Punctuated evolution of viscid silk in spider orb webs supported by mechanical behavior of wet cribellate silk

The origin of viscid capture silk in orb webs, from cribellate silk-spinning ancestors, is a key innovation correlated with significant diversification of web-building spiders. Ancestral cribellate silk consists of dry nanofibrils surrounding a stiff, axial fiber that adheres to prey through van der Waals interactions, capillary forces, and physical entanglement. In contrast, viscid silk uses chemically adhesive aqueous glue coated onto a highly compliant and extensible flagelliform core silk. The extensibility of the flagelliform fiber accounts for half of the total work of adhesion for viscid silk and is enabled by water in the aqueous coating. Recent cDNA libraries revealed the expression of flagelliform silk proteins in cribellate orb-weaving spiders. We hypothesized that the presence of flagelliform proteins in cribellate silk could have allowed for a gradual shift in mechanical performance of cribellate axial silk, whose effect was masked by the dry nature of its adhesive. We measured supercontraction and mechanical performance of cribellate axial silk, in wet and dry states, for two species of cribellate orb web-weaving spiders to see if water enabled flagelliform silk-like performance. We found that compliance and extensibility of wet cribellate silk increased compared to dry state as expected. However, when compared to other silk types, the response to water was more similar to other web silks, like major and minor ampullate silk, than to viscid silk. These findings support the punctuated evolution of viscid silk mechanical performance.     

Male Enchenopa treehoppers (Hemiptera: Membracidae) vary mate-searching behavior but not signaling behavior in response to spider silk

Finding and attracting mates can impose costs on males in terms of increased encounters with, and attraction of, predators. To decrease the likelihood of predation, males may modify mate-acquisition efforts in two main ways: they may reduce mate-searching efforts or they may reduce mate-attraction efforts. The specific behavior that males change in the presence of predator cues should depend upon the nature of risk imposed by the type of predator present in the environment. For example, sit-and-wait predators impose greater costs to males moving in search of mates. Here, we test whether cues of the presence of a sit-and-wait predator lead to a reduction in mate-searching but not mate-acquisition behavior. We used a member of the Enchenopa binotata complex of treehoppers—a clade of vibrationally communicating insects in which males fly in search of mates and produce mate-attraction signals when they land on plant stems. We tested for changes in mate-searching and signaling behaviors when silk from a web-building spider was present or absent. We found that males delayed flight when spider silk was present but only if they were actively searching for mates. These results suggest that males have been selected to reduce predation risk by adjusting how they move about their environment according to the cues of sit-and-wait predators.     

Sex-dependent immune response in a semelparous spider

Previous studies on arthropods showed that seasonality and parity in breeding considerably impact the direction of sex differences in immunocompetence, and it has been suggested that life span and the time window available for breeding play key roles in shaping sex-differences in immunity. One proposed mechanism behind this phenomenon is differential investment into life history traits in sexes. Here, we tested whether in a seasonally breeding semelparous arthropod sexes differ in their immunocompetence, predicting that females would show weaker immune response than males. We compared encapsulation efficiency (a well-established and widely used method for assessing immunocompetence) of freshly matured, virgin males and females of the lycosid spider Pardosa agrestis (Westring, 1861). On average, males mounted stronger immune response than females and the extent of encapsulation was positively associated with prosoma length in males, but not in females. Also, time until maturation was positively related to the extent of encapsulation in both sexes, but did not significantly affect adult prosoma length. We propose that sex-difference in encapsulation is likely shaped by combined effects of relatively higher costs of reproduction in females, narrow time window of reproductive activity, and the absence of trade-off between current and future reproduction.     

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.     

Host selection by a kleptobiotic spider

Why do kleptobiotic spiders of the genus Argyrodes seem to be associated with spiders of the genus Nephila worldwide? Observations following introduction of experimental insect prey of different sizes and weights on to host webs revealed that: (1) small prey are more effectively retained on the web of Nephila clavipes than on the web of another common host, Leucauge venusta. (2) N. clavipes did not consume small prey that accumulated on the web whereas larger, heavier prey were enveloped and stored. (3) We observed clear partitioning of prey items between N. clavipes and Argyrodes spp.; diet selection by Argyrodes did not overlap with that of N. clavipes but closely overlapped with that of L. venusta. (4) L. venusta responds very quickly to prey impact whereas N. clavipes is slower, offering a temporal window of opportunity for Argyrodes foraging. (5) The ability of L. venusta to detect and respond to small items also means that it acts aggressively to Argyrodes spp., whereas N. clavipes does not. Consequently, food-acquisition behaviours of Argyrodes were clearly less risky with N. clavipes compared with L. venusta. We conclude that when a kleptobiotic organism has a choice of various host species, it will opt for the least risky host that presents the highest rate of availability of food items. The fact that Nephila species present such characteristics explains the worldwide association with Argyrodes kleptobiotic spiders.     

