Ontogenetic shifts in competitive interactions and intra-guild predation between two wolf spider species

Abstract. 1. The wolf spiders (Araneae: Lycosidae) Hogna helluo (Walckenaer) and Pardosa milvina Hentz co-occur in soybean fields of south-west Ohio, U.S.A. As adults, Hogna is the larger species and has the competitive advantage in most interactions; due to differing phenologies, however, their size-classes frequently overlap and as such there is potential for shifts in competitive ability and intra-guild predation. The hypothesis that competitive interactions and intra-guild predation will favour Pardosa when Pardosa is similar-sized, or has a size advantage over Hogna , was tested in laboratory and field experiments.
2. Studies in laboratory arenas, pairing similar-sized individuals of these species and Hogna spiderlings with larger spiders of both species, revealed that intra-guild predation seldom occurs with similar-sized Hogna and Pardosa , however Pardosa will consume small Hogna individuals in laboratory arenas.
3. Field experiments involved stocking high densities (50 m^{– 2} ) of Pardosa and Hogna in enclosures placed in soybean fields. In experiments with spiders of similar size, no interspecific effects were uncovered, but an intraspecific effect was found for Pardosa as its survival and weight gain were lower in the presence of more conspecifics. Large Hogna or Pardosa had no effect on the survival or weight gain of Hogna spiderlings.
4. Although Hogna is a better competitor as an adult, it has no advantage over Pardosa when their size-classes overlap, and Pardosa effects on Hogna may be inconsequential under field conditions. Therefore, the co-existence of these species is fostered by the fact that there are few negative interspecific interactions during their ontogeny.     

Saline stress tolerance partly matches with habitat preference in ground‐living wolf spiders

Salinity interacts with many physiological functions and therefore probably influences the distribution of terrestrial fauna in tidal flooded salt marshes. The present study tests the hypothesis that the physiological tolerance of stenotopic wolf spiders for saline conditions at least partially determines their occurrence throughout salt‐marsh and nonsaline habitats. The duration of survival of three stenotopic wolf spider species (Araneae: Lycosidae) with different habitat preferences is compared in a controlled laboratory experiment. The forest‐dwelling Pardosa saltans, the salt‐marsh resident Pardosa purbeckensis and its sister species the inland‐living Pardosa agrestis are exposed to experimental conditions with different levels of salinity. Individuals (45 males and 20–45 females per treatment) are placed in individual air‐tight boxes filled with water‐saturated sand. Three levels of salinity are tested: nonsaline (0‰), medium saline (33–35‰) and highly saline (66–70‰). Contents of carbon, hydrogen and nitrogen and the molar ration carbon/nitrogen remain constant over time and do not differ among salinity treatments, indicating that starvation effects on survival, if any, are similar for all treatments. Conversely, body water significantly decreases over time and differs among salinity treatments, in accordance with patterns of survival. Conforming to their habitat preference, the survival of P. saltans and P. agrestis decreases quickly under highly saline conditions. Pardosa purbeckensis, however, has a high survival under both saline and nonsaline conditions. The duration of survival of females is significantly lower than that of males of P. saltans and P. purbeckensis. Durations of survival of ground‐living wolf spiders exposed to salinity partly match their habitat distribution but do not explain the restriction of salt‐marsh species to saline habitats.     

Experimental assessment of trophic ecology in a generalist spider predator: Implications for biocontrol in Uruguayan crops

