Subsocial behaviour and brood adoption in mixed-species colonies of two theridiid spiders

Cooperation and group living often evolves through kin selection. However, associations between unrelated organisms, such as different species, can evolve if both parties benefit from the interaction. Group living is rare in spiders, but occurs in cooperative, permanently social spiders, as well as in territorial, colonial spiders. Mixed species spider colonies, involving closely related species, have rarely been documented. We examined social interactions in newly discovered mixed-species colonies of theridiid spiders on Bali, Indonesia. Our aim was to test the degree of intra- and interspecific tolerance, aggression and cooperation through behavioural experiments and examine the potential for adoption of foreign brood. Morphological and genetic analyses confirmed that colonies consisted of two related species Chikunia nigra (O.P. Cambridge, 1880) new combination (previously Chrysso nigra) and a yet undescribed Chikunia sp. Females defended territories and did not engage in cooperative prey capture, but interestingly, both species seemed to provide extended maternal care of young and indiscriminate care for foreign brood. Future studies may reveal whether these species adopt only intra-specific young, or also inter-specifically. We classify both Chikunia species subsocial and intra- and interspecifically colonial, and discuss the evolutionary significance of a system where one or both species may potentially benefit from mutual tolerance and brood adoption.     

A comparative study of an innate immune response in Lamprologine cichlid fishes

Social interactions facilitate pathogen transmission and increase virulence. Therefore, species that live in social groups are predicted to suffer a higher pathogen burden, to invest more heavily in immune defence against pathogens, or both. However, there are few empirical tests of whether social species indeed invest more heavily in immune defence than non-social species. In the current study, we conducted a phylogenetically controlled comparison of innate immune response in Lamprologine cichlid fishes. We focused on three species of highly social cichlids that live in permanent groups and exhibit cooperative breeding (Julidochromis ornatus, Neolamprologus pulcher and Neolamprologus savoryi) and three species of non-social cichlids that exhibit neither grouping nor cooperative behaviour (Telmatochromis temporalis, Neolamprologus tetracanthus and Neolamprologus modestus). We quantified the innate immune response by injecting wild fishes with phytohaemagglutinin (PHA), a lectin that causes a cell-mediated immune response. We predicted that the three highly social species would show a greater immune reaction to the PHA treatment, indicating higher investment in immune defence against parasites relative to the three non-social species. We found significant species-level variation in immune response, but contrary to our prediction, this variation did not correspond to social system. However, we found that immune response was correlated with territory size across the six species. Our results indicate that the common assumption of a positive relationship between social system and investment in immune function may be overly simplistic. We suggest that factors such as rates of both in-group and out-group social interactions are likely to be important mediators of the relationship between sociality and immune function.     

David and Goliath: potent venom of an ant-eating spider (Araneae) enables capture of a giant prey

It is rare to find a true predator that repeatedly and routinely kills prey larger than itself. A solitary specialised ant-eating spider of the genus Zodarion can capture a relatively giant prey. We studied the trophic niche of this spider species and investigated its adaptations (behavioural and venomic) that are used to capture ants. We found that the spider captures mainly polymorphic Messor arenarius ants. Adult female spiders captured large morphs while tiny juveniles captured smaller morphs, yet in both cases ants were giant in comparison with spider size. All specimens used an effective prey capture strategy that protected them from ant retaliation. Juvenile and adult spiders were able to paralyse their prey using a single bite. The venom glands of adults were more than 50 times larger than those of juvenile spiders, but the paralysis latency of juveniles was 1.5 times longer. This suggests that this spider species possesses very potent venom already at the juvenile stage. Comparison of the venom composition between juvenile and adult spiders did not reveal significant differences. We discovered here that specialised capture combined with very effective venom enables the capture of giant prey.     

The role of perennial habitats for Central European farmland spiders

Spiders are important predators of insect pests. Some species invariably dominate spider communities in crop fields over large parts of Europe, and are therefore considered as “agrobionts”. It is however not clear if these species generally prefer arable habitats, or to what degree they utilize other habitats during times when crops are inhospitable. Here, spider abundances in 26 crop fields and 16 perennial habitats in Germany were compared during spring. Overall spider abundance in winter wheat was 75% lower than in perennial habitats, and species richness was reduced by 46%. Out of a total of 91 species, 73 had higher densities in perennial habitats, including the most dominant crop spiders Erigone atra and Tenuiphantes tenuis. Only one species (Oedothorax apicatus) was significantly more abundant in crops than in perennial habitats. Hence, the high dominance of most “agrobiont” species in crops was due to low densities of other species, and not to their own higher abundances. Therefore, perennial habitats adjacent to crop-fields could enhance spider populations and their potential for biological control in arable fields on a landscape level.     

