Jumping spiders attend to context during learned avoidance of aposematic prey

A large number of studies on both animals and humans have demonstratedthat learning is influenced by context or secondary cues thatare present when an association is formed. Few studies, however,have examined the functional value of attending to context.We first demonstrated that jumping spiders, Phidippus princeps,could be trained to avoid aposematic, distasteful milkweed bugs,Oncopeltus fasciatus. Spiders readily attacked bugs on firstexposure but were significantly less likely to do so after eighttrials, although they subsequently attacked and ate crickets.Spiders exposed to nontoxic milkweed bugs reared on sunflowerseeds did not show the same decline in attack rate. We nextexamined the effects of secondary contextual cues on spiderlearning by training spiders to avoid milkweed bugs in one oftwo environments. When spiders were tested in an environmentdifferent from the one in which they were trained, attack ratesincreased, and spiders no longer demonstrated retention of theassociation. Spiders tested in the same environment in whichthey were trained continued to avoid attacking the bugs. Theseresults have potential consequences for the evolution of bothpredator and prey and point to the importance of studying context-dependentlearning.     

Contrasting patch choice behaviour by immature ambush predators, a spider (Misumena vatia) and an insect (Phymata americana)

1. Spiderlings of the crab spider Misumena vatia and nymphs of the ambush bug Phymata americana normally ambush prey in flowers, as do the adults. The immatures of M. vatia are more mobile, by ballooning, than the ambulatory but largely sedentary adult female spiders, but the apterous immatures of P. americana are less mobile than the alate adults. The aim of the work reported here was to investigate how immatures, as compared with adults, select sites from which to ambush their prey. 2. Individual immatures of both species were released in experimental arenas with variously manipulated natural stems: normal, with and without leaves, and with and without flowers of three species of plants. 3. The spiderlings and nymphs showed discriminatory behaviour but not in accordance with optimal foraging in either case. The spiderlings chose more leafy stems than any other kind of stem. For the nymphs, the importance of leaves was less marked, food apparently being more important than shelter. 4. Neither of the predators discriminated among flowering stems of Solidago canadensis, Daucus carota, and Cirsium arvense. 5. In other experiments, in which flowered and deflowered stems were located in two separated groups in the arena and the individual was released in a bare area between the two groups, the spiderlings chose either group as predicted by chance but the nymphs showed a slight preference for the flowered sector. Nevertheless, once in either of the groups, the predators showed similar behaviour in choosing leafy stems more often than other stems. 6. Nymphs of P. americana are much more selective than adults but spiderlings of M. vatia are less selective than adult females. These results appear to accord with the time and energy costs of changing hunting sites, walking being slower than flying or ballooning. 7. The results indicate the need to include other parameters, such as enemy‐free space and time spent in not being able to forage, in the analysis of foraging behaviour.     

Presampling sensory information and prey density assessment by wolf spiders (Araneae, Lycosidae)

Decisions regarding foraging patch residence time and the assessmentof patch quality may be mediated by various sources of information.This study examined the use of sensory cues by hunting spidersto assess prey density in the absence of prey capture. Adultfemale wolf spiders [Schizocosa ocreata (Hentz); Lycosidae]had food withheld for 4 days and then were exposed to artificialforaging patches containing four densities of crickets (0, 3,10, 20) with different sensory stimuli (visual and vibratoryinformation, visual only, and vibratory only). The spiders werenot allowed to feed during trials, and patch residence timewas recorded. The spiders varied patch residence time basedon sensory cues alone and spent more time in patches with higherprey density. With visual information only, spiders could apparentlydistinguish among prey densities almost as well as with visualand vibratory cues combined, but residence time did not differamong prey densities when only vibratory information was presented.Measurements of vibration levels produced by cricket activityunder experimental sensory treatments conform to test results,suggesting that visual detection of crickets is important inpatch assessment used in determining patch residence time.     

