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Relationship Between Escape Speed and Flight Distance in a Wolf Spider, Hogna Carolinensis (Walckenaer 1805)

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Relationship Between Escape Speed and Flight Distance in a Wolf Spider, Hogna Carolinensis (Walckenaer 1805) 2005. The Journal of Arachnology 33:153±158 RELATIONSHIP BETWEEN ESCAPE SPEED AND FLIGHT DISTANCE IN A WOLF SPIDER, HOGNA CAROLINENSIS (WALCKENAER 1805) Matthew K. Nelson and Daniel R. Formanowicz Jr.: Department of Biology, University of Texas at Arlington, Arlington, Texas 76019. E-mail: [email protected] ABSTRACT. The relationship between running speed and ¯ight distance is an important one in terms of escape from predators, especially in species that may have multiple defensive strategies. In the wolf spider Hogna carolinensis, one important antipredator mechanism is ¯ight. We examined the relationship between sprint speed and ¯ight distance in wolf spiders by measuring sprint speed on a running track and, in a separate set of experiments with the same individual spiders, measured the distance at which they ¯ed from an advancing model predator. Sprint speed was not signi®cantly correlated with mass, size, or sex of the spiders. Sprint speed was positively correlated with ¯ight distance. This correlation may be the result of a trade-off between two competing modes of antipredator mechanisms: escape and crypsis. In individuals with higher sprint speeds, escape may be the more advantageous option. Slower individuals may have a greater chance of surviving an encounter with a predator simply by remaining still and relying on crypsis. Keywords: Antipredator strategy, risk, ¯ee, sprint speed Behavior patterns associated with predator made by Ydenberg & Dill's (1986) model is escape and avoidance are important to indi- that ``response'' does not necessarily equal vidual survival. These result in strong selec- ``detection,'' in that it can often be dif®cult to tive pressure favoring individuals that suc- assess whether or not the potential prey has cessfully avoid or escape from predators. detected a predator. In some cases, an individ- When an animal is approached by a potential ual may ignore an approaching predator until predator, it must evaluate the level of preda- it becomes necessary to initiate ¯ight. In other tion risk, and utilize the appropriate antipred- cases, certain ``alert behaviors'' may occur ator mechanism to neutralize the risk. The dis- that precede a ¯ight decision. The Ydenberg- tance from an approaching predator at which Dill model generally predicts that individuals an animal chooses to ¯ee has been referred to should delay ¯ight until the costs associated as ``¯ight distance'' (e.g. FernaÂndez-Juricic et with staying (e.g., increased predation risk) al. 2002), ``¯ight initiation distance'' (e.g. exceed the bene®ts associated with staying Bonenfant & Kramer 1996), ``approach dis- (e.g., time spent searching for food or mates). tance'' (e.g. Martin & Lopez 1999), and Several studies have examined the relation- ``¯ush distance'' (e.g. FernaÂndez-Juricic et al. ship between ¯ight distance and running abil- 2001). The latter two of these imply the per- ity. Rand (1964) found that body temperature spective of the predator. Since we will be dis- affected the distance at which Anolis lizards cussing the issue from the perspective of the ¯ed from approaching predators, attributing prey, ``¯ight distance'' seems the most appro- differences in escape distances among individ- priate and concise terminology. uals to lower body temperatures which re- Ydenberg & Dill (1986) discussed in detail duced sprint speeds. Cooler individuals tended the economics of escape from predators. They to ¯ee at greater distances because lower body suggested a cost-bene®t model of ¯ight dis- temperatures result in greater risk of capture. tance, incorporating the costs and bene®ts of Heatwole (1968) suggested that crypsis might continuing a particular behavior (such as for- also play a role in ¯ight distances. Cryptic aging) relative to the costs and bene®ts of species of Anolis may decrease their risk of ¯eeing. A concept critical to the predictions capture by remaining motionless and ¯eeing 153 154 THE JOURNAL OF ARACHNOLOGY at shorter distances than less cryptic species. usually ¯ee a meter or so, and then remain Species that rely on crypsis may not ¯ee from motionless (pers. obs.). an approaching predator until detection by the We examined the relationship between predator is certain. body size and running speed in male and fe- Schwarzkopf & Shine (1992) suggested male H. carolinensis, testing the hypothesis that ``vulnerability'' (risk of capture) of prey that larger individuals were faster. Since there should be evaluated in terms of the probability is a sexual dimorphism in body size in this of being detected by a predator. They found species, we also predicted that females and that gravid female water skinks, Eulamprus males should differ in running speed. Using tympanum, exhibited decreased running