Competition for a limited space in kleptoparasitic Argyrodes spiders revealed by field experiments

Most kleptoparasitic Argyrodes spiders rely exclusively on host spider webs for obtaining their food. Because their densities occasionally reach high levels within a restricted area, competitive interactions may be important for determining the density of these unique spiders. Here I used two Argyrodes species commonly found on webs of the large orb-web spider Nephila clavata to clarify whether inter- and intraspecific competition influences abundance and within-web distribution by using observational data and field experiment. Removing Argyrodes flavescens from the host webs induced a remarkably high immigration of that species while density on control webs remained almost at the same level, which is evidence for strong intraspecific competition. Larger individuals of A. flavescens were located more frequently at the capture area of the host webs where it is easy to access prey ignored by the host spider, and spiders immigrating into webs from which that species had been removed were smaller in body size, suggesting interference competition for space among conspecific kleptoparasites. Argyrodes bonadea increased in number on webs from which A. flavescens had been removed, and the increase was correlated with the number of A. flavescens removed. This finding is evidence for interspecific competition that is rarely reported in spiders. A multiple regression model including numbers of a conspecific parasite as well as web and body sizes of the host spider could not detect competitive interactions between species, suggesting the importance of experimental approaches. Received: May 22, 2000 / Accepted: December 1, 2000     

Evidence of exploitative competition among young stages of the wolf spider Schizocosa ocreata

Summary Previous research by many investigators has demonstrated food limitation in both web-building and wandering spiders. Field experiments have tested for exploitative competition for prey in web-building, but not wandering species. As a first step to examining the question of whether spiders without webs exhibit exploitative competition, we manipulated densities of young stages of a common wolf spider, Schizocos ocreata, and measured (1) spider growth rate and (2) numbers of Collembola, a potential prey organism. Replicate populations of recently hatched S. ocreata were established in 1-m² fenced plots at four levels: 0×, 0.25×, 1× and 4× natural density. Increasing spider density had a negative effect on spider growth rate, defined as increase in weight or cephalothorax width. Early in the experiment spider density had a weak negative effect on Collembola numbers [p(F)=0.08]. Taken together, this probable response by Collembola and the clear effect of spider density on growth rate constitute the first experimental evidence of intraspecific exploitative competition for prey in a species of wandering spider. We discuss (1) the strength of this evidence given the constraints of the experiment's design, and (2) the implications of the strong convergence in spider densities that had occurred after 2.5 months.     

Is different degree of individual specialization in three spider species caused by distinct selection pressures?

Individual specialization (IS), i.e. the relatively narrower niche of individuals compared to their population, can be caused by the interplay between intrinsic and extrinsic factors. We studied IS on prey type and size in three congeneric euryphagous sympatrically occurring species: Philodromus albidus, Philodromus aureolus, and Philodromus cespitum. We performed prey acceptance experiments in standardized laboratory conditions using adult females. We also studied possible factors affecting IS. As intrinsic factors, we measured aggressiveness toward prey and spider size as the measures of behavioral and morphological differences, respectively. As extrinsic factors, we studied the composition of the guild of actively hunting spiders in the area from where the philodromids were collected. We computed the degree of inter- and intraspecific crowding as a measure of inter- and intraspecific competition, respectively. Then we computed competition coefficients from census data in order to estimate the strength of interspecific competition among three species. There was no IS on prey type in any studied species. In terms of prey size, only P. albidus was found to be individually specialized, while there was no IS in the other two species. There was a significant effect of aggressiveness, but no significant effect of body size on IS in P. albidus. This species was the least aggressive of the studied species. The philodromids were the dominant actively hunting spiders in the study area. P. albidus was exposed to a relatively higher degree of interspecific than intraspecific competition, whereas in the other philodromids the situation was reversed. The presence of interspecific competition between P. albidus and the other two species was confirmed by the census data analysis. We documented context-dependent IS in Philodromus species. We argue that the higher degree of IS in P. albidus was influenced by interference competition mainly from the other two philodromid species. The niches of the other two species were influenced by food limitation, which decreased their IS.     