No discrimination against previous mates in a sexually cannibalistic spider

In several animal species, females discriminate against previous mates in subsequent mating decisions, increasing the potential for multiple paternity. In spiders, female choice may take the form of selective sexual cannibalism, which has been shown to bias paternity in favor of particular males. If cannibalistic attacks function to restrict a male's paternity, females may have little interest to remate with males having survived such an attack. We therefore studied the possibility of female discrimination against previous mates in sexually cannibalistic Argiope bruennichi, where females almost always attack their mate at the onset of copulation. We compared mating latency and copulation duration of males having experienced a previous copulation either with the same or with a different female, but found no evidence for discrimination against previous mates. However, males copulated significantly shorter when inserting into a used, compared to a previously unused, genital pore of the female.     

Influence of CO<Subscript>2</Subscript> on the micro-structural properties of spider dragline silk: X-ray microdiffraction results

The mechanical properties of spider dragline fibres are altered by CO₂ exposure under anaesthetizing conditions during the spinning process. In order to relate these macroscopic changes to a microscopic model, the extrusion of dragline silk was studied by synchrotron radiation microdiffraction. A brief exposure of a female Nephila senigalensis spider to CO₂ results after an incubation time of less than 7 min in the extrusion of a thread (two fibres) swollen with water. The data are interpreted for a model of crystalline -sheet domains containing nanofibrils, which reinforce a network of protein chains. The protein network absorbs water, leaving the nanofibrils unaffected. A continuous flow of CO₂ results in a co-extrusion of a dragline thread and an isotropic silk fraction, which probably has a glycine-rich sequence. Long CO₂ exposure reduces the axial alignment of nanofibrils, presumably due to a partial destruction of the amorphous network.     

Changes in element composition along the spinning duct in a Nephila spider

The silk gland of the golden orb spider Nephila edulis connects to the exit spigot through a long S-shaped duct that assists in the formation of the thread. Previous evidence suggests that the epithelium of the distal (last) part of the duct is specialized for ion transport and that a proton pump is involved in this process. Here, we present evidence from SEM (scanning electron microscope)-EDAX (energy dispersive X-ray) microanalysis of rapidly frozen material maintained at approximately -150°C and from the use of pH indicators that the element composition and pH change progressively as the dragline silk dope (spinning solution) passes down the duct to form the thread. Na⁺ and Cl^- composition decreased while K⁺ and P and S increased. Indicators suggested that the pH dropped from 6.9ǂ.1 to 6.3ǂ.1. These novel findings suggest that the absorption of Na⁺ and secretion of the more chaotropic K⁺ may help the silk protein molecules to refold while the secretion of H⁺ may assist in this process and reduce the repulsive charges on them. This in turn may allow the molecules to approach one another more closely to crystallize. Thus precise control of the ionic environment within the spider's spinning duct may be important in forming a tough insoluble thread and when devising mimetic processes to spin silk proteins industrially.     

Symbiotic bacteria in hornet pupal silk

The silk weave spun by hornet larvae before undergoing pupal metamorphosis is composed of fibers and sheets, both containing symbiotic bacteria. The bacteria are secreted from the silk gland and are glued to the secreted silk, which is made up of amino-acid polymers. In the dark, it possesses at first an electric current amounting to several hundred nanoamperes (nA) (i.e., a thermoelectric property), and a high electric capacitance of up to several milliFarads (mF). This electrical charge is used gradually by the developing pupa. The symbiotic bacteria penetrate through slits in the coat of the silk fibers to the core or into pockets in the sheets, where they gradually digest parts of the silk weave, thereby nullifying its mechanical properties and facilitating in due time the egress of the imago from the puparium.     

