Male-male competition and mating success in the orb-web spider,Nephila clavata,with reference to temporal factors

Seasonal occurrence patterns of adults of both sexes, intensity of male-male interactions, and mating success in the spider,Nephila clavata, were examined in the field. Adult males began to attend female webs about 2 weeks before female maturation. Large adult males were abundant in the early breeding season, but small males increased later in the season. From the distribution of males among female webs and size relationship of males within a web, male-male interactions seemed to be more intense when most females were still subadult. This was verified by a field experiment in which males were artificially introduced to female webs that were attended by other males. It was found that the probability of introduced males remaining on subadult female webs was lower than that on adult webs. As mating occurred mostly in the period shortly after the female final molt and first male sperm precedence was known in all spiders reported so far, intense male-male competition on subadult female webs seemed to be reasonable. Male longevity had an important influence on the mating success of males with just-molted females. Mating success was also affected by the relative body size of males present in a given period. Since larger males occupied the position closest to females within a web and stayed there longer, relative body size appeared to influence mating success through male-male competition. Female body size at maturation declined with time; hence, males that attained sexual maturity earlier had the advantage of mating with larger and more fecund females. Therefore, early maturation as well as larger size seem to be two important trairs influencing the reproductive success of males.     

The biology of Euryattus sp. indet., a web-building jumping spider (Araneae, Salticidae) from Queensland: utilization of silk, predatory behaviour and intraspecific interactions

The biology of an aberrant saltioid spider, Euryattus Thorell sp. indet., is described from observations in a Queensland rain forest and the laboratory. Pronounced morphological and behavioural changes occur during post-embryological development. Juveniles spin webs, but adult females make 'suspension nests' by suspending a curled-up leaf by heavy guylines from a rock ledge or vegetation. Adult males spin neither webs nor suspension nests. Females oviposit inside their suspension nests, but if denied access to leaves for suspension, they spin and oviposit in webs similar to those spun by juveniles. The flat, papery egg sacs of Euryattus are atypical for a salticid, being more like the egg sacs of many of the Gnaphosidae. Intraspecific display behaviour has characteristics in common with typical salticids, but also includes unique features. Male courtship includes vibratory displays performed on the surface of the suspended leaf. Mating occurs inside the curled-up leaf. Males co-habit with sub-adult females in suspension nests. Females take over suspension nests of other females and eat each other's eggs. Normal locomotion, intraspecific interactions, and predatory behaviour are characterized by frequent leaping. Euryattus routinely makes long and accurate leaps on to prey, including flying insects intercepted in mid-air and spiders and insects located in alien webs. The phylogenetic implications of the unusual characteristics of Euryattus are considered.     

Small Organic Solutes in Sticky Droplets from Orb Webs of the Spider Zygiella atrica (Araneae; Araneidae): β‐Alaninamide Is a Novel and Abundant Component

In northeastern North America, Zygiella atrica often build their orb webs near the ocean. We analyzed individual field‐built Z. atrica webs to determine if organic low‐molecular‐mass solutes (LMM) in their sticky droplets showed any unusual features not previously seen in orb webs of other species living in less salty environments. While two of the three most abundant organic LMM (putrescine (butane‐1,4‐diamine) and GABamide (4‐aminobutanamide)) are already well‐known from webs of inland spiders, the third major LMM, β‐alaninamide (3‐aminopropanamide), a homolog of GABamide, has not been detected in sticky droplets from any other araneoid spiders (27 species). It remains to be established, however, whether or not use of β‐alaninamide is related to proximity to saltwater. We observed variability in organic LMM composition in Z. atrica webs that appeared to be influenced more by an undetermined factor associated with different collecting locations and/or collection dates than by different genders or instars. Shifts in composition when adult females were transferred from the field to the laboratory were also observed. Structural similarities and inverse correlations among β‐alaninamide, GABamide, and N‐acetylputrescine suggest that they may form a series of LMM fulfilling essentially the same, as yet unknown, role in the webs of those species in which they occur.     

Multiple web-borne pheromones in a spider Frontinella pyramitela (Araneae: Linyphiidae)

The bowl-and-doily spider, Frontinella pyramitela, is a common inhabitant of low vegetation throughout most of temperate North America. All instars build concave-upward, bowl-shaped, nonviscid webs supported above and below by meshworks of silk. Previous studies of this species have revealed that chemical(s) on the silk of adult females elicit both courtship behaviour and positive geotaxis from adult males that contact the silk. This study demonstrates (1) that two different contact pheromones are responsible for the dual action of the silk of adult females and (2) that the webs of different age and sex classes of bowl-and-doily spiders (including the webs of adult males) contain functionally different mixtures of the two pheromones.     

