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Thomas Bilbo and Alex Benedick Wikipage: Pompilidae Family: Pompilidae Spider Wasps (Order: Hymenoptera, Suborder: Apocrita)

) The family Pompilidae are often referred to as spider wasps because of their distinctive behavior of using spiders as the living host for their larvae. This cosmopolitan family (meaning they occur throughout most of the world) of solitary wasps includes approximately 5000 species in 230 genera. Like other widespread insect families, they are most diverse in tropical regions of the world [1]. They belong to the superfamily Vespoidea which also includes the well known family Vespidae (paper wasps, hornets, and yellow jackets). Morphology Like other strong fliers, the thorax of Pompilidae is modified for efficient flight. The mesothorax is solidly fused to the pronotum and metathorax; Moreover, the prothorax is best developed in species such as Pompilidae and Scoliidae because they use their forelegs to dig (more in Behavior) [2]. Pompilids typically have a slender body with long, spiny legs and the hind femur is often long enough to reach past the tip of the abdomen. The tibiae of the rear legs usually have a conspicuous spine at their distal end. The first two segments of the abdomen being narrowed characterize the slender look of their body. The Pompilid body is typically dark (black or blue, sometimes with metallic reflections), but many brightly colored species exist. From a lateral view, its pronotum looks rectangular and it extends back to the tegulae, near the base of the wings. Most species are macropterous (having long wings), but a few brachypterous (short-winged) and apterous (no wings) are known [4]. Wings are typically tinted red/orange. An almost universal characteristic is the presence of a transverse suture on the mesopleuron (Fig. 1). This suture looks something like an indented ridge running across the mesopleuron. As hinted above, the dark body and bright wings (as well as a distinct odor in Hemipepsis and Pepsis sp.) are aposematicmeaning they are warning signals, specifically of its sting. Pompilids, especially the tarantula hawk species (Hemipepsis and Pepsis sp.) have very powerful stings (see Schmidt Pain Index below) and an enemy with the ability to learn will avoid pompilids. Sexual dimorphism is slight, so males look similar to females except for the presence of a stinger, which predators cannot notice. The similarity in coloring is an example of automimicry [2]. The most noticeable difference between males and females is the larger female body size in most species.

Image 1: Red arrow is pointing to the pompilids transverse suture on the mesopleuron.

Ecology and Behavior Currently, Pompilidae is split unequally into six subfamilies throughout most of the world. Ceropalinae and Notocyphinae contain two genera each and occur in Central and South America and the Oriental region. Epipompilinae contains a single genus and occurs in the Neotropics, the Australasian region, and the Afrotropical region. Ctenocerinae contains two genera in the Neotropics, four in Australia and 11 in Africa. Pepsinae and Pompilinae are the most diverse and the remaining genera are split between them [4]. As mentioned above, Pompilidae gets its common name, Spider Wasps, from the their notable behavior of hunting and killing spiders, often larger than themselves, as food for their larvae. They provide each of their larvae with a single prey item/ host, which must be large enough to serve as its food source throughout its development. It has been shown in Pepsis thisbe of the southwestern United States that there is a direct correlation between adult wasp body length and the weight of its host spider, Aphonopelma echina. Because the size of a P. thisbe adult is determined by the size of the host that was provided for it by its mother, it is implied that the seasonal frequency of host sizes will determine the size variation in adult wasps [5]. Because of the large body size of their prey, pompilids usually will either construct burrows near the site of attack or use the hosts own burrow or tunnel. The paralyzed spider is concealed in a burrow so that the pompilid larva can develop without disruption by other parasites or scavengers. Some pompilids only temporarily paralyze their host, which regains activity before being killed by the maturing wasp larva [2]. In another study on Pepsis thisbe [7], it was shown that chemosensory cues are used to detect its specific host. This study concluded that specific chemosensory cues attract the wasp to its prey,

Aphonopelma echina, despite other host spiders of the same size and frequency being present (see Image 2). In studies on Pepsis formosa [6], a pompilid of the southwestern United States, it was shown that the wasps had behavioral plasticity. Their hunting behavior concerning their host Rhechostica echina improved with experience. The time required completing all behavioral components decreased with each spider killed (see image 3). Concerning mating behavior, males acquire perch territories to scan for incoming receptive females. In studies on the tarantula wasp Hemipepsis ustulata [3], it was shown that larger males are more likely to acquire perch territories and that territorial males appear to increase their chances of mating because receptive females fly to perch sites held by said males (see image 4).

