2012 SOUTHEASTERN NATURALIST 11(Monograph 4):1–71 Hosts, Nesting Behavior, and Ecology of Some North American Spider Wasps (Hymenoptera: Pompilidae) Frank E. Kurczewski1,* and G.B. Edwards2 Abstract - This study broadens the range of host, behavioral, and ecological information for 65 pompilid taxa from North America: Pepsis sp., Entypus fulvicornis, E. unifasciatus californicus, E. u. unifasciatus, Priocnemis (Priocnemissus) minorata, Priocnemis (Priocnemis) cornica, P. (P.) germana, P. (P.) scitula relicta, Caliadurgus fasciatellus alienatus, Dipogon (Deuteragenia) c. calipterus, Dipogon (D.) s. sayi, Phanagenia bombycina, Auplopus a. architectus, A. architectus metallicus, A. bellus, A. caerulescens subcorticalis, A. carbonarius, A. m. mellipes, A. mellipes variitarsatus, A. nigrellus, Ageniella (Priophanes) arcuata, A. (Ageniella) accepta, A. (A.) coronata, A. (A.) euphorbiae, A. (Ameragenia) salti, Aporus (Plectraporus) hirsutus, Evagetes p. padrinus, Agenioideus (Agenioideus) humilis, Sericopompilus apicalis, S. neotropicalis, Episyron b. biguttatus, E. conterminus cressoni, E. oregon, E. quinquenotatus hurdi, E. q. quinquenotatus, E. snowi, Poecilopompilus a. algidus, P. algidus coquilletti, P. algidus willistoni, P. i. interruptus, Tachypompilus f. ferrugineus, T. u. unicolor, Anoplius (Lophopompilus) aethiops, A. (L.) atrox, A. (L.) cleora, A. (Notiochares) lepidus atramentarius, A. (Arachnophroctonus) americanus trifasciatus, A. (A.) insignis bahamas, A. (A.) nigritus, A. (A.) semicinctus, A. (A.) semirufus, A. (Pompilinus) insolens, A. (P.) marginatus complex, A. (P.) tenebrosus, A. (Anoplius) depressipes, A. (A.) imbellis, Ammosphex a. angularis, A. a. anomalus, A. s. solonus, Arachnospila arctus, A. fumipennis eureka, Aporinellus completus, A. medianus, A. t. taeniatus, and A. yucatanensis. New host spider families are introduced for Entypus unifasciatus (Pisauridae), Priocnemis minorata (Dysderidae), P. germana (Miturgidae), P. scitula (Miturgidae, Philodromidae), Auplopus architectus (Miturgidae), A. bellus (Anyphaenidae), A. mellipes (Anyphaenidae, Miturgidae), Ageniella coronata (Lycosidae), A. euphorbiae (Hahniidae, Anyphaenidae), A. salti (Anyphaenidae), Sericopompilus neotropicalis (Theridiidae), Tachypompilus unicolor (Sparassidae), Anoplius insolens (Thomisidae), A. imbellis (Pisauridae, Amaurobiidae), Ammosphex angularis (?Anyphaenidae, Clubionidae, Corinnidae, Philodromidae), A. anomalus (Lycosidae, Gnaphosidae), A. solonus (Gnaphosidae), Arachnospila arctus (Salticidae), A. fumipennis (?Philodromidae), and Aporinellus medianus (Miturgidae). The families Hahniidae and Sparassidae are reported as prey of North American Pompilidae for the first time. Numerous new host spider genera and species are recorded for many of the pompilid species. Introduction Pompilidae are long-legged, wing-flicking, highly active wasps that hunt spiders on the ground or on vegetation. The female spider wasp searches for and locates a suitable spider host, pursues it, rapidly stings it one or more times in the underside of its cephalothorax, usually transports and places the paralyzed 1PO Box 15251, Syracuse, NY 13215. 2Curator, Arachnida and Myriapoda, Florida State Collection of Arthropods, FDACS, Division of Plant Industry, Gainesville, FL 32614- 7100. *Corresponding author - fkurczewski@twcny.rr.com. 2 Southeastern Naturalist Vol. 11, Monograph No. 4 prey in a simple cell, lays an egg on its abdomen, and closes the nest. The larval spider wasp feeds and develops on the paralyzed spider over several days, spins a cocoon, eventually pupates, and emerges in several weeks or the following year. There is a positive correlation between pompilid size and host-spider size. Pompilids invariably provision their nests with a single relatively medium-sized or large spider. The size of the resultant adult wasp often depends on the amount of food stocked for its development. A small prey simply would not provide sufficient food for the developing wasp larva. Female spiders are the preferred host sex based on their larger biomass (especially the size of the abdomen), more sedentary ways, and seasonal availability (Kurczewski and Kurczewski 1968a, 1968b, 1972, 1973). Many species in the tribe Pepsini build multicellular nests underground or in series in plants or aboveground cavities. Many species in the tribe Ageniellini construct mud cells above ground and cut off the spider’s legs before placing it in a cell. Species of Pepsini and Ageniellini often prepare the depository before capturing the spider. Most species of Pompilini capture the spider before excavating a single-celled nest in the ground. The vast majority of North American Pompilini (85%) have a rake or comb on the foretarsus for excavating a burrow in friable soil (Evans 1950, 1951a, 1951b). Evans and Yoshimoto (1962) presented the pioneer paper on the ecology and nesting behavior of the North American Pompilidae. A number of papers have trickled out since then to enlarge this body of knowledge (Kurczewski 2010). Although nearly 2500 host records are available for the approximately 300 species of North American Pompilidae (Krombein 1979, Kurczewski et al. 1987, Powell 1985, Wasbauer 1982, Wilson and Pitts 2007), most species remain incompletely studied. Many pompilids are more polyphagous than once believed, especially those living in woodlands with two or more generations per year (Kurczewski 1999), and any recent host record has the potential to be new. Host type constitutes the most voluminous kind of biological information available for spider wasps, probably because the paralyzed spider is rather easily collected from the provisioning wasp. Methods This study presents a total of 339 host records for 65 North American pompilid taxa. The spider wasps with host spiders were hand-netted by the authors (F.E. Kurczewski [FEK], California, New York; G.B. Edwards [GBE], Florida), Lyle J. and Eileen Buss (LJB, Florida), Nancy B. Elliott (NBE; Bahamas, Idaho), and Keith E. Kurczewski (KEK, California). James R. Wiley located additional pompilids with prey in the Florida State Collection of Arthropods. Dick Walton provided his field observations on Anoplius semicinctus (Dahlbom) and A. tenebrosus (Cresson). Nick Fensler and J.C. Cokendolpher sent us previously unpublished prey records for some pompilid species. Julio A. Genaro sent us his notes on some Caribbean Pompilidae. Carol Davis sent us a seven-celled mud nest of Auplopus architectus metallicus (Banks) from Utah. Spider wasps and their prey spiders were identified from color photographs taken by Alice J. Abela 2012 F.E. Kurczewski and G.B. Edwards 3 (California), Ken H. Allen (Florida), Lyn S. Atherton (LSA, Florida), Sasha Azevedo (South Carolina), James W. Beck (JWB, Louisiana), Rollin E. Coville (California, Nevada), Linda F. Cooper (LFC, Texas), Carol Davis (Utah), Noah Elhardt (Florida), Wayne Hughes (Georgia), Lynette