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The wasp spider Argiope bruennichi (Arachnida, Araneidae): ballooning is not an obligate life history phase.

ABSTRACT. Aerial dispersal ("ballooning") of Argiopebruennichi spiderlings has been claimed being an obligate lifestyle historytrait along along with a prerequisite for spinning prey-capture webs. In case this weretrue, the ballooning stage could be crucial for just regarding any laboratory rearingof A. bruennichi creating rearing protocols particularly elaborate. Wetested your significance of ballooning regarding second-instar spiderlings inthe laboratory and also indicated that the actual ballooning behavior isn't essentialfor constructing prey-capture orb webs. Our outcomes furthermore give simply no evidencefor the particular hypothesis that will current all-natural assortment has evolved ballooningbehavior in newly founded field populations.

Keywords: Araneae, ballooning experiment, laboratory rearing, web-building behavior.


Ballooning is really a widespread dispersal mechanism for many modern spiders (Coyle 1983; Dean & Sterling 1985; Weyman 1993), which behavior is especially important regarding maintaining genetic cohesion among Argiope populations (Ramirez & Haakonsen 1999). The Actual - download terraria hack - life good popularity for Argiope is seen as an ballooning, the aerial transport on wind-blown silk threads. The excellent instance for the significance associated with ballooning with regard to range expansion will end up being the Palearctic wasp spider Argiope bruennichi (Scopoli 1772). The Particular spider is surely an r-strategist (Guttmann 1979), characterized by high aerial dispersal capability as well as an ongoing postglacial expansion of its geographical range in Europe (van Helsdingen 1982). Females involving A. bruennichi generate up to five cocoons within the field, frequently containing several hundred eggs (Crome & Crome 1961; Kohler & Schaller 1987). Your expansion in the species provides accelerated in the 2nd half of the last century possibly thanks to end up being able to aspects favoring dispersal by ballooning (Guttmann 1979; Levi 1983; Sacher & Bliss 1990; Scharff & Langemark 1997; Jonsson & Wilander 1999; Smithers 2000). The wasp spider prefers grassy or even herbaceous vegetation in open, ephemeral or shrubby sites (Wiehle 1931; Pasquet 1984; Malt 1996) in coarse-grained (patchy) landscapes (Gillandt & Martens 1980; Sacher & Bliss 1989) and contains regionally benefited through extra time of farming production and also urbanization (Lohmeyer & Pretscher 1979; Arnold 1986; Nyffeler & Benz 1987). River valleys get been identified as favored dispersal corridors further supporting the importance associated with ballooning for dispersion (Gauckler 1967; Puts 1988).

Follner & Klarenberg (1995) claimed ballooning to become an obligate phase in the progression of A. bruennichi. These types of authors monitored the actual pre-ballooning and ballooning behavior involving spiderlings in a grassland research site close to Munich (Germany). Since they in zero way found aggregations regarding orb webs in the neighborhood in the cocoons coming from which the over-wintering 2nd instar spiderlings eclosed and thus they only observed the creating of 1st prey-capture orb webs right after a ballooning trip, that they concluded "that aeronautic behaviour in Bavarian populations of A. bruennichi is actually obligatory". Moreover, these authors suggested that will spiderlings, that have hatched coming from the cocoon, will starve to death, unless they will carry out a ballooning trip. Ballooning ought to thus be an obligate cycle to swap coming from a non-predatory, passive cycle to certainly one of active predation by simply spinning prey-capture orbs. Follner & Klarenberg (1995) argued that the obligatory aerial dispersal might be a consequence of recent natural selection and stay the reason powering your swift expansion with the species. New populations that are established within a period involving expansion are always founded simply by individuals, which may have ballooned.