Leaf domatia reduce intraguild predation among predatory mites

1. Although theory suggests that intraguild predation destabilises food webs and may result in exclusion of species, empirical observations of food webs reveal that it is a common interaction. It has been proposed that habitat structure reduces the interaction strength of intraguild predation, thus facilitating the coexistence of species. 2. This was tested using acarodomatia, tiny structures on plant leaves, and predatory mites, which usually reside in these domatia. Sweet pepper plants (Capsicum annuum L.) were used, which possess domatia consisting of tufts of hair, and coffee plants (Coffea arabica L.) with pit‐shaped domatia. 3. On sweet pepper, the predatory mites Neoseiulus cucumeris Oudemans and Iphiseius degenerans Berl. feed on each other's juveniles. Larvae of each of the species were therefore used as intraguild prey with adult females of the other species as intraguild predators. On coffee, a similar set‐up was used, with larvae and adult females of Amblyseius herbicolus Chant and Iphiseiodes zuluagai Denmark & Muma as intraguild prey and intraguild predators, respectively. 4. Domatia on detached, isolated sweet pepper and coffee leaves were either closed with glue or left open, after which larvae and adult predators were released. As a control, larvae were released on leaves with open or closed domatia without an adult predator. 5. Survival of larvae was high in the absence of the adult (intraguild) predator. In the presence of the intraguild predator, survival was significantly higher on leaves with open domatia than on leaves with closed domatia. 6. This shows that even such tiny structures as plant domatia may significantly affect the interaction strength of intraguild predation.     

Impact of predation by spiders on a territorial damselfly (Odonata: Calopterygidae)

Summary Predation by orb-weaving spiders and crab spiders on the damselfly Calopteryx haemorrhoidalis was studied at a small stream in Southern France. One species of orb-weaving spider, Larinioides folium, caught 76% of the damselflies which fell prey to spiders. Displacement experiments on spiders on sections of bank and the positioning of webs in male territories show that the density and distribution of damselflies is not influenced by orbweb density or by the position of webs. Predation rates corresponded to orb-web density, but neither for sex nor for stage was there a relationship with damselfly density. Mean daily predation rates ranged between 0.9% for females and 4.1% for adult males. Predation risk to adult damselflies by orb-weaving spiders was male biased, whereas among tenerals there was no bias. Males were captured more frequently at territories near the water. Captures show a maximum at noon when territorial disputes of adult males were most frequent. After orbwebs were placed within territories predation rate of males was strongly increased. Predation risk to adult females in the direct vicinity of the stream was less than in the bank vegetation where they perch close to orbwebs. The risk of predation by crab spiders, which catch damselflies at their perching sites, was not sex-biased.     

Ontogenetic Changes in Web Design in Two Orb‐Web Spiders

The first orb web built by newly hatched spiders resembles the adult web in its overall form and structure. However, many details show ontogenetic changes. One possible explanation for these changes is that the tiny early‐instar spiders with their minute brains will make more mistakes and build less ‘perfect’ orb webs than older and larger juveniles and adults. To test this hypothesis, known as the size limitation hypothesis, I analysed orb webs from three developmental stages, spiderlings, juveniles and adult females, in two neotropical orb‐web spiders, the araneid Eustala illicita and the nephilid Nephila clavipes. Neither species showed clear signs of being behaviourally limited or more prone to committing errors as spiderlings than were older juveniles or adults. These findings therefore do not support the size limitation hypothesis in either species. Finally, I looked for evidence of the ‘biogenetic law’, which predicts that juveniles should build less derived orb webs than the adults. Evidence for this was found in E. illicita, but not in N. clavipes.     

Stabilimenta of Philoponella vicina (Araneae: Uloboridae) and Gasteracantha cancriformis (Araneae: Araneidae): Evidence Against a Prey Attractant Function

Both the uloborid Philoponella vicina and the araneid Gasteracantha cancriformis spiders sometimes placed silk stabilimenta on non-orb "resting webs" that consisted of only one or a few lines. These webs completely lacked sticky silk, so their stabilimenta could not function to attract prey. Some non-orbs were built by spiders when their orb webs are damaged. These observations contradict the prey attraction camouflage hypothesis for stabilimentum function, but are compatible with the spider camouflage and web advertisement to avoid web destruction hypotheses.     