Schimming (Washington), Marie L. Schmidt (Pennsylvania), Edward Trammel (Arkansas), Troy R. Zurovec (TRZ, Texas), and various other contributors to open online sites. The excellent photographs and detailed observations of these persons were instrumental in allowing us to identify the spider wasps and their host spiders. The spider wasps were identified by F.E. Kurczewski or James P. Pitts (California specimens plus opinions on other species) and arranged taxonomically according to the Catalog of Hymenoptera in America North of Mexico (Krombein 1979). The spiders were identified by: G.B. Edwards; Rick West (Theraphosidae); David Bixler, Allen Brady, Tom Prentice, Rick Vetter, and Margaret Hodge (Lycosidae); Kelly Kissane (Pisauridae); Darrell Ubick (Gnaphosidae); and Wayne Maddison and Marshall Hedin (Salticidae); with nomenclature following Platnick (2012). Body lengths of the wasps and spiders, if known, are given along with species identifications. Many wasps and spiders were measured in the field before they were pinned or preserved in 70% alcohol, respectively. Many observations and collections were made on Onondaga Hill, 3.2 km SW of Syracuse, Onondaga County, NY (NY). California observations and collections were made at Montana de Oro State Park, San Luis Obispo County (CA), unless indicated otherwise. Date and time (EDT, CDT, CST, PDT, PST) of observation or collection, air temperature (at), non-sand ground surface temperature (gt) or sand surface temperature (st) in degrees centigrade, and behavioral descriptions are given. Wasp specimens are deposited at Cornell University, Florida State Collection of Arthropods, and Utah State University. Spider specimens are deposited at Florida State Collection of Arthropods and State University of New York College of Environmental Science and Forestry. In this study, we used for the first time online media images and videos as an aid in spider wasp and spider identification and description of associated predatory behavior. Such an innovative investigative tool enhanced the scope and quality of this paper and broadened the known host-spider species spectrum for some pompilids. Descriptions of hunting, prey transport, and nest construction in certain spider wasps are included to extend variation in species behavorial components. Nonetheless, despite this and other recent studies, there is still much to learn about host selection and nesting behavior in the North American Pompilidae. Results FAMILY POMPILIDAE Subfamily PEPSINAE Tribe Pepsini Pepsis sp. TX: Hidalgo County, 7.2 km SW Mission, National Butterfly Center; 18 November 2011; TRZ; 1700; at = 27–29°. Host: Aphonopelma anax (Chamberlin) (Theraphosidae), subadult. The wasp dragged the spider backwards on the ground, grasping it by the base of its right forecoxa with her mandibles (Fig. 1). 4 Southeastern Naturalist Vol. 11, Monograph No. 4 She transported it a distance of 9.1 m for nearly 10 minutes, eventually entering a ravine and disappearing in the ground beneath the base of a fallen tree (T.R. Zurovec, San Benito, TX, 2011 pers. comm.). Entypus fulvicornis (Cresson) NJ: Cape May County, Cape May Point, Delaware Bayshore; September 2007; R.K. Walton. Host: Hogna sp., adult or subadult female (Walton 2007). SC: Horry County, Myrtle Beach; 2 August 2007; T. Fish. Host: Schizocosa avida (Walckenaer) (Lycosidae), adult male. This wasp walked backwards across a lawn, grasping the spider by the base of its left foreleg or pedipalp with her mandibles (Tracyyxx 2007). Entypus unifasciatus californicus (Townes) CA: Alameda County, Fremont, Lake Elizabeth; 1 July 2009; A. Cavette. Host: Schizocosa mccooki Montgomery or S. maxima Dondale & Redner, adult or subadult female. This wasp dragged her spider backwards, grasping it by the right pedipalp or base of foreleg with her mandibles, fought off three conspecific females, and finally pulled her prey down an open hole (Cavette 2009). CA: Solano County, Suisun/Fairfield; 21 June 2010; El Cobrador. Host: Schizocosa mccooki or S. maxima, adult or subadult female. This wasp captured the spider on the ground, stinging and paralyzing it. She then dragged it backwards to her previously prepared nest, grasping a pedipalp with her mandibles (El Cobrador 2010). Entypus unifasciatus unifasciatus (Say) AR: Sharp County, Evening Shade; 8 July 2010; Dawn419. Host: Dolomedes albineus Hentz (Pisauridae), adult or subadult female. The wasp dragged the spider backwards over rocks, grasping a pedipalp with her mandibles, and eventually pulled it into a hole among the rocks (Dawn419 2010). Figure 1. Pepsis sp. female dragging Aphonopelma anax subadult backwards on ground, grasping the base of its right forecoxa with her mandibles (T.R. Zurovec, pers. comm.). 2012 F.E. Kurczewski and G.B. Edwards 5 IL: Wayne County, Fairfield; 28 July 2007; S. Scott. Host: Rabidosa rabida (Walckenaer) (Lycosidae), adult female. This wasp dragged her spider backwards, grasping a pedipalp with the mandibles. She attacked and drove off a Tachypompilus f. ferrugineus female attempting to steal the spider (Scott 2007). KS: Sedgwick County, Wichita; 3 September 2009; J. Frisch. Host: Rabidosa rabida. The wasp stung and paralyzed the spider and then flew away (Frisch 2009). KS: Wyandotte County, Kansas City; ?2004; D. Murrow. Host: Hogna sp. (Lycosidae), adult or subadult female. This wasp grasped the spider by its left pedipalp with her mandibles and dragged it backwards across a backyard (Murrow 2006). LA: Lafayette Parish, Lafayette, along Bayou Vermilion; 7 August 2008; JWB. Host: Dolomedes albineus, adult female. The interaction between this spider wasp and its prey spider occurred in hardwoods-bottomland. The spider fell from a large, mature Liquidambar styraciflua L. (Hamamelidaceae) (Sweet Gum) in a semi-conscious state suspended by a silken thread, having apparently already been stung by the wasp. The E. unifasciatus female flew in and immediately stung the spider again in mid-air (Fig. 2). The immobile spider fell to the ground, a distance of 3.65 m, followed by the wasp, which examined it with her antennae. The wasp began dragging the spider backwards, dorsal side upward, grasping it Figure 2. Entypus u. unifasciatus female stinging Dolomedes albineus adult female in its cephalothorax in mid-air as it dangles from a silken thread emitted from its spinnerets (Beck 2009). 