Selective olfactory attention of a specialised predator to intraspecific chemical signals of its prey

Prey-specialised predators have evolved specific cognitive adaptations that increase their prey searching efficiency. In particular, when the prey is social, selection probably favours the use of prey intraspecific chemical signals by predatory arthropods. Using a specialised ant-eating zodariid spider, Zodarion rubidum, which is known to prey on several ant species and possesses capture and venom adaptations more effective on Formicinae ants, we tested its ability to recognise chemical cues produced by several ant species. Using an olfactometer, we tested the response of Z. rubidum towards air with chemical cues from six different ant species: Camponotus ligniperda, Lasius platythorax and Formica rufibarbis (all Formicinae); and Messor structor, Myrmica scabrinodis and Tetramorium caespitum (all Myrmicinae). Z. rubidum was attracted to air carrying chemical cues only from F. rufibarbis and L. platythorax. Then, we identified that the spiders were attracted to airborne cues coming from the F. rufibarbis gaster and Dufour's gland, in particular. Finally, we found that among several synthetic blends, the decyl acetate and undecane mixture produced significant attraction of spiders. These chemicals are produced only by three Formicine genera. Furthermore, we investigated the role of these chemical cues in the communication of F. rufibarbis and found that this blend reduces their movement. This study demonstrates the chemical cognitive capacity of Z. rubidum to locate its ant prey using chemical signals produced by the ants. The innate capacity of Z. rubidum to olfactory detect different ant species is narrow, as it includes only two ant genera, confirming trophic specialisation at lower than subfamily level. The olfactory cue detected by Zodarion spiders is probably a component of the recruitment or trail pheromone.     

Context-dependent crypsis: a prey’s perspective of a color polymorphic predator

Many animals use body coloration as a strategy to communicate with conspecifics, prey, and predators. Color is a trade-off for some species, since they should be visible to conspecifics but cryptic to predators and prey. Some flower-dwelling predators, such as crab spiders, are capable of choosing the color of flowers where they ambush flower visitors and pollinators. In order to avoid being captured, visitors evaluate flowers visually before landing. The crab spider Mecaphesa dubia is a polymorphic species (white/purple color morphs), which inhabits the flower heads of a dune plant, Palafoxia lindenii. Using full-spectrum photography of spiders and flowers, we evaluated how honeybees perceived the spiders at different distances. Using visual modeling, we obtained the chromatic and achromatic contrasts of the spiders on flower heads as perceived by honeybees. Purple morphs were found mainly on the receptacle area and white morphs were equally likely to be found in the flowers and receptacle. According to theoretical modeling, white morphs were visible to honeybees from a distance of 10 cm in receptacle area but appeared to be cryptic in the flower area. Purple morphs were cryptic on the receptacle and less so when they were on the flowers. Spiders on flower heads are predicted to be more easily detected by honeybees using chromatic contrast. Our study shows that the conspicuousness of flower dwelling spiders to honeybees depends on the color morph, the distance of observation, and the position of spider on the flower head.     

降水波动对呼伦贝尔草甸草原初级生产力年际动态影响

降水是控制草原植被生产力最关键的因素. 为探明降水量在不同时间尺度上的波动及其对草原植被的响应机制,以呼伦贝尔草甸草原为研究对象,选用反映年际降水量波动的降水集中度和偏离期2个因子,将以地面光谱生物量模型计算获得的草甸草原植被NPP(net primary production,净初级生产力)与不同周期降水量和降水波动因子建立回归模型,分析年际降水量波动对草甸草原植被NPP的影响. 结果表明:①在呼伦贝尔草甸草原区,以年内4—7月为关键期,P_k(关键期累积降水量)对草甸草原植被NPP的影响最大,C_dk(关键期降水集中度)平均值为0.439±0.182,d_k(关键期降水偏离期)平均值为31.6 d,变幅为-3.6~94.2 d. ②以P_k和以旬为单位的C_dk、d_k构建的草甸草原植被NPP估算模型,y=-52.11+88.957C_dk+0.724d_k+0.953P_k,能较好地反映草甸草原植被NPP与降水波动之间的关系,模型估测精度可达91.0%. 因此,在半干旱草原区,利用基于遥感植被反射光谱构建的NPP模型计算草甸草原植被NPP具有较高的可信度,并且与样方调查结果有极高的关联性.      