Developmental changes in barrier web structure under different levels of predation risk inNephila clavipes (Araneae: Tetragnathidae)

Individuals of the orb-weaving spider Nephila clavipesbuild complex webs with a region used for prey capture, the orb, and tangle webs opposite either face, the barrier webs. Barrier webs have been hypothesized to serve a variety of functions, including predator defense, and the primary function of the barrier web should be reflected in the relative size of the barrier to the orb under varying conditions of foraging success and predation risk. To investigate the effects of predation pressure and foraging success on barrier web structure, I conducted a comparative study in three disjunct populations that differed in predation risk and foraging success. Although both the orb web and the barrier webs are silk, there was no indication of a foraging-defense trade-off. Barrier web structure did not change during seasonal shifts in orb web size related to changes in preycapture rate, and barrier web silk density and orb radius were positively correlated. The hypothesis that the construction of barrier webs is in part a response to predation pressure was supported. Barrier webs do deflect attacks by some predators, and barrier webs built by small spiders, suffering frequent predation attempts, had a higher silk density than barrier webs built by larger individuals. Additionally, barrier web complexity decreased at a later age in areas with higher predation risk.     

Effects of an experimental increase in prey abundance upon the reproductive rates of two orb-weaving spider species (Araneae: Araneidae)

Summary A field experiment was performed to determine if food is a limited resource for adult females of two species of orb-weaving spiders, Mecynogea lemniscata and Metepeira labyrinthea. Spiders built webs after being added to open experimental units located in a mixed deciduous-pine forest in Maryland, USA. Each unit was a frame supporting dead branches of the type used by both species for anchoring webs. Spiders on half the units were exposed to natural prey densities only, while each spider on the other units was given laboratory-reared flies in order to increase prey availability above natural levels. Supplemental feeding continued for 2.5 months. At the end of the experiment all egg sacs were removed from the units.Providing additional prey did not increase the survival rate on the units (net effect of mortality, emigration and immigration). However, both species responded to additional prey by significantly increasing the number of eggs produced per female, indicating that food was a limited resource for these species. Median egg production per female increased from 34 to 62 for Mecynogea lemniscata and from 65 to 145 for Metepeira labyrinthea. Egg weight was not affected.Feeding rates and nearest neighbor distances were determined for spiders in non-experimental populations, which permitted evaluation of the experiment's naturalness. The effects of food supply upon the reproductive rates of the two species are discussed in relation to their numerical response and population dynamics.     

Trail-following behavior by males of the wolf spider, <Emphasis Type="Italic">Schizocosa ocreata</Emphasis> (Hentz)

Male spiders are able to detect and respond to chemical cues deposited by females in the environment. In many species, detection of these chemicals may be the first indication a male has to the presence of a nearby female. In wolf spiders (Lycosidae), which do not produce webs, females leave a trail of silk and chemical cues as they move through the leaf-litter habitat. Males could increase encounter rates with receptive females if they were able to follow these trails. We used behavioral assays to determine whether male Schizocosa ocreata (Hentz) wolf spiders are able to detect and respond to cues resulting from a single-pass trail by a female, and whether they are able to determine the direction of female travel. Our focal males responded to virgin adult female trails with following behavior, but showed no propensity to follow trails from other conspecifics (subadult females or males). While males were able to follow a female trail, our observations and analysis indicates that they are not able to determine trail directionality.     

Nest-building by the virgin female mouse exposed to ultrasound from inaccessible pups

When a naïve virgin female mouse is exposed to an inaccessible litter so that it perceives the odour and the ultrasounds from the pups, its nest-building behaviour undergoes changes that are different from changes occurring in a control condition providing exposure to an inaccessible adult female. Virgin females exposed to a litter built their nest as near as possible to the pups. Other characteristics of their nest-building behaviour were influenced differently according to the type of ultrasound that the pups were emitting. In a condition where pups were mainly calling in response to cold, virgin females built heavier nests than control females. When mainly calls in response to handling were given by the pups, the females built lighter nests than control females. Finally, there was also some suggestion that the females exposed to ultrasounds from handled pups tended to chew the nest-material more than either control females or females exposed to ultrasound from cooled pups.     

Mate guarding, competition and variation in size in male orb-web spiders, Metellina segmentata: a field experiment

In the common orb-web spider Metellina segmentata, males are more powerfully built with longer legs, although females are heavier because of their egg load. Males guard females before attempting to mate, and there is considerable male-male competition because of the male-biased operational sex ratio. We used a field removal experiment to examine (1) seasonal changes in the average morphology of guarding males and (2) whether there is a pool of small males that is excluded from the webs of females. Morphological measures were subjected to a principal components analysis and changes in PC scores were examined for seasonal effects and the effects of previous removal of males. The size of guarding males (PC1) increased over the season, suggesting that smaller males were increasingly excluded from webs, but the condition of guarding males (PC2) decreased, indicating that energy reserves are depleted because the males gain little access to food during the reproductive season. When guarding males were removed, smaller males were able to take up residence. Our results show that large males have a clear advantage in monopolizing females. We discuss the manner in which selection acts to maintain large male size in this spider.Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.      