Conservative biological control promotes the use of native natural enemies to limit the size and growth of pest populations. Although spiders constitute one of the most important groups of native predators in several crops, their trophic ecology remains largely unknown, especially for several generalist taxa. In laboratory, we assessed the predatory behaviour of a wandering spider (the wolf spider Lycosa thorelli (Keyserling, 1877) against several arthropods varying in size and trophic positions, all found in South American soybean and rice crops. As prey we used the bug Piezodorus guildinii (Westwood, 1837) as well as larvae and adults of the moth Spodoptera frugiperda (Smith, 1797), both being considered important pests in Uruguayan crops. We also used several non-pest arthropods as prey, sarcophagid flies, carabid beetles and wolf spiders. All prey were attacked in more or less high, although not statistically differing, proportions. However, carabids were not consumed, and bugs were consumed in significantly lower proportions than flies. A negative correlation was found between prey size and acceptance rate. Immobilization times were longer against larvae when compared to moths and flies, while predatory sequences were longer for bugs when compared to flies, moths and spiders. In addition, we found a positive effect of prey size on predatory sequence length and complexity. Our results confirm the ability of spiders to attack and feed upon prey with different morphologies, included well-defended arthropods, and their potential use as natural enemies of several pests in South American crops.     

农田狼蛛亚群落结构的研究

本文对南宁市郊农田九种生境狼蛛亚群落结构进行了讨论。群落有7属37种狼蛛。文中揭示了农田各生境区间狼蛛亚群落的主要成分及其主要属性差异的原因。以狼蛛类群为指示动物,用群落相似性比例和聚类分析法,能较好地区分幼豹蛛(Pardosa pusiola)与奇异獾蛛(Trochosa ruricola)和拟环纹豹蛛(Pardosa pseudoannulata)与拟水狼蛛(Pirata subpiraticus)是构成农田区旱地和水田生境型具有代表性的两个狼蛛亚群落主要成分。     

拟环纹豹蛛的生物生态学研究

运用田间观察和室内外饲养相结合的方法,对稻田蜘蛛优势种拟环纹豹蛛Pardosa pseudoannulata的生物生态学特性进行了研究。拟环纹豹蛛在湘西北1年发生2～3代,第3代不完整,以第2代历期最短,第3代（越冬代）历期最长;以成蛛、亚成蛛或幼蛛越冬,其分布图式为聚集分布;4月下旬即由田埂向稻田内迁移,在水稻生育期有3次数量高峰; 属游猎型蜘蛛,可步行在水稻、水面、陆地等处捕食飞虱、叶蝉等多种目标害虫; 受惊时潜入水中或潜伏。雌雄蛛均可多次交配,雌蛛一生最多可产卵5次,实验种群卵囊含卵量76～337粒,平均156粒。孵化的幼蛛,先群集在雌蛛背面,3～5天后开始扩散,6～7天扩散到高峰。幼蛛期47～158天;成蛛期121～236天,雌蛛较雄蛛长21～62天。性比除第2代外的各代均为雄性多于雌性。本文详细记述了拟环纹豹蛛的求偶与交配行为,产卵与护卵习性,孵化与携幼行为,幼蛛生长、蜕皮、各龄期形态特征及耐饥、耐旱能力等。     

Offspring recognition by mother crab spiders with extreme maternal care

Maternal care is provided by several spider species, but there are no reports of mother spiders recognizing their young, which suggests that maternal care can be exploited by unrelated individuals. Diaea ergandros, a crab spider with extreme, sacrificial maternal care, does accept unrelated spiderlings (ca. 43.9% of spiderlings) into its nest in areas of high nest density. However, a field and a laboratory experiment with mother spiders and natural and adoptive spiderlings demonstrated that mothers did recognize their own offspring. Recognition was not expressed in survival as adopted (unrelated) spiderlings had similar survival rate to that of natural offspring. Instead it was displayed in growth; mother D. ergandros caught large prey items for their own offspring, but not for adopted spiderlings, and so natural offspring grew more than adopted spiderlings. Also, mothers produced trophic oocytes, which are important for the sacrificial care that influences spiderling survival, only when they lived with their own offspring.     

