2010 NORTHEASTERN NATURALIST 17(1):115–124 Prey and Nesting Behavior of Some North American Spider Wasps (Hymenoptera: Pompilidae) Frank E. Kurczewski* Abstract - This study presents prey records and nesting information for 13 species of spider wasps: Hemipepsis ustulata, Priocnessus nebulosus, and Priocnemis cornica (Tribe Pepsini); Auplopus architectus, A. caerulescens (Tribe Auplopodini); and Tachypompilus ferrugineus, T. unicolor, Anoplius (Pompilinus) marginatus complex, A. (P.) splendens, A. (Anoplius) nigerrimus, A. (A.) virginiensis, Arachnospila arctus, and A. scelestus (Tribe Pompilini). Cumulative prey records for Priocnessus nebulosus, Tachypompilus ferrugineus, T. unicolor, and Arachnospila scelestus indicate limited prey selection. Cursorial hunting spiders were more susceptible to predation by the spider wasps than web-spinning spiders. Wasp nesting sites varied among natural and man-made microhabitats. Introduction The family Pompilidae has approximately 300 species in North America north of Mexico (Krombein 1979). Although more than 2000 prey records have been published for this group, there is much to learn about prey selection of the individual species (Kurczewski et al. 1987). A summary of prey spiders for the North American species of Pompilidae is presented in the Catalog of Hymenoptera in America North of Mexico (Krombein 1979). Wasbauer (1982) added prey records for some spider wasps from the western United States. Kurczewski et al. (1987) supplemented prey records and nesting behavior for some eastern pompilids. Evans and Yoshimoto (1962) reported on the ecology and nesting behavior of some Pompilidae from the northeastern United States. More recent additions to the ecology and nesting behavior of this fauna include studies on Calicurgus hyalinatus (Fabricius) (Kurczewski and Spofford 1985), Auplopus caerulescens (Dahlbom) (Kurczewski 1989a), Ageniella fulgifrons (Cresson) (Kurczewski and Kurczewski 1987), Episyron quinquenotatus (Say) (Kurczewski 2001), Tachypompilus ferrugineus (Say) (Kurczewski 1989b), Anoplius relativus (Fox) (McQueen 1978), Anoplius cylindricus (Cresson) (Gwynne 1979), Anoplius tenebrosus (Cresson) (Alm and Kurczewski 1984), Anoplius depressipes Banks (Roble 1985), and Aporinellus wheeleri Bequaert and A. taeniolatus (Dalla Torre) (Kurczewski et al. 1988). Despite these reports, there is still much to learn about the prey and nesting behavior of the North American species of Pompilidae. The nesting behavior of many species remains unstudied. Certain species are highly secretive in their habits and difficult to study because of their cryptic nesting *PO Box 15251, Syracuse, NY 13215; Fkurczewski@twcny.rr.com. 116 Northeastern Naturalist Vol. 17, No. 1 situations. Some species, such as Anoplius bengtssoni (Regan), are evidently crepuscular or nocturnal in their nesting behavior (Kurczewski 1999), which further adds to the difficulties of their study. This study adds previously unknown spider prey and presents typical and atypical nesting behavior for 13 pompilid species. The species of Pompilidae are presented according to their arrangement in the Catalog of Hymenoptera in America North of Mexico (Krombein 1979). The spider names follow the current classification of Platnick (2009). Wet weights or body lengths of the wasps and spiders are given along with the species identifications. Localities, dates of collection, and observations on nesting behavior are also recorded. The wasps are placed in the Cornell University insect collection. The spiders have been deposited in the State University of New York College of Environmental Science and Forestry invertebrate collection. SUBFAMILY PEPSINAE Tribe Pepsini Hemipepsis ustulata Dahlbom CA: Along 17-Mile Drive near Pescadero Point, Pebble Beach, Monterey County; 19 August 2007; 1525–1630 (PDT); 18 °C (air temperature). This female, 24 mm long, was excavating her burrow in sandy clay-loam atop an ocean-side cliff, 140 cm from where she deposited her spider, an immature Aphonopelma sp. (Theraphosidae), 31 mm long. She removed soil from her burrow by walking backward and using the forelegs alternately. She held her bright orange wings on the dorsum except for occasionally flicking them when on the surface. She backed from her burrow with soil 14 times in 45 minutes, excluding the duration she was absent from the nest area. The soil