Spider Mimicry in Fruit Flies (Diptera: Tephritidae): Further Experiments
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Spider Mimicry in Fruit Flies (Diptera: Tephritidae): Further Experiments on the Deterrence of Jumping Spiders (Araneae: Salticidae) by Zonosemata vittigera (Coquillett) DOUGLAS W. WHITMAN,1 LARRY ORSAK,2 AND ERICK GREENE3 Department of Entomology, University of Georgia, Athens, Georgia 30602 Ann. Entomol. Soc. Am. 81(3): 532-536 (1988) Downloaded from https://academic.oup.com/aesa/article/81/3/532/60805 by guest on 29 September 2021 ABSTRACT Zonosemata vittigera (Coquillett) possesses dark transverse wing bands and abdominal eye spots, which cause it to resemble a salticid spider. Jerky motion, wing flicking behavior, and sudden short flights to nearby foliage enhance this resemblance. Here we provide observations on the behavioral interactions between jumping spiders and Z. vittigera. In the laboratory, sympatric jumping spiders (Salticidae) attacked and captured more house flies (Musca domestica L.) or wing-painted Z. vittigera (wing markings disguised by green paint) than unpainted Z. vittigera. Furthermore, salticids backed away from, or gave threat or courtship displays to, normal Z. vittigera but not to house flies or wing-painted Z. vittigera. These results provide further evidence that Z. vittigera mimic jumping spiders and that this resemblance reduces jumping spider predation. KEY WORDS Insecta, Araneae, mimicry, predation 3 NUMEROUS SPIDERS mimic insects such as ants search station. Flies were kept in a 2-m cage and (Reiskind 1970, 1977, Englehardt 1971, Edmunds provided with water, sugar water, honey, dry pow- 1978, Foelix 1982, Jackson 1982a). However, Hin- dered milk, sliced fruit, and potted S. elaeagni- ton (1976) noted that virtually no insect seemed to folium plants. Salticid spiders were collected from mimic spiders; the few postulated cases were with- the desert flats and foothills near Portal on S. out empirical evidence. Flies of the family Te- elaeagnifolium or adjacent vegetation. Zonose- phritidae offer a fascinating exception. Monteith mata vittigera also occurred in this area. The spi- (1972) noted the uncanny resemblance of apple ders were maintained in 6-ml vials and starved 1- maggot, Rhagoletis pomonella (Walsh) adults, to 6 d prior to testing. sympatric jumping spiders (Salticidae), as did Eis- Experiment 1: Salticid Spider Response to ner (1984) with the southwestern tephritid Zono- House Flies Versus Fruit Flies. On 14 July, 19 semata vittigera (Coquillett). Although these mim- spiders were isolated in 1-liter glass jars containing icry patterns were very apparent, the significance two dry sticks and a S. elaeagnifolium stem (13 of this phenomenon remained untested until Greene cm long). After a 30-min acclimation period, each et al. (1987) and Mather & Roitberg (1987) pro- of 10 jars received a specimen of M. domestica; vided evidence that spider mimicry in the Te- the other nine were given one of Z. vittigera. Fol- phritidae serves to reduce predation by salticid spi- lowing fly introduction, spider-fly interactions were ders. Here we present additional confirmation of monitored for 50 min or until the spider caught this hypothesis derived from wing-painting exper- the fly. Captured flies were immediately taken from iments and speculate on the conditions that led to the spider and removed from the jar to prevent the evolution of this mimicry. spider feeding. A second 30-min acclimation pe- riod was followed by the introduction of the alter- nate fly species and a second 50-min observation Materials and Methods period. The S. elaeagnifolium was periodically re- Experiments were conducted at the Southwest- placed with fresh material. Ambient temperatures ern Research Station (Portal, Cochise County, Ariz.) during tests ranged from 27 to 29°C. during July 1986. Zonosemata vittigera were ob- Experiment 2: Salticid Spider Response to Nor- served in the field and collected from their food mal and Disguised Fruit Flies. On 15 July, the plant, Solanum elaeagnifolium Cavanilles, in and same 19 spiders were tested as in Experiment 1, near Rodeo, N. Mex. Wild house flies (Musca do- except that they received either normal Z. vittigera mestica L.) were collected near refuse at the re- or flies that had been painted on the upper and lower wing surfaces with a green felt-tip marker. 1 Department of Entomology, University of Georgia, Athens, This painting completely obscured the dark wing Ga. 30602. markings, destroying the spider-leg image. Spider- 2 Institute of Ecology, University of Georgia, Athens, Ga. 30602. 