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

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

Effects of chronic exposure to a toxic prey in a generalist predator

Abstract.  Generalist predators experience large differences in prey quality, and eating toxic prey may affect their fitness even if the toxic prey constitutes only a small fraction of a mixed diet. Feeding and life history parameters were examined in the wolf spider Pardosa prativaga during continuous exposure to the toxic collembolan Folsomia candida . Spiderlings were divided into a control group fed adult Drosophila melanogaster , a group fed only F. candida , and a mixed diet group fed both types of prey. Folsomia candida reduced survival, developmental stage attained and growth rate of all exposed spiders. Spiders chronically exposed to F . candida increased their consumption of D. melanogaster compared to control spiders. Folsomia candida thus inhibited the utilization of the high-quality food and/or increased the respiration rate. The intake of F . candida remained at a constant low level throughout the experiment, indicating that P. prativaga was unable to develop an absolute aversion against this prey. Half of the control group was also given the mixed diet after a large juvenile instar was attained. These spiders were not affected to the same extent as the newly hatched spiderlings, indicating an ontogenetically increased tolerance. Comparison of laboratory-raised and field-caught spiders confirmed that tolerance to F. candida was size-dependent and thus not an induced response. A differential survival in the mixed diet group, based partly on maternal effects, indicated possible genetic variability in the physiological tolerance to F. candida .     

Physiological costs during the first maternal care in the wolf spider Pardosa saltans (Araneae,Lycosidae)

Many arachnids like other terrestrial arthropods, provide extensive maternal care. Few studies have quantified the underlying physiological costs of maternal care. We investigated how maternal care affects the free-moving wolf spider’s (Pardosa saltans) energy requirements. We described in detail their basic reproduction biology (i.e. carrying cocoon and young) and we evaluated the variation in the females’ energy reserves during maternal care. Our results show that mothers guard eggs until hatching and then guard their spiderlings for 27–30 more days. Laboratory observations indicated that spiderlings start leaving the maternal abdomen gradually 5–7 days after hatching. Females carry an egg sac (cocoon) that can weigh up to 77% of their post-reproduction weight and carry young that weigh 87–100% of their body mass. Females lost weight over time despite regular food intake, while carrying cocoon and young; but their weights increased gradually during the dispersal of young. The contributions of proteins, glucose and triglycerides to maintain females’ energy were calculated. Their energetic state varied during maternal care, in particular lipid levels declined, during the care of spiderlings when the females’ predatory behaviour was inhibited. Our results show that the maternal care provided by P. saltans females is particularly costly physiologically, during the 30 days following egg sac formation and development of spiderlings, even when food is available.     

Effects of glufosinate‐ammonium herbicide and pod sealant on spider Pardosa agrestis

Herbicides based upon glufosinate‐ammonium (GLA) are among the world's most widely used. They also are applied on the most prominent oil crops as desiccants in combination with pod sealants to prevent pod shatter and seed loss close to harvest. Even though these crops occupy a significant part of the world's agroecosystems, the effects of GLA herbicides on non‐target arthropods, and in particular natural enemies of pests, have been studied very rarely, and such effects of pod sealants have never been studied. We studied in our laboratory mortality as well as prey capture efficiency of the common GLA herbicide and desiccant Basta 15^®, pod sealant Arrest^®, and a mixture of both on the wolf spider Pardosa agrestis. We found that Basta 15^® and the mixture had lethal effect on spiders. We also found significant, short‐term effect on predatory activity of spiders after all treatments. Basta 15^® significantly influenced the amount of captured prey also in the long term. This is the first study showing lethal effect on spiders of the herbicide and herbicide plus pod sealant mixture. This is also the first study examining the effects of pod sealant on the mortality and predatory activity of a pest antagonist. More studies regarding the effects of agricultural chemical mixes are needed to uncover their effects on beneficial organisms existing within agroecosystems.     

用ELISA方法研究稻田节肢动物的食物关系

应用酶联免疫吸附试验双抗体夹心法,研究了稻田节肢动物,包括19种捕食性天敌、4种主要水稻害虫及1种中性昆虫之间的食物关系。在检测的19种捕食者中,有15种捕食了白背飞虱,11种捕食了褐飞虱。其中食虫沟瘤蛛、拟水狼蛛、八斑鞘蛛、粽管巢蛛、四斑锯螯蛛和拟环纹豹蛛对两种稻飞虱的捕食阳性率较高。有7种捕食了稻纵卷叶螟幼虫,其中粽管巢蛛、褶管巢蛛、拟环纹豹蛛、青翅蚁形隐翅虫和印度细颈步甲的捕食阳性率较高。拟环纹豹蛛、浙江豹蛛和拟水狼蛛对稻蝗若虫有较高的阳性率。13种捕食者捕食了摇蚊,摇蚊作为替代或补充猎物,对稻田捕食性节肢动物亚群落的重建和发展具有重要意义。在实验检测的基础上,构建了被检测节肢动物猎物与捕食者间的食物关系图。     

