Widespread reliance on olfactory sex and species identification by lyssomanine and spartaeine jumping spiders

Jumping spiders (Salticidae) are renowned for their exceptional vision, but this does not preclude use of other senses. Here we provide evidence that olfactory pheromones are widespread in the Spartaeinae and Lyssomaninae, two subfamilies regarded as basal clades within the Salticidae. Pheromone use by salticids was tested in a series of experiments: males were tested with the odour of conspecific females, heterospecific females, and conspecific males, and females were tested with the odour of conspecific males. With seven of the 29 species tested, we also tested males using the draglines of conspecific females (spider absent) as the odour source. Males of all species tested were attracted to the odour of conspecific females and to the odour of the draglines of conspecific females. There was no evidence of males responding to the odour of heterospecific females or conspecific males, or of females responding to the odour of conspecific males. These findings suggest that it is primarily males that respond to olfactory sex pheromones, consistent with the apparent trend within spiders of males more actively searching for females and females placing greater emphasis on mate‐choice decisions. Compared with most salticid groups, lyssomanines and spartaeines are unusually sedentary and this lifestyle may favour olfactory mate searching. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 107 , 664–677.     

A specialized araneophagic predator's short‐term nutrient utilization depends on the macronutrient content of prey rather than on prey taxonomic affiliation

A specialist predator that has a specialized diet, prey‐specific prey‐capture behaviour and a preference for a particular type of prey may or may not be specialized metabolically. Previous studies have shown that jumping spiders of the genus Portia prey on other spiders using prey‐specific prey‐capture behaviour, prefer spiders as prey to insects and gain long‐term benefits in terms of higher survival and growth rates on spider diets than on insect diets. However, it is unclear whether there are substances uniquely present in spiders on which Portia depends, or, alternatively, spiders and insects all contain more or less the same nutrients but the relative amounts of these substances are such that Portia perform better on a spider diet. These questions are addressed by testing the hypothesis that prey specialization includes metabolic adaptations that allow Portia an enhanced nutrient extraction or nutrient utilization efficiency when feeding on spider prey compared with insect prey. Three groups of Portia quei Zabka are fed either their preferred spider prey or one of two types of flies (Drosophila melanogaster Meigen) that differ in nitrogen and lipid content. Portia quei shows a higher feeding rate of high‐protein flies than of high‐lipid flies and spiders but, after 5 days of feeding, there is no significant difference in growth between treatments, and the diets lead to significant changes in the macronutrient composition of P. quei as a result of variable extraction and utilization of the prey. The short‐term utilization of spider prey is similar to that of high‐lipid flies and both differ in several respects from the utilization of high‐protein flies. Thus, the short‐term nutrient utilization is better explained by prey macronutrient content than by whether the prey is a spider or not. The results suggest that spider prey may have a more optimal macronutrient composition for P. quei and that P. quei does not depend on spider‐specific substances.     

Love is in the air: olfaction-based mate-odour identification by jumping spiders from the genus Cyrba

Jumping spiders (Salticidae) are known for having good eyesight, but the extent to which they rely on olfaction is poorly understood. Here we demonstrate for the first time that olfactory pheromones are used by two species from the salticid genus Cyrba (C. algerina and C. ocellata). Using a Y-shape olfactometer, we investigated the ability of adult males and females of both species to discriminate between mate and non-mate odour. A hidden spider or a spider’s draglines (no spider present) were used as odour sources. There was no evident response by females of either Cyrba species to any tested odour. Males of both species chose odour from conspecific females, or their draglines, significantly more often than the no-odour control, but there was no evident response by males to any of the other odours (conspecific male and heterospecific female). Our findings demonstrate that C. algerina and C. ocellata males can make sex- and species-specific discriminations even when restricted to using olfaction alone. Also, by showing that draglines can be a source of olfactory pheromones, our findings illustrate the difficulty of ruling out olfaction when attempting to test for chemotactile cues.     