Adjustment of web-building initiation to high humidity: a constraint by humidity-dependent thread stickiness in the spider Cyrtarachne

Cyrtarachne is an orb-weaving spider belonging to the subfamily Cyrtarachninae (Araneidae) which includes triangular-web-building Pasilobus and bolas spiders. The Cyrtarachninae is a group of spiders specialized in catching moths, which is thought to have evolved from ordinary orb-weaving araneids. Although the web-building time of nocturnal spiders is in general related to the time of sunset, anecdotal evidence has suggested variability of web-building time in Cyrtarachne and its closely related genera. This study has examined the effects of temperature, humidity, moonlight intensity, and prey (moths) availability on web-building time of Cyrtarachne bufo, Cyrtarachne akirai, and Cyrtarachne nagasakiensis. Generalized linear mixed model (GLMM) have revealed that humidity, and not prey availability, was the essential variable that explained the daily variability of web-building time. Experiments measuring thread stickiness under different humidities showed that, although the thread of Cyrtarachne was found to have strong stickiness under high humidity, low humidity caused a marked decrease of thread stickiness. By contrast, no obvious change in stickiness was seen in an ordinary orb-weaving spider, Larinia argiopiformis. These findings suggest that Cyrtarachne adjusts its web-building time to favorable conditions of high humidity maintaining strong stickiness, which enables the threads to work efficiently for capturing prey.     

Biphasic activity of a jumping spider

Individual variation is a ubiquitous and important factor that affects ecological dynamics. This study examined individual variation in the nest-use pattern of the jumping spider Phidippus audax. Although the jumping spider is a diurnal species, field observations in this study revealed that the majority of individuals remained in their nests during the day. An accompanying examination of the hunger level of the spiders revealed that spiders that remained in nests were more starved than those observed outside nests. If spiders actively forage when they are starved, as has been suggested by previous studies, one would expect to see the opposite trend (i.e., spiders that remained in nests are more satiated). Thus, the pattern observed in the field contradicts the known behavioral pattern of the spiders. An individual-based model was used to investigate the behavioral mechanism of the spider and the discrepancy found in the observations. A basic assumption of the model is that spiders possess distinct inactive and active phases (biphasic activity pattern), and transitions between the two phases are regulated by the hunger level of the spider. Data from a laboratory experiment were used to examine the assumptions of the model partially. The model was able to capture patterns observed in the data, suggesting that the pattern of transitions in biphasic activity is an important trait of the foraging behavior of the jumping spider.     

Armoured spiderman: morphological and behavioural adaptations of a specialised araneophagous predator (Araneae: Palpimanidae)

In a predator–prey system where both intervenients come from the same taxon, one can expect a strong selection on behavioural and morphological traits involved in prey capture. For example, in specialised snake-eating snakes, the predator is unaffetced by the venom of the prey. We predicted that similar adaptations should have evolved in spider-eating (araneophagous) spiders. We investigated potential and actual prey of two Palpimanus spiders (P. gibbulus, P. orientalis) to support the prediction that these are araneophagous predators. Specific behavioural adaptations were investigated using a high-speed camera during staged encounters with prey, while morphological adaptations were investigated using electron microscopy. Both Palpimanus species captured a wide assortment of spider species from various guilds but also a few insect species. Analysis of the potential prey suggested that Palpimanus is a retreat-invading predator that actively searches for spiders that hide in a retreat. Behavioural capture adaptations include a slow, stealthy approach to the prey followed by a very fast attack. Morphological capture adaptations include scopulae on forelegs used in grabbing prey body parts, stout forelegs to hold the prey firmly, and an extremely thick cuticle all over the body preventing injury from a counter bite of the prey. Palpimanus overwhelmed prey that was more than 200% larger than itself. In trials with another araneophagous spider, Cyrba algerina (Salticidae), Palpimanus captured C. algerina in more than 90% of cases independent of the size ratio between the spiders. Evidence indicates that both Palpimanus species possesses remarkable adaptations that increase its efficiency in capturing spider prey.     

An estimated 400–800 million tons of prey are annually killed by the global spider community

Spiders have been suspected to be one of the most important groups of natural enemies of insects worldwide. To document the impact of the global spider community as insect predators, we present estimates of the biomass of annually killed insect prey. Our estimates assessed with two different methods suggest that the annual prey kill of the global spider community is in the range of 400–800 million metric tons (fresh weight), with insects and collembolans composing >90% of the captured prey. This equals approximately 1‰ of the global terrestrial net primary production. Spiders associated with forests and grasslands account for >95% of the annual prey kill of the global spider community, whereas spiders in other habitats are rather insignificant contributors over a full year. The spider communities associated with annual crops contribute less than 2% to the global annual prey kill. This, however, can be partly explained by the fact that annual crop fields are “disturbed habitats” with a low buildup of spider biomass and that agrobiont spiders often only kill prey over short time periods in a year. Our estimates are supported by the published results of exclusion experiments, showing that the number of herbivorous/detritivorous insects and collembolans increased significantly after spider removal from experimental plots. The presented estimates of the global annual prey kill and the relative contribution of spider predation in different biomes improve the general understanding of spider ecology and provide a first assessment of the global impact of this very important predator group.     