she removed from the burrow formed a tumulus, 75 mm wide and 125 mm long, in front of the opening. This wasp left her burrow and flew to examine her spider 3 times at intervals of 11–18 minutes. She returned to her burrow each time walking to recommence excavation. I collected the wasp and prey before she finished the burrow. Priocnessus nebulosus (Dahlbom) PA: Wintergreen Gorge Cemetery, 1.6 km SE of Erie, Erie County; 28 July 1987. This wasp (weight = 63 mg) was observed carrying a female Agelenopsis utahana (Chamberlin & Ivie) (Agelenidae) (weight = 94 mg) forward on the ground. The female straddled the spider ventral side upward, grasping it by the base of a chelicera (?). The wasp released her prey on a gravelly bank in the shade, ventral side upward, and flew back and forth in front of a Marmota monax (L.) (Woodchuck) hole, at which time the pair was collected. Priocnemis cornica (Say) NY: Auburn, Cayuga County; 2 September 1991. This female (weight = 5 mg) was walking backward, grasping a male Cheiracanthium inclusum (Hentz) (Miturgidae) (weight = 9 mg) by a hindcoxa. The wasp held the spider in an almost perpendicular position during transport. 2010 F.E. Kurczewski 117 Tribe Auplopodini Auplopus architectus (Say) NY: Onondaga Hill, 3.2 km SW of Syracuse, Onondaga County; 16 August 1996; 1615 (EDT); 25 °C (air temperature). This wasp was transporting a female Phidippus sp. (Salticidae) across a concrete step on an intermittently rainy day. Each leg of the spider had been amputated at the coxa-trochanter joint. With her mandibles, the female held the spider ventral side upward by its spinnerets. Maintaining the spider in this position, the wasp walked up the side of a house and entered a crack in a shingle. Another female or the same one (weight = 24 mg) was collected the next day at the same location with a female Phidippus whitmani Peckham & Peckham (weight = 59 mg). The wasp was carrying the spider, which had all its legs amputated at the coxa-trochanter joints, in the same manner. Auplopus caerulescens (Dahlbom) NY: Auburn, Cayuga County; 12 August 1991. This female (weight = 6.5 mg) was seen carrying a female Habronattus decorus (Blackwall) (Salticidae) (weight = 11.5 mg), which had all its legs amputated at the coxatrochanter joints, up the outside wall of a basement foundation. Maintaining the prey underneath in a ventral side upward position and grasping it by the spinnerets with the mandibles, the wasp proceeded toward a hole in the mortar between two concrete blocks, at which time the pair was collected. NY: Onondaga Hill, 3.2 km SW of Syracuse, Onondaga County; 8–27 July 2007. Two females were observed nesting between a window screen and windowpane of a house. They periodically entered an opening with small immature Pellenes and Sitticus spp. (Salticidae). The wasps occupied this site for approximately three weeks. SUBFAMILY POMPILINAE Tribe Pompilini Tachypompilus ferrugineus (Say) NY: Clark Reservation State Park, 2.1 km W of Jamesville, Onondaga County; 19 August 1992; 1935 (EDT); 19 °C (air temperature). This wasp (weight = 186 mg) was nesting beneath a rusted truck tailgate lying atop soft pulverized shale mixed with deciduous twigs, pine needles, and other dried vegetation. A partly paralyzed female Rabidosa rabida (Walckenaer) (Lycosidae) (weight = 428 mg), with legs tucked in but moving, was found dorsal side upward in a concave depression in the shale beneath the tailgate. The depression was about 30 mm deep and 45 mm in diameter at the top. The wasp continued to walk around the area with raised wings as I removed the spider from the concavity. If present, the wasp’s egg was dislodged from the spider’s abdomen during the prey’s removal from the nest. Tachypompilus unicolor (Banks) CA: Montana de Oro State Park along Pecho Valley Road, 4.0 km S of Los Osos, San Luis Obispo County; 9, 11 August 2009; 1510–1525, 1423– 118 Northeastern Naturalist Vol. 17, No. 1 1445 (PDT); 20 °C (air temperature). Two (possibly three) females, 14–15 mm long, were seen searching for prey in the chaparral-covered foothills in an area of sandy clay-loam, ca. 10 m2. The wasps searched on the ground primarily under dried vegetation, in crevices, and in holes up to 3 cm in diameter. Females remained inside some holes for 0.5–1.5 minutes. The wasps ran rapidly in an almost straight line, but occasionally zigzagged while