Ecosystem linkages revealed by experimental lake-derived isotope signal in heathland food webs

Cross-ecosystem movement of nutrients and biomass can have important effects on recipient systems. Emerging aquatic insects are subsidies to terrestrial ecosystems and can influence foodweb interactions in riparian systems. In a 2-year field experiment, we simulated aquatic insect deposition by adding adult midge carcasses (150?g dry mass m^?2 year^?1) to 1-m² heathland plots at a site with low natural midge deposition. We established four levels of midge-addition treatments and measured stable isotopes (??¹³C and ??¹⁵N) in plants and arthropods within each treatment. We used a multiple-source isotope Bayesian mixing model to estimate the terrestrial versus aquatic contribution to the diets of arthropods. Aquatic resources were incorporated into plant, detritivore, and predator biomass. Detritivorous Collembola showed the greatest difference in isotope values (+3??? ??¹⁵N and +4??? ??¹³C) between midge-addition and reference treatments. Isotope values of small spiders followed the same trend of enrichment as Collembola while other arthropods (mites and large spiders) were only enriched after 2?years of midge addition. Although predator diets did not change, they became isotopically enriched via their likely prey (Collembola). Plants also had elevated ??¹⁵N (+1???) in midge-addition treatments. The time required and amount of midge-derived C and N detected varied and depended on trophic position. Midge-derived nutrients were no longer present in arthropod biomass in the year following midge addition. Aquatic insect carcasses can be rapidly incorporated into terrestrial food webs in nearshore habitats, and repeated inputs can be detected at multiple trophic levels, thus highlighting the importance of the detrital pathway for aquatic to terrestrial cross-ecosystem subsidies.     

Early season natural control of the brown planthopper, Nilaparvata lugens: the contribution and interaction of two spider species and a predatory bug

Nilaparvata lugens St?l, the Brown Planthopper (BPH), is a major pest in rice. The lycosid Pardosa pseudoannulata (B?senberg & Strand) and the linyphiid Atypena formosana (Oi) are the early dominant predators in irrigated rice. Later, predatory bugs, including the mirid Cyrtorhinus lividipennis Reuter, become dominant. In unsprayed rice, BPH numbers normally remain low. While P. pseudoannulata is known to be a key natural enemy of BPH, the contribution to BPH reduction by the smaller A. formosana is less well known. Due to the size difference, A. formosana may also be an intraguild prey of P. pseudoannulata. To investigate predation on BPH by the two spider species alone, together and in combination with C. lividipennis, two cage experiments were conducted, the first with adult spiders, the second with immature/unmated spiders. In the latter, spiders were introduced with a delay to allow a better establishment of C. lividipennis. In both experiments, BPH numbers were lowest in treatments with more than one predator species present. Intermediate BPH reduction was obtained with P. pseudoannulata followed by A. formosana and C. lividipennis. While P. pseudoannulata can prey upon large nymphs and adult BPH, A. formosana favour early instar BPH, leading to a more even control across instars in treatments with both spider species. Pardosa pseudoannulata numbers increased more with A. formosana or A. formosana and C. lividipennis present. In contrast, P. pseudoannulata reduced A. formosana and C. lividipennis numbers. The presence of C. lividipennis apparently lessened intraguild predation on A. formosana and may be an intraguild prey of A. formosana. Competition for prey may have added to the antagonistic interactions found. From a biological control perspective, the presence of both spiders in early rice is an advantage for the biological control of BPH in rice. In early rice, results show that A. formosana is an important predator of BPH and, like C. lividipennis, can complement control by P. pseudoannulata and serve as intraguild prey for the latter species.     