Intraspecific variation in responses to aposematic prey in a jumping spider (Phidippus regius)

Aposematic signals often allow chemically defended prey to avoid attack from generalist predators, including jumping spiders. However, not all individual predators in a population behave in the same way. Here, in laboratory trials, we document that most individual Phidippus regius jumping spiders attack and reject chemically defended milkweed bugs (Oncopeltus fasciatus), immediately releasing them unharmed. However, a small number of individuals within the population kill and completely consume these presumably toxic prey items. This phenomenon was infrequent with only 14% of our sample (17/122) consuming the milkweed bugs over the course of the study. Individuals that killed and consumed bugs often did so repeatedly; specifically, individuals that consumed a bug in their first test were more likely to kill a bug in their second test and also tended to consume them again. We explored what might drive some (but not all) individuals to consume these bugs and found that neither sex, sexual maturity, body size, laboratory housing type, nor being wild-caught or being laboratory-reared, predicted milkweed bug consumption. Consuming bugs had no negative effects on spider mass or body condition; contrary to expectations, individuals that consumed milkweed bugs actually gained more body mass and increased in body condition. We discuss potential behavioural and physiological variation between individuals that may drive these rare behaviours and the implications for the evolution of prey defences.     

Choice of hunting site as a consequence of experience in late-instar crab spiders

Earlier experiences may play an important role in the choice of hunting sites, but their effects on the foraging repertoire of most animals remain poorly understood. I tested the role of previous flower choices (hunting sites) by penultimate-instar female crab spiders Misumena vatia in making subsequent patch-choice decisions. M. vatia is a sit-and-wait predator, and the two flower species used, ox-eye daisy Chrysanthemum leucanthemum and common buttercup Ranunculus acris, are important hunting sites. Spiders with different immediate experience showed similar short-term (<1 day) giving-up times on the two flower species, independent of their previous substrate. However, four-fifths of the individuals that remained a day or longer tended to leave buttercups sooner than daisies, especially if they had previously occupied daisies. Thus they may directly assess the quality of a potential hunting site, perhaps in response to prey abundance, but previous experience may play a minor role as well. Of spiders that made several consecutive choices of hunting sites, those on daisies often confined these runs to daisies (one of two years); those on buttercups did not exhibit comparable fidelity. Spiders molting into the adult stage almost always subsequently chose the same flower species (either daisy or buttercup) as the one on which they molted. Thus, juvenile experiences may influence adults, the critical stage when virtually all of the spiders' reproductive resources are gathered, even if this resulted from imprinting on their molt sites rather than carrying information over the molt. Received: 26 December 1998 / Accepted: 21 April 1999     

Risk Sensitivity and the Paradox of Colonial Web-Building in Spiders

Group foraging is rare in spiders, occurring only where preyavailability is high. If colonial web-building increases individualprey capture rates as shown, why does group foraging not occurmore often where prey are scarce? Risk sensitivity may explainthis paradox, as variance in prey capture is reduced in groups;risk-averse spiders should join groups only when prey exceeda threshold level. Field studies show that group foraging variesas predicted between species, between populations of a singlespecies, and between sites within a population. However, recentmodels suggest the necessity of examining variance within individualsover time rather than between individuals within populations.Additionally, mechanisms responsible for variance reductionin colonial webs may be less effective than previously assumed.New field data suggest that while prey variance over time maybe somewhat less for individual spiders in groups than for solitaries,the relationship between colonial web-building and variancein prey capture is far more complex than originally thought.The influence of risk sensitivity on reproductive success andthe evolution of colonial web-building is discussed.     

Flower choice by naïve young crab spiders and the effect of subsequent experience

Initial responses of na?ve individuals to critical environmental stimuli provide important information about the innate contribution to behaviour, and subsequent responses to the same stimuli may show the role of experience in mediating those initial responses. To test the role of these factors, I measured initial patch choices and giving-up responses of just-emerged, na?ve, second-instar crab spiders, Misumena vatia, on several hunting sites they encountered after leaving their natal nests. In follow-up tests I measured the effects of these experiences on subsequent patch choice decisions. The choice of hunting sites is a vital decision at all stages of the life cycle for sit-and-wait predators such as Misumena. In their initial tests these spiderlings remained more frequently on goldenrod (Solidago spp.) flowers than on green or yellow goldenrod buds, a preference they retained through tests run on 5 consecutive days. Individuals on green and yellow buds shifted sites more quickly and frequently than those from flowers, and made most of these moves to flowers, which attracted many more prey than did buds. These differences were not affected by age, energetic condition, or loss of information over the period of the experiment. Once spiderlings moved from buds, they showed a high, increasing tendency to move from buds in subsequent runs, those from flowers showed a consistently low tendency. These results suggest that spiderlings retain their innate behavioural patterns through the second instar, but that experience also plays a modest role in patch choice at this stage. Copyright 2000 The Association for the Study of Animal Behaviour.     