6 Southeastern Naturalist Vol. 11, Monograph No. 4 by a pedipalp with her mandibles (Fig. 3). She paused to examine it again with her antennae and mouthparts (Fig. 4). The wasp resumed pulling the spider backwards, dorsal side upward, grasping it by a pedipalp with her mandibles (Beck 2009; J.W. Beck, Broussard, LA, 2011 pers. comm.). MO: Greene County, Springfield; 27 August 2008; K. Firth. Host: Rabidosa rabida, adult female (Firth 2008). NY: 25 July 2010; FEK; at = 27°. Host: Rabidosa rabida, adult female, 20 mm (wasp, 21 mm). The spider was dragged backwards through an overgrown field by a pedipalp or base of foreleg being grasped with the wasp’s mandibles. OH: Gallia County; 9 August 2009; J. McCormac. Host: Hogna sp. This wasp captured her spider in leaf litter by stinging it. She then dragged it backwards through open woodland, over branches, twigs, and leaves, grasping it by a pedipalp with her mandibles. She periodically released the spider on the ground to reconnoiter ahead. Her 40-m-long transport of the prey took 15 minutes. She arrived at a small hole amidst entangled rootlets at the base of a sloping hill and immediately pulled the spider inside, probably by its spinnerets with her mandibles (McCormac 2009). OH: Montgomery County, Vandalia; 6 August 2011; S. Lessig. Host: ?Hogna sp., subadult female or juvenile. The wasp examined the paralyzed spider with her antennae among thick weeds and grasses (Lessig 2011). Figure 3. Entypus u. unifasciatus female dragging Dolomedes albineus adult female backwards on ground, dorsal side upward, grasping its right pedipalp with her mandibles (Beck 2009). 2012 F.E. Kurczewski and G.B. Edwards 7 OK: Cleveland County, Norman; 20 August 2009; J.A. Grzybowski. Host: Rabidosa sp. The wasp pulled her spider backwards on the ground, grasping its right pedipalp with her mandibles (JAGpix 2009). PA: Adams County; 16 August 2008; B. Moul. Host: Rabidosa rabida, adult or subadult female. The wasp grasped the spider by its left pedipalp or chelicera with her mandibles and dragged it backwards on the ground (Moul 2008). SC: Greenwood County; 5 September 2004; M. Boone. Host: Rabidosa rabida, adult female. This wasp dragged her spider backwards, grasping a pedipalp with her mandibles (Boone 2004). WV: Nicholas County, Kesslers Cross Lanes; 16 August 2009; E.K. Bell. Host: Dolomedes tenebrosus Hentz, adult female. This spider was dragged backwards across a road by its pedipalp with the wasp’s mandibles (E.K. Bell 2009). WV: Upshur County, near Ruraldale; 25 August 1989; S. Creswell. Host: Rabidosa rabida, adult female. This wasp dragged her spider backwards, grasping a pedipalp with her mandibles (Creswell 1989). Priocnemis (Priocnemissus) minorata Banks AR: Newton County, Buffalo National River; 23 April 2007; E. Trammel. Host Pisurina mira (Walckenaer) (Pisauridae), adult or subadult female, 15.0 mm (wasp, 12.0 mm) (Trammel 2007a). Figure 4. Entypus u. unifasciatus female examining Dolomedes albineus adult female with her mouthparts and antennae before attempting further transport (Beck 2009). 8 Southeastern Naturalist Vol. 11, Monograph No. 4 MA: Franklin County, Colrain; 30 April 2004; C. Eiseman. Host: Coras sp. or Wadotes sp. (Amaurobiidae), adult female. The wasp pulled the spider across dried fallen leaves, grasping it with her mandibles by the base of its 3rd or 4th left leg (Eiseman 2004). NY: 17, 18 May 2010, 22, 27 May 2011, 24 May 2012; FEK; at = 21–28°. Hosts: Dysdera crocata C.L. Koch (Dysderidae), adult female, 13.0, 12.0, 12.5 mm; juvenile, 11.0 mm (wasps, 13.5, 12.5, 12.0, 13.0 mm); Trochosa ruricola (De Geer) (Lycosidae) (Rustic Wolf Spider), adult female, 10.0 mm (wasp, 12.0 mm); Trochosa sp., probably ruricola, penultimate female, 9.5, 10.0 mm (wasps, 12.0, 12.5 mm); Pisaurina mira, adult male, 13.0 mm, 70.0 mm leg span (wasp, 12.5 mm); Agelenopsis pennsylvanica (C.L. Koch) (Aglelenidae), adult female, 11.0 mm (wasp, 12.5 mm); Amaurobius ferox (Walckenaer) (Amaurobiidae), adult female, 10.5, 10.0 mm (wasps, 11.5, 12.0 mm). The spiders were dragged backwards on the ground in deciduous woodland, the wasps grasping a hindleg with the mandibles. Priocnemis (Priocnemis) cornica (Say) (all-black color variant) NY: 15–16, 18 June 2010; FEK; at = 25–28°, gt = 43–48°. Hosts: Lycosidae sp., not Hogna, juvenile, 4.5 mm (wasp, 6.0 mm); Hogna sp., juvenile, 5.0 mm (wasp, 6.5 mm); Pardosa milvina (Hentz) (Lycosidae), adult male, 5.5 mm (wasp, 6.5 mm). The wasps dragged their spiders backwards on the ground, grasping a hindleg with the mandibles. They nested in preexisting burrows in sparsely vegetated, gravelly loam. Priocnemis (Priocnemis) germana (Cresson) NY: 23 July 2011; FEK; at = 28°. Host: Cheiracanthium mildei L. Koch (Miturgidae), adult female, 8.5 mm (wasp, 9.0 mm), subadult female, 8.0 mm (wasp, 8.0 mm). The wasps carried the spiders backwards across low vegetation in deciduous woodland, grasping them by a hindleg with their mandibles. They released the spiders on leaves and walked ahead, at which time they were collected. Priocnemis (Priocnemis) scitula relicta Banks NY: 25 July, 11, 22 September 2010, 10–11, 21, 29 July, 13, 17–18 August, 18 September 2011; FEK; at = 25–27°, gt = 38–39°. Hosts: Cheiracanthium ?mildei, juvenile, 6.0, 6.5 mm (wasps, 6.0, 6.5 mm); Philodromus sp. (Philodromidae), adult male, 5.5 mm (wasp, 6.0 mm), with right foreleg missing beyond coxatrochanter joint; Platycryptus undatus (De Geer) (Salticidae), juvenile, 5.5, 6.0, 6.0, 4.0, 5.5 mm (wasps, 6.0, 6.0, 6.5, 5.5, 6.5 mm); Phidippus audax (Hentz) (Salticidae), juvenile, 4.8 mm (wasp, 5.5 mm); Naphrys pulex (Hentz) (Salticidae), adult female, 4.5 mm; adult male, 4.5 mm (wasps, 5.5, 5.5 mm); Sitticus fasciger (Simon) (Salticidae), adult male, 4.5, 4.5 mm; juvenile, 4.0 mm (wasps, 5.5, 5.5, 6.0 mm). Prey transport involved the wasps walking backwards, grasping their spiders by a hindleg with the mandibles. 2012 F.E. Kurczewski and G.B. Edwards 9 Caliadurgus fasciatellus alienatus (Smith) MA: Worcester County, Harvard; 7 August 2004; T. Murray. Host: Neoscona sp. (Araneidae), juvenile. The wasp grasped the base of the spider’s 3rd or 4th right leg and, holding her prey above ground, walked forward (Murray 2010a). NY: 5, 10 July 2011; FEK; at = 27–28°. Hosts: Araneus sp. (Araneidae), juvenile, 4.5 mm (wasp, 7.0 mm); Cyclosa conica (Pallas) (Araneidae), adult female, 5.5 mm (wasp, 7.0 mm); Neoscona sp. (Araneidae), subadult female, 5.0 mm (wasp, 7.0 mm). Dipogon (Deuteragenia) calipterus calipterus (Say) IL: Champaign County, Urbana; 23 July 2011; A. Wild. Host: Trachelas tranquillus (Hentz) (Corinnidae), penultimate male. The wasp dragged the paralyzed spider backwards across a porch, grasping its spinnerets with her mandibles (Wild 2011). Dipogon (Deuteragenia) sayi sayi Banks NY: 19 June 2011; FEK; at = 26–27°. Hosts: Bassaniana versicolor (Keyserling) (Thomisidae), adult female, 6.0 mm (wasp, 6.5 mm); Xysticus sp. (Thomisidae), penultimate female, 5.5 mm (wasp, 6.5 mm). This wasp nested in a preexisting empty bamboo tubule, one of about 75 different-sized bamboo tubules comprising an artificial mason bee house. PA: Northumberland County, Elysburg; 7 July 2010; J.M. Yuschock. Host: Xysticus sp., juvenile. This wasp grasped the spider by its spinnerets with her mandibles and walked backwards (Yuschock 2010). Tribe Ageniellini Phanagenia bombycina (Cresson) AR : Faulkner County, Bell Slough Wildlife Management Area; 9 October 2007; H. Raney. Host: ?Schizocosa sp., adult or penultimate male. The wasp carried the spider face forward and dorsal side upward, straddling it and grasping its right pedipalp with her mandibles. She had amputated all of its legs at the coxa-trochanter joints (Raney 2007). FL: Indian River County, St. Sebastian River Preserve; 14–20 June 2007; J. Hollenbeck. Host: Rabidosa hentzi (Banks), penultimate female, 11 mm long, with 3rd and 4th left and right legs amputated at coxa-trochanter joints. The wasp was reared from a mud cell (Hollenbeck 2007). NY: 10 July 2011; FEK; 1306; at = 28°. Host: Platycryptus undatus, adult female, 13.5 mm (wasp, 13.0 mm), with all legs amputated at coxa-trochanter joints. The wasp transported her spider face foward and dorsal side upward, grasping its right pedipalp with the mandibles. Auplopus architectus architectus (Say) MA: Middlesex County, Carlisle; 17 June 2007; D. Walton. Host : Eris sp. (Salticidae), adult female, with all legs amputated at coxa-trochanter joints. The wasp transported the spider forward, straddling it ventral side upward and 10 Southeastern Naturalist Vol. 11, Monograph No. 4 grasping its spinnerets with her mandibles as well as dorsal side upward and grasping it by a pedipalp (Walton 2007). Auplopus architectus metallicus (Banks) UT: Salt Lake County, Taylorsville; 30 September 2011; C. Davis. Host: Cheiracanthium inclusum (Hentz), adult or penultimate male, with all but left hindleg amputated at coxa-trochanter joints. Four photographs show the wasp amputating the spider’s legs at the coxa-trochanter joints, using her mandibles. A fifth photograph shows the wasp transporting the spider on the ground. She grasped the spider by its spinnerets with her mandibles, ventral side upward, and walked forward, eventually disappearing from view (Davis 2011). Two other photographs (dorsal, ventral views) depict a group of seven mud cells from a prior year affixed together beneath a bottom screen-door hinge (Fig. 5). Each cell has an emergence hole the diameter of the wasp’s head (2.5– 3.0 mm). The composite group of cells is 32 mm long, 18 mm wide, and 9 mm high. The individual cells average 5.9 (5.0–6.5) mm wide and 11.8 (11.0–12.5) mm long (C. Davis, Taylorsville, UT, 2012 pers. comm.). Auplopus bellus (Cresson) BAHAMAS: San Salvador Island, Dump; 31 January 1982; NBE. Host: Hibana sp. (Anyphaenidae), juvenile, 6.5 mm (wasp, 6.5 mm), with all legs amputated at coxa-trochanter joints. The wasp carried the spider forward, straddling it ventral side upward and grasping its spinnerets with her mandibles (N.B. Elliott, Siena College, Loudonville, NY, 2010 pers. comm). Auplopus caerulescens subcorticalis (Walsh) NY: 8 July 2011; FEK; 1336; at = 28°. Host: Sitticus fasciger, adult male, 4.5 mm (wasp, 6.5 mm), with all legs amputated at coxa-trochanter joints. The wasp transported the spider forward, straddling it ventral side upward and grasping its spinnerets with her mandibles. Auplopus carbonarius (Scopoli) MI: Kalamazoo County, Kalamazoo; June–August 1999; M.C. Cassino. Host: Cheiracanthium ?mildei, adult female. The wasp straddled the spider ventral side upward, grasping its spinnerets with her mandibles. She carried it across a concrete driveway and disappeared inside a large planter beside a house (Cassino 1999; M.C. Cassino, Kalamazoo, MI, 2012 pers. comm.). ONTARIO: Ottawa; 12 August 2007; S. Kassam. Host: Cheiracanthium mildei, adult female, with only the hindlegs cut off at the coxa-trochanter joints. This wasp recently captured the spider and landed on the wood siding of a house, straddling it ventral side upward and grasping its spinnerets with her mandibles (Kassam 2007). WA: Pierce County, Puyallup, 1805 35th Street PL SE; 21 July 2012; L. Schimming. Host: Eris militaris (Hentz) (Salticidae), penultimate female. The wasp was photographed on the side of a plastic tool cabinet, 1.2 m above ground 2012 F.E. Kurczewski and G.B. Edwards 11 level, in the process of amputating the spider’s legs with her mandibles at the coxa-trochanter joints (Figs. 6, 7). After amputating all of the spider’s legs, she grasped her prey ventral side upward by its spinnerets with her mandibles and took flight (L. Schimming, Puyallup, WA, 2012 pers. comm.). Auplopus mellipes mellipes (Say) OK: Creek County; 3 July 2007; J.J. Cross. Host: Paraphidippus aurantius (Lucas) (Salticidae), adult or subadult female with all legs amputated at Figure 5. Seven mud cells of Auplopus architectus metallicus from a prior year’s nesting season affixed together from beneath a bottom screen door hinge. The spider wasp emergence holes are visible on many of the cells (C. Davis, pers. comm.). A. Dorsal view of nest; B. ventral view. 12 Southeastern Naturalist Vol. 11, Monograph No. 4 Figure 6. Auplopus carbonarius female using her mandibles to amputate the 3rd and 4th left legs of Eris militaris adult female. She eventually amputated all eight legs of her prey spider (L. Schimming, pers. comm.). Figure 7. Auplopus carbonarius female grasping an Eris militaris adult female by its spinnerets with her mandibles before flying to her previously prepared mud cell (L. Schimming 2012). 2012 F.E. Kurczewski and G.B. Edwards 13 coxa-trochanter joints. The wasp grasped the spider by its spinnerets with her mandibles, straddled it ventral side upward, and proceeded forward (Cross 2007). VA: Fairfax County, McLean, Turkey Run Park; 23 June 2009; T. Bentley. Host: Hibana ?gracilis (Hentz), adult female. This wasp was photographed in the process of amputating the spider’s leg at a coxa-trochanter joint. It appears that all but three of the legs had been cut off (Bentley 2009). VA: Independent City of Virginia Beach; 21 May 2010; no collector. Host: Hibana ?gracilis, adult female. The wasp amputated all of the spider’s legs at the coxa-trochanter joints and walked forward, straddling it ventral side upward and grasping its spinnerets with her mandibles (w00dy4012 2010). Auplopus mellipes variitarsatus (Dalla Torre) NY: 30 June–5 July 2009, 15 July 2010; FEK; at = 21–27°. Host: Platycryptus undatus, adult female, 10, 10 mm; adult male, 8.5, 8.5 mm; juvenile, 8.5, 9.5 mm (wasp, 9.5 mm); all spiders’ legs amputated at coxa-trochanter joints. One female constructed five mud cells over a six-day period applying the mud in single threads using her mouthparts and trowel-shaped abdominal apex. The cells were affixed vertically in single file to the edge of a closed basement window (Fig. 8). Wasp entry was through a narrow space where the window closed against its casing. The cells were 7–8 mm wide and 12–13 mm long on the exterior and 5–6 mm wide and 10–11 mm long on the interior. Each cell had a flattened end facing inward and a convex end facing outward (Fig. 8). One female, 9.5 mm long, emerged from the middle cell on 15 July 2010 at 1610. The contents of the other four cells never emerged, each cell containing a desiccated cocoon with dried wasp