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Disturbance Behaviors in the Spider Uloborus Glomosus (Araneae

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Disturbance Behaviors in the Spider Uloborus Glomosus (Araneae 1090 Disturbancebehaviors in the spider Uloborusglomosus (Araneae, Uloboridae): possiblepredator avoidancestrategies Pluln E. CusHrNG AND BnrNr D. Oprll Department of Biology, Virginia PolytechnicInstitute and State University, Blacksburg, VA 24061, U.S.A. ReceivedMay 25, 1989 CusHtNc, P. E., and OpEu-, B.D. 1990. Disturbancebehaviors in the spider Uloborus glomosus(Araneae, Uloboridae): possiblepredator avoidancestrategies. Can. J. Zool. 68:1090-1097. When disturbed, Uloborus glomosuseither remain in position at the hub of their orb webs,jump from the web, move to the edgeof the web, or shakethe web. Juvenilesmore frequentlyexhibited moving andjumping responses,whereas the majority of adultsjumped from the web or remainedin position. Adults with linear stabilimentatended to shaketheir webs in the morning and to jump from their websin the evening. Juvenileswith linear stabilimentashook their webs in the afternoonand evening, whereas thosewith circular stabilimentatended to move to the edgeof the web or shakethe web during all times of day. Adult femaleswith egg-sacchains were usually alignedwith the egg sacsand tendedto remain in position when disturbed.Both adults andjuveniles lacking stabilimentaor egg-sacchains tendedto jump from their webs or move to the edge of their webs. Spiders filmed in a natural settingresponded to contactby insects.Predatory wasps repeatedly flew near the spiderswithout eliciting a response.A tetheredpredatory wasp held aboveboth juveniles and adultsin the laboratoryalso failed to elicit responsesfrom the spiders.The data agree with the hypothesisthat moving to the edge of, shaking, and jumping from the web may be predator avoidance strategieselicited by direct contact. CustttNc, P. 8., et OPELL,B. D. 1990.Disturbance behaviors in the spider Uloborus glomosus(Araneae, Uloboridae): possible predatoravoidance strategies. Can. J. Zool.68: 1090-1097. En casd'aletle, Uloborus glomosuss'accroche au centrede sa toile, sautehors de satoile, seddplace vers sa bordureou la fait vibrer. Les araign6esjuv6niles ont surtoutdes r6actions de d6placementou de saut, alorsque la majoritd des adultessautent hors de leur toile ou ne bougentpas. Les adultesi stabilimenturrislin6aires ont tendanced faire vibrer leur toile le matin et h sauterhors de leur toile le soir. Les juvdniles h stabilimentums lin6aires font vibrer leur toile I'aprds-midi, alors que les juv6niles h stabilimentumscirculaires ont tendancei se d6placervers la bordureou i faire vibrer leur toile en tout temps de la journ6e. Les femellesadultes qui portentdes cocons ovigdres se tiennent ordinairement en ligne avecleurs cocons et ont tendanced garderleur position en casd'alerte. Les adulteset lesjuv6niles sansstabilimentums ou sanscocons ont tendanced sauterhors de leur toile ou i se d6placer vers sa bordure. Des araign6esfilm6es dans leur habitat naturel ont r6agi au contact d'insectes. Des gu6pes prddatricesont vol6 plusieursfois vers les araigndessans susciter de r6action. En laboratoire,les araigndesjuv6niles et adultes sontrest6es sans rdaction en pr6senced'une gu6pepr6datrice attachde et maintenueau-dessus de la toile. Les donn6esconoborent I'hypothEseselon laquelle les d6placementsvers la bordure de la toile, le tremblementde la toile ou les sautshors de la toile constituentprobablement des stratdgiesde fuite en r6action i un contact direct avec des pr6dateurs. lTraduit par la revue] Introduction web'shub (Fig. lB). The legsof thesespiders display disruptive Although spiders are important prey items of some wasps, coloration and the first legs are beset with a setal brush birds, and other spiders (Bristowe l94l; Krombein 1967; (Fie. 1A). Robinson and Valerio 1977: Pyke 1980; Johnsgard 1983; Mature female U. glomosas and juvenile spidersoften add Coville 1987), few studiesinvestigate the defensivemecha- linear bands of silk stabilimentato the hubs of their orb webs nisms they use to avoid predation. This study has four objec- (Fig. lB). Somejuveniles add a circularstabilimentum platform tives: (f) to define more precisely and describethe responsesto to the hubs of their webs (Fig. lD) (Cushing 1988). Mature disturbanceshown by an orb-weaving species, (ii) to survey female U. glomosusdeposit their stellateegg sacsin a chainthat adult and juvenile populations of the species to determine extendsalong a radius from the web's perimetertoward its hub. what factors influence the expression of the disturbancebe- When an egg-sacchain is present,a femaletypically positions haviors, (iii) to document on film actual encountersbetween herself adjacentto the most recently deposited(most centripe- the orb weaver and its natural predators,and (iv) to determine