If ballooning were a new genuinely obligate phase, it might not only be important regarding natural selection nevertheless also be essential for almost any rearing protocol for A. bruennichi. allowing pertaining to ballooning inside a rearing procedure might simply render laboratory breeding unfeasible since it could prove being too time-consuming along with laborious. However, an obligate ballooning cycle has never been observed before, neither throughout other Argiope nor inside the typically well studied A. bruennichi. Tolbert (1976, 1977) studied ballooning behavioral elements of A. trifasciata (Forska[degrees]l 1775) and also A. aurantia Lucas 1833. He concluded from field and laboratory observations that will "it is actually unnecessary for spiderlings associated with possibly Argiope species to be able to engage throughout aerial dispersal before constructing an orb web" (Tolbert 1977), which is an obvious discrepancy to Follner's and Klarenberg's (1995) claims. We here test your significance associated with ballooning for your construction associated with the first preycapture web within the laboratory simply by evaluating spiderlings reared under a couple of experimental conditions, 1 along with and something without having ballooning.

We collected cocoons associated with A. bruennichi (n = 6) within dry and also semi-dry grasslands northeast associated with Halle (Saale) throughout late April 2002 (Germany, 160 m a.s.l., 51[degrees]33'31" N, 011[degrees]52'49" E). They had been maintained in the lab within individual glass vials (9 cm diameter, 13 cm height, coated with fine gauze) in 23 [+ or even -] 2 [degrees]C along with mist-sprayed using water every a pair of times to be able to avoid desiccation. the vial bottom was covered with initially wet cellulose wadding (1 cm). Second-instar spiderlings hatched from your cocoons in early May.

One day right after hatching we simulated individual ballooning with regard to 60spiderlings (10 from each and also every cocoon) simply by exposing the actual spiderling in aspatula to an air stream generated by a heat supply along using a fan (see Figs.1-4 for details of the actual experimental design). We observed behavioralelements within the pre-ballooning cycle in more detail and also noticed its mode.When the spiderling became airborne, we tracked it and also retrieved it atthe "landing strip" (Figs. 3, 4). The Actual ballooning experimentwas repeated immediately (re-ballooning) for each individual to satisfya achievable "ballooning drive" (see Tolbert 1977). Thespiderlings needed to actively participate within this experiment through showingthe entire sequence of pre-ballooning along with ballooning behavior (Figs.1-4).


Following the actual experiments, the "ballooners" were held in the same unheated indoor space with windows admitting indirect natural light. These People were housed within teams (n = 20) throughout three gauze covered glass terraria (50 x 30 x 31 cm; 25 [+ or perhaps -] three [degrees]C; 65 [+ or even -] 10% RH) and fed ad libitum 45-50 live Drosophila melanogaster as quickly as any day. Every two times we sprinkled the within surfaces with the terraria using water. This prevented desiccation as well as allowed with regard to typical drinking behavior of the spiderlings. The Particular bottoms in the terraria were covered using a layer of commercial, pasteurized potting soil (3 cm) along with grass tufts, some dry twigs along with wooden skewers in order to enhance your number of potential attachment points with regard to web building.

A control group regarding spiderlings (n = 60) ended up being treated in the same way, nevertheless without the particular ballooning procedure ("non-ballooners"). In both teams (ballooners vs. non-ballooners) spiderlings and also orb webs were noted three times day-to-day from 6 a.m., 12 p.m. and six p.m. to end up being able to ensure individual based data sets. The Particular rearing time period has been cut off after 19 days when all the surviving people had spun their own initial preycapture orb-webs.

Voucher specimens tend to be deposited in the Entomological Collection of the Martin-Luther-University Halle-Wittenberg (Zoological Institute), Germany (identification amount 2568).

The web-building exercise of the spiderlings elevated in each the ballooners as well as the non-ballooners over time as well as reached 90 [+ or -] 5% for ballooners (n = 54, three terraria) as well as 95 [+ or even -] 5% for non-ballooners (n = 57, 3 terraria) within a period regarding 19d (Fig. 5). The differences in the web-building activity (Fig. 5) were not statistically significant in between both sets of spiderlings (Kruskal-Wallis test, P = 0.7515; tested for every day built-first webs). The mean latency moment pertaining to web-building (time through hatching through the cocoon to the construction involving the first prey-capture web) has been 8.61 [+ or -] 4.28 days as well as 8.18 [+ or perhaps -] 3.60 days regarding ballooners (n = 54) and non-ballooners (n = 57) respectively. This difference was not statistically significant (t-test, P = 0.56).


