Age-related sequential web building in the colonial spider Metepeira incrassata (Araneidae): an adaptive spacing strategy

Colonial orb-weaving spiders provide insight into the proximate mechanisms by which social animals space themselves within a group. We examined mechanisms for the temporal patterns of web building that determine individual positions in Metepeira incrassata (Araneidae) colonies. The spiders display a characteristic age-related sequence of daily web building, with larger spiders completing their webs significantly earlier than smaller ones. We used data on behavioural interactions, web building, prey capture and predator attacks to evaluate four hypotheses. (1) Larger spiders are better competitors and pre-empt optimal spatial positions. (2) Smaller spiders reduce competition with larger individuals by building webs later. (3) Prey captured by different size classes is available at different times. (4) Differential predation risk determines web-building times. Large individuals dominated behavioural interactions. Disturbances by larger spiders during web construction significantly delayed the completion of smaller individuals' webs and precipitated movements to new web sites. One prediction of the first hypothesis, that spatial needs translate into earlier building, was confirmed by significantly earlier web building by mature females with egg sacs (which are unable to move their egg sacs) compared with same-sized females without eggs (which can change locations freely). Experiments to determine whether the presence of large spiders inhibited the web building of smaller individuals were equivocal. Prey availability and risk of predation are not factors affecting web-building patterns. Sequential web building appears to be a result of both larger spiders competing to pre-empt space from one another and smaller individuals attempting to reduce conflict during web construction. Sequential web building is a proximate mechanism that influences spacing among colonial orb-weaving spiders and helps shape the typical hierarchical size distribution of spiders within the colony. Similar spacing mechanisms may be seen in colonial birds and marine invertebrates. Copyright 2000 The Association for the Study of Animal Behaviour.     

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.     

Subsocial spiders in space and time: A fine scale approach to the dynamics of dispersal

Less than 0.2% of all spider species live in close associations with conspecifics. Among these, subsocial spiders show characteristics of both solitary spiders (e.g., individuals disperse for breeding) and social spiders (e.g., prolonged cooperative behaviours at least prior to independent reproduction). Dispersing individuals build small webs, usually with one inhabitant, whereas colonies are large webs with plant debris and harbouring multiple females. We studied the spatiotemporal dynamics of dispersal in the subsocial spider Anelosimus baeza. We followed the occupancy of all colonies and dispersal webs over the breeding season by mapping the number and sex of spiders with respect to their location in three dimensions. We studied the settlement patterns of new webs and fluctuation in web occupancy through movement between occupied and abandoned webs of colonies and dispersal webs. The occupancy of webs was highly dynamic with changes occurring at small time scales. The similarity in the patterns of web occupancy by females among dispersal webs was partially explained by their spatial and their temporal proximity. Our results suggest that dispersal webs may be used by spiders as a temporary refuge by both sexes during the breeding season. Patterns described here suggest new approaches to dispersal studies in group living spiders.     

The Effect of Wind Exposure on the Web Characteristics of a Tetragnathid Orb Spider

Studies on spiders in their natural habitats are necessary for determining the full range of plasticity in their web-building behaviour. Plasticity in web design is hypothesised to be important for spiders building in habitats where environmental conditions cause considerable web damage. Here we compared web characteristics of the orb spider Metellina mengei (Araneae, Tetragnathidae) in two different forest habitats differing in their wind exposure. We found a notable lack of differences in web geometry, orientation and inclination between webs built along an exposed forest edge and those built inside the forest, despite marked differences in wind speed. This suggests that M. mengei did not exhibit web-building plasticity in response to wind in the field, contrasting with the findings of laboratory studies on other species of orb spiders. Instead, differences in prey capture and wind damage trade-offs between habitats may provide an explanation for our results, indicating that different species employ different strategies to cope with environmental constraints.     

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.     

Female body size, fecundity parameters and foundation of new colonies in Anelosimus jabaquara (Araneae, Theridiidae)

Summary. As in other social spider species, subadult Anelosimus jabaquara females found new colonies after solitary dispersal. Some individuals, however, usually remain and reproduce in their natal nests. To test the hypothesis that large females disperse more often than smaller ones, we compared the body size of A. jabaquara females that remained in their natal colonies with those that left to build solitary webs. We also compared clutch size, egg diameter, total egg volume and spiderling size in both conditions. Emigrating females were significantly larger and laid larger clutches. The smaller females that had not dispersed laid significantly larger eggs, although their total egg volume was lower. Spiderlings of solitary females were smaller and had a smaller range of size variation than those from colonies. We discuss the implications of these results in terms of costs and benefits of dispersion for spiders in different nutritional conditions.     