Do plant cues influence the oviposition behavior of Japanese beetles?

A bioassay arena was developed to test the response of the Japanese beetle, Popillia japonica Newman (Coleoptera: Scarabaeidae), to various plant‐derived factors that may influence the selection of oviposition sites. In choice experiments with natural and artificial plant cues, females preferred to lay eggs in ryegrass rather than in artificial resources. Artificial grass stems received more eggs on average than bare ground, indicating the role of vertical objects on the soil surface as oviposition cues. Popillia japonica females were able to discriminate between the quality and quantity of the artificial stems, as more eggs were laid in soil blocks that had larger diameter stems or a higher number of stems per soil block. Beetles were capable of evaluating plant characteristics on the soil surface and adjusted their investment accordingly; more eggs were laid in resources that were chosen more frequently. Observations of females in choice arenas revealed how surface cues influence digging behavior and the time spent under the soil surface. The number of digging events was not significantly different among the three treatments with surface cues (ryegrass, grass roots, and artificial grass), but beetles spent significantly more time in the soil under ryegrass than any of the other treatments. These findings suggest that oviposition by P. japonica is selective and is influenced by plant‐derived cues evaluated before and after digging into the soil.     

Uganda kob prefer high-visibility leks and territories

In lekking species, where males provide estrous females withlittle more than sperm, it has been widely supposed that theonly possible benefits to females of mate choice are genetic.We studied female choice of leks and territories in a reduncineantelope, the Uganda kob (Kobus kob thomasi), and found thatfemales consistently preferred high-visibility mating sites.Leks were elevated and had shorter grass and fewer thicketsthan the surrounding areas. Changes in the number of male andfemale kob on 10 leks were correlated with changes in surroundinggrass height, and both females and males preferred leks withexperimentally reduced grass height over neighboring controls.Within a lek, territory popularity was the primary determinantof male daily mating success, and females preferred territoriesrelatively far from thickets, but removal of thickets did notaffect female territory preferences. Because lion hunting successon kob increases with grass height and thicket density, femalesmay benefit directly from these preferences by reducing therisk of predation.     

Fitness costs and benefits of antipredator behavior mediated by chemotactile cues in the wolf spider Pardosa milvina (Araneae: Lycosidae)

Animals may exhibit a variety of defensive behaviors in thepresence of indirect predator cues. Such behavior offers immediatefitness benefits but may also incur substantial foraging andreproductive costs. We measured shifts in space use (verticalclimbing) by the wolf spider Pardosa milvina induced by chemotactilecues (silk and excreta) from a co-occurring predatory wolfspider Hogna helluo. We then measured foraging and reproductive costs, as well as survival benefits, of this behavior. For 2weeks, we maintained mated adult female Pardosa in plasticcontainers with one of three treated peat moss substrates:a container previously occupied by a conspecific for 3 days,a container previously occupied by an adult Hogna for 3 days,and a container devoid of either cue (control). We measuredprey capture efficiency, body condition, egg sac production,egg sac weight, and egg number for individuals in each treatment.We also counted the number of Pardosa that survived and exhibitedclimbing behavior in the presence of a live Hogna with andwithout silk and excreta cues. Pardosa climbed container wallssignificantly more often in the presence of Hogna silk andexcreta relative to other treatments. Pardosa exposed to Hognacues coupled with live Hogna survived significantly longerthan spiders that had no predator cues available. Pardosa placedin containers with Hogna cues, but no Hogna, lost weight morequickly, ate fewer prey, were in poorer body condition, producedlighter egg sacs, and produced fewer eggs than spiders in controlor conspecific treatments.     

