensatory responses, suggesting that bottom-up factors mediate effects of spiders.     

Seasonality of spiders (Araneae) in Mediterranean ecosystems and its implications in the optimum sampling period

Abstract.  1. Fields such as ecology, macroecology, and conservation biology rely on accurate and comparable data. This is especially important for mostly unknown and megadiverse taxa such as spiders and regions such as the Mediterranean. Short-term sampling programmes are increasingly seen as the best option for sampling spiders. Comparability of results, however, demands standard procedures both in methodology and in sampling period. Cost-efficiency dictates that this period should be the most species rich.
2. Pitfall trapping was conducted in 23 sites from north to south Portugal, comprising three large-scale environmental zones and many different habitat types, during 10 months in each site. The annual richness pattern, differences in this pattern between areas and habitats, the complementarity between sampling periods and possible environmental correlates of richness were studied.
3. May and June present the optimal time for collecting spiders in Mediterranean areas. Northern areas have a later peak in richness and dense tree-cover sites offer more flexibility for sampling, with a higher proportion of species present at each period throughout the year.
4. Day length is the environmental factor most correlated with species richness. Maximum daily temperature may reduce richness, especially in southernmost areas, where summer temperatures can be extremely harsh.
5. It is recommended that short-term sampling programmes, intended to give a reasonable picture of spider communities in Portugal and in the Iberian Peninsula (and possibly extending to all the Mediterranean), should be conducted during May or June, with variable flexibility according to area and habitat. The proposed suggestions should appeal to everyone working in the field, given the cost-efficiency and comparability of results by adopting a common standardised approach.     

Kin recognition and cannibalism in a subsocial spider

Evolution of cooperation and group living in spiders from subsocial family groups may be constrained by their cannibalistic nature. A tendency to avoid cannibalizing kin may facilitate tolerance among spiders and implies the ability to identify relatives. We investigated whether the subsocial spider Stegodyphus lineatus discriminates kin by recording cannibalism among juveniles in experiments during which amount of food and size difference among spiders in groups were varied. We hypothesized that family groups should be less cannibalistic than groups of mixed‐parental origin. Further, we tested whether food‐stress would influence cannibalism rates differently in kin and nonkin groups and the effect of relatedness on cannibalism within groups of spiders of variable size compared with those of homogenous size. In groups of six spiders, more spiders were cannibalized in nonsib groups than in sib groups under low food conditions. A tendency for nonkin biased cannibalism in starved spider pairs supported that kin recognition in S. lineatus is expressed when food is limited. Size variance of individuals within well‐fed groups of siblings and unrelated spiders had no influence on cannibalism rates. Apparently, both hunger and high density are important promoters of cannibalism. In addition to inclusive fitness benefits, we suggest that an ability to avoid cannibalizing kin will favour the evolution of cooperation and group living in phylogenetically pre‐adapted solitary species.     

Spider diversity in a tropical habitat gradient in Chiapas, Mexico

This paper presents an assessment of spider diversity in a complex landscape of southern Mexico. Eighteen different habitats were identified, measured and mapped across this fragmented landscape. Habitat types were characterized by measuring various features, including number of plant forms, tree cover and litter depth. Each month from February to April (dry season) and from June to August 2002 (wet season), spiders were sampled on each habitat by using pitfall traps and direct collection. Correlations between spider diversity and habitat characteristics were carried out to explore the relative contribution of each habitat variable as related to changes in spider composition and richness. In total, 115 spider species were recorded in 18 habitat types, and the mean number and density of species per habitat were 21 (± 3, standard error of mean) and 57 (± 9), respectively. The species recorded represent 41% of the fauna recorded in the Mexican state of Chiapas and 4% of the fauna recorded in Mexico. Relatively pristine habitats (e.g. deciduous forest) contained an important proportion of spider diversity in this fragmented landscape. Epigean spider diversity was significantly correlated with tree cover and with the diversity of plant forms during the rainy season. No correlation was detected between soil spider diversity and the habitat variables measured for any season. The results of this work suggest that in highly fragmented tropical landscapes, some habitat types (e.g. coffee plantations, hedgerows) might play an important role for the persistence of spider populations. The prevalence of relatively stable conditions in some of these habitats can allow spiders species (e.g. Nephila clavipes ) to overcome adverse conditions such as a decrease in humidity and dramatic changes in temperature and wind exposure, allowing them to recolonize when favourable conditions return.     