Contrasting effects of habitat fragmentation,population density,and prey availability on body condition of two orb‐weaving spiders

1. Habitat fragmentation is a major threat to biodiversity because it disrupts movement between habitat patches. In addition, arthropod fitness may be reduced in fragmented habitats, e.g. due to reduced prey availability. 2. We studied the relationship of spider body condition with habitat fragmentation, population density, and prey availability. We expected that prey availability and population density of spiders would be affected by landscape composition and patch isolation. Body condition should be enhanced by high prey availability, but negatively affected by population density due to competition. 3. We sampled spiders on 30 groups of cherry trees that varied independently in the level of isolation from other woody habitats and in the percentage of woody habitat within 500 m radius. As a measure of body condition, we used residuals of the relationship between individual body mass/opisthosoma width and prosoma width of the two most common orb‐weaving spider species, Nuctenea umbratica Clerck and Araniella opisthographa Kulczynski. 4. Body condition of A. opisthographa was positively correlated with the abundance of flies, which increased with the percentage of forest in the landscape. In contrast, body condition of N. umbratica was reduced at high population densities, presumably due to intraspecific competition. In addition, body condition and population density of A. opisthographa was lower at isolated sites. 5. Our study suggests that effects of landscape fragmentation on body condition vary strongly between spider species, depending on the relative role of food limitation and intraspecific competition.     

Resource partitioning and competition for shells between intertidal hermit crabs on the outer coast of Washington

Summary Resource partitioning was quantified for 6 species of intertidal hermit crabs in the genus Pagurus, that occur on the outer coast of Washington. This, together with field evidence of shell shortage and with laboratory experiments to quantify the mechanism of interactions for shells, allowed estimation of the relative intensities of inter-and intraspecific competition between these species. The findings were that: (1) the magnitude of intraspecific competition was greater than any single interspecific competitive effect for all of the species; and (2) the relative proportion of intraspecific competition was greater for the middle and upper intertidal species than for the lower intertidal species. Studies at several outer coast sites supported these generalizations. Both of these findings are consistent with the hypothesis that competitive divergence has occurred in the past. The structure of the outer coast hermit crab assemblage is compared with that of the San Juan Archipelago hermit crab assemblage. Differences between the two do not seem to be the result of adaptive responses to the presence of more competing species in the former group.     

荒漠草原三种蝗虫成虫种内和种间竞争的研究

荒漠草原三种蝗虫成虫种内和种间竞争的研究贺达汉方成郑哲民（宁夏农学院农学系,永宁７５０１０５）（陕西师范大学动物研究所,西安７１００６２）Ｉｎｔｒａ＿ａｎｄＩｎｔｅｒｓｐｅｃｉｆｉｃＣｏｍｐｅｔｉｔｉｏｎａｍｏｎｇｔｈｅＡｄｕｌｔｓｏｆＴｈｒｅｅＧｒ...     

Combined versus Single Species Release of Predaceous Mites: Predator–Predator Interactions and Pest Suppression

Interactions such as competition, intraguild predation (IGP), and cannibalism affect the development and coexistence of predator populations and can have significance for biological control of commonly exploited pest organisms. We studied the consequences of combined versus single release of two predaceous mite species (Phytoseiidae), with differing degrees of diet specialization, on their population dynamics and the suppression of the carmine spider mite, Tetranychus cinnabarinus Boisduval (Tetranychidae), on greenhouse-grown gerbera. Population growth of the specialist predator Phytoseiulus persimilis Athias-Henriot was greater and population decline steeper when released in combination with the generalist Neoseiulus californicus McGregor than when released alone. In contrast, the N. californicus population grew and declined more gradually when released in combination with P. persimilis, compared to the single species release. The differential impact on each other's population dynamics can be primarily attributed to contrasting properties in competition, IGP, and cannibalism. At the same overall predator density and as long as prey was abundant, the specialist P. persimilis was more strongly affected by intraspecific competition than by interspecific competition with the generalist N. californicus. In contrast, interspecific competition with P. persimilis had a greater impact on N. californicus than intraspecific competition. After prey depletion, the generalist predator N. californicus was more likely to engage in IGP than was the specialist predator P. persimilis. Overall, the study demonstrates that prey specificity has significance for the quality and intensity of predator–predator interactions and indicates potential implications for biological control of spider mites. All predator releases (i.e., either species alone and both species in combination) resulted in reduction of the spider mite population to zero density. Individual release of the specialist P. persimilis led to the most rapid spider mite suppression. Nonetheless, in perennial greenhouse-grown crops P. persimilis and N. californicus could have complementary effects and a combination of the two predators could enhance long-term biological control of spider mites. The potential risks and benefits associated with the release of both species are discussed.     