Aggregation and Mate Location in the Red Milkweed Beetle (Coleoptera: Cerambycidae)

We examined behavioral mechanisms underlying aggregation and mate location in the red milkweed beetle, Tetraopes tetrophthalmus (Forster) (Coleoptera: Cerambycidae). Larvae of this species feed on rhizomes of common milkweed, Asclepias syriaca L., and adults feed on the flowers and foliage, aggregating on individual stems within milkweed patches. Adults preferred to aggregate on milkweeds that had multiple, large inflorescences. Males actively searched for females, often flying between host plants. Mate location did not appear to involve long-range pheromones or vision, but rather males landed on milkweed stems arbitrarily, whether or not females were present. Males remained for longer periods, and so tended to accumulate, on milkweed stems that had female-biased sex ratios. We conclude that aggregation of T. tetrophthalmus is cued by host plant characteristics but dynamically influenced by the sex ratio of conspecifics present on individual stems.     

Perception of ultraviolet light by crab spiders and its role in selection of hunting sites

The perception of ultraviolet (UV) light by spiders has so far been only demonstrated in salticids. Crab spiders (Thomisidae) hunt mostly on flowers and need to find appropriate hunting sites. Previous studies have shown that some crab spiders that reflect UV light use UV contrast to enhance prey capture. The high UV contrast can be obtained either by modulation of body colouration or active selection of appropriate backgrounds for foraging. We show that crab spiders (Thomisus sp.) hunting on Spathiphyllum plants use chromatic contrast, especially UV contrast, to make themselves attractive to hymenopteran prey. Apart from that, they are able to achieve high UV contrast by active selection of non-UV reflecting surfaces when given a choice of UV-reflecting and non-UV reflecting surfaces in the absence of odour cues. Honeybees (Apis cerana) approached Spathiphyllum plants bearing crab spiders on which the spiders were high UV-contrast targets with greater frequency than those plants on which the UV contrast of the spiders was low. Thus, crab spiders can perceive UV and may use it to choose appropriate backgrounds to enhance prey capture, by exploiting the attraction of prey such as honeybees to UV.     

Monarch butterfly host plant (milkweed Asclepias spp.) abundance varies by habitat type across 98 prairies

The decline in migratory monarch butterflies (Danaus plexippus) over the past 20 years has been attributed to several drivers, including loss of their host plants (milkweeds Asclepias spp.). This has sparked widespread interest in milkweed ecology and restoration. We developed a model on environmental and habitat‐type variables to predict milkweed abundance by sampling 93 prairie plantings (47 conservation plantings and 46 roadsides) and 5 unplowed prairie remnants throughout the state of Iowa, United States. Milkweeds were censused in 10–25 random locations within each site, and data on plant diversity, age of planting, soil characteristics, and management were tested as predictors of abundance. Milkweed densities of all species combined were highest in remnant prairies (8,705 stems/ha), intermediate in roadside plantings (1,274 stems/ha), and lowest in conservation plantings (212 stems/ha). Most milkweeds were common milkweeds Asclepias syriaca, which were more abundant in roadside than conservation plantings. Remnants contained the most milkweed species. Total milkweed and common milkweed abundance were both predicted by higher soil pH, a more linear site shape, and lower soil bulk density across restorations. Our results indicate that common milkweed is maintained by disturbance, and establishes readily in rural roadside habitat. Remnants are important as reservoirs for multiple milkweed species and should be protected.     

The reactions of two species of aphid to chemotactile cues from a wolf spider match their vulnerability to predation

Various strategies have evolved to protect animals from predators. We explored the activity and predation risk experienced by two species of aphid. Both species will drop from plants when disturbed and face a suite of predators, including wolf spiders, when they reach the ground. We focused on Aphis fabae Scopoli and Aphis nerii Boyer de Fonscolombe (Hemiptera: Aphididae, Aphidini); A. nerii sequesters cardiac glycosides when it feeds on milkweed. We explored the interactions between these aphids and the wolf spider Pardosa milvina Hentz (Araneae: Lycosidae) that is likely a predator they encounter when they are not on their host plants. We hypothesized that there would be differences in the susceptibility of the two species to predation and that the more vulnerable species would react more strongly to substrate-borne cues deposited by the spider. We predicted that any behavioral reactions that the aphid displayed in response to predator cues would be effective in reducing risk. We documented the activity of each aphid species on chemotactile cues from P. milvina and measured predation rate in arenas with and without those same cues. Aphis fabae altered their activity in the presence of P. milvina cues but A. nerii did not. Likewise, A. fabae was more susceptible to predation by P. milvina when no cues were present, but when cues were present, predatory success was much lower. Aphis nerii, the less desirable prey for this predator, moved less and had a different locomotory pattern than A. fabae in control trials with no spider cues and so we cannot determine whether its chemical protection or activity were more important in reducing predation levels. These results provide insight into the risks faced by aphids when they are off of their host plant and in a barren environment.     