取食转基因抗虫棉上的棉蚜对粉舞蛛存活和发育的影响

本文以转双价基因棉花SGK321、棉蚜Aphis gossypii和粉舞蛛Alopecosa pulverulenta为对象,研究了捕食转基因植物上的植食性害虫对多食性捕食性天敌的影响。结果表明,粉舞蛛可以猎食棉蚜,但单独捕食棉蚜不足以长期维持若蛛的生存和发育; 与果蝇混合饲养,能显著提高若蛛存活率和体重。在猎物过量或数量不足的情况下,单独捕食转基因棉或常规棉上的棉蚜,若蛛的生存曲线和体重差异不显著。在猎物过量的条件下,用转基因棉上的棉蚜与果蝇混合饲养,若蛛的存活率显著高于用常规棉上的棉蚜与果蝇混合的处理; 但这两种处理下,若蛛的体重差异不显著。在猎物数量不足的情况下,用转基因棉或常规棉上的棉蚜与果蝇混合饲养,若蛛的存活率和体重差异都不显著。可见,转双价基因棉花SGK321上的棉蚜对粉舞蛛的存活和发育没有显著的不利影响。     

A mass rearing method for the linyphiid spider species Erigone atra (Blackwall) (Araneae: Linyphiidae)

Abstract:  A mass rearing method for Erigone atra (Blackwall) (Araneae: Linyphiidae) allowing continuous laboratory rearing is described. Twenty 1–2-day old spiderlings were kept together in plastic boxes, which were filled with soil containing a culture of the Collembola species, Lepidocyrtus lanuginosus (Gmelin) (Entomobryidae), and serving the spiders as a continuous available prey source. Once per week vestigial-wing fruit flies of Drosophila melanogaster Meigen (Diptera: Drosophilidae) were provided as additional prey. In addition, the rearing boxes were filled with wood-wool serving spiders as points of contact for their webs. After 3–5 weeks most of the spiderlings developed to adults, which were separated individually into glass tubes filled with soil and Collembola until all of them became adult. To produce a new generation of spiders 20–40 adult/subadult spiders originating from different mass rearing boxes were brought together and kept and fed in the same way as the spiderlings. Within a few days females started to produce eggsacs. The eggsacs were transferred into glass tubes filled with a layer of moist plaster of Paris until the spiderlings hatched, which were then bred as described above. Erigone atra was bred over 12 generations within a period of 2 years. The mean rearing success (from 1 to 2-day-old spiderlings to adults) was 59.3%. Decreasing rearing success, decrease of fecundity or decrease of adult spider size were not observed. Advantages and use of the mass rearing method are discussed in relation to rearing methods for other spiders.     

The Nutritional Content of Prey Affects the Foraging of a Generalist Arthropod Predator

While foraging theory predicts that predatory responses should be determined by the energy content and size of prey, it is becoming increasingly clear that carnivores regulate their intake of specific nutrients. We tested the hypothesis that prey nutrient composition and predator nutritional history affects foraging intensity, consumption, and prey selection by the wolf spider, Pardosa milvina. By altering the rearing environment for fruit flies, Drosophila melanogaster, we produced high quality flies containing more nitrogen and protein and less lipid than low quality fruit flies. In one experiment, we quantified the proportion of flies taken and consumption across a range of densities of either high or low quality flies and, in a second experiment, we determined the prey capture and consumption of spiders that had been maintained on contrasting diets prior to testing. In both cases, the proportion of prey captured declined with increasing prey density, which characterizes the Type II functional response that is typical of wolf spiders. Spiders with similar nutritional histories killed similar numbers of each prey type but consumed more of the low quality prey. Spiders provided high quality prey in the weeks prior to testing killed more prey than those on the low quality diet but there was no effect of prior diet on consumption. In the third experiment, spiders were maintained on contrasting diets for three weeks and then allowed to select from a mixture of high and low quality prey. Interestingly, feeding history affected prey preferences: spiders that had been on a low quality diet showed no preference but those on the high quality diet selected high quality flies from the mixture. Our results suggest that, even when prey size and species identity are controlled, the nutritional experience of the predator as well as the specific content of the prey shapes predator-prey interactions.     