Dynamics of spider colonization of apple orchards from adjacent deciduous forest

The colonization of foliage-dwelling spiders into insecticide-free “micro” apple orchards at two distances (10 and 50 m) from an adjacent source habitat of deciduous forest was quantified in southern Québec, Canada. Both colonization rate and composition of colonizing spider assemblages were affected by distance at this small spatial scale: the samples from the micro-orchards 10 m from the forest were intermediate in composition between the assemblages in the deciduous forest and those in the micro-orchards 50 m from the forest. Spider species found in micro-orchards were the same as those documented from collections from older, insecticide-free orchards although species evenness was higher in the older orchards indicating that species composition shifts over time within the habitat. Maintaining deciduous forest near to orchards will provide a source of annual colonists for spider populations within the orchard.     

Seasonal changes in burrow geometry of the common mole rat (Rodentia: Bathyergidae)

Sociality in mole rats has been suggested to have evolved as a response to the widely dispersed food resources and the limited burrowing opportunities that result from sporadic rainfall events. In the most arid regions, individual foraging efficiency is reduced, and energetic constraints increase. In this study, we investigate seasonal differences in burrow architecture of the social Cryptomys hottentotus hottentotus in a mesic region. We describe burrow geometry in response to seasonal weather conditions for two seasons (wet and dry). Interactions occurred between seasons and colony size for the size of the burrow systems, but not the shape of the burrow systems. The fractal dimension values of the burrow systems did not differ between seasons. Thus, the burrow complexity was dependent upon the number of mole rats present in the social group.     

Design and use of a time-sorting pitfall trap for predatory arthropods

An electronically-controlled time-sorting pitfall trap was designed which improves upon earlier models in having a 24-h rotation time, and in being capable of running continuously for at least one week without attention. Four prototypes were built, using readily available, inexpensive components, and used from June to October, 1995, to collect carabid beetles and linyphiid spiders in clover-undersown and weeded cabbages. The traps operated without malfunction during the hottest weather ever recorded in Aberdeen, when soil surface temperatures exceeded 40°C, and also during periods of torrential rain. Three spider and two carabid species dominated the catch, their nocturnal and diurnal patterns of activity being in general agreement with other published data. Some predators were apparently able to adapt to the contrasting vegetation in the two treatments, two species of Erigone spiders being nocturnal in weeded cabbages but tending to be diurnal in undersown crops.     

Supercooling ability in the house spider, Achaearanea tepidariorum: effect of field-collected and laboratory-reared prey

The influence of prey animals on the supercooling ability of the house spider, Achaearanea tepidariorum, was studied by feeding spiders with field-collected and laboratory-cultured prey animals. Irrespective of the prey species supplied, spiders given field-collected prey had a higher supercooling point than those given laboratory-cultured counterparts. This means that (1) the field prey animals contained some efficient ice nucleators, whereas the laboratory animals were free from such substances and (2) the ice nucleators must be of external origin. Several lines of evidence also suggest that, under natural conditions, potential prey animals for the house spider are highly contaminated with ice nucleators.     

Prey detection without successful capture affects spider’s orb-web building behaviour

Animals obtain information from past foraging experience to adjust their foraging activity according to their environment. The ability of spiders to obtain information from unsuccessful predation experiences was investigated by examining the effects on web building, a significant foraging investment, of prey detection without successful capture in the orb-web spider Cyclosa octotuberculata. Four treatments were employed: (1) successful capture and feeding: one syrphid fly was allowed to be captured and consumed by the spider on the web; (2) single prey-item detection: a syrphid fly was placed on the web to lure the spider, but was removed before capture; (3) five prey-item detection: above prey-item detection stimulus was given five times; and, (4) control: neither prey nor feeding on the web. While control spiders decreased the total thread length and capture area of their webs, prey-item detection spiders in both conditions increased them, indicating that the spider obtained information from unsuccessful predation experience to adjust their foraging investment. The fed spiders exhibited a significantly greater increase than the prey-detection-only spiders, suggesting that prey detection alone and prey detection with consumption had different informational effects. Total thread length did not differ between single and five prey-item detection spiders, but distance between two adjacent sticky spirals increased only in the former spiders, possibly because five times unsuccessful predations prevented spiders to reduce web stickiness. It suggests that the spider changed web morphology according to the number of prey detection.     