Evidence for Mass Flow in Flowering Individuals of the Submersed Vascular Plant Myriophyllum heterophyllum

Myriophyllum heterophyllum Michx. is a rhizomatous submersed aquatic plant that produces a short, emergent floral spike. We hypothesized that lacunar pressures in emergent spikes should be at or near atmospheric pressure and that a mass flow of gases from submersed stems through the rhizome to emergent stems may occur as lacunar O2 concentrations and pressures in submersed stems increase during photosynthesis. We examined the potential for a pressure gradient ([delta]P) to develop along this pathway by measuring diurnal changes in lacunar gas composition and pressure in submersed stems of nonflowering plants and emergent stems of flowering individuals. Methane release from emergent spikes was also monitored during three diurnal cycles to evaluate the hypothesis that the [delta]P is maintained by the release of lacunar gases to the atmosphere. Lacunar O2 concentrations and pressures in submersed stems increased at sunrise and reached maximum levels by midday. Although O2 fluctuated similarly in emergent stems, lacunar pressures remained at or near atmospheric pressure, indicating that a [delta]P is generated between submersed and emergent stems during photosynthesis. Methane release from emergent spikes increased as lacunar pressures increased, indicating that rhizome gases are transported through emergent stems by mass flow and the [delta]P is maintained by venting lacunar gases from emergent spikes. The potential for mass flow in both flowering and nonflowering individuals is discussed.     

REPRODUCTIVE BIOLOGY OF ASCLEPIAS EXALTATA

An analysis of the relationships between plant size and survivorship and reproductive success was carried out by sampling four populations of the herbaceous perennial milkweed Asclepias exaltata in Virginia from 1980 to 1982. The annual survivorship rate (about 65%) is the lowest measured for any species of Asclepias. Survivorship was strongly size-dependent but showed no clear relationship with previous history of fruit production. Non-flowering plants were significantly smaller than flowering plants and showed very strong (r > 0.87) correlations between root dry weight and stem or leaf dry weight. Flowering plants were similar to nonflowering plants in root: shoot ratio (approximately 1:1) but differed in that root dry weight was not strongly correlated with stem or leaf dry weight. Components of inflorescence size were strongly correlated within a given level of comparison (e.g., stems per plant with flowers per plant) but less strongly correlated between levels (e.g., stems per plant with flowers per stem). Number of fruits per plant and percentage fruit-set were positively correlated with every component of inflorescence size. Although overall fruit-set was low (about 2%), fruits that were initiated had a high probability of surviving to maturity. There was no evidence of an early period of high fruit abortion: a relatively constant proportion of fruits aborted between each age class.     

Mechanisms governing sex-ratio variation in dioecious Rumex nivalis

Sex ratios of flowering individuals in dioecious plant populations are often close to unity, or are male biased owing to gender-specific differences in flowering or mortality. Female-biased sex ratios, although infrequent, are often reported in species with heteromorphic sex chromosomes. Two main hypotheses have been proposed to account for female bias: (1) selective fertilization resulting from differential pollen-tube growth of female- versus male-determining microgametophytes (certation); (2) differences in the performance and viability of the sexes after parental investment. Here we investigate these hypotheses in Rumex nivalis (Polygonaceae), a European alpine herb with female-biased sex ratios in which females possess XX, and males XY1Y2, sex chromosomes. Using field surveys and a glasshouse experiment we investigated the relation between sex ratios and life-history stage in 18 populations from contrasting elevations and snowbed microsites and used a male-specific SCAR-marker to determine the sex of nonflowering individuals. Female bias among flowering individuals was one of the highest reported for populations of a dioecious species (mean female frequency = 0.87), but males increased in frequency at higher elevations and in the center of snowbeds. Female bias was also evident in nonflowering individuals (mean 0.78) and in seeds from open-pollinated flowers (mean 0.59). The female bias in seeds was weakly associated with the frequency of male flowering individuals in populations in the direction predicted when certation occurs. Under glasshouse conditions, females outperformed males at several life-history stages, although male seeds were heavier than female seeds. Poor performance of Y1Y2 gametophytes and male sporophytes in R. nivalis may be a consequence of the accumulation of deleterious mutations on Y-sex chromosomes.     