Comparative biology of Portia africana,P. albimana,P. fimbriata,P. labiata,and P. shultzi,araneophagic, web-building jumping spiders (Araneae: Salticidae): Utilisation of webs,predatory versatility,and intraspecific interactions

Abstract

Portia is a behaviourally complex and aberrant salticid genus. The genus is of unusual importance because it is morphologically primitive. Five species were studied in nature (Australia, Kenya, Malaysia, Sri Lanka) and in the laboratory in an effort to clarify the origins of the salticids and of their unique, complex eyes. All the species of Portia studied were both web builders and cursorial. Portia was also an araneophagic web invader, and it was a highly effective predator on diverse types of alien webs. Portia was an aggressive mimic, using a complex repertoire of vibratory behaviour to deceive the host spiders on which it fed. The venom of Portia was unusually potent to other spiders; its easily autotomised legs may have helped Portia escape if attacked by its frequently dangerous prey. Portia was also kleptoparasitic and oophagic when occupying alien webs. P. fimbriata from Queensland, where cursorial salticids were superabundant, used a unique manner of stalking and capturing other salticids. The display repertoires used during intraspecific interactions were complex and varied between species. Both visual (typical of other salticids) and vibratory (typical of other web spiders) displays were used. Portia copulated both on and away from webs and frequently with the female hanging from a dragline. Males cohabited with subadult females on webs, mating after the female matured. Adult and subadult females sometimes used specialised predatory attacks against courting or mating males. Sperm induction in Portia was similar to that in other cursorial spiders. Portia mimicked detritus in shape and colour, and its slow, mechanical locomotion preserved concealment. Portia occasionally used a special defensive behaviour (wild leaping) if disturbed by a potential predator. Two types of webs were spun by all species (Type 1, small resting platforms; Type 2, large prey-capture webs). Two types of egg sacs were made, both of which were highly aberrant for a salticid. Responses of different species and both sexes of Portia were quantitatively compared for different types of prey. Many of the trends in behaviour within the genus, including quantitative differences in predatory behaviour, seemed to be related to differences in the effectiveness of the cryptic morphology of Portia in concealing the spider in its natural habitat (‘effective crypsis’). The results of the study supported, in general, Jackson & Blest’s (1982a) hypothesis of salticid evolution which, in part, proposes that salticid ancestors were web builders with poorly developed vision and that acute vision evolved in conjunction with the ancestral spiders becoming proficient as araneophagic invaders of diverse types of webs.     

Hatching responses of subsocial spitting spiders to predation risk

The carrying of eggs often renders parents vulnerable to predators due to increased conspicuousness or decreased mobility. Nonetheless, egg-carrying parents can escape from the predators to which they are vulnerable. Previous studies have demonstrated heavy predation by spider-eating jumping spiders (Portia labiata) on egg-carrying spitting spider (Scytodes pallida) females, but little predation on eggless females. If the timing of hatching is phenotypically plastic, then both S. pallida females and their eggs could reduce the risk of predation by hatching early. Hence, this study examines the hatching responses of S. pallida to chemical cues from P. labiata, both in the laboratory and in the field, and addresses the following questions. (i) Do cues from predatory P. labiata influence the hatching traits of S. pallida? (ii) Are the olfactory cues from predators sufficient for predator detection by S. pallida ? (iii) Are hatching responses to predatory P. labiata controlled by egg-carrying S. pallida females, or directly by their embryos? The study provides evidence of hatching as a life-history switch point, which shows an adaptive plasticity in response to predation risk in egg-carrying S. pallida. Egg-carrying S. pallida females, but not unattended eggs, adjust egg-hatching time (the interval between oviposition and hatching) in response to the threat of predation on both the female and her eggs by P. labiata. In the presence of P. labiata, eggs that are carried by females hatch sooner; the hatchlings of these eggs are therefore smaller than hatchlings born in the absence of P. labiata. Chemical cues that are released from the draglines of P. labiata are sufficient to elicit changes in the egg-hatching traits of S. pallida. Hatching early in response to this predator may benefit both females and their offspring. To my knowledge, this is the first direct experimental study to demonstrate predator-induced hatching plasticity in spiders and, in particular, in animals with parental care.     

