Speculative Hunting by an Araneophagic Salticid Spider
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SPECULATIVE HUNTINGBY ANARANEOPHAGICSAL TICID SPIDER by ROBERT J.CLARK , DUANE P.HARLAND and ROBERT R.JACKSON 1,2) (Departmentof Zoology,University of Canterbury, Private Bag 4800, Christchurch, New Zealand) (Acc.3-VII-2000) Summary Portia mbriata ,anaraneophagic jumping spider ( Salticidae),makes undirected leaps ( er- raticleaping with no particulartarget being evident) in the presence of chemicalcues from Jacksonoidesqueenslandicus ,anothersalticid and a commonprey of P. mbriata. Whether undirectedleaping by P. mbriata functionsas hunting by speculation is investigatedexperi- mentally.Our rsthypothesis, that undirected leaps provoke movement by J.queenslandicus , wasinvestigated using living P. mbriata andthree types of luresmade from dead, dry arthro- pods (P. mbriata, J.queenslandicus and Muscadomestica ).When a living P. mbriata made undirectedleaps or aspring-drivendevice made the lures suddenly move up and down, sim- ulatingundirected leaping, J.queenslandicus respondedby wavingits palps and starting to walk.There was no statisticalevidence that the species from which the lure was made in u- enced J.queenslandicus ’responsein these tests. Our second hypothesis, that J.queenslandi- cus revealsits location to P. mbriata bymoving, was investigated by recording P. mbriata’s reaction to J.queenslandicus when J.queenslandicus reactedto luressimulating undirected leaping.In these tests, P. mbriata respondedby turning toward J.queenslandicus and waving its palps. Keywords: Portia mbriata , Jacksonoidesqueenslandicus ,jumpingspiders, predation, spec- ulativehunting. 1) Correspondingauthor; e-mail address: [email protected] 2) WethankPhil T aylorand David Blest for useful discussion and valuable comments on the manuscript.Financial support was provided by theNational Science Foundation ( GrantBNS 861078)and the Marsden Fund of New Zealand(Grant UOC512). c KoninklijkeBrill NV ,Leiden,2000 Behaviour137, 1601-1612 ® 1602 CLARK, HARLAND&JACKSON Introduction Ageneralproblem facing predators is howto locate prey(Curio, 1976). Whenvision is relied on,being out of the predator’sline ofsight orbeing camouaged will interfere with detection (Edmunds,1974). ‘ Huntingby speculation’(directing attacks at refugeswhere prey tend to befound or probingareas in whichprey normally hide) is apotential solution (Curio, 1976).Envisaged not as prey-capturebehaviour, but instead as atactic forlocating prey,hunting by speculation might functionfor a predator byprovoking a responsethat reveals the prey’slocation. W oodstorks,for example,may probe submerged vegetation even when no prey is visible (Kahl& Peacock,1963), Octopuscyanea Grayattacks holes in coral even in the absenceof prey(Y arnell, 1969)and lions mayrun to the topof hills, apparentlyin anticipation ofstartling unwaryprey on the otherside (Schaller, 1972).However, examples fromarthropods, and experimental studies onany predator,have been scarce. Inthe present paper,we investigate huntingby speculation in Portia m- briata (Doleschall) fromQueensland, Australia, anaraneophagic jumping spider (Salticidae) that preysespecially oftenon other salticids (Jackson& Blest, 1982). Jacksonoidesqueenslandicus Wanless (Salticidae) is especially abundantin the same habitat as P. mbriata (Jackson,1988 )andis probably the salticid species onwhich P. mbriata most oftenpreys. Preliminary stud- ies showthat chemical cues from J.queenslandicus ,evenin the absence of J.queenslandicus , prepare P. mbriata forpredation by stimulating the adoptionof a special palpposture (retracted palps) characteristic ofstalk- ingsequences against salticids as preyand by heightening P. mbriata’s at- tention to visual cues from J.queenslandicus .Preliminary studies indicate that chemical cues from J.queenslandicus also elicit intermittent undirected leaping(erratic leapingwith noparticular target beingevident) by P. mbri- ata.Theabsence of anapparent target suggests that this behaviourfunctions as speculative hunting.T wohypotheses are consideredhere: (1)undirected leaps by P. mbriata stimulate J.queenslandicus to move;(2 )bymoving, J.queenslandicus gives awayits location to P. mbriata. Methods General Standardmaintenance procedures in a controlled-environmentlaboratory (light-dark cycle, 12L:12D;lights on at0800 h) wereadopted, as detailedelsewhere (Jackson & Hallas,1986). SPECULATIVE HUNTINGBYSPIDER 1603 Adultfemales of P. mbriata and J.queenslandicus (bodylength: P. mbriata 10-11 mm; J.queenslandicus 5-6mm), from laboratory cultures, were used. Experiment1: J. queenslandicusviewing P .mbriata Thesetests were used to get baseline information on how J.queenslandicus reactedwhen P. mbriata madeundirected leaps. For a testchamber, we used a