A comparative study of larval cannibalism in three species of ladybird

Abstract. 1. Cannibalism was studied under laboratory conditions in three species of Coccinellidae (Coleoptera): Cycloneda sanguinea , Olla v-nigrum , and Harmonia axyridis . Larval cannibalism varied among species, C. sanguinea  > H. axyridis  ≥  O. v-nigrum .
2. Larvae of all species cannibalised more in response to reduced food availability (14 h starvation daily) than in response to reduced food quality (dry Ephestia eggs).
3. Larvae of H. axyridis cannibalised siblings at lower rates than non-siblings, but larvae of C. sanguinea and O. v-nigrum did not. Rates of cannibalism increased significantly with increasing size disparity among larvae of all three species.
4. Cannibalism in C. sanguinea and H. axyridis , but not in O. v-nigrum , increased with larval density, suggesting that not all attacks on conspecifics are driven by hunger.
5. Costs, not benefits, were the predominant effects of exclusively and partially cannibalistic diets. A diet of conspecific eggs yielded survival equivalent to the Ephestia egg diet for H. axyridis and O. v-nigrum , but developmental time was extended and adults were smaller. Larvae of C. sanguinea had superior survival and faster development on conspecific eggs than on the Ephestia egg diet. Only H. axyridis larvae survived as well on diets comprised exclusively or partially of conspecific larvae as on Ephestia eggs, although they developed more slowly.     

Movement patterns of the bush cricket Platycleis albopunctata in different types of habitat: matrix is not always matrix

Abstract.  1. Inter-patch movement is usually assumed to be homogeneous across a landscape. As the intervening area between suitable patches is usually richly textured, it cannot be assumed to be uniform in real landscapes.
2. In an experimental mark-and-resight study, the movement behaviour of the bush cricket Platycleis albopunctata in four habitat types as well as at the border between two of these habitat types was observed.
3. Analysis of recapture data indicated differences in mortality risk (or emigration rates) between habitat types.
4. When released at the border between suitable habitat and a crop field, P. albopunctata did not show a consistent preference for the suitable habitat. This suggests that the crop field is at least temporarily attractive for P. albopunctata .
5. Movement in suitable habitat was not always different from movement in the matrix, and movement between different types of matrix also differed.
6. The results indicate that the movement behaviour of P. albopunctata is influenced not only by suitability for breeding but also by structural resistance and other factors (e.g. food availability or habitat-specific mortality risk).     

Influence of prey availability and conspecifics on patch quality for a cannibalistic forager: laboratory experiments with the wolf spider Schizocosa

 Because cannibals are potentially both predator and prey, the presence of conspecifics and alternative prey may act together to influence the rate at which cannibals prey upon each other or emigrate from a habitat patch. Wolf spiders (Lycosidae) are cannibalistic-generalist predators that hunt for prey with a sit-and-wait strategy characterized by changes in foraging site. Little information is available on how both prey abundance and the presence of conspecifics influence patch quality for these cursorial, non-web-building spiders. To address this question, laboratory experiments were conducted with spiderlings and older juveniles of the lycosid genus Schizocosa. The presence of insect prey consistently reduced rates of spider emigration when spiders were housed either alone or in groups. Solitary juvenile Schizocosa that had been recently collected from the field exhibited a median giving-up time (GUT) of 10 h in the absence of prey (Collembola); providing Collembola increased the median GUT to 64 h. For solitary spiders, the absence of prey increased by about fourfold the rate of emigration during the first 24 h. In contrast, for spiders in patches with a high density of conspecifics, the absence of prey increased the 24-h emigration rate by only 1.6-fold. For successful cannibals in the no-prey patches, the presence of conspecifics improved patch quality by providing a source of food. Mortality by cannibalism was affected by both prey availability and openness of the patch to net emigration. In patches with no net emigration, the presence of prey reduced rates of cannibalism from 79% to 57%. Spiders in patches open to emigration but not immigration experienced a rate of cannibalism (16%) that was independent of prey availability. The results of these experiments indicate that for a cannibalistic forager such as the wolf spider Schizocosa, (1) the presence of conspecifics can improve average patch quality when prey are absent, and (2) cannibalism has the potential to be a significant mortality factor under natural field conditions because cannibalism persisted in prey patches that were open to emigration. Received: 12 April 1996 / Accepted: 14 August 1996     

Nutritional value of cannibalism and the role of starvation and nutrient imbalance for cannibalistic tendencies in a generalist predator