Effect of arbuscular mycorrhiza on inter- and intraspecific competition of two grassland species

We were interested in the role of arbuscular mycorrhiza (AM) in the competition between plants of different sizes. A pot experiment of factorial design was established, in which AM root colonization and competition were used as treatments. Five-week-old Prunella vulgaris seedlings were chosen as target plants (i.e. plants whose response to competition was studied) and the following (13 replicates of each) were used as neighbours: (1) a large, 10-week-old P. vulgaris, (2) two P. vulgaris seedlings, and (3) a large, 10-week-old Fragaria vesca. In the experiment where small neighbours were grown together with small target plants, competition did not reduce target plant weight significantly, compared to the other two treatments. The competitive effects of large neighbours were significant, regardless of species (both older neighbours reduced the weights of target plants similarly), but there was a clear difference between intra- and interspecific competition when plants were mycorrhizal. In intraspecific competition with a large neighbour, the target plant shoot weight was reduced 24% when inoculated with AM. Thus, AM amplified rather than balanced intraspecific competition. In interspecific competition with old F. vesca, the shoot weights of target plants were 22% greater when inoculated with AM than when non-mycorrhizal. The results showed that, for given soil condition, AM might increase species diversity by increasing competitive intraspecific suppression and decreasing the interspecific suppression of small plants by larger neighbours.     

Seasonal reversal of competitive advantage between two spider species

Summary To test for inter- and intraspecific competition between two common orb-weaving spiders, Metepeira grinnelli and Cyclosa turbinata, I conducted separate field experiments in the spring and summer. During the spring experiment, the prey-consumption rate of Metepeira was reduced at higher inter- and intraspecific densities, but Cyclosa was only affected by intraspecific density. During the summer experiment, Metepeira prey-consumption was not affected by inter- or intraspecific densities, while Cyclosa was affected by both inter- and intraspecific densities. In addition to competition for food, during the spring experiment Cyclosa was observed displacing Metepeira but not vice versa, whereas during the summer experiment Metepeira displaced Cyclosa but not vice versa. The seasonal reversal in the competitive advantage between the species was linked to their asynchronous phenologies; in the spring most Cyclosa were larger than Metepeira, whereas in the summer most Metepeira were larger than Cyclosa. I suggest that in this system, temporal variability in the competitive abilities may promote species coexistence.     

Food limitation of population density in the orb-wed spider,Nephila clavata

Field studies were conducted to clarify whether variation in food availability among habitats influences population density, and whether population density has a negative effect on foraging success in the orb-web spider, Nephila clavata. Lifetime food consumption per individual (i.e., foraging success) strongly correlated with mean body size of adult females and mean fecundity in populations. Also, there was a positive correlation between foraging success and population density. Since foraging success reflected potential prey availability in the habitat, food resource appeared to be a limiting factor for populations in this spider. Mean fecundity per individual correlated with population density of the following year, suggesting that decreased reproduction is a major component of food limitation on population density. Consistent defferences in mean body size between particular sites were observed over years, while such difference was less obvious in density. Thus, ranking of food abundance among habitats seems to be predictable between years. A field experiment revealed that an artificial increase in population density had no negative effect on the feeding rate of individuals, suggesting that intraspecific competition for food is not important in this species.     