transparentperspex box (length ´ width ´ height: 211 ´ 144 ´ 44mm). Three regions of the box were de ned: region1 extended50 mm outfrom one end of thebox; region 2 extended50 mm outfrom theopposite end; region 3 wasthe space between the other two regions. There was a 5-mm widehole ( keptplugged with a cork)in the bottom of thebox in the centre of region 1 and anotherin the centre of region 2. Testspider 1 wasput in thebox 24-28 h beforetesting started and had free access to the threeregions during this pre-test interval. T estingbegan between 0800 and 1000 h. When test spider1 waswithin 50 mm ofoneend of the box, a partition(partition 1) wasput into place, therebyclosing this region ( region1) offfrom the rest of the box. At thesame time, another partition(partition 2) wasput into place, closing off region 2. Testspider 2 wasintroduced 10minlater into region 2 throughthe hole in the bottom of the cage. For transfer, rsttest spider2 wasenticed into a plastictube ( diameter5 mm),then the tube was positioned with oneend against the hole in thetest chamber. When gently prodded by inserting a softbrush throughthe other end of thetube, test spider 2 walkedslowly out into the test chamber. Testingbegan only if J.queenslandicus hadremained quiescent for the previous 5 sfacing region2. With J.queenslandicus quiescent,partition 2 followedby partition1 wasremoved. Thebehaviour of thetwo spiders was observed for the next 10 min. Control tests were the sameas experimental tests except that partition 2 remainedin place ( i.e.J. queenslandicus couldnot see P. mbriata duringthese tests). Beinginterested speci cally in how J.queenslandicus reactedto undirected leaping by P. mbriata,weconsider only those experimental and control tests in which: (1) P. mbriata madea singleundirected leap during the 10-min test interval and (2) J.queenslandicus remainedquiescent for the entire period prior to P. mbriata makingits undirected leap. The testwas aborted if: ( 1) P. mbriata movedduring an experimental test into region 3 before makingan undirectedleap or (2) P. mbriata madea secondundirected leap before the 10- mintest interval elapsed. No individual J.queenslandicus or P. mbriata wasused in more thanone successful test. Different individuals were used in experimentaland control tests. Experiment2: J. queenslandicusviewing a lure Ourobjective was to test simultaneously how J.queenslandicus reactedto a moving lure( asimulationof undirected leaps by P. mbriata) and how P. mbriata reacted to J.queenslandicus ’reactionto thelure. The test chamber ( Fig.1) wasa rectangularperspex box (length ´ width ´ height: 147 ´ 51 ´ 51mm) withtwo tubes. Its design permitted viewingof a lureby J.queenslandicus andviewing of J.queenslandicus by P. mbriata. Tube1 (internaldiameter 13 mm), made of transparent glass, tinsidethe box at one end.Except when introducing J.queenslandicus ,thehole opening to the outside was kept stoppered.Initially, a holeat the opposite end of thetube was blocked by anopaquemetal screen( partition1). Partition 1, which tintoa slitin the box, could be moved from side toside( indicatedby dotted lines in Fig.1). Tube 2 (internaldiameter 13 mm), situated on 1604 CLARK, HARLAND&JACKSON Fig.1. Apparatus used in experiment 2. Jacksonoidesqueenslandicus (inTube 1), but not Portia mbriata (inTube 2), can view the lure. P. mbriata canview the reaction of J.queenslandicus tothelure. T opand bottom of the box are opaque perspex and the sides transparentperspex. Tubes are transparent glass. Before testing begins, the opening of each tubeinto the box is blockedby aslidingpartition, as indicated by dottedlines. During testing bothtubes are unblocked by aligning the hole in thepartition with the tube opening. The lure ison top of a woodenrod connected to anelectric motor ‘ leapgenerator’ ( notshown). thetop of thechamber, housed P. mbriata.Thedistal opening of tube 2 waskept stoppered exceptwhen introducing P. mbriata.Theproximal opening of tube 2 openedinto a wider tube( internaldiameter 20 mm) whichin turn opened into the box. An opaque metal screen (partition2) coveredthe opening between the narrow and wide tube. Partition 2 tintoa slit inthewider tube, and it couldbe movedfrom side to side( indicatedby dotted lines in Fig. 1). Thewide tube was positioned so that its centre was directly above the opening of tube 1 intothe interior of thebox. The top of thebox was opaque. This meant that the only part of thebox’ sinteriorvisible to P. mbriata duringa testwas around the opening of tube 1. At thefar end of the box, positioned in front of tube 1, there was a holein the bottom ofthe box through which a lurecould move. Each lure was made by positioning a dead J.queenslandicus , P. mbriata or house y (Muscadomestica L.)in a lifelikeposture on a corkdisk (diameter 17 mm; height22