Image 2: Pepsis thisbe

Image 3: Pepsis formosa

Image 4: Hemipepsis ustulata >>>>VIDEO http://www.youtube.com/watch?v=ctAXcBxHdCg&feature=player_embedded

Life Cycle and Diet Unlike many other families in the order hymenoptera, wasps in this family are solitary and nest alone [8]. A female wasp will search the ground and tree trunks for a spider and upon finding the spider will sting it, rendering the spider paralyzed. Once the spider is paralyzed, the female wasp will make a burrow or take the spider to a previously made burrow [9]. The female wasp will lay one single egg on the abdomen of the spider using its ovipositor and then enclose the spider in the burrow [9]. The egg will hatch and the larva will feed on the spider, breaking through the integument with its mandibles [5]. The larva has 5 instar stages before it pupates, where there are no major morphological differences between the first 4 instars, with the exception of size [5]. At the conclusion of the final instar, the larva will spin a silky cocoon where it will emerge as an adult either later in the same summer season or will overwinter, depending on the species and the time of year the larva pupates [5]. Adult Pompilidae are nectar-feeding insects and feed on a variety of plants [8, 10]. The female wasps search for a variety of spiders for their larva to feed on, including wolf spiders (Lycosidae), huntsman spiders (Sparassidae), and baboon spiders (Harpactirinae) [8]. As the larva feeds on its host, it saves the vital organs, such as the heart and central nervous system, for last. By waiting until the final larva instar, it ensures that the spider will not decompose before the larva has fully developed [10].

Image 4: Pompilidae larva feeding on tarantula

Schmidt Pain Index In 1984, Joseph O. Schmidt, a researcher from Arizona, developed a Hymonpeteran sting pain scale; what is now known as the Schmidt Sting Pain Index. This index runs from 0 to 4, where a 0 is given to a sting from an insect that can not break through human skin, a 2 is given for intermediate pain, and a 4 is given for intense pain. The scale rates stings from 78 different species in 42 different genera [11]. Spider Wasps of the genera Pepsis, also known as Tarantula Hawks, have a sting rating of 4.0. The sting is described as "blinding, fierce, shockingly electric. A running hair drier has been dropped into your bubble bath" [12]. Only the bite of the Bullet Ant, Paraponera clavata, is ranked higher, with a 4.0+ rating. References [1] Wasbauer, M.S. 1995 Pompilidae. In: P.E. Hanson & I.D. Gauld (Eds) The Hymenoptera of Costa Rica (pp.522-539). Oxford: Oxford University Press. [2] Daly, Howell V., John T. Doyen, and Alexander H. Purcell. Introduction to Insect Biology and Diversity. Oxford: Oxford UP, 1998. [3] Alcock, J. & Kemp, J. 2006. The behavioral significance of male body size in the Tarantula Hawk Wasp Hemipepsis ustulata (Hymenoptera: Pompilidae). Ethology 112: 691-698. [4] Pitts, J., Wasbauer, M. S., and Dohlen, C. D. 2005. Preliminary morphological analysis of relationships between the spider wasp subfamilies (Hymenoptera: Pompilidae): revisiting an old problem. Zoologica Scripta 35: 63-84. [5] Punzo, F. 1994. The biology of the spider wasp, Pepsis thisbe (Hymenoptera: Pompilidae) from Trans Pecos, Texas. I. Adult morphometrics, larval development and the ontogeny of larval feeding patterns. Psyche 101: 229-242.

[6] Punzo, F. and Garman, B. 1989. Effects of encounter experience on the hunting behavior of the spider wasp, Pepsis Formosa (Say) (Hymenoptera: Pompilidae). The Southwestern Naturalist 34: 513-518. [7] Punzo, F. and Ludwig, L. 2005. Behavioral Responses of Pepsis thisbe (Hymenoptera: Pompilidae) to Chemosensory Cues associated with host spiders. Journal of Insect Behavior 18: 757766. [8] CSIRO Fact Sheet-http://www.csiro.au/resources/Spider-Wasp-Factsheet.html [9] Australian Museum-http://australianmuseum.net.au/Spider-wasps [10] Punzo, F. 2005. Studies on the natural history, ecology, and behavior of Pepsis Cerberus and p. mexicana (Hymenoptera: Pompilidae) from Big Bend National Park, Texas. Journal of the New York Entomological Society, 113(1): 84-95.

[11] Berenbaum, M. 2003. A Stinging Commentary. American Entomologist,49(2): 68-69.

[12] http://tucsoncitizen.com/wryheat/2010/07/16/tarantula-hawks-deliver-the-big-sting/ Image 1:Image by Troup1 on Flickr. Spider wasp in Prachuap Khiri Khan, Thailand. >http://www.flickr.com/photos/23271361@N06/6061812016/in/photostream/< Inner Image: >http://quizlet.com/3395420/insect-ordersfamilies-with-images-flash-cards/< Image 2: http://images.dpchallenge.com/images_challenge/10001999/1419/800/Copyrighted_Image_Reuse_Prohibited_960970.jpg Image 3: http://tucsoncitizen.com/wryheat/files/2010/07/TarantulaHawk-large3-550x420.jpg Image 4: http://farm4.static.flickr.com/3111/2733563768_071cd73127.jpg Image 5: http://www.qm.qld.gov.au/Find+out+about/Animals+of+Queensland/Insects/Wasps+and+bees/Comm on+species/Spider+Wasps Video: http://www.youtube.com/watch?v=ctAXcBxHdCg&feature=player_embedded

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