The effect of colour variation in predators on the behaviour of pollinators: Australian crab spiders and native bees

1. Australian crab spiders exploit the plant–pollinator mutualism by reflecting UV light that attracts pollinators to the flowers where they sit. However, spider UV reflection seems to vary broadly within and between individuals and species, and we are still lacking any comparative studies of prey and/or predator behaviour towards spider colour variation. 2. Here we looked at the natural variation in the coloration of two species of Australian crab spiders, Thomisus spectabilis and Diaea evanida, collected from the field. Furthermore, we examined how two species of native bees responded to variation in colour contrast generated by spiders sitting in flowers compared with vacant flowers. We used data from a bee choice experiment with D. evanida spiders and Trigona carbonaria bees and also published data on T. spectabilis spiders and Austroplebeia australis bees. 3. In the field both spider species were always achromatically (from a distance) undetectable but chromatically (at closer range) detectable for bees. Experimentally, we showed species‐specific differences in bee behaviour towards particular spider colour variation: T. carbonaria bees did not show any preference for any colour contrasts generated by D. evanida spiders but A. australis bees were more likely to reject flowers with more contrasting T. spectabilis spiders. 4. Our study suggests that some of the spider colour variation that we encounter in the field may be partly explained by the spider's ability to adjust the reflectance properties of its colour relative to the behaviour of the species of prey available.     

Role of interspecific competition in the coexistence ofApodemus argenteus andA. speciosus (Rodentia: Muridae)

Reciprocal-removal experiments with two replicates were conducted to test for the role of interspecific competition in the coexistence ofApodemus argenteus andA. speciosus. Population density, rate of appearance of new (unmarked) individuals, reproduction, survival rate and habitat use were monitored during pre- and removal periods. In both removal experiments, the removal ofA. argenteus had little effect onA. speciosus, while that ofA. speciosus affected several population characteristics ofA. argenteus. Namely, the removal ofA. speciosus shifted the distribution ofA. argenteus to the habitat with a denser shrub cover in one experiment. Also, the removal increased the population densities and appearance rates of new individuals ofA. argenteus in another experiment. Interspecific interactions between the two species appeared to be a one-way action fromA. speciosus toA. argenteus. In removal periods in both experiments, the rates of appearance of new individuals in each species were the highest on the grid where that species was removed. These results suggest that, though interspecific competition occurred between the two species, intraspecific competition had greater effects than interspecific competition on the abundance and habitat use ofA. argenteus andA. speciosus. This implies that the fundamental niches ofA. argenteus andA. speciosus differ potentially, which may play an important role in the coexistence of the two species.     

Interactions amongst species in a guild of subtidal benthic herbivores

Summary The subtidal coralline flats of northeastern New Zealand support a characteristic guild of grazing herbivores. The most important members of this guild are an echinometrid echinoid, patellid, turbinid and trochid gastropods. Densities of these herbivores fluctuate through time. Interactions within and among the different species of echinoids and gastropods were investigated experimentally. Different combinations of species were caged at densities up to 5 times that of ambient for a 24 week period in an experiment designed to differentiate between intra- and interspecific competition.The echinoidEvechinus chloroticus and the turbinid gastropodCookia sulcata exhibited reduced mean dry weight with increasing intraspecific densities. There was little evidence of density-related mortality in these species. The limpetCellana stellifera showed comparatively large losses of weight and enhanced mortalities in intraspecific experimental treatments but this was not related to density.Investigation of interspecific interactions amongstEvechinus andCookia revealed no evidence of a negative influence of one species on the other. In terms of dry weight,Cookia was indifferent to the presence ofEvechinus, andEvechinus benefited in the reciprocal interaction.Cookia also enjoyed an enhanced mean dry weight when in the presence ofCellana compared to the equivalent intraspecific treatments. There were no coherent trends in proportional mortality in any treatments with enhanced interspecific densities. Cellana, in the presence ofCookia, exhibited a dramatic decrease in mortality rate and increase in mean dry weight. The presence of the turbinid gastropod was clearly beneficial to the limpet when compared to the intraspecific treatments with enhanced intraspecific densities and the control cages containingCellana at ambient density. We suggest that subtidal areas constitute poor habitats for limpets in the absence of agents such asCookia which may provide or maintain suitable sites for attachment and grazing.For the combinations of densities and species investigated there was a consistent trend towards positive interspecific interactions. It seems unlikely that at the sites investigated interspecific competition could act to restrict distributions, or limit abundances of species.     