hunting. Their antenna tips constantly tapped the substrate. They held their wings upward at a 45° or greater angle and intermittently flicked them once or twice per second. The females occasionally made low, rapid flights up to 1 m in length to new hunting areas. Their hunting activity ceased abruptly at the onset of coastal marine fog and disappearance of full sunlit conditions. Anoplius (Pompilinus) marginatus (Say) complex NY: Auburn, Cayuga County; 13 September 1991. This wasp (weight = 27 mg) was captured dragging a female Schizocosa crassipes (Walckenaer) (Lycosidae) (weight = 52 mg) backward across the edge of a lawn. The female grasped the spider by the base of a hindleg with the mandibles. MI: 1.5 km N of Grayling, Crawford County; 9 August 1992. A female (weight = 16 mg), with very frayed wings, was noted transporting a female Thanatus formicinus (Clerck) (Philodromidae) (weight = 53 mg) on sand, grasping the spider as described above. Another wasp (weight = 17 mg) at that locality on the same day was observed dragging an immature Xysticus sp. (Thomisidae) (weight = 23 mg) across sand in the same manner. Anoplius (Pompilinus) splendens (Dreisbach) MI: Allegan State Game Area, Fennville, Allegan County; 13 June 1993. This wasp (weight = 22 mg) was collected dragging an immature Thanatus sp. (weight = 23 mg) rapidly backward across sand. The female grasped the spider with the mandibles as described for the Anoplius marginatus complex. MI: 1.5 km N of Grayling, Crawford County; 9 August 1992. This female (weight = 23 mg) was captured dragging an immature Schizocosa sp. (weight = 31 mg) backward on sand. The 2nd right leg of the prey was missing beyond the coxa-trochanter joint. The female grasped the spider with the mandibles as described above. Anoplius (Anoplius) nigerrimus (Scopoli) NY: Onondaga Hill, 3.2 km SW of Syracuse, Onondaga County; 24–25 September 2009. Three females were seen hunting during two afternoons (1344–1651, EDT; air temperatures = 18–21 ºC; ground surface temperatures = 33–41 ºC). The wasps searched for spiders around and under stones, in cavities in the soil, beneath decumbent vegetation, and in grasses at the edge of a lawn. One female searched unsuccessfully in an almost straight line for 13 m during 35 minutes. One wasp with moderately frayed fore- and hindwings, 11 mm long, entered a patch of decumbent vegetation on bare soil. Several seconds later, a small lycosid ran from the vegetation followed rapidly by the wasp. The female caught the spider after 15 cm, bent her abdomen underneath, and inserted her sting into its ventral cephalothorax for 1–2 seconds. She 2010 F.E. Kurczewski 119 dismounted and, using her mandibles, manipulated the prey’s mouthparts (?) and forelegs for 3–4 seconds. She then grasped the spider by the base of a 3rd leg with her mandibles and, holding it above ground, walked rapidly backward across the bare soil. The female released the prey dorsal side upward beside a small hole, entered, reappeared in the opening, and, grasping the spider by its spinnerets with the mandibles, pulled it backward into the burrow. The wasp reappeared in the entrance 4 minutes later and began pulling soil and debris into the burrow with her mandibles and forelegs. I collected the female before she finished filling the burrow. The entrance and burrow were only 5 mm wide. The burrow sloped downward almost vertically, ending in a rough circular cell, 6.5 x 12 mm, 34 mm below the surface. A female Trochosa ruricola (DeGeer), 11 mm long, was placed cephalothorax outward and dorsal side upward in the cell. The wasp’s egg, ca. 2.5 mm long, was affixed to the left side of the spider’s abdomen almost midway from the base. A female with severely frayed fore- and hindwings, 11 mm long, was observed beginning a burrow at 1259, EDT. She used her mandibles to loosen the loamy soil and her mid- and hindlegs to push the loosened soil to the surface. When inspected, this wasp’s left mid- and hindtibiae were caked with dried mud. Every several minutes, the female exited backward and distributed the loose soil around the opening, resulting eventually in a shallow tumulus 55 mm wide and 75 mm long. The wasp removed soil through the entrance 13 times during the 37 minutes of burrow excavation. At 1336, after removing soil from the burrow, she turned in front of the entrance and walked slowly away in a