Colony size and individual fitness in the social spider Anelosimus eximius

ABSTRACT The effects of colony size on individual fitness and its components were investigated in artificially established and natural colonies of the social spider Anelosimus eximius (Araneae: Theridiidae). In the tropical rain forest understory at a site in eastern Ecuador, females in colonies containing between 23-107 females had india significantly higher lifetime reproductive success than females in smaller colonies. Among larger colonies, this trend apparently reversed. This overall fitness function was a result of the conflicting effects of colony size on different components of fitness. In particular, the probability of offspring survival to maturity increased with colony size while the probability of a female reproducing within the colonies decreased with colony size. Average clutch size increased with colony size when few or no wasp parasitoids were present in the egg sacs. With a high incidence of egg sac parasitoids, this effect disappeared because larger colonies were more likely to be infected. The product of the three fitness components measured-probability of female reproduction, average clutch size, and offspring survival-produced a function that is consistent with direct estimates of the average female lifetime reproductive success obtained by dividing the total number of offspring maturing in a colony by the number of females in the parental generation. Selection, therefore, should favor group living and itermediate colony sizes in this social spider.     

Maternal defensive behaviors of <Emphasis Type="Italic">Uloborus</Emphasis> sp. (Araneae,Uloboridae): behavioral repertoire and influence of clutch size and female size on female aggressiveness

Maternal care in spiders often involves behaviors associated with the protection of eggs and spiderlings against parasitoids and predators (including conspecifics). The females of several species have been documented to move their egg sacs away from natural enemies or to invest in active defense behaviors against web invaders, such as parasitoid wasps or araneophagic spider species, to protect their brood. In this study, we present observations of protective behavior by Uloborus sp. females carrying egg sacs. We also investigated whether brood size and female size influence female aggressive behaviors and response time against an artificial source of disturbance. Females carrying egg sacs almost immediately perceived and reacted aggressively against the artificial stimulus, whereas females without egg sacs moved away or ran to the web margins, avoiding the source of disturbance. The aggressive response was independent of clutch size and female body size, indicating that all females will risk interacting with potential agents of egg mortality. This systematic response by all females with egg sacs may be important for reducing the incidence of attack by the egg predator wasp Bathyzonus sp. (Ichneumonidae).     

Variable dependence on detrital and grazing food webs by generalist predators: aerial insects and web spiders

Recent studies have shown that organisms from the detritus food web subsidize generalist predators in aboveground food webs, but its significance in space and time is largely unknown. Here we report seasonal dynamics of aerial insects from grazing and detritus food webs in both forest and grassland habitats, and show how these patterns influence the dependence of web spiders on the detritus food web. Detrital insects were more abundant in spring, decreased in summer, and then increased slightly in autumn. This pattern was most conspicuous in Nematocera. Due to different seasonal activity patterns of grazing and detrital insects, the proportion of detrital insects was greater in spring and autumn. Detrital insects were relatively more abundant in the forest than in the grassland. Prey captured by web spiders generally reflected seasonal and spatial patterns of aerial insect abundance. In particular, Leucauge spiders reversed their dependence on the two food webs seasonally. Body size of spiders was negatively correlated with the proportion of detrital prey, suggesting that the detrital subsidy is essential for relatively small predators. This size effect probably resulted from interaction of the following two factors: 1) the maximum body size of prey that can be caught increased with spider body size, 2) larger body size classes of aerial insects included a higher proportion of insects from the grazing food web.     

Forest fragmentation and colony performance of forest tent caterpillar

We monitored survival and egg production in colonies of Malacosoma disstria in the understorey to examine relationships between colony performance and forest structure at several spatial scales. As forest cover increased, fewer moths survived. Increased mortality from a virus and a parasitoid assemblage were the main causes of this result. Forest cover measured at the smallest spatial scale (53 × 53 m) was the best predictor of colony performance. We attribute these effects of fragmentation to fine-scale differences in microclimate betweeen edge and interior habitats (edge effects).     

Individual decisions and group dynamics: why pholcid spiders join and leave groups

To examine the role of individual variation in the dynamics of group formation, I conducted a mark-recapture study and a series of laboratory and field experiments with Holocnemus pluchei spiders (Araneae: Pholcidae). These spiders can either share webs or live alone, and individuals shift frequently between these strategies. Spiders' decisions were influenced by size and recent feeding success. In the laboratory, small hungry spiders introduced into a web that held a larger conspecific resident were more likely than small well-fed spiders to abandon the web and build their own web. This behaviour pattern gradually reversed as spiders grew: large hungry spiders were more likely than large well-fed spiders to stay in the shared web. When I introduced spiders into empty webs, they were more likely to stay compared with spiders tested with conspecifics. However, hunger level also influenced behaviour even when conspecifics were not present. Food-deprived spiders were more likely to abandon webs and build their own, consistent with the idea that spiders were following a win-stay/lose-shift strategy. In the field, spiders were more likely to stay in webs overnight when they were given supplemental food. In another experiment, spiders that were found building webs in cleared areas were smaller and thinner than average. Finally, I tested whether the size of the intruder or the resident affected whether spiders joined webs. Large intruders were more successful at remaining in webs than smaller intruders, although spiders of all sizes had some success in joining groups. Additional synthetic theoretical work is needed to integrate the complex processes underlying the formation and persistence of groups.     

The effect of a single colony of the red wood ant,Formica polyctena,on the spider fauna (Araneae) of a beech forest floor

Summary The predation on spiders in a forest ecosystem by a colony of red wood ants, Formica polyctena, was estimated using a barrier to isolate the colony. Of the ants' total prey, 4.6% were spiders. In order to estimate the effect of F. polyctena within their hunting area on the spider population, the spiders' population density was studied inside and outside the hunting area. Samples of the forest floor were taken, spider webs were counted and pitfall traps were used. No significant difference was found in density or composition of the spider fauna inside and outside the hunting area.     