Although mortality increased in the second half of the actual observation period (Fig. 6), it didn't exceed 22% following the particular experiment (ballooners: 21.7 [+ or perhaps -] 2.89%, n = 13, non-ballooners: 20.0 [+ or -] 8.66%, n = 12, distinction not really significant, t-test, P = 0.77). The surviving animals caught prey in their orb webs and confirmed normal development along with approximately four molts inside your experimental time.


Using our protocol, we're able to initiate the actual complete sequence of ballooning behavior promptly in every experiment. the A. bruennichi spiderlings often showed an identical sequence involving pre-ballooning and ballooning behavior (Fig. 1-4). Any time exposed to the heat in the lamp, they displayed the "ballooning drive" behavior. Individuals walked for the margin with the spatula, spooled out a dragline and dropped down hanging from your line. Whilst suspended along with holding upon towards the drag line, they will let loose an additional series regarding 50-100 cm ballooning silk (Fig. 1). Any time this is lifted by the breeze generated from the fan as well as the heat source, the actual spiderlings cut the particular dragline along with became airborne (Figs. 2, 3). Following landing (Fig. 4) they will hauled in the ballooning line, formed it with the legs into a silk blob and lastly ate the actual silk, bringing the ballooning behavioral sequence to end up being able to completion.

Tolbert (1977) observed a pair of modes associated with preparation pertaining to ballooning in sympatric field populations of A. trifasciata and also A. aurantia. A spiderling attempting to end up being able to grow to become able to be airborne climbed for the surface of some blade of grass or virtually any other structures as well as adopted the typical "tiptoe" posture simply by depressing the particular cephalothorax and elevating the opisthosoma. Multiple silk lines were then exuded coming from the spinnerets. Any time moving air generated sufficient silk, the actual spiderling became a new "ballooner" (Nielsen 1932; Richter 1970; Eberhard 1987). Alternatively, the particular spiderling could grow for you to be airborne by dropping and hanging from a dragline, spinning the ballooning thread, which in turn then gradually lifted along with lengthened within the breeze. The Particular ballooner then cut the dragline as well as floated off to the air (Nielsen 1932; Bristowe 1939).

Argiope bruennichi can display both pre-ballooning modes. However, the drop and dragline mediated ballooning seems being more frequent (Follner & Klarenberg 1995). Throughout the particular field, second-instar spiderlings usually attach the actual draglines to guidelines associated with grass blades or even they'll use silk threads which in turn connect the actual suggestions of grass haulms as attaching points (Follner & Klarenberg 1995). in our experiments, we offered individual spiderlings optimal starting conditions, and which we never observed the particular tip-toe ballooning mode. Follner (1994) suggested that "tip-toe" might be the tactical alternative for people in unfavorable beginning points (e.g., overcrowded suggestions associated with grass blades).

Our outcomes show that it isn't essential for spiderlings associated with A. bruennichi to engage inside aerial dispersal before building any prey-capture web. Whilst ballooning will be frequent in the area (Follner & Klarenberg 1995), it actually is clearly not necessarily an obligate part in the growth and development of this species. Throughout spite of the rapid expansion with the species more than the past decades and the possible significance involving aerial dispersal regarding colonizing new habitats, the role of ballooning in A. bruennichi doesn't differ from A. trifasciata as well as A. aurantia exactly where this cycle in daily life history is also not obligate (Tolbert 1977).