Responses of an araneophagic assassin bug,Stenolemus bituberus,to spider draglines

Abstract 1. Animals, as they move through their environment, leave traces of their passage that can be informative to others and convey significant advantages to the animal producing them. However, such traces may also reveal presence, location or identity to enemies. 2. We studied an araneophagic (‘spider‐eating’) assassin bug, Stenolemus bituberus (Heteroptera, Reduviidae), testing whether it associated with areas containing chemotactile traces (e.g. draglines, excreta) left behind by nine sympatric spider species. Stenolemus bituberus were presented with a choice between a substrate containing draglines and a clean substrate. Each hour, for a duration of 12 h, we recorded which substrate was occupied. 3. Stenolemus bituberus tended to associate especially with draglines left by spiders from the genus Achaearanea, their most common prey in nature. 4. These results suggest that S. bituberus exploits draglines from these spiders as cues for indicating prey presence. We also found an increasing tendency to associate with draglines from some spider species through the day, which may be related to circadian patterns or slower response times of some individuals.     

Male-specific (Z)-9-tricosene stimulates female mating behaviour in the spider Pholcus beijingensis

Chemical signals play an important role in spider sexual communication, yet the chemistry of spider sex pheromones remains poorly understood. Chemical identification of male-produced pheromone-mediating sexual behaviour in spiders has also, to our knowledge, not been reported before. This study aimed to examine whether chemically mediated strategies are used by males of the spider Pholcus beijingensis for increasing the probability of copulation. Based on data from gas chromatography–mass spectrometry analysis, electroantennography assay and a series of behavioural tests, we verified that (Z)-9-tricosene is a male-specific compound in the spider P. beijingensis. This compound acts as an aphrodisiac: it increases the likelihood that a female will mate. Mate-searching males release (Z)-9-tricosene to stimulate sexual behaviour of conspecific females. In the two-choice assay, however, sexually receptive females show no preference to the chambers containing (Z)-9-tricosene. This indicates that the male pheromone of P. beijingensis is not an attractant per se to the conspecific females. This is, to our knowledge, the first identification of a male-produced aphrodisiac pheromone in spiders.     

Silk‐ and volatile‐based male mate choice in the genital plug‐producing spider

Female mating history can have a strong effect on male fertilization success. Although males often prefer to mate with virgin females, they often also engage with mated females. As the intensity of sperm competition can differ among mated females, males are expected to evolve means to identify their status. In spiders, males often use female silk to gather information about female quality. Males of many spider species deposit mating plugs into female genitalia to hinder further copulations. We tested whether males of the foliage‐dwelling, plug‐producing spider Philodromus cespitum, which is an important natural enemy of pests, discriminate between females of different mating status and whether they can determine the extent of genital plugging in mated females solely on the basis of cues gained from deposited female silk. We presented males with draglines of females that varied in either mating status (virgin vs. mated), the extent of plugging (small vs. big plug), or the age of the plug (fresh vs. old plug) and examined their mate preferences. Additionally, we tested whether males were attracted to volatile cues produced by female bodies. Our experiments revealed that males preferred draglines of virgin females to those of mated females, and mated females with small plugs to those with large plugs. They were also attracted to female volatile cues. This study suggests that males are able to extract fine‐scale information on mating status from female draglines.     

Secondary host choice by the autoparasitoid Encarsia pergandiella

Autoparasitoids are species of parasitic wasps in the family Aphelinidae which produce females as solitary primary endoparasitoids of homopterans such as whitefly and scale insects (primary hosts), and males as solitary hyperparasitoids. Males generally develop on immature conspecific females or on individuals of other primary parasitoid species (secondary hosts). Encarsia pergandiella is an autoparasitoid that has been introduced to Italy for control of greenhouse whitefly Trialeurodes vaporariorum, in greenhouses and field crops. In this study we examined the secondary host selection behaviour of this species with regard to conspecific females and females of two thelytokous species, E. formosa and E. meritoria. Encarsia formosa has been used successfully for greenhouse whitefly control in Northern Europe, but has not been effective in Southern Italy in winter crops in unheated greenhouses. E. meritoria has recently spread in Italy, and may have potential for biological control of whitefly in the greenhouse environment. In the first experiment, female E. pergandiella were exposed to one of three pair-wise combinations of the three species in petri dish arenas. Parasitism was determined by dissection of the hosts. The number of hosts parasitized by E. pergandiella females did not differ with host species. However, significantly greater numbers of eggs were laid in E. meritoria in both treatments in which it was present; these hosts were more likely to be superparasitized. In a second experiment, observations of females in arenas with equal numbers of all three host species indicated that females encountered and parasitized all host species with approximately equal frequency, although the length of time females spent in the oviposition posture differed with host species.     

