Relating the Cost of Spinning Silk to the Tendency to Share It for Three Embiids

BEHAVIOR Relating the Cost of Spinning Silk to the Tendency to Share It for Three Embiids with Different Lifestyles (Order Embiidina: Clothodidae, Notoligotomidae, and Australembiidae) 1 J. S. EDGERLY, S. M. SHENOY, AND V. G. WERNER Department of Biology, Santa Clara University, Santa Clara, CA 95053 Environ. Entomol. 35(2): 448Ð457 (2006) ABSTRACT Although adult female embiids (Order Embiidina) superÞcially lack morphological diversity, their variety of habitats may impose distinct selective pressures on behavior, such as their use of silk and their tendency to aggregate. For example, where silk serves as a primary defense from environmental threats, coloniality might be adaptive. The cost of production or spinning might also prompt them to share silk. These ideas were tested in laboratory trials involving three species of embiids with different lifestyles: an arboreal species (Antipaluria urichi (Saussure) from a neotropical rain forest, a species (Notoligotoma hardyi (Frederichs) that dwells on surfaces of granite outcrops in Australia, and another Australian species (Australembia incompta Ross) that stitches leaf litter together. The cost of spinning silk was analyzed by recording CO2 output in the short term during spinning and by measuring performance in long-term trials where embiids were forced to repeatedly replace their silk. Their subsequent development or reproductive output was scored. Overall, the cost of spinning was relatively low. However, the tendencies to spin and to aggregate varied in a manner related to how silk is used in the Þeld. As such, the more exposed the embiid is to the elements, as for the two species that spin on surfaces, the more silk they spun and the more likely they settled near a neighbor. In contrast, the embiid that used dead leaves (not silk) as walls for their abodes produced scant silk and showed little tendency to aggregate. KEY WORDS Embioptera, webspinner, silk, social evolution, metabolic cost LIVING IN SILKEN ABODES deÞnes lifestyles for many study, we will brießy review relevant research on arthropods, especially for masterful weavers such as arthropod silks. insects in the order Embiidina. Silk serves as nesting Generalizations cannot easily be made about the material, retreats, protection for eggs, and coverings cost of silk production. In some cases, spinning is not over foraging zones. Because silk-using arthropods costly materially, metabolically, or behaviorally, range from solitary to colonial to communal, research- whereas in others, direct impacts on performance ers have been able to apply comparative methods to have been shown. For example, caddisßy larvae that determine how silk use relates to social evolution (as spin extra silk lose more weight than starved individ- in social spiders; see references in Avile´s 1997, Uetz uals, suggesting a high cost of spinning for these insects and Hieber 1997). An argument with some support is (Dudgeon 1987). Behavioral mechanisms may have that coloniality may be favored when the costs of silk evolved in response to costs associated with spinning. production are high for any individual and beneÞts of For example, Jakob (1991) showed that pholcid spi- sharing are great. The purpose of this report is to derlings beneÞted by sharing silk with older conspe- summarize results of a comparative study studying silk ciÞcs because of the relatively high cost of spinning, spinning in the context of natural history in the order Embiidina (or Embiodea, see Grimaldi and Engel despite a loss of food to competitors. Tietjen (1986) 2005). Through a series of laboratory experiments, we showed that as colony size increased for the social evaluated possible costs of spinning and explored spider Mallos gregalis (Simon) individuals beneÞted whether embiids can reduce costs by sharing silk. by living in a group because each contributed less silk Embiids that construct complex, extensive silk abodes to the shared nest and spent more time sitting still. In may be more likely to form groups; those relying more their review of spider evolution, Uetz and Hieber on building materials in the environment may be less (1997) emphasized that a common element promot- prone to do so. Before describing methods for this ing coloniality for some spider species is the three- dimensionality of their aggregations and considerable energetic savings for individuals sharing silk. In con- 1 Corresponding author, e-mail: [email protected]. trast, Avile´s (1997), in a review of permanent- sociality 0046-225X/06/0448Ð0457$04.00/0 ᭧ 2006 Entomological Society of America April 2006 EDGERLY ET AL.: SILK SPINNING DIVERSITY IN EMBIIDS 449 in spiders, did not identify silk as a catalyst for social ing sites (domiciles) to foraging zones. Generally, em- evolution. biids live hidden within silk in motherÐoffspring as- While it seems that costs can promote group- living, sociations. In the humid tropics, large-bodied embiids alternatively, dependence on