3 Department of Biology, Princeton University, Princeton, N.J. fly interactions were observed for 10 min followed 08544. by a 30-min fly-free acclimation period, and then 0013-8746/88/0532-0536$02.00/0 © 1988 Entomological Society of America May 1988 WHITMAN ET AL.: SPIDER MIMICRY IN FRUIT FLIES 533 Table 1. Response of sympatric jumping spiders when presented with house flies (M. domestica) or tephritid flies (Z. vittigera) for 50 min Capture latency (min) Spider Species M. domestica Z. vittigera 1 Sassacus papenhoei Peck ham 2 Phidippus sp. A — — 3" Metaphidippus arizo- 3.9 — nensis (Peckham) 4 Habronattus sp. 1.0 10.9 5 Phidippus apacheanus 13.8 — Fig. 1. Posterior aspect of Zonosemata vittigera. Note Chamberlin & Downloaded from https://academic.oup.com/aesa/article/81/3/532/60805 by guest on 29 September 2021 leglike patterns on wings, and abdominal eyespots ar- Gertsch 6 Metaphidippus sp. A — — ranged in a spiderlike pattern. 7 Phidippus sp. A 1.2 7.0 8 Thiodina sp. 4.3 — 9* Habronattus sp. 4.5 44.4 a final 10-min period with the alternate fly species. 10 Eris sp. 0.5 21.0 Temperatures during tests ranged from 20 to 23°C. 11 Phidippus apacheanus 5.9 — The resulting data were analyzed using the Mann- Chamberlin & Whitney U test and a binomial test for changes Gertsch 12 Eris sp. (prob. aurantia — — with small expected frequencies (Siegel 1956). (Lucas)) 13 Thiodina sp. — — 14 Metaphidippus sp. A — — Results 15 Phidippus apacheanus 0.3 8.6 Chamberlin & Appearance and Behavior of Z. vittigera. Eis- Gertsch ner's (1984) observations were quickly confirmed. 16" Phidippus sp. B — _ In nature, Z. vittigera resembled jumping spiders. 17 Phidippus sp. B 2.3 — 18C Phidippus apacheanus 0.2 19.0 The wings possessed dark lines (Fig. 1), which, Chamberlin & because of the translucence of the rest of the wing, Gertsch appeared as legs. This salticid image was further 19 Phidippus sp. B 23.8 15.3 enhanced by dark abdominal spots (arranged in a x ± SD 5.14 ± 7.00 18.03 ± 12.74 pattern resembling spider eyes), and a proclivity a to rotate the wings outward and upward. This ro- Spider backed away from wing-rotating Z. vittigera. b Spider responded to Z. vittigera with a threat display then tation caused the dark wing lines to move up and backed away. down in a manner resembling the foreleg threat c Spider responded to Z. vittigera by repeatedly signaling with or courtship displays common to salticids (Foelix first one leg then the other. 1982, Jackson 1982b,c,d, Witt & Rovner 1982). The flies were in nearly constant motion on both to Z. vittigera wing flicks, some spiders performed the upper and lower S. elaeagnifolium vegetation behaviors similar to salticid sexual or aggressive surfaces, exhibiting short, jerky locomotory move- displays (Tables 1 and 2); the front legs were jerked ments reminiscent of the stereotypic movements upward, or moved in a repetitious manner, and of jumping spiders. They generally remained with- the whole prosoma was lifted up, exposing the che- in the S. elaeagnifolium vegetation, making sud- licerae. Such displays were never elicited by house den short flights from adjacent leaves or stems and flies or wing-painted Z. vittigera. Courting of spi- occasionally to adjacent plants. ders by male fruit flies did not result in predation. In the laboratory both sexes, but particularly In the two cases when male flies landed on spiders, males, increased wing flicking in response to the the spiders became highly agitated and attempted presence of other small objects such as other te- to escape. phritids, house flies, or spiders. During the exper- Experiment 1: House Flies Versus Normal Fruit iments, male Z. vittigera sometimes "courted" Flies. During the 50-min observation period, 12 jumping spiders by facing them and moving back out of 19 M. domestica were captured, whereas and forth sidewise while rotating their wings. Twice, significantly fewer (7 out of 19; binomial test for male flies actually flew onto the backs of spiders changes, P = 0.031) Z. vittigera were taken (Table but immediately decamped and flew away. 1). In addition, the salticids required a significantly Salticid Behavior in the Laboratory. During longer time (18.0 ± 12.7 min versus 5.1 ± 7.0 min; successful prey capture, salticids oriented toward Mann-Whitney U = 10, P < 0.01) to make the Z. and stalked flies, then at a distance of 1-6 cm, made vittigera captures, indicating a reluctance to at- a short leap that ended in prey capture. This se- tack. Half the M. domestica captures took place quence was typical of house fly encounters. How- in less than 2.5 min, whereas none of the Z. vittigera ever, most salticids failed to attack normal Z. vit- captures occurred before 7 min had elapsed. In tigera, and instead backed up or rotated and faced, response to Z. vittigera, two spiders displayed and but did not advance toward, the prey. In response three backed away (Table 1). 534 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 81, no. 3 Table 2. Jumping spider response to unpainted (nor- and that this mimicry reduces attack from this mal) and wing-painted Z. vittigera during 10-min trials predacious group. Capture latency (min) We feel that spiders mistook the tephritid flies for other jumping spiders. Salticid spiders are well Spider Species Normal Wing-painted known to use vision in both hunting and courtship Z.