northeasterly direction. Staying entirely on the ground and holding her wings on the dorsum, she disappeared into dense vegetation. At 1351, the female reappeared walking across the bare ground toward her entrance. She examined the opening and, walking slowly, retraced her path into the dense vegetation. At 1422, she flew back to her entrance, examined it, and walked away much more rapidly than before, perhaps because the ground surface temperature had increased from 33 to 38 ºC. At 1448 (air temperature = 18 ºC, ground surface temperature = 38 ºC), more than an hour after finishing her excavation, the wasp returned walking rapidly backward holding a paralyzed wolf spider with her mandibles above ground by the base of a 3rd or 4th leg. Her return with prey was from the opposite direction (southwest) in which she left, implying that she had dug her burrow before hunting. The burrow, 5.5 mm wide, sloped almost vertically, and the cell, 6.5 x 13 mm, was placed at a right angle at the bottom. The distance from the entrance to the cell was 29 mm. The spider, a female Trochosa ruricola, 10 mm long, was positioned in the cell as above and the egg, 2.5 mm long, was affixed similarly. The spider recovered from the effect of the wasp’s venom within 24 hours and was able to move its legs fully, possibly because the wasp was very old. Anoplius (Anoplius) virginiensis (Cresson) NY: Onondaga Hill, 3.2 km SW of Syracuse, Onondaga County; 31 August– 17 September 2009. Females were seen hunting in mid-late afternoon 120 Northeastern Naturalist Vol. 17, No. 1 (1410–1655, EDT; air temperatures = 19.5–25.0 ºC; ground surface temperatures = 31–42 ºC). They searched for prey in subterranean burrows and cavities, around stones, beneath decumbent vegetation, and under the bark of trees at ground level. Most females hunted apparently randomly but rather thoroughly through 1–2 m2 areas for 5–10 minutes before searching elsewhere. Hunting wasps walked or ran forward tapping their antennae on the surface, occasionally flicked their wings, and made long (>5 m) winding flights to new areas. One female, 11 mm long, entered dense vegetation at ground level. Less than 1 minute later, a small wolf spider hurriedly exited the vegetation and ran across the stony ground with the wasp flying in rapid pursuit. The pompilid twice landed just behind the spider, but failed to capture it. Upon landing a 3rd time, apparently atop the wolf spider, the wasp rapidly inserted her sting into the underside of the spider’s cephalothorax stopping its movement. From the time the spider exited the vegetation until it was subdued by the wasp, only 5–6 seconds elapsed. The site of the capture was 0.9 m from the vegetation. The wasp repositioned the spider’s legs with her mandibles, turned it over, and applied her mouthparts to the spider’s ventral cephalothorax for 7 seconds. She then grasped the spider by the base of a hindleg with her mandibles and began dragging it backward toward a decaying pine stump 1.6 m away. During transport, the wasp released the spider three times, walked straight to the stump, and then returned to the prey walking in a straight line. She grasped the spider as before, dragged it farther, released it ventral side upward at the base of the stump, and entered a crevice between two large bare roots. The wasp exited headfirst, grasped the spider by its spinnerets with the mandibles, and pulled it into the opening. After waiting a few minutes, I excavated the area of entry. The paralyzed spider was located, positioned on its side with its cephalothorax outward, 54 mm from the mouth of the crevice in a rough cell, 7 mm wide and 12 mm long. The female Trochosa ruricola, 10.5 mm long, was completely motionless. It had the wasp’s egg attached to the upper left side of its abdomen, about two-fifths of the way from the base. Excavating deeper, I found two or three old A. virginiensis cells from prior generations containing dried spider body fragments and coarse sawdust partitions. Another female, 13.5 mm long, was observed dragging a funnel-webweaving spider backward, grasping it in the same manner as described above. She, too, released her prey three or four times during transport, walked straight to another decaying pine stump, then returned to the spider on the ground walking in a straight line. This wasp pulled the spider into an opening in the base of the pine