The strange occurrence of male cannibalism on adult females in snakes

Cannibalism is widespread in the animal kingdom. The most common, well-known cases are found in some arthropod systems where the larger females occasionally consume the smaller males before, during, or after mating (i.e., sexual cannibalism). The killing and consumption of females by males are, on the other hand, seemingly much rarer and theoretically more difficult to explain, at least among adults, because the consumption of females would result in the loss of potential mates for males. Here, we use observations submitted to a community science reporting system to describe three instances of adult males feeding on adult females in the Montpellier snake, Malpolon monspessulanus, in the south of France. We discuss our observations in the theoretical framework of cannibalism to provide potential explanations for these observations and briefly review empirical findings of cannibalism in vertebrates, which confirms that this sex-reversed form of sexual cannibalism is rare in nature.     

Behaviour of colonial and solitary spiders of the theridiid species Anelosimus eximius

Most Anelosimus eximius live in colonies, but a few females emigrate short distances and establish small, individual webs. Field studies were conducted on one colony and nearby smaller webs in order to describe communal activities and division of labour, and to note costs and benefits of remaining in the parental colony and emigrating. Adult and juvenile females repaired the web and captured prey. Adult females, rather than juveniles, cared for egg sacs and fed young. When colonial egg sacs were abundant, females moved from sac to sac. Several colonial females regurgitated food to spiderlings which had recently emerged from a particular sac. In smaller webs inhabited by two or three females, the mother cared for the sac but all females fed the young by regurgitation. Males rarely participated in communal activities. Advantages of colony living include protection from predators, the availability of large prey and, in the event of a female's death, the care of her egg sac and feeding of her young. Cannibalism of the egg sac is a potential cost of coloniality, affecting about 10–20% of colonial sacs. Cannibalism was not observed in the smaller webs. However, the costs of emigrating are very high: most of the solitary females disappeared, leading to interspecific predation on their eggs and young.     

Aggression with mixed age-sex groups of bolivian squirrel monkeys following single animal introductions and new group formations

The social organization of the Bolivian squirrel monkey (Saimiri boliviensis boliviensis) is thought to be sexually segregated, with males and females forming separate social groups during the nonbreeding season. To investigate the influence of this social order on patterns of aggression, controlled single animal introductions within established groups and establishment of new groups were studied in a systematic manner. Behavioral observations were made using an all-occurrences technique to sample all agonistic interactions. In study I, 4 animals of each age-sex class (adult males and females, juvenile males and females) were introduced one at a time into 4 different social groups composed of 1 adult male and 8 to 10 adult females. Behavioral observations were made prior to and after the introductions. Newly introduced adult males received significantly more contact aggression than other age classes. Only the adult females increased aggression after the introduction of new animals. In study II, new social groups were formed and behavioral observations were made following formation. One group was formed from 7 pairs of familiar females and an unfamiliar adult male. This group had a high frequency of aggression during the first half hour, with contact aggression rising to peaks at 3 and 5 h. The second group consisted of 10 familiar females, 2 pairs of familiar females and an unfamiliar male. There was a significant peak in contact aggression 3 h into the observation.     

The presence of webbing affects the oviposition rate of two-spotted spider mites, <Emphasis Type="Italic">Tetranychus urticae</Emphasis> (Acari: Tetranychidae)

Several species of tetranychid mites including Tetranychus urticae Koch (Acari: Tetranychidae) construct complicated three-dimensional webs on plant leaves. These webs provide protection against biotic and abiotic stress. As producing web is likely to entail a cost, mites that arrive on a leaf with web are expected to refrain from producing it, because they will gain the benefit of protection from the existing web. Mites that produce less web may then allocate resources that are not spent on web construction to other fitness-enhancing activities, such as laying eggs. To test this, the oviposition rate of T. urticae adult females was examined on leaves with web. As a control, we used leaves where the web had been removed, hence both types of leaves had been exposed to conspecifics previously and were thus damaged. On leaves with web, the oviposition rate of T. urticae females was higher than on leaves where the web had been removed. Therefore, the presence of web constructed by conspecifics enhanced the oviposition rate of T. urticae females. This provides indirect evidence that mites use the web constructed by conspecifics and thereby save resources that can be allocated to other traits that enhance reproductive success.     