fragments. NY: 19–25, 28 July, 17, 18 August, 2 September 2011; FEK; 1203–1451; at = 22–30°. Hosts: Cheiracanthium mildei, adult female (gravid), 9.5 mm (wasp, 9.5 mm), with all but left pedipalp and right foreleg amputated at coxa-trochanter joints (collected while attempting to cut off remaining leg), 9.0 mm (wasp, 9.0 mm), with all legs amputated at coxa-trochanter joints; Platycryptus undatus, adult male, 9.5 mm; juvenile, 8.0, 9.0, 8.0, 9.0, 8.0, 9.5 mm (wasp, 9.0 mm); all spiders’ legs amputated at coxa-trochanter joints. One wasp constructed seven mud cells over an eight-day period. The cells were affixed longitudinally in single file to the inside of a horizontally positioned, 10-mm-wide, 14-cm-long bamboo tubule, part of an artificial mason bee house. The cells were 7–8 mm wide and 11–12 mm long on the outside. Another female constructed her mud cells beneath the back deck on a house, 26 cm above ground level. She flew in every 1–2 minutes holding a mud pellet with her mouthparts, disappeared between two horizontal planks (floor boards), exited in 60–90 seconds without the mud pellet, and flew straight away in the direction from which she came. She entered with prey through a 5-mm space between two planks of the deck, straddling the spider ventral side upward and grasping its spinnerets with her mandibles. Females were observed hunting for P. undatus 5 July 2009 (cloudy, at = 21– 24°), 20 July 2010 (sunny, at = 29°), and 8 August 2010 (overcast, at = 27–28°) on gray cedar siding of a house and gray, unstained cedar post-and-rail fence. 14 Southeastern Naturalist Vol. 11, Monograph No. 4 They ran across the wood flicking their wings and tapped the surface with their antennae, interspersed with short flights. The gray body color of P. undatus may have served as camouflage to avian predators but not to the hunt ing wasps. PA: Delaware County, Norwood; 25 May 2011; M.L. Schmidt. Host: Hibana ?gracilis, adult female (gravid), 10.3 mm (wasp, 9.1 mm), with all legs amputated Figure 8. Five fully provisioned mud cells of Auplopus mellipes variitarsatus affixed vertically to edge of basement window. Flattened ends of cells face inward and convex ends face outward (J.W. Stoll, Syracuse University, Syracuse, NY, 2010 pers. comm.). 2012 F.E. Kurczewski and G.B. Edwards 15 at coxa-trochanter joints. The wasp carried the spider forward, straddling it ventral side upward and grasping its spinnerets with her mandibles (Fig. 9, Schmidt 2011; M.L. Schmidt, Norwood, PA, 2011 pers. comm.). VA: Accomack County, Belle Haven; 1 June 2008; A. Bradford. Host: Anyphaena sp. (Anyphaenidae), adult female, 5.5 mm (wasp, 8.8 mm), with all but two or three front legs missing. This wasp transported the spider forward, straddling it ventral side upward and grasping its spinnerets with her mandibles. She flew from leaf to leaf, then to the ground, ran across a patio and garden, up an oak tree, then to the garden and disappeared (zxgirl 2008). Auplopus nigrellus (Banks) TX: Lubbock County; Lubbock, 21 July 1991; J.C. Cokendolpher. Host: Hibana incursa (Chamberlin), adult male, adult female, both sexes with all legs amputated at the coxa-trochanter joints (J.C. Cokendolpher, Lubbock, TX, 2012 pers. comm.). Ageniella (Priophanes) arcuata (Banks) AR: Washington County, Fayetteville; 30 July 2006; E. Trammel. Host: Oxyopes salticus Hentz (Oxyopidae), adult female, with all legs amputated at coxa-trochanter joints. The wasp walked forward with the spider on grass blades, straddling it ventral side upward and grasping its spinnerets with her Figure 9. Auplopus mellipes variitarsatus female transporting Hibana ?gracilis adult female, with all legs amputated at coxa-trochanter joints, forward in a ventral side upward position, grasping its spinnerets with her mandibles (Schmidt 2011). 16 Southeastern Naturalist Vol. 11, Monograph No. 4 mandibles (Fig. 10). She flew from this position to the ground, after which the pair disappeared underground (Diogmites 2009a; E. Trammel, Fayetteville, AR, 2011 pers. comm.). Ageniella (Ageniella) accepta (Cresson)* TX: Williamson County, Balcones Canyonlands National Wildlife Refuge, Gainer Tract; 2 September 2010; P. Lenhart. Host: Rabidosa punctulata (Hentz), juvenile, with all legs amputated at coxa-trochanter joints. This wasp transported her spider forward on the ground, dorsal side upward and face forward, grasping its chelicera or pedipalp with her mandibles (Lenhart 2010). *See Waichert et al. (2011). Ageniella (Ageniella) coronata Banks CA: Los Angeles County, Los Angeles, Placerita Canyon State Park; 26 June 2010; G. Park. Host: Lycosidae sp., penultimate male. This wasp carried the spider face forward and dorsal side upward after cutting off its legs at the coxatrochanter joints with her mandibles. She grasped the spider with her mandibles by its left chelicera or pedipalp, straddled it, and ran forward using all legs or attempted to fly with frayed wings (Park 2010). Ageniella (Ageniella) euphorbiae (Viereck) CA: 29 August, 25 September 2010; KEK; at = 27–28°, st = 50–53°. Hosts: Hibana ?incursa, juvenile, 6.0 mm (wasp, 7.5 mm); Calymmaria lora (Chamberlin & Ivie) (Hahniidae), adult male, 6.5 mm (wasp, 8.2 mm). One wasp walked Figure 10. Ageniella arcuata female transporting Oxyopes salticus adult female, with all legs cut off at coxa-trochanter joints, forward on grass blade. The wasp is grasping the spider’s spinnerets with her mandibles and holding it in a ventral side upward position (Diogmites 2009a). 2012 F.E. Kurczewski and G.B. Edwards 17 forward on perennial veldt grass (Ehrharta calycina J.E. Smith) (Poaceae), 34 cm above ground, holding her spider underneath, flew down to the sand surface, and released her prey ventral side upward. She entered her burrow, 30 cm away, and was collected as she returned to retrieve her prey (K.E. Kurczewski, Los Osos, CA, 2010 pers. comm.). Ageniella (Ameragenia) salti (Banks) FL: Orange County, Orlando; 14 February 2011; J. Reben. Host: Hibana velox (Becker), adult female. The wasp amputated all of the spider’s legs at the coxatrochanter joints. She grasped the spider by its spinnerets with her mandibles, straddled it ventral side upward, and walked forward (Reben 2011). Subfamily POMPILINAE Tribe Aporini Aporus (Plectraporus) hirsutus (Banks) CA: San Luis Obispo County, Baywood-Los Osos, Sweet Springs Nature Preserve; 6 April 2011; FEK; 1448–1510; at = 15°. This female, 8.1 mm, walked slowly on damp sand, flicked her wings, and searched among dried leaves and plant litter with her antennae. CA: San Luis Obispo County, Los Osos, El Moro Elfin Forest; 13 April 2011, 17 April 2012; FEK; 1050–1145; at = 14–16°. Two wasps, 9.5 and 11.7 mm, stayed entirely on the ground, flicking their wings and searching with their antennae in dried plant litter beneath shrubs in the shade. They made no attempt to fly under cool, moderately windy (13–24 mph) conditions. CA: San Luis Obispo