Cross-habitat variation in the phenology of a colonial spider: insights from a reciprocal transplant study

In species with widespread distribution, populations found in markedly different environments can show differences in developmental traits. This, in time, can have an effect on reproductive success. Sources of variation in developmental traits can be genetic or environmentally induced. I examined the relationship between environmental and genetic influences on juvenile development in populations of the colonial spider, Parawixia bistriata, located at sites with different moisture regimes and associated environmental variables (e.g., prey availability). It was expected that individuals from different populations would show differences in developmental traits and that those differences will be associated with lower reproductive success at dry sites. I recorded the phenology and developmental traits of native and transplanted individuals in the field and estimated reproductive success based on clutch size. Colonies from wet versus dry sites showed different phenologies, with individuals at dry sites maturing later. Transplant results suggest plasticity in instar duration caused by environmental effects. Despite differences in resources and spider phenology, clutch sizes of native dry and wet populations were similar. Transplanted individuals, however, were differentially affected. Transplants from wet to dry sites (WD) showed lower growth rates and smaller clutches, whereas transplants from dry to wet sites had larger clutch sizes than in native habitat. Delayed maturation and failure to reproduce in WD individuals is associated with a lower tendency to capture prey in groups and less aggressive interactions during prey capture. Thus, despite negative environmental effects on development, dry native individuals have evolved non-developmental traits that allow successful reproduction.     

Geographical variation in sexual behavior and body traits in a sex role reversed wolf spider

Mating partners need to recognize, assess each other, and exchange information through behavioral events that occur before, during, and after mating. Sexual signals, as well as life history traits, are influenced by selective pressures and environmental factors that can vary across distant geographical areas. Allocosa senex is a sand-dwelling wolf spider which constructs burrows along the sandy coasts of Argentina, Brazil, and Uruguay. Females are the mobile sex that searches for males and initiates courtship. They prefer males which construct longer burrows, and males prefer virgin females in good body condition. The objective of this study was to compare sexual behavior patterns, as well as body characteristics and burrow dimensions, between two geographically distant locations of A. senex, one in Uruguay (Uruguayan location) and the other from central Argentina (Argentinean location). We found differences in the number of male abdominal vibrations, male and female touches during mating, and number of erections of male leg spines, which all were higher in matings of Argentinean pairs. On the other hand, male body mass and female body condition were higher in Uruguayan individuals. The wide distribution of A. senex could be determining variations in the biotic and abiotic features that affect the species, generating differences in the strength of selective forces acting on individuals from the two studied locations.     

Temporal variation in size-assortative mating and male mate choice in a spider with amphisexual care

Males should be more selective when they have a high investment in reproduction, especially in species with biparental or paternal care. In this context, male mate choice can promote size-assortative mating (SAM) when (1) large males win intrasexual disputes, (2) large females are more fecund, and (3) males prefer larger females to smaller ones. In the spider Manogea porracea, males exhibit high reproductive investment by building their webs above those of females and exhibiting extended care of offspring in the absence of females. Under these circumstances, we expect the occurrence of SAM and male preference for large females. Herein, we performed observations and experiments in the field to evaluate the hypotheses that (1) M. porracea mates assortatively by size and (2) SAM is influenced by male mate choice. Furthermore, we measured variables that could affect mating patterns, the sex ratios, and densities of both sexes. Pairing in M. porracea was positively size-assortative in 2012, but not in 2013. Large males won most disputes for mates and preferred larger females, which produced more eggs. The inconsistency in detection of SAM was due to population dynamics, namely variations in sex ratio and population density across the breeding season. Furthermore, we found that the significance of male mate choice on sexual selection of body size in M. porracea strongly depends on the competition intensity for mating opportunities. The traditional sexual selection hypothesis of SAM needs to be reviewed and must include measures of competition intensity.     

Remote monitoring of vibrational information in spider webs

Spiders are fascinating model species to study information-acquisition strategies, with the web acting as an extension of the animal’s body. Here, we compare the strategies of two orb-weaving spiders that acquire information through vibrations transmitted and filtered in the web. Whereas Araneus diadematus monitors web vibration directly on the web, Zygiella x-notata uses a signal thread to remotely monitor web vibration from a retreat, which gives added protection. We assess the implications of these two information-acquisition strategies on the quality of vibration information transfer, using laser Doppler vibrometry to measure vibrations of real webs and finite element analysis in computer models of webs. We observed that the signal thread imposed no biologically relevant time penalty for vibration propagation. However, loss of energy (attenuation) was a cost associated with remote monitoring via a signal thread. The findings have implications for the biological use of vibrations by spiders, including the mechanisms to locate and discriminate between vibration sources. We show that orb-weaver spiders are fascinating examples of organisms that modify their physical environment to shape their information-acquisition strategy.