stump, after releasing it on the ground in a dorsal side upward position, entering the opening for a minute, and exiting. After a few minutes, the wasp was seen pulling pieces of moist, decaying wood from the sides of the opening with her mandibles and placing them farther down the cavity, at which time I excavated the nest. A female Agelenopsis pen[n]sylvanica (C.L. Koch) (Agelenidae), 14 mm long, was located on its side with its cephalothorax outward in a rough cell, about 7 mm wide and 15 mm long, about 40 mm from the opening. The wasp’s egg was affixed 2010 F.E. Kurczewski 121 to the upper right side of the spider’s abdomen, about one-third of the way from the base. Two other wasps, 8 and 9 mm long, were collected dragging backward on the ground incompletely paralyzed immature Agelenopsis pen[n]sylvanica, 9.5 and 10 mm long. These or other females were twice seen walking across and entering funnel webs in low vegetation without getting entangled. The spiders must have escaped through the opening of the funnel web as both searches proved fruitless. Arachnospila arctus (Cresson) NY: Auburn, Cayuga County; 7 September 1988. This wasp (weight = 28 mg) was collected slowly dragging a male Arctosa rubicunda (Keyserling) (Lycosidae) (weight, 81 mg) backward at the edge of a backyard patio. The female grasped the spider by the base of a hindleg with the mandibles. NY: Onondaga Hill, 3.2 km SW of Syracuse, Onondaga County; 9–17 October 2008. Two females were observed for more than a week nesting among pebbles and loamy soil near a house foundation. They hunted at the bases of shrubbery by running rapidly on the ground, flicking their wings, and making quick short flights to new areas. Prey transport involved grasping the spider by the base of a hindleg with the mandibles and dragging it backward on the ground. The spider was pulled into the nest entrance by its spinnerets with the wasp’s mandibles. A female nesting beneath an outdoor ground light entered with prey through a notch in the base of the light! A nest in an open area comprised an oblique burrow, 75 mm long, that ended in a round-shaped cell. The paralyzed spider was placed in the cell dorsal side upward and cephalothorax outward. The wasp’s egg was affixed transversely to the underside of the spider’s abdomen about midway from the base. Three spiders collected from wasps or their nests were identified as female Schizocosa crassipes (weights = 44, 47, 41 mg). One wasp weighed 28 mg. Arachnospila scelestus (Cresson) MI: Allegan State Game Area, Fennville, Allegan County; 13 June 1993. This wasp (weight = 52 mg) was observed slowly dragging a female Schizocosa avida (Walckenaer) (Lycosidae) (weight = 254 mg) backward on sand. The 4th right leg of the spider had been amputated at the coxa-trochanter joint. The wasp grasped the base of the prey’s hindleg with the mandibles during transport. Discussion Hemipepsis ustulata preys upon the trap-door spider Brachythele longitarsis (Simon) (Nemesiidae) and tarantulas of the genus Aphonopelma (Williams 1956), as observed in this paper. All known prey records for Priocnessus nebulosus are for the genus Agelenopsis (Krombein 1979). Virtually nothing is known about the nesting behavior of this species (Evans and Yoshimoto 1962). Priocnemis cornica is very unselective of its prey, capturing nine families of cursorial hunting spiders (Krombein 1979, Kurczewski 1999, Kurczewski 122 Northeastern Naturalist Vol. 17, No. 1 et al. 1987, this study). This species nests in pre-existing holes in the ground (Evans and Yoshimoto 1962). Auplopus architectus builds mud nests in sheltered places and provisions the nests with spiders belonging to the families Gnaphosidae, Clubionidae, Thomisidae, and Salticidae (Kurczewski 1999). Several prey records comprise the genus Phidippus (Krombein 1979). Auplopus caerulescens also builds mud nests in sheltered spots and preys on cursorial hunting spiders of the families Clubionidae, Anyphaenidae, Thomisidae, and Salticidae (Krombein 1979, Kurczewski 1989a). Tachypompilus ferrugineus is rather selective of its prey, capturing spiders of the families Pisauridae and Lycosidae in the eastern United States and Ctenidae in the western United States (Krombein 1979, Kurczewski 1989b, Wasbauer 1982). This species is reported to nest in weathered mortar of an abandoned building foundation (Rau and Rau 1918), in