silk may lead to aggres- can form conspicuous and extensive colonies that sion. For example, in laboratory experiments, parsnip cover entire tree trunks where they graze on algae and webworm larvae defended webs from potential usurp- lichens (see references in Edgerly 1997, Ross 2000). In ers and, when given a choice, adopted empty webs arid zones, they retreat underground in crevices, rather than spun new ones (Berenbaum et al. 1993, emerging (especially after rain) at the surface, par- Green et al. 1998). Comparative work on spider spe- ticularly at rockÐsoil boundaries, to feed on vegetation cies that range in how much silk they use showed that or detritus. the more individuals invested in silk for construction, To test our ideas, we selected three species that vary the less likely they were to relocate where they had to in their use of silk in the Þeld. The focal species were spin anew (Tanaka 1989). Apparently, the function of as follows: the relatively large arboreal embiid Anti- silk can inßuence dispersion of individuals. paluria urichi (Saussure) (Clothodidae) of the Trin- The degree of reliance on silk and the tendency to idad rain forest, a facultatively colonial species that share it may depend on challenges imposed by envi- often forms large colonies (Edgerly 1987b, 1994); N. ronmental factors. In a search for function, Seibt and hardyi of subtropical Queensland, Australia, which Wickler (1990) found that silk nests of two social lives mostly on granite outcrops, but also on bark, species of Stegodyphus spiders in South Africa did not where they spin dense silk and feed on lichens; and protect occupants from potentially serious threatsÑ Australembia incompta Ross (Australembiidae), also wind, hail, direct sun radiation, desiccation, or over- of Queensland, which stitches dead leaves together heating. The authors concluded that predation on the and feeds on detritus on the forest ßoor (Edgerly and spiders was more signiÞcant in promoting sharing silk Rooks 2004). We evaluated short- and long-term costs and sociality (Seibt and Wickler 1988). Likewise, of spinning and tested whether sharing silk may func- Edgerly et al. (2005) discovered that embiid silk in tion to reduce such costs. We also tested how readily Queensland, Australia, did not protect against over- adult females spun silk and whether they tended to heating in colonies exposed to direct sun. The embiids aggregate. We related these behaviors to their appar- in that study [Notoligotoma hardyi (Friederichs): No- ent reliance on silk in the Þeld. Our prediction was that toligotomidae] relied instead on internal heat-shock embiids living exposed on rock-faces or trees are more proteins and microhabitat choice (see also Edgerly dependent on silk, which serves as primary protection and Rooks 2004). In contrast, silken abodes of cater- against abiotic and biotic threats. These species should pillars do play a signiÞcant role in behavioral thermo- regulation, offering warmth or shade depending on spin more and aggregate more than the leaf-litter where the larvae sit in their three-dimensional tents dweller, which should be less reliant on silk which (Carlberg 1980, Joos et al. 1988, Fitzgerald and Un- functions more as stitching than as primary cover. derwood 2000). Reliance on silk is fundamental to embiids, an order of Ϸ2,000 species, many still undescribed (Ross 2000). Materials and Methods Individuals of all ages and both sexes spin with their front tarsi by releasing numerous strands as their silk Laboratory Cultures and General Methods. Embi- Њ ejectors touch the substratum (Edgerly et al. 2002). ids were reared at Santa Clara University at 27 C with Females are wingless, elongate, ßexible, and juvenile 12 h of overhead ßuorescent lighting each day. A. in form, traits that are well suited to life in narrow silk urichi and A. incompta fed on romaine lettuce and passages. The few detailed behavioral studies show lived in a mulch of dead oak leaves, typical rearing that adult females contribute the most silk to the medium for embiids (Ross 2000), whereas N. hardyi colony, and indeed, spinning silk is a key component would only feed on lichens scraped from oak bark. All of maternal care (Edgerly 1988). Adult males spin lichens and leaves were from local California Live Oak little silkÑin most species, they ßy off to mate and die (Quercus agrifolia). Adult females were used in all soon thereafter without feeding or contributing much experiments unless noted. Their mean weights ranged to the silk covering. The differential contributions to from 0.042 g for A. urichi (n ϭ 20) to 0.021 g for A. spinning made by males and females still needs study, incompta (n ϭ 14), and 0.012 g for N. hardyi (n ϭ 20). especially for those species where males are wingless Body length follows the same relationship: A. urichi at and more likely to spend time in silk galleries, partic- 1.60 Ϯ 0.02 cm (Edgerly 1987a), A. incompta at 1.18 Ϯ ularly true for embiids in arid environments.

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