County, Los Osos; 18 September 2011; KEK. Host: Aptostichus atomarius Simon (Cyrtaucheniidae), adult male, 14.5 mm (wasp, 11.5 mm) (K.E. Kurczewski, 2011 pers. comm.). Tribe Pompilini Evagetes padrinus padrinus (Viereck) CA: 3–4, 9, 11, 14, 25 April 2011, 28 May 2012; KEK. Six females (6.1, 4.5, 6.2, 4.7, 5.9, 7.2 mm) were observed and collected in an area where several females of Episyron quinquenotatus hurdi Evans were nesting gregariously. In addition to entering burrows of E. quinquenotatus hurdi, females of this species were seen grappling with and entering burrows of Ammosphex a. angularis and Aporinellus t. taeniatus (K.E. Kurczewski, 2011–2012 pers. comm.). Agenioideus (Agenioideus) humilis (Cresson) MI: Calhoun County, Springfield; 28 July 2009; S. Warner. Host: Araneus sp., subadult female, 5.6 mm (wasp, 7.7 mm). This wasp dragged the spider backwards, grasping it with her mandibles (Warner 2009). QUEBEC: Brossard; 20 July 2010; A. Bateman. Host: Araneus ?diadematus Clerck, subadult female, 7.2 mm (wasp, 8.0 mm). The wasp pulled the spider up a wall, grasping the base of its 2nd left leg with her mandibles (Bateman 2010). 18 Southeastern Naturalist Vol. 11, Monograph No. 4 Sericopompilus apicalis (Say) CA: 9 May 2011; KEK. Hosts: Phidippus sp. probably johnsoni Peckham and Peckham (Salticidae), penultimate female, 10.5 mm (wasp, 12.5 mm); Phidippus sp., juvenile, 10.0 (wasp, 12.0 mm) (K.E. Kurczewski, 2011 pers. comm.). Sericopompilus neotropicalis (Cameron) FL: Lee County, North Fort Myers; 22 June 2010; N. Elhardt. Host: Latrodectus mactans (Fabricius) (Theridiidae), adult female. This wasp carried her spider from place to place, grasping it by a pedipalp, chelicera, or base of foreleg with her mandibles, as she attempted to escape an aggregation of small reddish-yellow ants or re-excavated a previously existing burrow with her forelegs in a patch of bare sandy soil. The spider dangled downward as it was straddled and carried forward on vegetation, being held by a chelicera or pedipalp with the wasp’s mandibles, or was transported backward on the ground (Fig. 11; Elhardt 2010; N. Elhardt, South San Francisco, CA, 2011 pers. comm.). Episyron biguttatus biguttatus (Fabricius) AR: Washington County, Fayetteville; 25 August 2006; E. Trammel. Host: Neoscona crucifera (Lucas), subadult female (Diogmites 2009b) . Figure 11. Sericopompilus neotropicalis female transporting Latrodectus mactans adult female forward on vegetation, grasping its pedipalp with her mandibles (Elhardt 2010). 2012 F.E. Kurczewski and G.B. Edwards 19 CN: New Haven County, Southbury; 23 September 2010; T.S. Crider. Host: Araneus marmoreus Clerck, adult or subadult female. The wasp grasped the spider by the base of its 3rd right leg with her mandibles and dragged it backwards (Crider 2010). MA: Barnstable County, Hyannis; no date or collector. Host: Araneus bicentarius McCook, adult female (Bugman 2011a). MA: Middlesex County, Groton; 8 July 2008; T. Murray. Host: Araneus sp., subadult female. This wasp grasped the base of the spider’s right hindleg with her mandibles and pulled it backwards on the ground (Murray 2010b). ME: Hancock County, Acadia National Park; 23 July 2005; P. Moniotte. Host: Araneus nordmanni (Thorell), ?adult female (Moniotte 2005). NY: Ulster County, New Paltz; 20 August 2011; K. Andreas. Host: Araneus diadematus, ?subadult female. The wasp grasped the spider by the base of its right foreleg and pulled it backwards (Futureman1 2011). Figure 12. Episyron b. biguttatus female pulling Eriophora ravilla adult female up pine tree trunk, grasping the base of its 2nd left leg with her mandibles (Azevedo 2009). 20 Southeastern Naturalist Vol. 11, Monograph No. 4 SC: Charleston County, Charleston; 26 August 2009; S. Azevedo. Host: Eriophora ravilla (C.L. Koch) (Araneidae), adult female. The wasp pulled the spider backwards up a pine tree trunk, grasping it with her mandibles by the base of its 2nd left leg (Fig. 12, Azevedo 2009; S. Azevedo, Charleston, SC, 2011 pers. comm.). VT: Orange County, Topsham; 8 July 2008; T. Murray. Host: Araneus nordmanni, adult or subadult female. The wasp grasped the spider by the base of its 3rd right leg with her mandibles and dragged it backwards (Murray 2008). ONTARIO: Presqu’ile Provincial Park; 6 August 2007; J. Illingworth. Host: Neoscona sp., penultimate female. The wasp pulled the spider up tall grasses near the Lake Ontario shore, grasping the base of its right foreleg with her mandibles. Presumably, she would have cached her prey in the grasses prior to excavating her burrow (Jamuudsen 2007). Episyron conterminus cressoni (Dewitz) FL: Highlands County, Highlands Hammock State Park; 14 April 2010; LSA. Host: Gea heptagon (Hentz) (Araneidae), subadult female. The wasp inspected the spider with her antennae before pulling it into her burrow by its spinnerets with her mandibles (L.S. Atherton, Tierra Verde, FL, 2011 pers. comm.). FL: Okeechobee County, Kissimmee Prairie Preserve State Park; 15 March 2011; LSA. Host: Eustala anastera (Walckenaer) (Araneidae), subadult female. The wasp cached her spider on vegetation above ground level while she excavated her burrow. She then carried the prey to the ground, grasping it with her mandibles, and inspected it before pulling it into her burrow by its spinnerets (Fig. 13; L.S. Atherton, 2011 pers. comm.). Figure 13. Episyron conterminus cressoni female pulling a Eustala anastera subadult female backwards across a dried leaf, grasping the base of its 2nd right leg with her mandibles (L.S. Atherton, pers. comm.). 2012 F.E. Kurczewski and G.B. Edwards 21 Episyron oregon Evans CA: 20 June, 8 August 2011; KEK; 1431–1529; at = 17–23°, st = 30–57°. Host: Araneus montereyensis (Archer), adult female, 5.0, 5.0, 5.5 mm (wasps, 8.0, 7.5, 9.0 mm). One spider not cached on a plant but left on the ground was pilfered by scavenging ants which dismembered and carried off the pedipalps, legs, and carapace. One wasp interrupted burrow excavation to inspect her spider four times, each time returning to her entrance to enlarge the opening. Another female, after excavating her burrow, retrieved her spider from a plant and dragged it backwards on the sand, ventral side upward, grasping it by a leg with her mandibles. During transport she released the spider temporarily on the sand several times, walked to and examined her entrance, and eventually pulled it into her burrow by its spinnerets with her mandibles. A Hedychridium coruscum Bohart (Chrysididae) female entered one E. oregon burrow twice while the female was away examining her cached spider (K.E. Kurczewski, 2011 pers. comm.). Episyron quinquenotatus hurdi Evans CA: 2, 9, 22 May 2011; KEK; 1032–1730; at = 16–29°, st = 32–60°. Host: Araneus montereyensis, adult female, 5.0, 5.0, 5.5, 5.5, 5.0 mm (wasps, 8.5, 7.5, 8.5, 9.0, 8.0 mm). All wasp burrows were located in bare sand, away from vegetation. In each case, the female cached her paralyzed spider on a plant above ground some distance from her burrow. Four females were observed nesting 17 October 2011 (1135–1245; at = 20–23°, st = 40–44°) (K.E. Kurczewski, 2011 pers. comm.). Figure 14. Episyron quinquenotatus hurdi female dragging paralyzed Araneus montereyensis backwards across sand, alternately grasping the spider by the base(s) of its 2nd, 3rd, and/or 4th legs with her mandibles (A.J. Abela, pers. comm.). 