1. Cannibalism is considered an adaptive foraging strategy for animals of various trophic positions, including carnivores. However, previous studies on wolf spiders have questioned the high nutritional value of cannibalism. We therefore analysed two different aspects of nutritional quality of conspecifics in the wolf spider Pardosaprativaga: their value for survival, growth and development; and the growth efficiency of feeding on conspecifics. We also measured the propensity for cannibalistic attacks and the consumption rate of conspecifics in an experiment where hunger level and nutrient balance were manipulated. In all experiments, cannibalism was compared with predation on fruit flies as control prey. 2. The growth experiment gave ambiguous results regarding the nutritional quality of conspecifics. Spiders on pure cannibalistic diets split into two distinct groups, one performing much better and the other much worse than spiders on fruit fly diets. We discuss the possibility that the population is dimorphic in its cannibalistic propensity, with the latter group of individuals showing a high level of inhibition against cannibalistic attacks in spite of a high nutritional value of cannibalism. 3. The food utilization experiment confirmed the high nutritional quality of conspecifics, as cannibalistic spiders had the same growth rate as spiders fed insect prey in spite of a much lower consumption rate. 4. Inhibition against cannibalistic attacks was demonstrated in medium-sized juveniles: only half of the spiders attacked a prescribed victim of 50% the size of their opponents, and the latency for those that did attack was more than half an hour, compared with a few minutes for spiders fed fruit flies. 5. Nutrient-imbalanced spiders utilized an alternative insect diet less efficiently than balanced spiders, whereas no difference was present in efficiency of utilizing conspecifics. This result indicates that spiders can remedy at least part of a nutrient imbalance through cannibalism. 6. As spiders can escape nutritional imbalance as well as restore energy reserves through cannibalism, we predicted both nutrient imbalance and hunger to stimulate cannibalism. This prediction was confirmed only with respect to hunger. Nutrient-imbalanced spiders had reduced cannibalistic consumption, perhaps due to lowered predatory aggressiveness as a result of bad condition.     

Physiological bases of genetic differences in cannibalism behavior of the confused flour beetle Tribolium confusum

Physiological causes of genetic differences in cannibalism were examined to gain a better understanding of constraints on behavior evolution. Cannibalism has complex population level consequences in Tribolium confusum, including dramatic effects on population size. Laboratory strains with low and high cannibalism rates, obtained through inbreeding, have maintained distinct levels of cannibalism for over two decades even in the absence of artificial selection to maintain the differences. Why strains differ in their cannibalism rates was examined by measuring: (1) the nutritional benefit from cannibalism in both nutritionally good and poor environments, and (2) the possibility that eggs are an important source of water. How strains achieve differences in cannibalism was examined by testing for differences between strains in their ability to find eggs and in their tendency to eat eggs. Beetles from both strains survive equally well in a nutritionally good environment, but they accomplish this in different ways. The low cannibalism strain has high survivorship with and without cannibalism. The high cannibalism strain has low survivorship when not fed eggs and survivorship equivalent to the low cannibalism strain when fed eggs, suggesting it compensates for poor nutritional adaptation by eating eggs. The strains also differ in feeding behavior; beetles from the high cannibalism strain have a higher appetite for eggs. Beetles from the two strains did not differ in locomotor activity, search efficiency, or need for water. The observed behavioral and nutritional differences may contribute to the maintenance of different levels of cannibalism.     

Territoriality mediates atypical size‐symmetric cannibalism in the Amblypygi Phrynus longipes

Cannibalism can have important demographic and ecological effects on populations. Typically, cannibalism is size‐structured, where larger individuals eat smaller conspecifics. Initial cursory observations of the whip spider, Phrynus longipes, however, suggested that cannibalism might not be size‐structured in this species, perhaps because cannibalism is often a by‐product of territory contests. We staged paired interactions and recorded latency to escalate to physical aggression or cannibalize to understand the dynamics of cannibalism and resource contests. We employed a multimodel comparative approach to tease apart the contest characteristics that best predicted cannibalism during behavioral trials between P. longipes opponents. We found that, while armament size symmetry predicted escalation of contests, cannibalism was best predicted by body mass asymmetry. Further, cannibalism was most likely to occur among individuals similar in armament, but dissimilar in body mass. This suggests a discrepancy between phenotypes that may have evolved to communicate resource holding potential (e.g., armaments which benefit individuals if dishonest), and body mass as a cue of resource holding potential.     