Predator–Prey Interactions Between Aggressive-Mimic Jumping Spiders (Salticidae) and Araneophagic Spitting Spiders (Scytodidae) from the Philippines

In Los Baños in the Philippines (Laguna, Luzon), Scytodes sp. indet. is a web-building spitting spider (Scytodidae) that preys primarily on jumping spiders (Salticidae) and Portia labiata is an aggressive-mimic jumping spider that preys especially frequently on Scytodes. Tactics by which three species of Portia (P. africana, P. fimbriata, and P. labiata) and, for Portia labiata, three disjunct populations (Sri Lanka and, in the Philippines, Sagada and Los Baños) capture this especially dangerous prey are compared. Local adaptation to Scytodes by the Los Baños P. labiata is discussed. The Los Baños P. labiata uniquely made consistent use of tactics (soft plucking with palps and signal–detour–leap sequences) that were apparently responsible for greater prey-capture success and more effective avoidance of being spat on. Inter- and intraspecific differences were evident despite having used Portia that were reared in the laboratory with no prioir experience with scytodids.     

Behavioural evidence of UV sensitivity in jumping spiders (Araneae: Salticidae)

Jumping spiders are known to possess ultraviolet (UV) receptors in the retinas of their large-principal eyes. The existence of UV visual cells, however, does not prove that jumping spiders can see into the UV part of spectrum (300–400 nm) or whether such an ability plays any role in salticid intra-specific interactions. In the study reported herein, we performed behavioural experiments to test whether a UV−reflecting jumping spider, Cosmophasis umbratica, is sensitive to UV wavelengths and whether UV cues are important in intra-specific communication. The absence of UV cues not only affected intra-specific behaviour by significantly reducing the frequency of agonistic displays, but also elicited unprecedented courtship displays in males towards their own mirror images and conspecific opponents. Furthermore, C. umbratica males were able to respond rapidly to changes in UV cues of conspecific mirror images by switching between agonistic and courtship displays. These findings clearly demonstrate that C. umbratica males are capable of seeing UV wavelengths and that UV cues are necessary and sufficient for this species to enable the agonistic displays. Hence, UV light may have an important role to play in intra-specific communication in jumping spiders.     

Reliance on trial and error signal derivation by <Emphasis Type="Italic">Portia africana</Emphasis>, an araneophagic jumping spider from East Africa

All species from the jumping spider (Salticidae) genus Portia appear to be predators that specialize at preying on other spiders by invading webs and, through aggressive mimicry, gaining dynamic fine control over the resident spider’s behavior. From previous research, there is evidence that P. fimbriata, P. labiata and P. schultzi derive signals by trial and error. Here, we demonstrate that P. africana is another species that uses a trial and error, or generate and test, algorithm when deriving the aggressive-mimicry signals that will be appropriate in different predator–prey encounters. We discuss the implications of these new findings and the findings from previous work in order to understand the selection factors that drive the evolution of flexibility in aggressive-mimicry strategies.     

The Trochidae and Turbinidae of the Kermadec Ridge (Mollusca: Gastropoda)

Abstract

Taieria erebus (Gnaphosidae) was found to be a versatile predator: it captured insects both cursorially (away from webs) and kleptopar-asitically (on alien webs); it captured spiders in both the presence and absence of webs; and it also ate the eggs of host spiders (oophagy). When T. erebus invaded webs, it was as an aggressive mimic — it performed a repertoire of vibratory behaviours to lure the host spider. Although T. erebus pursued and captured spiders on diverse web-types, it was more effective as a predator when invading densely (rather than sparsely) woven cribellate and non-sticky webs, and was especially effective on non-cribellate sticky webs. Gnaphosids are traditionally referred to as hunting spiders, but T. erebus built a small prey-capture web. T. erebus also preyed on segestriid spiders, then used their webs to catch more prey, this being an unusual example of a spider using, as a tool for predation, the spinning-work of another species from an unrelated family. T. erebus used specialised behaviours to prey on nesting cursorial spiders. Prey was either grasped or stabbed; the venom of T. erebus was highly potent against spiders. Experiments indicated that vision was of little or no importance in the predatory behaviour of T. erebus. The behaviour of T. erebus is compared to that of Portia, a web-building salticid spider which is very versatile in its predatory behaviour and has acute vision. T. erebus is discussed in relation to hypotheses concerning gnaphosid and salticid evolution.     