22 Southeastern Naturalist Vol. 11, Monograph No. 4 CA: San Luis Obispo County, Los Osos, Middle School Trail; 18 September 2011; KEK; 1442–1506; at = 18°, st = 56°. Host: Larinioides cornutus (Clerck) or L. patagiatus (Clerck) (Araneidae), penultimate male, 9.0 mm (wasp, 11.0 mm, no wing fray). This wasp excavated her burrow using her forelegs alternately. She interrupted burrow excavation four times to examine her prey cached 1.1 m away on Eriogonum fasciculatum Bentham (Polygonaceae) (California Buckwheat), and twice to obtain nectar from an adjacent plant of the same species. The wasp eventually went to the cachment site, grasped the spider by the base of its 2nd left leg, and flew to the ground. She then dragged the prey backwards across the sand toward her open burrow, at first grasping it with her mandibles by the base of a 2nd or 3rd leg but, later, by the base of a hindleg. Another female nested at this site on 10 October 2011 (KEK, 1230, at = 19°) (K.E. Kurczewski, 2011 pers. comm.). CA: Santa Barbara County, Vandenberg Air Force Base, 26 July 2012; A.J. Abela. Host: Araneus montereyensis, adult female. This wasp was photographed dragging her paralyzed spider backwards across the sand surface, grasping it alternately by the base(s) of its 2nd, 3rd and/or 4th legs (Fig. 14). The wasp eventually cached her prey atop a raised sand ledge and began excavating her burrow (Abela 2012; A. J. Abela, Santa Maria, CA, 2012 pers. comm.). Episyron quinquenotatus quinquenotatus (Say) ONTARIO: Essex County; 10 July 2006; no collector. Host: Larinioides cornutus, adult female, 10.0 mm long (wasp, 12.0 mm). The wasp dragged the spider to a bare sandy area near her nest entrance, grasping a pedipalp with the mandibles. She then pulled it inside the burrow by its spinnerets (Jackie 2006). Episyron snowi (Viereck) UT: Cache County, Wasatch/Cache National Forest; 24 August 2008; Algidus. Host: Araneus sp. or Neoscona sp., adult female. This wasp dragged her spider backwards, grasping the base of its 2nd right leg with her mandibles (Algidus 2008). Poecilopompilus algidus algidus (Smith) AR: Johnson County, Hurricane Creek Wilderness Area; 4 August 2007; E. Trammel. Host: Neoscona crucifera, subadult female (Trammel 2007b). AR: Pulaski County, Jacksonville; no date; A.D. Tetkoskie. Host: Argiope aurantia Lucas (Araneidae), adult female. The wasp grasped the spider by the base of its left hindleg and pulled it backwards on the ground (Bugman 2011b). FL: Palm Beach County, Belle Glade, Everglades Experiment Station; 7 October 1971; C.E. Seiler. Host: Eriophora ravilla, adult female, 15.5 mm (wasp 19.0 mm) (J.R. Wiley, Florida State Collection of Arthropods, Gainesville, FL, 2011 pers. comm.). FL: Pinellas County, Brooker Creek Preserve near Education Center; 16 September 2008; M. Deep. Host: Nephila clavipes (L.) (Nephilidae), adult female (small). The wasp chased the spider from its orb web onto the ground, stung it, and, after examining it, dragged it off by grasping a front right leg 2012 F.E. Kurczewski and G.B. Edwards 23 with her mandibles. During transport, the prey was stolen by an anole nearby (Deep 2008). FL: Pinellas County, Fort De Soto Park; 24 September 2008; LSA. Host: Nephila clavipes, adult female (small). The wasp dragged the spider backwards through vegetation, holding it with her mandibles by the base of a leg (Fig. 15; L.S. Atherton, 2011 pers. comm.). TX: Fayette County, La Grange; 6 September 2005; C. Vannoy. Host: Argiope aurantia, adult female. The wasp grasped the spider by the base of its left hindleg with her mandibles and pulled it backwards on the ground (danielj 2005b). Poecilopompilus algidus coquilletti (Provancher) CO: Boulder County, Boulder; 21 August 2011; S.E. Severs. Host: Argiope trifasciata (Forskal), adult female. The spider was cached atop a plant stem while the wasp excavated her burrow nearby. During excavation, she periodically paused and raised her wings (Severs 2011). UT: Cache County; 6 August 2006; Algidus. Host: Araneus sp., adult or subadult female (Algidus 2006). Poecilopompilus algidus willistoni (Patton) AZ: Cochise County, Benson, Mescal Road; 7 September 2007; M. Brummermann. Host: Neoscona ?oaxacensis (Keyserling), adult female. The wasp dragged the spider backwards, grasping it by the base of its right hindleg with her mandibles (Brummermann 2007). Figure 15. Poecilopompilus a. algidus female dragging Nephila clavipes adult female backwards on ground, grasping the base of a right leg with her mandibles (L.S. Atherton, pers. comm.). 24 Southeastern Naturalist Vol. 11, Monograph No. 4 Poecilopompilus interruptus interruptus (Say) AR: Faulkner County, Conway; 18 July 2005; H.D. Raney. Host: Neoscona crucifera, subadult female. The wasp pulled the spider backwards up several stems and flower stalks, finally wedging it in the axile of a plant atop a leaf. She returned periodically to examine her cached prey, interspersed with periods of self cleaning. She then proceeded with burrow excavation (Raney 2005). FL: Alachua County, S9 T10S R18E; 17 May 1974; W.H. Pierce. Host: Acanthepeira ?stellata (Walckenaer) (Araneidae), adult female, 8.0 mm (wasp, 14.0 mm) (J.R. Wiley, Florida State Collection of Arthropods, Gainesville, FL, 2011 pers. comm). FL: Okeechobee County, Kissimmee Prairie Preserve State Park; l0 June 2009, 16 June 2011; LSA. Hosts: Eriophora ravilla, subadult female; Neoscona crucifera, subadult female. One wasp with prey was photographed prior to burrow entry (Fig. 16; L.S. Atherton, 2011 pers. comm.). TX: Bexar County, S of San Antonio, Medina River Park; 18 June 2007; L. Elliott. Host: Neoscona crucifera, adult female. The wasp stung and then examined the spider as it laid ventral side upward on the ground (Elliott 2007). Tachypompilus ferrugineus ferrugineus (Say) AR: Faulkner County, Conway; 7 July 2009; H. Raney. Host: Dolomedes albineus, adult female. This wasp dragged her spider backwards, grasping its left pedipalp with her mandibles (Raney 2009). AR: Searcy County, Ozark Mountains; 7 July, 1 October 2008; M. Smith. Hosts: Rabidosa rabida, adult female; unidentified spider. One wasp (7 July) pulled her Figure 16. Poecilopompilus i. interruptus female inspecting Neoscona crucifera subadult female with her mouthparts and antennae prior to dragging it into open burrow (L.S. Atherton, pers. comm.). 