Web spider's dilemma: a risky move or site dependent growth

Summary Nephila clavipes inhabiting adjacent habitats showed differences in rate and ratio of growth. Spiders inside the forest fed less, grew slower and stayed smaller than those at the forest edge, a richer environment. The mortality rate in this habitat was about 1% per day, the mortality of vagrant males was even higher. Experimentally fed spiders did not shift their webs readily, even starving individuals stayed much longer than expected. The spiders avoided frequent moves and in a poor site grew more slowly and in smaller size increments. It seems that the costs of small adult size are less than the risks of active foraging.     

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

Abstract

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

Behavior of Colonial Orb-weaving Spiders during a Solar Eclipse

The behavior of colonial orb-weaving spiders (Metepeira incrassata) in tropical Veracruz, Mexico was studied during the total solar eclipse on July 11, 1991. Spiders behaved in a manner typical of daily activity until totality, when many began taking down webs. After solar reappearance, most spiders that had begun taking down webs rebuilt them. There was no significant difference in the overall activity patterns of spiders during totality across a range of colony sizes. Experimental illumination of part of a colony during totality altered web takedown behavior. While spiders in the darkness of totality began to take down webs, those spiders which were artificially illuminated did not. These observations suggest that the primary environmental cue responsible for the daily rhythm of web building behavior in this species is light level.     

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.     

Population studies of two colonial orb-weaving spiders

Colonial spiders have individual capture webs (territories) within a communally shared web structure. I describe here the life histories and colony population dynamics of two communal species, Ctrtophora moluccensis (Doleschall) (Araneidae) in Papua New Guinea and Philoponella republicana (Simon) (Uloboridae) in the Panama Canal Zone. In both species, dispersal and foundation of new colonies are primarily by groups of immatures. Population growth of new-colonies was rapid during the first generation, but then colony population size decreased markedly. Colonies of P. republicana rarely lasted more than one generation, whereas those of C. moluccensis attained an equilibrium population size and often persisted for many generations at the same site. Reproduction occurred during the wet season in P. republicana colonies and year-round in colonies of C. moluccensis. Reproduction was synchronized in widely separated colonies of P. republicana. Factors controlling population growth and survival of colonies are discussed. Cyrtophora moluccensis colonies were probably regulated by density dependant factors, especially predation and parasitism, and perhaps a shortage of flying insects due to colony visibility. Philoponella republicana colonies were most likely limited by climatic conditions and instability of the habitat (i.e. density independent factors). Colonial social organization influences both dispersal and colony population growth. Coloniality is, however, compatible with various life history strategies.     

Patch shape alters spider community structure: links between microhabitat choice and sensitivity to increased edge habitat

Increased edge effects in fragmented habitats can affect the abundance of edge-dwelling organisms, but these impacts may depend on the biological attributes of species. Microhabitat choice, a species characteristic that reflects combinations of biological traits, may affect the ability of peripheral species to take advantage of increased edge habitat in the presence of edge effects. In this field study, we built artificial shrub modules designed to encourage web spiders to build webs on the periphery. While modules were identical in volume, they differed in shape (cubic and elongated), so that elongated modules had more edge habitat and were subject to enhanced edge effects. Given that the tangle-web spiders Theridion and Dictyna built webs on module edges and strongly differed in terms of concealment and substrate generalization, two habitat characteristics associated with lower vulnerability to habitat modification, we tested the hypothesis that Theridion, which built webs in more concealed locations and on a greater diversity of substrate configurations in the modules compared to Dictyna, would take better advantage of increased edge habitat. As predicted, Theridion was significantly more abundant on elongated modules whereas the abundance of Dictyna did not respond to shape, even though the change in module shape entailed a similar increase in favored substrate for both spider groups. Our results suggest that the microhabitat associations of organisms may be linked to their propensity to be sensitive to edges, and that a better understanding of these links can improve our ability to predict the effects of habitat modification on biodiversity.     

Female morphology,web design,and the potential for multiple mating in Nephila clavipes: do fat‐bottomed girls make the spider world go round?

In animal species where females mate with multiple males, female mating success might be expected to covary with aspects of female morphology, such as size or shape. Spiders are especially interesting in this regard, as the females of several spider groups weave intricate webs that often accommodate multiple male spiders, all of whom are potential mates. Because web design is likely to be dependent on female size/shape, we use multivariate methods to assess the relationships among female morphology, web design, and reproductive ecology over a range of body sizes in the orb-weaving spider Nephila clavipes . Of the measured variables, only abdomen size explained a significant amount of the variation in number of males on a web, and this relationship holds even after statistically accounting for body size. Because abdomen size is an indicator of body condition in spiders, we suggest that condition is likely to be an important factor relating to potential mating success in female spiders. We found no evidence for an association between web design and number of males on a web, although our data indicate that larger females build webs that are both larger and further from the ground than those of smaller females.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 87 , 95−102.     

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.