The mortality regarding with regards to 20% after 19 days inside both experimental groups (difference statistically not really significant) points too rearing of A. bruennichi spiderlings to adulthood could end up being challenging. Our rearing method based on an consuming plan along with Drosophila melanogaster, similar to Mu"ller & Westheide (1993), worked well regarding our purpose, where we just tested the actual outcomes of ballooning inside second-instar spiderlings on their ability to produce his or her initial web.

On average, more then eight days elapsed before A. bruennichi spiderlings began to build their initial prey-capture web. This specific appears to be a surprisingly long period, since the animals is only in the position to feed once the initial web is built. We cannot exclude this can become a laboratory artifact, pertaining to instance because of in order to unattractive websites pertaining to web construction. However, the particular lengthy latency did not interfere using the rearing regime. The animals appeared being well adapted for you to temporary starvation since the mortality was reduced in this stage (Fig. 6). Furthermore within the field, the spiderlings do not immediately start with prey-capture web construction (Follner & Klarenberg 1995) as well as endure extended durations of starvation. Argiope spiderlings effortlessly survive a few days neighborhood their cocoons, sometimes with communal meshworks regarding interlocking dragline threads ("communal tangles") (Tolbert 1976, 1977; Follner & Klarenberg 1995) where they will discover shelter until favorable weather or microclimate circumstances allow for ballooning (Tolbert 1977; Follner & Klarenberg 1995; notice additionally Suter 1999 pertaining to physics associated with ballooning).

Argiope spiderlings actively select suitable sites by ballooning, re-ballooning or perhaps walking (Enders 1973; Tolbert 1977; Follner & Klarenberg 1995). In Addition in this nonpredatory stage the actual spiderlings must steer clear of starvation. Tolbert (1976) kept A. aurantia spiderlings in the laboratory with out meals as well as water. Mortality remained reasonable in these experiments for several days in assistance of elevated distinctly about two weeks after hatching.

The behavioral ballooning sequence could be effortlessly triggered under artificial circumstances inside our study, suggesting that will it will furthermore occur in your field whenever environmental conditions allow. Therefore dispersal as well as population composition will be primarily driven by microclimatic circumstances inside the neighborhood habitats. The Actual local persistence of non-emigrants (non-ballooners as well as short-distance ballooners) inside A. bruennichi populations may facilitate aggregated dispersion patterns, just as with weather phases that are unfavorable with regard to aerial dispersal. Given ballooning is truly a less capable method of long distance dispersal than formerly thought (Roff 1981; Decae 1987; Sensible 1993; Bonte et al. 2003), this could furthermore explain the particular genetic differentiation among habitat patches within some other Argiope species (Ramirez & Haakonsen 1999).

The role associated with all-natural choice inside range expansion provides just lately been discussed for insects within the context regarding climatic change (e.g., Pimm 2001; Thomas et al. 2001). However, improving environmental circumstances at range margins could initiate range extensions purely on the basis of ecological, physiological as well as population-dynamic processes not requiring any evolutionary alter (Thomas et al. 2001; discover in addition Coope 1995; Williamson 1996). Our outcomes will always be in range with 1 of these views as well as reject the hypothesis involving Follner & Klarenberg (1995) that will evolutionary processes get changed ballooning behavior within newly founded populations.


We are usually grateful for you to Peter Neumann, Gail E. Stratton and two anonymous reviewers regarding helpful feedback on prior drafts of the manuscript. We thank Christian W.W. Pirk for statistical advice, Vlastimil Ruzicka along with Theo Blick with regard to providing literature. This kind of study was supported by way of a analysis fellowship to be able to A.W. in the State of Saxony-Anhalt along with by the Deutsche Forschungsgemeinschaft (DFG) grant to P.B. (BL 776/1-1).

Manuscript received 17 September 2004, revised 10 August 2005.


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Andre Walter, Peter Bliss (1) as well as Robin F.A. Moritz: Institut fur Zoologie, Martin-Luther-Universitat Halle-Wittenberg, Hoher Weg 4, D-06120 Halle (Saale), Germany. E-mail:

(1) Corresponding author.

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