我国狼蛛科5种记述

记述我国蜘蛛目狼蛛科5种,包括1种豹蛛(罩豹蛛Pardosa vulvitecta)雄性新发现,1种豹蛛(意大利豹蛛Pardosa italica)中国新纪录,以及我国已记载的2种豹蛛和1种舞蛛的修订。     

Collembola as alternative prey sustaining spiders in arable ecosystems: prey detection within predators using molecular markers

Collembola comprise a major source of alternative prey to linyphiid spiders in arable fields, helping to sustain and retain these predators as aphid control agents within the crop. Polymerase chain reaction primers were developed for the amplification, from spider gut samples, of DNA from three of the most abundant species of Collembola in wheat crops in Europe, namely Isotoma anglicana, Lepidocyrtus cyaneus and Entomobrya multifasciata. The primers amplified fragments of the mitochondrial cytochrome oxidase subunit I (COI) gene and were designed following alignment of comparable sequences for a range of predator and prey species. Each of the primer pairs proved to be species-specific to a Collembola species, amplifying DNA fragments from 211 to 276 base pairs in length. Following consumption of a single collembolan, prey DNA was detectable in 100% of spiders after 24 h of digestion. We report the first use of DNA-based techniques to detect predation by arthropods on natural populations of prey in the field. All three species of Collembola were consumed by the spiders. By comparing the ratios of the Collembola species in the field with the numbers of spiders that gave positive results for each of those species, it was possible to demonstrate that the spiders were exercising prey choice. Overall, a single target species of Collembola was eaten by 48% of spiders while a further 16% of spiders contained DNA from two different species of Collembola. Preference was particularly evident for I. anglicana, the species most frequently found in spider guts yet the least numerous of the three target species in the field.     

The life-cycle of the Wolf spider Pardosa lugubris in Holland

Pardosa lugubris in Scotland has a 2-year life-cycle. In Holland the situation is less clear-cut: spiderlings emerging from the summer egg sac can reach sexual maturity by the following spring. These spiders, with a 1-year life-cycle, have one instar fewer than the Scottish spiders. The Dutch spiderlings from the autumn egg sac, like the Scottish spiders, have a 2-year life-cycle and have the same number of instars as the Scottish spiders.
The shorter life-cycle of some of the Dutch spiders is probably due to the fact that summer temperatures are higher in Holland than in Scotland. It is suggested that the length of the life-cycle may be an important factor in spider distribution and that this may account for the relative scarcity of large spider species in high latitudes.     

Herbivore release through cascading risk effects

Predators influence prey through consumption, and through trait-mediated effects such as emigration in response to predation risk (risk effects). We studied top-down effects of (sub-) adult wolf spiders (Lycosidae) on arthropods in a meadow. We compared risk effects with the overall top-down effect (including consumption) by gluing the chelicers of wolf spiders to prevent them from killing the prey. In a field experiment, we created three treatments that included either: (i) intact (‘predation’) wolf spiders; (ii) wolf spiders with glued chelicers (‘risk spiders’); or (iii) no (sub-) adult wolf spiders. Young wolf spiders were reduced by their (sub-) adult congeners. Densities of sheetweb spiders (Linyphiidae), a known intraguild prey of wolf spiders, were equally reduced by the presence of risk and predation wolf spiders. Plant- and leafhoppers (Auchenorrhyncha) showed the inverse pattern of higher densities in the presence of both risk and predation wolf spiders. We conclude that (sub-) adult wolf spiders acted as top predators, which reduced densities of intermediate predators and thereby enhanced herbivores. Complementary to earlier studies that found trait-mediated herbivore suppression, our results demonstrate that herbivores can be enhanced through cascading risk effects by top predators.