A spider population in flux: selection and abandonment of artificial web-sites and the importance of intraspecific interactions in Lephthyphantes tenuis (Araneae: Linyphiidae) in wheat

Lepthyphantes tenuis, a small sheet-webbuilding linyphiid spider is one of the most abundant spider species of cereal fields in Europe. In the present study we examined the process of web-site selection and web-site tenacity by adult females of this species in a winter wheat field. Spiders were selective in their choice of web-site. Different immigration rates into various manipulated web-sites, in field and laboratory, suggested that structural support and suitable micro-climate (high humidity) are the most important factors in the selection. Small holes dug in the ground were the most favoured web-sites. Web-site occupation was influenced by the presence of other conspecific spiders. Territorial contests occurred between spiders attempting to occupy the same web, these almost invariably led to the take-over of the web when the intruder was heavier. Interference, but also a certain level of tolerance, between spiders within the same web-site but in different webs was suggested by direct and indirect evidence. Many holes supported two or even three spiders in vertically stratified webs. Leaving probability of marked spiders was significantly higher in multiply occupied holes than in holes with a single web. Comparison with the results of a no-interference stochastic model showed that multiple occupancy in nature is less frequent than predicted by the model. There was further evidence for weak extra-web-interference between spiders in that multiple occupancy was even less frequent and overall occupancy was lower in web-sites which were packed close to each other. However, a level of tolerance for crowding is shown by the fact that closely packed hole colonies supported a spider density 13 times higher than in natural web-sites in the field. A marking experiment was carried out to gain information on web-site tenacity (i.e. the length of time a spider spends in a web-site) and abandonment. The average duration of tenacity was less than 2 days. A random loss function gave a good fit to the tenacity distribution and suggested that spiders abandoned web-sites randomly with a fixed leaving probability of c. 0.5. Individual webs were often used consecutively by more than one spider, and some spiders built more than one web in the same web-site. Calculations showed that abandonment is the most frequent leaving mode, while take-over by contest between spiders and disappearance due to destruction were some-what less frequent and equally likely modes of ending tenacity. It is suggested that the apparent contradiction between the selectiveness and competitiveness of spiders for web-sites and the relatively short tenacity observed can be resolved by hypothesising that spiders leave websites soon because they apply the strategy of spreading risk: spiders by frequently moving from one web-site to another distribute their reproductive efforts across several localities. This hypothesis is further supported by changes in web-site preference and ballooning behaviour at the onset of the reproductive stage in L. tenuis.     

The spinning apparatus of Polenecia producta (Araneae,Uloboridae): Structure and histochemistry

Summary The spinning apparatus of the uloborid spider Polenecia producta was studied to complete previous studies on the same family of spiders. The structure of spinnerets and spigots, under scanning electron microscopy, and the main anatomical and histochemical characteristics of the spinning glands of adult females and males are described. In addition some observations on the spinning apparatus at three successive stages of development are made. There are nine kinds of silk glands in Polenecia, i.e. one more (aciniform — B glands) than found in other uloborids. The spinning apparatus of Polenecia is, therefore, the most complex so far known. It is also more complex than that presently known of Araneoidea. The characteristics of the spinning glands of Polenecia are compared with those of other uloborids. Present knowledge of the spinning apparatus of uloborids leads to a renewed discussion of the origin of the orb web in this family and in araneids. It is concluded that these two types of orb webs emerged from independent evolutionary processes.     

Is Male Cohabitation Costly for Females of the Spider Stegodyphus lineatus (Eresidae)?

Reproductive interests of females and males can vary over a number of issues, including the number of matings and the occurrence and duration of mate guarding and cohabitation. The mating system of the spider Stegodyphus lineatus (Eresidae) is characterized by an exceptional sexual conflict where males induce females to remate by committing infanticide. Females experience high costs because of this male strategy, while males benefit. During the mating season, males tend to stay with females for several days. We examined whether this male strategy of cohabitation is also disadvantageous for females of S. lineatus. A field experiment revealed that male presence in a female's nest negatively affected her body condition, whereas cohabiting males gained weight because they fed on prey caught in webs of females. As a response, females did not renew their webs when males were present. Thus, females with a cohabiting male experienced the combined cost of prey loss and loss of time available for foraging. These costs are expected to increase with every additional cohabiting male. Mated females behaved more aggressively toward males than did virgin females. This behavior may be interpreted as an adaptive response to reduce mating costs.