abil- the same spiders, we examined the distance at ity and shorter ¯ight distances relative to non- which they ¯ed from an approaching model gravid females. They suggested that gravid fe- predator. We used the data collected on run- males switched antipredator tactics from ning speed and ¯ight distance to test the fol- escape to crypsis because of the decrease in lowing hypothesis based on Ydenberg & running speed. They interpreted these results Dill's (1986) model: faster individuals would as evidence that escape speed may not always be expected to ¯ee at shorter distances from be the most important element involved in de- the predator. termining when to ¯ee. Formanowicz et al. METHODS (1990) investigated similar effects in the liz- 5 ard Scincella lateralis with differences in Hogna carolinensis (n 77; 44 males, 33 sprint speed related to tail autotomy. Skinks females) used in this study were collected on with autotomized tails were found to exhibit 26 March and 11 April, 1997 at the Texas Na- slower running speeds and relatively shorter ture Conservancy's Independence Creek Pre- ¯ight distances. They suggested that individ- serve, approximately 37 km south of Shef- ®eld, Terrell County, Texas, on the uals that had lost their tails switched to a cryp- northeastern edge of the Chihuahuan desert. tic antipredator strategy to compensate for re- Voucher specimens have been deposited at the duction in sprint speed. Denver Museum of Nature & Science. Most In this study, we examined the relationship spiders were collected at night by using head- between running speed and ¯ight distance in lamps to produce eyeshine; a few were col- the wolf spider Hogna carolinensis (Walcken- lected by turning rocks during the day. Spiders aer 1805). Hogna carolinensis is a large, bur- were not found to be active during the day. rowing wolf spider that is active on the sur- Females were often found near the mouth of face of the ground from dusk until dawn and a burrow, and sometimes removed from a bur- is distributed from southernmost Maine and row. In almost every case, females were found Ontario throughout the southeastern U.S. and within a meter of the burrow. Males, however, west to Baja California (Dondale & Redner were only occasionally found near a burrow 1990). Very little has been published concern- but never in a burrow. Spiders were housed ing the life history of this species. Individuals individually, in clear plastic containers (18.5 of Hogna carolinensis construct a burrow 3 7.5 3 9 cm) with a sand substrate (approx. with a turret of sticks and grass surrounding 1 cm deep). Each spider was fed one adult the mouth of the burrow. Or, in some cases, cricket/week, and water was available ad li- these spiders may inhabit a deserted rodent bitum. Temperatures in the housing and test- burrow (Shook 1978). The depth of the bur- ing area ranged from 25±26 8C. row likely varies between geographical re- Escape Speed.ÐSpider escape speeds were gions and possibly with the substrate. In west measured on a wooden track 9 cm wide and Texas, where this study was conducted, I have 2 m long, with sheet-metal side walls approx- found burrows as deep as 25 cm (pers. obs.). imately 21 cm high. All trials were conducted Likely predators include lizards, centipedes during daylight hours. A start box was sepa- (pers. obs.), scorpions, coyotes, owls and var- rated from the track by a removable metal di- ious predacious insects (Shook 1978). If a vider (21 3 9 cm). A spider was placed in the burrow is near, individuals will retreat to a start box, allowed to acclimate for 15 minutes, burrow when disturbed (Kuenzler 1958); how- the divider was raised, and the spider was ever, if a burrow is not near, the animal will prodded on the posterior end of the abdomen NELSON & FORMANOWICZÐESCAPE DISTANCE OF WOLF SPIDER 155 with a ®berglass rod until it ran. Using a stop- 0.8640) indicating that the speed of the model watch, we recorded the time that the spider predator from trial to trial was not signi®- crossed each 50 cm segment of track. Each cantly different. Another set of trials was run spider was run twice with 24 hours between using only the motor without the model pred- trials, after which time its mass was recorded. ator to rule out the possibility of cues from Spider cephalothorax lengths were measured the sound and vibration of the motor. In these at the end of the study with a caliper. trials, none of the spiders responded to the The fastest 50 cm speed (cm/s) for each spi- activation of the motor (n 5 10). The response der was used for statistical analyses. T-tests of the spider to the approaching model pred- were used to determine whether mass, ceph- ator was viewed through the observation holes alothorax length and speed differed between in the wall of the spider chamber. Escape was the sexes. Normality was evaluated using Sha- operationally de®ned as a spider turning and piro-Wilk's W, and none of the groups violat- running in the opposite direction from the ed this assumption at 0.05 level.
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