Floral density and co-occurring congeners alter patterns of selection in annual plant communities*

Although the evolution and diversification of flowers is often attributed to pollinator-mediated selection, interactions between co-occurring plant species can alter patterns of selection mediated by pollinators and other agents. The extent to which both floral density and congeneric species richness affect patterns of net and pollinator-mediated selection on multiple co-occurring species in a community is unknown and is likely to depend on whether co-occurring plants experience competition or facilitation for reproduction. We conducted an observational study of selection on four species of Clarkia (Onagraceae) and tested for pollinator-mediated selection on two Clarkia species in communities differing in congeneric species richness and local floral density. When selection varied with community context, selection was generally stronger in communities with fewer species, where local conspecific floral density was higher, and where local heterospecific floral density was lower. These patterns suggest that intraspecific competition at high densities and interspecific competition at low densities may affect the evolution of floral traits. However, selection on floral traits was not pollinator mediated in Clarkia cylindrica or Clarkia xantiana, despite variation in pollinator visitation and the extent of pollen limitation across communities for C. cylindrica. As such, interactions between co-occurring species may alter patterns of selection mediated by abiotic agents of selection.     

COMPETITION AND THE EVOLUTION OF AGGRESSIVE BEHAVIOR IN TWO SPECIES OF TERRESTRIAL SALAMANDERS

The effects of competition on the evolution of interspecific interference mechanisms were studied by comparing the aggressive behavior of two terrestrial salamander species from two localities that differ in the intensity of interspecific competition. Plethodon jordani and P. glutinosus are closely related, ecologically similar species that are sympatric at intermediate elevations in the southern Appalachian Mountains. Previous removal and transplant experiments showed that interspecific competition is more intense in the northeastern Great Smoky Mountains, where the species are narrowly sympatric, than in the nearby Balsam Mountains, where sympatry is broader. In laboratory encounters, P. glutinosus from the Great Smoky Mountains were more aggressive to heterospecific and conspecific intruders than were P. glutinosus from the Balsam Mountains. For P. jordani, however, the variation in interspecific and intraspecific aggressive behavior among individuals within populations was as great as the variation between populations. Alpha-selection (i.e., improved competitive ability by the acquisition of interspecific interference mechanisms) has occurred in populations of P. glutinosus under conditions of intense interspecific competition. The evolution of aggressive behavior appears to have been influenced by the intensity of intraspecific competition as well.     

Interspecific competition among urban cockroach species

The aim of this study was to quantify and to compare the effects of intraspecific and interspecific competition among pairs of urban cockroaches [Blatta orientalis L., Periplaneta americana (L.), and Periplaneta australasiae Fabricius (Dictyoptera: Blattodea)] in relation to the limitation of resources such as shelter or food. Our approach was to assess whether the presence of one species affected the resource exploitation of another. A reduction in access to shelters or to food for one species revealed dominant/subordination relationships among species and induced spatial segregation in shelters and temporal segregation during food exploitation. The fragmentation of available resources facilitated spatial segregation and the access of more individuals of the subordinate species. Individuals of each species were aware of the presence of non‐conspecifics. The outcome of interspecific competition under laboratory conditions should help us to understand how segregated spatial distributions occur under natural conditions in urban areas.     