The cannibalistic behaviour of two Gammarus species (Crustacea: Amphipoda)

This study examines the cannibalistic behaviour of the freshwater amphipods Gammarus duebeni celticus Stock & Pinkster, 1970 and G. pulex (L., 1758). In the first experiment, interactions were staged among all combinations of single adult males, single adult females, adults in the precopulatory mate-guarding phase and juveniles. Cannibalism by inter-moult individuals on newly moulted conspecifics occurred in all interaction categories in both species. Gammarus d. celticus , however, were significantly more cannibalistic than G. pulex. Cannibalism between and within sex and size categories (males > females > juveniles) was facilitated by the vulnerability of individuals at moult. Individuals of smaller size categories, however, did not cannibalize newly moulted conspecifics of larger size categories. Males were less cannibalistic on newly moulted females than on newly moulted males and juveniles and, when in the precopulatory condition, appeared to defend females from cannibalistic attacks. In a second experiment, stream conditions were simulated in the laboratory and replicated populations monitored for nine weeks. High levels of cannibalism, and the species and sex differences in cannibalism identified in the first experiment, were confirmed under these heterogeneous conditions. Cannibalism by males on their newly moulted female mating partners, termed 'reversed' sexual cannibalism, was further investigated. When males were deprived of foraging opportunities, cannibalism of precopulatory partners was significantly more frequent. The occurrence of 'reversed' sexual cannibalism is thus interpreted as a conflict between motivation to feed and motivation to mate.     

THE GENETICS AND EVOLUTION OF CANNIBALISM IN FLOUR BEETLES (GENUS TRIBOLIUM)

Cannibalism plays a major role in population regulation in Tribolium confusum, accounting for up to tenfold differences in population size between different genetic strains. I characterized the within- and between-strain genetic variation for cannibalism using standard quantitative-genetic methods. The four laboratory strains studied have similar birth and death rates but differ in their strain-specific cannibalistic tendencies. The cannibalism rates of the strains were stable for more than 60 generations of laboratory husbandry. I found considerable genetic variation for cannibalism within each strain. A genetic analysis of the between-strain differences in each of three types of cannibalism (larvae eating eggs, adults eating eggs, and adults eating pupae) showed that all three cannibalism pathways are autosomally inherited and exhibit minor degrees of dominance. Adult cannibalism of eggs and larval cannibalism of eggs appear to be genetically correlated. The differences between the “high” and “low” cannibalism strains appear to be polygenic for two kinds of cannibalism, larvae eating eggs and adults eating pupae. However, strain differences in adult cannibalism of eggs may be due to only two loci. The stability of the between-strain differences for more than 60 generations, the additive nature of inheritance, and the demonstration of considerable within-strain genetic variation suggest that cannibalism may be selectively neutral or under stabilizing selection with many adaptive peaks.     

Habitat preferences of the urban wall spider Oecobius navus (Araneae, Oecobiidae)

Abstract  To assess the relative impact of a range of habitat variables on spider abundance, field and laboratory experiments were conducted on populations of the urban wall spider Oecobius navus in suburban Perth, Western Australia. Habitat characteristics investigated were: substrate type, wind speed, rainfall, sunlight exposure, relative humidity, air temperature, substrate temperature, artificial lighting and prey type/abundance. In the field, O. navus was found to be associated with high humidity, low air temperature and shelter from sunlight and rainfall. Oecobius navus was more abundant at sites with greater prey abundance. The most common prey item was the red meat ant Iridomyrmex chasei . Juvenile spiders were more abundant than adult spiders; however, patterns between spider abundance and habitat variables were similar for both adults and juveniles. Laboratory experiments showed that O. navus preferred to build webs on wooden substrates, and pitted limestone walls. These findings indicate that O. navus may be vulnerable to desiccation and/or thermal stress, and thus survives better on sheltered walls.     

Intra-guild predation relaxes natural enemy impacts on herbivore populations

Abstract. 1. To investigate the role of intra-guild predation in mediating the impact of the natural enemy complex on herbivore populations, a manipulative field experiment was conducted using uncaged plots (islets of Spartina cordgrass) on a North American salt marsh. The densities (moderate or low) of two invertebrate predators, the generalist wolf spider Pardosa littoralis and the specialist mirid bug Tytthus vagus , were manipulated in a 2 × 2 factorial design, and the resulting treatment effects on the population growth of their herbivorous prey, Prokelisia planthoppers, were assessed.
2. The abundance of wolf spiders on experimental islets was unaffected by the presence of mirid bugs, however the density of mirid bugs was influenced very negatively by the presence of the wolf spider.
3. The negative effect of the wolf spider on mirid bugs most probably resulted from the intra-guild predation of mirids by spiders because planthopper limitation by the wolf spider alone was significantly greater than when both predators were present.
4. As a result of intra-guild predation, planthopper population growth was positive in the presence of both predators, despite the fact that each predator alone promoted a decrease in planthopper population growth.
5. Notably, the occurrence of intra-guild predation diminished top-down impacts on planthopper populations in a relatively simple food web where strong top-down effects were expected. This result, however, was limited to habitats on the marsh with simply structured vegetation lacking leaf litter.     