tal) egg sac where her linear posture makes it very difficult to what stimuli (vibratory, visual, or contacQelicit the disturbance distinguishspider from egg-sacchain (Fig. 1C). During this behaviors. time, femalescontinue to feed and maintain their webs. The We studiedthe orb weaverUloborus glomosus (Walckenaer), linear and circular stabilimentaand the linear chain of egg sacs family Uloboridae. These horizontal orb weaving spiders are may function as cryptic devices, concealing the spider from common throughoutthe easternUnited Statesand are abundant visually hunting predators(Hingston 1927; Bristowe l94l; on the shrubberyof the Virginia PolytechnicInstitute and State Marson 1947a, 1947b; Marples 1969; Ewer 1972; Eberhard University(vptsu) campus (Cushing 1988). Spiderlings emerge 1973;Tolbert 1975;Levi 1977;Lubin 1980). from egg sacsas 2nd instarsand reachmaturity in the 6th instar. We formulated three hypotheses:(i) the disturbancebehaviors Mature females have a cephalothorax-abdomen length of shown by U. glomosus are predator avoidancestrategies and 10-12 mm. Uloborids build their webs at night, but monitor will be expressedat different frequenciesthroughout the day, them and capture prey during both day and night. During the reflecting changesin predatorypressure; (ii) the stabilimentr day, both juvenile and adult Uloborus hang from the underside and egg-sac chains are functioning as cryptic devices anci of their orb webs in what appearsto be a twiglike cryptic posture spidersthat have thesestructures and are aligned with them are (Opell and Eberhard 1984) (Fig. lA). They remain in this lesslikely to show activeavoidance behaviors when disturbed; posture excep: when wrapping prey and when feeding at the and (ni) becausethese disturbancebehaviors protect spiders Prinrcd in Canada / Imprimd au Canada CUSHING AND OPELL l09l Frc. l. (A) Adult female U. glomosush^cryptic posture.(B) Femalefeeding at the hub ofa web alignedwith a linear silk stabilimentum.(C) Femalealigned with an egg-sacchain (anow points to the spider). (D) Circular stabilimentumplatform of a juvenile web. Bar, 5 mm. from hunting wasps, they are elicited by the vibratory stimuli of the observer's presence.The rubber bulb was consideredan approxi- a flying wasp. mately standardizedstimulus, similar in size to the rubberpencil eraser used by Tolbert (1975) in his study of predator avoidancebehaviors. Methods This stimulus was assumedto simulate the sudden contact of an attacking predator. The data were analyzedusing a log-linear analysisdesigned Description of the behaviors for multivariate categoricaldata (Bishop et al. 1975;Fienberg 1987). To more preciselydefine and describethe disturbancebehaviors, we collectedadult femaleson campusand establishedthem on framesin a ofjuveniles greenhousewhere we filmed them with a Super 8 mm movie cameraas Survey During August and September1987, we conducteda similar survey they jumped from and shook their webs in responseto contact stimuli ofjuvenile U . glomosu.sin the field. To determinethe stadiaofjuvenile (the tip of a finger). Since uloborids often shaketheir webs during prey spiderswe collected every l0th individual and measuredthe length of capture,we also filmed the shakingbehavior of an adult female when its carapace and first femur. We compared these data with the values fruit flies were placed in her web. usedby Opell (1987) to assessthe stadiaof U. glomosasfrom the same Survey of adults population. Adult spiders living in shrubbery at three sites on the vplsu campus This survey differed from the adult survey in four ways. (i) As were studiedduring June and July, 1986. We made 222 obsewations juvenile U. glomosas males spin typical capture webs, this survey during the morning (08:00-10:00), 227 dunng the afternoon (12:00- includedboth malesand females.As male and femaleuloborids cannot 14:00), and 146 during the evening (17:00-19:00) for a total of 595 be easily distinguisheduntil they enter the penultimate(5th) instar, the observations. Since adult male U. glomosas do not build typical sex of the individuals tested could not be determined. (li) Being capturewebs (Opell1979), we usedonly adult femalesin this survey. juveniles, thesespiderlings never had egg-sacchains in their webs and Temperature was inadvertently recorded during only one of three some individuals had circular rather than linear silk stabilimenta.(iii) mornings during which observationswere made (25.9'C) and two of The stimulus usedfor this juvenile survey was a 4 mm diameterplastic three afternoons (mean : 35.6oC, SD : 2.62). The mean evening sphere affixed to the end of a 15 cm long wooden applicator stick. temperaturewas 28.5oC(SD : 0.38, n : 3). Relative to spider size, this stimulus was approximatelythe samesize Fourvariableswere recordedduring the adult survey:(i) time of day, as that used in the adult survey.
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