Does ontogenetic change in orb web asymmetry reflect biogenetic law?

Most orb web spiders face downward on the web hub, and their webs are vertically asymmetrical, that is, the lower part of the web is larger than the upper part and the ratio of the lower part to the whole web area increases as the spider grows. This phenomenon may reflect biogenetic law such that young animals exhibit a general ancestral trait whereas adults exhibit specific and derived traits. An alternative explanation is that vertical asymmetry may arise from the difference in time required by spiders to move up or down the web to capture prey. The present study tested these two hypotheses for Eriophora sagana. Subadults of this species build their webs with reverse asymmetry in that the upper part of the web area is larger than the lower part. In both subadults and adults, the upper proportion decreased with spider weight, and adult spiders built more symmetric webs. These results support the capture time difference hypothesis.     

Reverse positional orientation in a neotropical orb-web spider, <Emphasis Type="Italic">Verrucosa arenata</Emphasis>

Most orb-web spiders face downwards in the web. A downward orientation has been proposed to be the optimal strategy because spiders run faster downwards and thus can catch prey quicker. Consequently, orb-web spiders also extend their web in the lower part, leading to top-down web asymmetry. Since the majority of orb-web spiders face downwards, it has been difficult to test the effect of orientation on prey capture and web asymmetry. In this study, we explored the influence of reverse orientation on foraging efficiency and web asymmetry in Verrucosa arenata, a neotropical orb-web spider that faces upwards in the web. We show that reverse orientation does not imply reverse web asymmetry in this species. V. arenata spiders captured more prey in the lower part of the web but more prey per area on the upper part. The average running speeds of spiders did not differ between upward and downward running, but heavier spiders took longer to capture prey while running upwards. We discuss these findings in the context of foraging efficiency and web asymmetry.     

Flux characteristics of total dissolved iron and its species during extreme rainfall event in the midstream of the Heilongjiang River

The occurrence of extreme rainfall events and associated flooding has been enhanced due to climate changes, and is thought to influence the flux of total dissolved iron(TDI) in rivers considerably. Since TDI is a controlling factor in primary productivity in marine ecosystems, alteration of riverine TDI input to the ocean may lead to climate change via its effect on biological productivity. During an extreme rainfall event that arose in northeastern China in 2013, water samples were collected in the midstream of the Heilongjiang River to analyze the concentration and species of TDI as well as other basic parameters. The speciation of TDI was surveyed by filtration and ultrafiltration methods.Compared with data monitored from 2007 to 2012, the concentration of TDI increased significantly during this event, with an average concentration of 1.11 mg/L, and the estimated TDI flux reached 1.2 × 105 tons, equaling the average annual TDI flux level.Species analysis revealed that low-molecular-weight complexed iron was the dominant species, and the impulse of TDI flux could probably be attributed to the hydrological connection to riparian wetlands and iron-rich terrestrial runoff. Moreover, dissolved organic matter played a key role in the flux, species and bioavailability of TDI. In addition,there is a possibility that the rising TDI flux could further influence the transport and cycling of nutrients and related ecological processes in the river, estuary coupled with the coastal ecosystems, which merits closer attention in the future.     

Grassy field margins and arthropod diversity: a case study on ground beetles and spiders in eastern Austria (Coleoptera: Carabidae; Arachnida: Aranei, Opiliones)

In 1983, the species composition and diversity of carabids, spiders and harvestmen was investigated by pitfall traps in a grassy field margin and in an adjacent, biologically farmed wheat field at Obere Lobau, Vienna, Austria.The carabid fauna of both habitats was diverse; it showed a high degree of similarity, the abundant species in both habitats being represented by typical field species. Only Metophonus spp. predominantly lived in the margin. Numerous species of different habitat origin occurred in the margin at low abundances indicating transitional stay rather than permanent populations.In contrast, the spider fauna showed a low degree of similarity, with few agrobiotic species strongly dominating field fauna. In the field margin, a rich spider fauna containing rare elements indicates the general importance of grassy margins for nature conservation. The apparently low faunal exchange of spiders between the two habitat types is discussed. Harvestmen occurred mainly in the field margin; only two thermophilic species invaded the field.Seasonal fluctuations in the trapping numbers of Bembidion lampros, Platynus dorsalis and Brachinus explodens are interpreted as indicating migrations between field margin and field. Poecilus cupreus and certain other field species, however, apparently spend their entire life cycle in the field. The possible function of field margins for restoration of carabid losses due to impacts of intensive cultivation is discussed.It is concluded that both grassy field margins and sustainable farming methods, in particular biological farming, are necessary to maintain a diverse predatory arthropod fauna in agroecosystems.