Ontogenetic niche shifts in two populations of juvenile threespine stickleback, Gasterosteus aculeatus, that differ in pelvic spine morphology

Most research on ontogenetic niche shifts has focused on changes in habitat or resource use related to food resource distribution and heterospecific size-limited predation. Cannibalism, an intraspecific interaction, can also affect habitat selection or resource use by vulnerable size classes. Morphological defenses, such as spines, increase the effective size of an individual, making it more difficult to consume. The importance of such defense structures in affecting niche shifts in early life history stages is unclear. Using a combination of field observations and experiments in aquaria and wading pools, we examined the relative roles of cannibalism and morphology in determining juvenile habitat use in two populations of threespine stickleback that differ in pelvic spine morphology. Juveniles were categorized into three size classes: small (5–10 mm), medium (11–15 mm), and large (15–25 mm). In experiments assessing the relative vulnerability of juveniles to cannibalism by adults, we documented a significant difference among size classes in the number of juveniles eaten such that more large juveniles were eaten from the population lacking pelvic spines. The natural distribution of small and large juveniles in two distinct littoral microhabitats, open water and vegetation, was determined in each lake. In both populations, small juveniles were more abundant in vegetation. In the population with pelvic spines, a greater proportion of large juveniles was observed in open water than in vegetation. In the population without pelvic spines, the proportion of large juveniles did not differ between the two habitats. Experiments comparing juvenile habitat use in the presence or absence of adult conspecifics suggest that differences in habitat use may not only depend on the size of the individual, or the size of the individual relative to the size of the adult predator, but also on the degree of development or expression of defensive structures.     

Consequences for a host–parasitoid interaction of host-plant aggregation, isolation, and phenology

Abstract.  1. Spatial habitat structure can influence the likelihood of patch colonisation by dispersing individuals, and this likelihood may differ according to trophic position, potentially leading to a refuge from parasitism for hosts.
2. Whether habitat patch size, isolation, and host-plant heterogeneity differentially affected host and parasitoid abundance, and parasitism rates was tested using a tri-trophic thistle–herbivore–parasitoid system.
3.  Cirsium palustre thistles ( n = 240) were transplanted in 24 blocks replicated in two sites, creating a range of habitat patch sizes at increasing distance from a pre-existing source population. Plant architecture and phenological stage were measured for each plant and the numbers of the herbivore Tephritis conura and parasitoid Pteromalus elevatus recorded.
4. Mean herbivore numbers per plant increased with host-plant density per patch, but parasitoid numbers and parasitism rates were unaffected. Patch distance from the source population did not influence insect abundance or parasitism rates. Parasitoid abundance was positively correlated with host insect number, and parasitism rates were negatively density dependent. Host-plant phenological stage was positively correlated with herbivore and parasitoid abundance, and parasitism rates at both patch and host-plant scales.
5. The differential response between herbivore and parasitoid to host-plant density did not lead to a spatial refuge but may have contributed to the observed parasitism rates being negatively density dependent. Heterogeneity in patch quality, mediated by variation in host-plant phenology, was more important than spatial habitat structure for both the herbivore and parasitoid populations, and for parasitism rates.     

The impact of altered precipitation on spatial stratification and activity-densities of springtails (Collembola) and spiders (Araneae)

Abstract.  1. A field experiment was conducted to determine how short-term changes in moisture can alter activity-densities of spiders and springtails.
2. In a Kentucky forest 10 unfenced 4-m² plots were divided into two rainfall treatments. A clear roof over five plots excluded rainfall to simulate severe drought conditions ( drought treatment). Water was sprayed on the five uncovered plots at a rate equal to two times the long-term mean in order to establish the high-rainfall treatment. Activity-densities of Collembola and spiders were measured using pitfall traps designed to sample the top, middle, and bottom layers of leaf litter. The experiment ran from 20 July to 23 September 2001.
3. Overall (i.e. litter layers pooled) activity-density (mean number trapped each sampling date) of Collembola was ≈ 60% lower in drought plots than in plots receiving increased precipitation. Surprisingly, overall spider activity-density was ≈ 1.6 times greater in the drought plots.
4. Differences in rainfall affected the spatial stratification of Collembola and spiders in strikingly different ways. Activity-densities of neither group differed between drought and high-rainfall treatments in the bottom litter layer. Collembola activity-density was three times greater in the top and middle litter layers in high-rainfall plots than in drought plots. In contrast, spider activity-density did not differ between treatments in the top layer, but activity-density was decreased by 50% in the middle layer of high-rainfall plots compared with drought plots.
5. Three Collembola families (Sminthuridae, Tomoceridae, and Entomobryidae) accounted for most of the Collembola pattern. The spider response was due to altered activity-density of one family of wandering spider, the Gnaphosidae.     

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.