Sexual Selection in the Water Spider: Female Choice and Male–Male Competition

The water spider Argyroneta aquatica is the only spider spending its whole life under water, and one of the few spider species in which males are larger than females. Previous studies indicated that males can cannibalize females, which is uncommon among spiders. Here we aimed to further test for a potential influence of sexual selection on male body size. We examined the importance of female choice by testing whether females prefer the larger of two simultaneously presented males as mating partners. Further, we examined the influence of male–male competition by comparing the fighting behaviour between large and small males when alone or when together with a female, and we determined the outcome of fights. We found that females approach and choose large males as mating partners, despite the risk of male cannibalism. Additionally, males intensively compete for females, and large males clearly win against smaller ones. Hence sexual selection seems to be important for the evolution of the peculiar sexual size dimorphism of water spiders, as large size is beneficial for males in both the intra‐ and intersexual context. Previous studies have suggested an important role of natural selection in the sex‐specific body size of water spiders, but natural and sexual selection mechanisms apparently work in the same direction, favouring large male size.     

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     

Acoustic communication in the trilling field cricket,Gryllus rubens (Orthoptera: Gryllidae)

Male field crickets produce calling songs that are used for attracting conspecific females for mating. Acoustic communication was studied in the trilling field cricket, Gryllus rubens.A trilled calling song consists of a continuous train of sound pulses, each of which was repeated at a stereotyped rate. Singing males were recorded at different temperatures. The pulse period of the calling song decreased with increasing temperature; there was less effect of temperature on pulse duration. Female phonotaxis was studied on a noncompensating, spherical treadmill. In two-stimulus (choice) playback experiments, females preferred the conspecific trilled song over the chirped calling song of a sympatric species, Gryllus fultoni.This preference persisted even when the song of G. fultoniwas 6 dB louder. Females also discriminated between synthetic trills having different pulse periods; females chose trills with the conspecific pulse period over trills having lower and higher pulse periods.     

Retinal mosaics of a primitive jumping spider,Spartaeus (Salticidae: Araneae): A transition between principal retinae serving low and high spatial acuities

Summary Previous papers (Blest andPrice 1984,Blest andSigmund 1984) have described the receptor mosaics in the tiered principal retinae of jumping spiders with high and low visual acuities, respectively. The latter paper offers a phylogenetic model for the later stages in the evolution of high acuity retinae in which layer I, farthest from the dioptrics, is alone required to sustain fine visual discriminations. Nevertheless, current evidence does not establish that the low acuity principal retinae of primitive jumping spiders were modified to yield the high acuity retinae of advanced forms: each might have evolved independently from a common precursor.We now show that the principal retina ofSpartaeus (the type genus of a primitive subfamily, theSpartaeinae) is a true intermediate between the low acuity retina ofYaginumanis and the high acuity retina ofPortia (also in theSpartaeinae).InSpartaeus, foveal layer I receptors at the anatomical outer side of the retina each bear a single rhabdomere designed as a light guide. Inner layer I receptors each have two rhabdomeres, and rhabdomeres of horizontally adjacent receptors are contiguous, so that optical pooling limits their acuity. Horizontal staircasing of the tips of layer I receptors was shown byBlest et al. (1981) to cause outer receptors to receive focused images from remote planes in object space, and inner receptors from nearer planes. In the high-acuity retina ofPortia all foveal layer I receptors are light-guides, and in the low-acuity retina ofYaginumanis none are. Layer I inSpartaeus, therefore, is a true intermediate step in the evolution of a high-acuity retina, and the local region of the foveal layer I mosaic modified to that end is concerned, appropriately, with the resolution of distant objects.The layer II mosaic is degraded, although less so than inPortia. Layer III is conservative throughout theSalticidae. Foveal layer IV exhibits greater homogeneity inSpartaeus than in other forms.     