loose dry soil beneath buildings (Strandtmann 1953), and in mixed loose soil, powdered mortar, and debris underneath the base of a cemetery monument (Kurczewski 1989b). Nothing has been published on the nesting or prey of Tachypompilus unicolor. However, three snippets on this species were found on internet photograph- and video-sharing sites (Flickr, YouTube, pics.WW.com; Anonymous 2009, Bleon 2006, Williamson 2007 ). A video shows a female T. unicolor carrying a female lycosid backward in a dorsal side upward position, grasping it with her mandibles by a chelicera or pedipalp and, later, ventral side upward by a 2nd (?) leg. All prey were large female wolf spiders (Lycosidae) that appeared to weigh several times the wasp’s weight. The images indicate that T. unicolor hunts lycosids near suburban homes in the southwestern United States, although I found females searching for prey in chaparral-covered foothills. The Anoplius marginatus complex consists of five species recognizable in the male sex by their genitalia. The females are indistinguishable from one another (Evans and Yoshimoto 1962). The species identified as “Anoplius marginatus” is reported to capture 11 families of spiders and 1 family of harvestman. This species complex excavates its own nest from the soil surface or uses pre-existing holes or burrows in the ground (Evans and Yoshimoto 1962). Anoplius splendens has been collected with 10 families of cursorial hunting spiders, but some of the records may pertain to the closely related A. rectangularis (Dreisbach), which is a species in the Anoplius marginatus complex. Anoplius splendens excavates its nest from the soil surface or uses pre-existing burrows in the ground. Anoplius nigerrimus is holarctic, inhabiting Europe and North America (Evans 1951, Krombein 1979). In Europe, A. nigerrimus nests chiefly in pre-existing cavities and preys mainly on Lycosidae but also Pisauridae and Gnaphosidae (Richards and Hamm 1939). The only prey record for this species from North America is another holarctic species, Trochosa terricola Thorell (Lycosidae) (Kurczewski et al. 1987). Trochosa ruricola, the prey reported in this paper, is also holarctic. My observations imply that this spider wasp may excavate a burrow before capturing the spider. 2010 F.E. Kurczewski 123 Anoplius virginiensis preys on Agelenidae, Amaurobiidae, and Lycosidae in open woodland (Krombein 1979, Kurczewski and Kurczewski 1973). This species nests in pre-existing crevices and cavities in decaying stumps and logs (Evans and Yoshimoto 1962). Arachnospila arctus preys on cursorial hunting spiders of the families Lycosidae, Gnaphosidae, Amaurobiidae, and Clubionidae (Krombein 1979). Little is known about its nesting behavior (Evans and Yoshimoto 1962). Prey records for Arachnospila scelestus include mainly Lycosidae, and, rarely, Pisauridae and Salticidae (Krombein 1979). Prey species of Lycosidae are often considerably larger than the wasp, sometimes weighing 5–10 times as much, and transported with some difficulty (Kurczewski and Kurczewski 1973, Kurczewski et al. 1987). This species excavates nests primarily in sandy soil, beginning from the surface. Acknowledgments David McCorquodale, Cape Breton University, kindly agreed to serve as Guest Editor and reviewed the manuscript in detail. John Lerg provided the facilities of the Allegan State Game Area, Michigan. My father, Edmund Kurczewski (deceased), and Teresa Doherty, State University of New York College of Environmental Science and Forestry, assisted with some of the field studies. My son, Keith Kurczewski, and Esther Clark assisted with the observations on Hemipepsis ustulata and Tachypompilus unicolor. Bonnie Brierton and Roy Norton, both of the State University of New York College of Environmental Science and Forestry, identified many of the prey spiders. James Pitts, Utah State University, named the spider prey of Tachypompilus unicolor from photographs and a video. Partial funding for the field research was provided by the Michigan Chapter of The Nature Conservancy and New York State United University Professions. Literature Cited Alm, S.R., and F.E. Kurczewski. 1984. Ethology of Anoplius tenebrosus (Cresson) (Hymenoptera: Pompilidae). Proceedings of the Entomological Society of Washington 86:110–119. Anonymous. 2009. Redwasp-spider 675. 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