Temperature modulates the effects of predation and competition on mosquito larvae

1. Predation risk affects interspecific competition by decreasing foraging activity and relative competitive ability. Predation risk is determined by predators' prey choice and prey responses, both of which can be influenced by temperature. Temperature is especially important for larval prey and can result in a trade‐off between predator‐induced decreases in foraging activity and growth. Interspecific competition must also be examined in relation to intraspecific density‐dependent competition; weaker interspecific competition leads to coexistence of competitors. 2. This study explored how temperature (15 and 25 °C) could affect a focal species, larvae of the mosquito Culex quinquefasciatus, by examining prey choice in a shared predator (mosquitofish; Gambusia holbrooki) and the effects of predation risk on interspecific competition with Limnodynastes peronii tadpoles. Intraspecific density‐dependent competition in C. quinquefasciatus at these temperatures was also examined. 3. At 25 °C, G. holbrooki consumption of both C. quinquefasciatus and L. peronii increased; however, the effects of interspecific competition on mosquito survival did not decrease with L. peronii exposure to predation risk. The relationship between intraspecific density‐dependent competition and interspecific competition was temperature‐dependent, with competitive dominance of L. peronii at 25 °C. Male and female mosquitoes had different temperature‐dependent responses, indicating sex‐specific intrinsic responses to starvation and differential selection pressures. At 25 °C, females were susceptible to interspecific competition by L. peronii, while males were susceptible to intraspecific competition. 4. The use of competitors as biological controls has implications for mosquito disease transmission, and these results suggest that control effectiveness may be modified by climate change.     

Competition among the adults of three grasshoppers (Orthop., Acrididae) on an alpine grassland

Abstract:  The intraspecific and interspecific competition among the adults of three common grasshoppers, Angaracris rhodopa (Fischer-Waldheim), Chorthippus dubius (Zubovsky) and Chorthippus fallax (Zubovsky) was investigated on Ganjia High Mountain Grassland in north-west China. The results indicated a strong intraspecific competition within species at high densities. The interspecific competition between the larger-sized species, A. rhodopa , and the smaller-sized species, C. dubius or C. fallax , was weak, while competition between C. dubius and C. fallax was strong. An asymmetric competitive interaction in mortality and fecundity associated with the diet and size was observed among the three species. Competition was strong between the two species that had a similar diet. The large-sized species had higher mortality rates and lower fecundity than the smaller-sized species. Intraspecific competition between the three species was mainly affected by food limitation, while interspecific competition was mainly associated with food limitation as well as the grasshoppers' adaptation to the environment.     

Survival of transplanted nests of the red wood ant Formica aquilonia （Hymenoptera： Formicidae）： The effects of intraspecific competition and forest clear-cutting

The fitness and survival of ant colonies depend on the resources near their nests. These resources may be limited due to poor habitat quality or by intra- and interspecific competitions, which in extreme cases may cause the ant colony to perish. We tested the effect of intraspecific competition and habitat degradation （forest clear-cutting） on colony survival by transplanting 26 nests of the red wood ant （Formica aquilonia Yarrow, 1955） in 26 different forest areas that contained 0-11 conspecific alien nests per hectare. F. aquilonia is highly dependent on canopy-dwelling aphids, thus the removal of trees should cause food limitation. During the course of the 4-year experiment, 9 of the forests were partially clear- cut. We found that while forest clear-cutting significantly decreased the colonies＇ survival, intraspecific competition did not. As a highly polygynous and polydomous species, E aquilonia seems to tolerate the presence of alien conspecific colonies to a certain extent.     

Clubiona spider (Araneae: Clubionidae) visiting flowers of nectariferous orchid Neottianthe cucullata

Spiders are usually thought to obtain their energy for growth, maintenance and reproduction from captured prey. However, not all spiders should be assumed to receive all of their energy from prey. Accumulated studies have documented nectarivory in a wide variety of hunting spiders. We report a Clubiona spider visiting flowers of the nectariferous orchid Neottianthe cucullata as an example of spider nectarivory. In addition, during the visitations by Clubiona spider, pollinia removal was observed. The impact of the Clubiona spider on the N. cucullata reproductive success is also discussed.