Aspects of the ecological energetics of the wolf spider Pardosa (Lycosa) lugubris (Walckenaer)

Summary Feeding experiments were carried out on various stages of the wolf spider Pardosa lugubris in order to construct an energy budget for the spider. Under the rearing conditions the spiders developed at a similar rate to those in the field. P. lugubris females ingest 308 cal during their life. 73.0% of this energy, goes into respiration and 25.7% into production. Males ingest 98 cal of which 81.3% goes into respiration and 16.1% into production. P. lugubris ingests a large proportion of the food it kills and it resembles other arthropod carnivores in having high growth efficiencies.The annual rate of energy flow through a wolf spider population was calculated. 1.39 Kcal/m²/year are killed of which 81.2% is ingested. Of this ingested energy 71.1% goes into respiration and 26.4% into production.The difficulties involved in the assessment of the results of single-species energetics studies are discussed.The annual population energy budget of P. lugubris is compared with that of Pyrrhosoma nymphula, an aquatic arthropod carnivore with a similar lifecycle to P. lugubris. Although the magnitudes of the various components of the energy budget are larger in P. nymphula, there are similarities between the two species. Thus, the amount of energy killed per unit of predator biomass is similar and a similar proportion of this energy is returned to the ecosystem, although proportionally more goes into decomposers in P. lugubris.     

Age and Growth of the Whiskery Shark, Furgaleus macki, from Southwestern Australia

Age and growth of the whiskery shark, Furgaleus macki, from southwestern Australia were examined using vertebral ageing and tag-recapture data. The readability of bands on the vertebral centra varied markedly between individuals. Four readers were used to make band counts, with the most experienced reader having the lowest index of average percent error and the highest level of agreement with final counts. Marginal increment analysis indicated that opaque bands form in January. With parturition occurring from August to October, size data suggests that the first band is probably formed 15–17 months after birth. The age at maturity was estimated to be 4.5 years for males, and 6.5 years for females. The oldest male was 10.5 years, and oldest female was 11.5 years. Von Bertalanffy growth parameters for males were L =121.5cm fork length, K=0.423 year^–1, t ₀=–0.472 years, were L =120.7cm fork length, K=0.369 year^–1, t ₀=–0.544 years for females, and were L =118.1cm fork length, K=0.420 year^–1, t ₀=–0.491 years for combined sexes. Data from a tag recapture study were analysed using a maximum likelihood method to verify the estimates of growth parameters from vertebral ageing. Von Bertalanffy growth parameters from the tag recapture study were L =128.2cm fork length, K=0.288 year^–1, t ₀=–0.654 years. The two methods of estimating growth parameters produced similar results, with rapid growth until approximately 5 years of age, after which there was little increase in length.     

Interspecific predation as a mortality factor among overwintering spiders

Summary Results from field experiments indicate that predation occurs among spruce-living spiders during winter in SW Sweden. Field observations of natural activity showed that Philodromus spp and Pityohyphantes phrygianus together make up 80% of the spiders active on spruce in winter. They are therefore potential predators on other overwintering spiders. Laboratory experiments were performed at +4° C to assess the importance of such predation between spiders. Small spiders (length <2.5 mm) had a mean winter mortality of 58% when kept together with large spiders (2.5 mm) which had a mean mortality of 3% only. Among the small spiders the Erigninae spp seemed to be more vulnerable to predation than other taxonomic groups. Predation also occurred when large P. phrygianus were kept together, but such predation caused mortality of less importance to the spider populations than the mortality among small spiders. Differences in spider density and food availability did not change this pattern. Considerable weight increase occurred in subadult P. phrygianus when fed during winter. This suggests that winter foraging specimens increase their fitness. Interspecific predation among spiders is suggested to be an important mortality factor in natural populations at high spider densities in November and December, when the ambient temperature often is above 0° C and when the density of large spiders is not yet substantially reduced by bird predation.