Visual preference of flower-visiting crab spiders (Ebrechtella tricuspidata) for host flowers

1. Crab spiders (Thomisidae) are common flower-visiting spiders that ambush prey on inflorescences. As such, they require specific flowers or substrates for hunting, which are most often selected using sensory cues (e.g. vision). However, few studies have examined the visual preference of crab spiders for particular flowers. In this study, the visual preferences of the crab spider Ebrechtella tricuspidata for different inflorescence characteristics (e.g. colour and shape) were investigated. 2. The results showed that adult spiders explored all colours and shapes, whereas juvenile spiders displayed an overall preference for long (red) and short (purple) wavelength colours. Thus, differences in colour were not particularly important for E. tricuspidata with regard to visual attractiveness and selection. 3. However, inflorescence shape (e.g. tulip) was found to be a more desirable trait for selection, which was probably due to the provision of shelter. 4. These results also suggest that male preference for female spiders depended somewhat on the background colour (wavelength) of the flower on which the female was located.     

Signaling by decorating webs: luring prey or deterring predators?

Many organisms convey false signals to mislead their prey orpredators. Some orb-weaving spiders build conspicuous structureson webs called decorations. Web decorations and spider colorationsare both suggested to be important signals involved in interactionsbetween spiders and other organisms. There are several hypothesesabout the functions of signaling by decorations, among whichprey attraction had received much support, but empirical evidenceregarding predator defense is controversial. In this study,we conducted field experiments to investigate the effects ofspider decoration and coloration on insect interception ratesof webs built by Argiope aemula and to evaluate whether presenceof decorations may decrease predation risk of spiders. Decoratedwebs with spiders present had the highest prey interceptionrate, followed by undecorated webs with spiders, and then undecoratedwebs without spiders. Such results indicated that decorationsof Argiope spiders functioned as visual lures, and so did spiders'bright body colorations. In the field, almost all wasp attackevents occurred on medium-sized spiders rather than on largeones. Moreover, medium-sized Arg. aemula on decorated webs receivedfar more attacks than those on undecorated webs. Results ofthis study thus show that the signals conveyed by decorationscan visually lure prey but at the cost of an increased predationrisk. Received 20 March 2007; revised 3 August 2007; accepted 5 August 2007.     

Bird predation affects diurnal and nocturnal web-building spiders in a Mediterranean citrus grove

Spiders and birds can greatly decrease insect populations, but birds also limit spider densities in some habitats. Bird predation is thought to be one of the causes behind nocturnal activity in spiders, so night-active spiders that hide in retreats during the day should be less affected by bird foraging than day-active spiders. However, this hypothesis has not yet been tested. We investigated the importance of bird predation on the spider community of a Mediterranean organic citrus grove. We excluded birds by placing net cages over the trees and we conducted visual searches in the canopies to sample web-building spiders. As there are many nocturnal species in the family Araneidae, we conducted searches both by day and by night to compare the abundance of active araneids in these two time periods. We sampled the tree trunks with cardboard bands to collect hunting spiders. In bird-excluded canopies there were more spiders of the families Araneidae and Theridiidae. There were higher numbers of active Araneidae at night, but these were just as negatively affected by bird predation as day-active Araneidae, so there was no evidence of nocturnal activity serving as an anti-predator strategy. We did not find any negative impact of birds on hunting spiders. Our results contrast with other studies reporting a negative effect of birds on hunting but not on web-building spiders.