2012 F.E. Kurczewski and G.B. Edwards 25 spider backwards through grasses and a garden for over 15 m, grasping its right pedipalp with her mandibles (Smith 2008a). The other wasp (1 October) pulled her prey up a vertical wall, grasping a pedipalp with her mandibles (Smith 2008b). DE: New Castle County, Bear; 14 September 2008; T. Kropiewnicki. Host: Hogna sp., adult or subadult female, 17.5 mm. The wasp, 17.5 mm, dragged the spider backwards across a patio, up concrete steps, across a wall, and into an open space between the wall and a basement window—a distance of 9.1 m that took 15 minutes. She held the spider by its left pedipalp with her mandibles (Kropiewnicki 2008; T. Kropiewnicki, Bear, DE, 2011 pers. comm.). FL: Alachua County, Gainesville, 10014 SW 87th Terrace; LJB. On 30 July 2005, a T. ferrugineus (14.0 mm) dragged a Dolomedes albineus, adult female (19.0 mm), backwards in a yard, grasping its right pedipalp with her mandibles. A D. albineus adult female (22.5 mm) was buried in a children’s sandbox by another T. ferrugineus on 26 September 2009. A 3rd T. ferrugineus (19.0 mm) dragged a D. albineus, adult female (21.0 mm) backwards across an outside house wall on 25 July 2010, grasping its left pedipalp with her mandibles. Ten large adult females of D. albineus and Hogna spp., including lenta (Hentz), with egg or larva affixed on the anterior dorsum of the abdomen, were uncovered in Figure 17. Tachypompilus f. ferrugineus egg (arrow) affixed to left anterior latero-dorsal surface of abdomen of Dolomedes albineus adult female (L.J. Buss, pers. comm.). 26 Southeastern Naturalist Vol. 11, Monograph No. 4 the sandbox on 16 July 2010. This lot included a D. albineus (21.0 mm) with wasp’s egg attached to its abdomen and an H. lenta (17.0 mm) with wasp’s larva attached to its abdomen (Figs. 17, 18). Two T. ferrugineus ?females, 11.0 and 8.5 mm, eclosed from other paralyzed D. albineus in late August 2010. A female and possible parent T. ferrugineus (17.0 mm) was captured in the sandbox at night on 23 July 2010. Numerous spiders, wasp larvae, and wasp pupae in cocoons were sifted and collected from the sandbox on 25–26 July 2010; eight adults of T. ferrugineus (males 7.0, 10.5, 11.0 mm; females 13.0, 13.5, 14.0, 14.5, 16.0 mm) emerged from this material 18 August to 9 September 2010. Another cocoon unearthed from the sandbox 19 October 2010 produced a male of T. ferrugineus (12.0 mm) in January 2011 (L.J. and E. Buss, University of Florida, Gainesville, FL, 2011 pers. comm.). Live-caught T. ferrugineus females from northern Florida were significantly larger (P = 0.003) than hand-reared females from the same area: live-caught, Figure 18. Tachypompilus f. ferrugineus larva feeding at anterior dorsal surface of abdomen of Hogna lenta adult female (L.J. Buss, pers. comm.). 2012 F.E. Kurczewski and G.B. Edwards 27 mean body length = 17.9 mm, range = 14.0–20.0 mm, n = 6; hand-reared, mean body length = 12.9 mm, range = 8.5–16.0 mm, n = 7. The 95% confidence interval about the difference of mean body length in the two samples is (2.13, 7.85). FL: Levy County, Cedar Keys National Wildlife Refuge, Seahorse Key; 3 August (year, collector unknown); 17 September 2010; GBE; 1800. Host: Isohogna timuqua (Wallace), adult female, 20.0, 19.0, 18.5 mm (wasps, 19.0, 17.5, 15.0 mm). Two T. ferrugineus females with prey were observed on a sandy, partly mowed lawn in front of the research facility (lighthouse). One female dragged her spider slowly backwards across the lawn by a ?leg, grasping it with the mandibles. Another wasp softened her spider’s anterior abdomen with her mandibles, probably in preparation for oviposition. FL: Manatee County, Sun City; 23 September 2010; M. Keokuk. Host: Hogna sp., adult or subadult female. The wasp pulled the spider backwards, grasping its right pedipalp with her mandibles (Keokuk 2010). FL: Monroe County, Lignumvitae Key, near Islamorada; 12 March 2010; S. Dunn. Host: Hogna sp., adult or subadult female. The wasp pulled the spider backwards up a wall, grasping its left pedipalp with her mandibles (Bugman 2010). FL: Okeechobee County, Okeechobee; 14 July 2009; no collector. Host: Hogna sp., adult or subadult female (Eaton 2009). GA: Clarke County, Athens, Sandy Creek Nature Center; 5 July 2009; C. Champagne. Host: Dolomedes albineus, adult female (Champagne 2009). GA: Oglethorpe County, Wolfskin District, 16 km SE Athens; 4 August 2008 (at dusk), 17 July 2009 (afternoon); W. Hughes. Hosts: Rabidosa rabida, subadult; Dolomedes albineus, adult or subadult female. Both wasps dragged their spider backwards across a front deck of a house, grasping the left pedipalp with the mandibles (Fig. 19). The D. albineus was twice dragged up and down a wall Figure 19. Tachypompilus f. ferrugineus female dragging Dolomedes albineus adult or subadult female across front deck of house, grasping its left pedipalp with her mandibles (W. Hughes, pers. comm.). 28 Southeastern Naturalist Vol. 11, Monograph No. 4 and then over a raised threshold into the kitchen (Hughes 2008–2009; W. Hughes, Wolfskin District, GA, 2012 pers. comm.)! IL: White County, Norris City; 27 September 2006; K.D. Harrelson. Host: Hogna helluo (Walckenaer), adult female (Harrelson 2006). KS: Gray County; 30 August 2006; Drakken. Host: ?Geolycosa sp. (Lycosidae), subadult female. The wasp pulled the spider up the side of a house, grasping its pedipalp with her mandibles (Drakken 2006). LA: Jefferson Davis Parish, China; 4 August 2010; jciv. Host: Rabidosa rabida, adult female. The wasp dragged the spider backwards across dried leaves and live vegetation, grasping a pedipalp with her mandibles. She traveled over 15 m in almost 10 minutes (jciv 2010). MD: Harford County, Bel Air; 20 August 2011; Greg; 1600; 31°. Host: Hogna sp., adult female (Bugman 2011c). MD: Wicomico County, Salisbury; 1 August 2008; M.M. Beckey. Host: Rabidosa punctulata, adult female. The wasp grasped the spider by its left pedipalp with her mandibles and dragged it backwards up a concrete wall (Beckey 2008). NE: Douglas County, Omaha; 21 July 2009; B. Peterson. Host: Dolomedes tenebrosus, juvenile. The wasp dragged the spider backwards, alternately grasping its left and right pedipalps with her mandibles (Peterson 2009). NE: Sarpy County, Bellevue; 17 July 2008; L. and B. Padelford. Host: Dolomedes tenebrosus, subadult female. The wasp grasped the spider by its right pedipalp with her mandibles and dragged it backwards across a lawn to its nest in a rock wall (Padelford and Padelford 2008). NJ (central): Locality not listed; 18 September 2005, 30 July 2006; danielj. Hosts: Hogna sp., adult female; Rabidosa rabida, adult female. The two wasps were observed transporting their spider backwards across a driveway and backyard, respectively, grasping a pedipalp with the mandibles (danielj 2005a, 2006). NC: Guilford County, Oak Ridge; 15 August 2009; S. Gilliam. Host: Rabidosa rabida, adult female. The wasp dragged the spider backwards, grasping its right pedipalp with her mandibles (Gilliam 2009). OH: Hamilton County, Cincinnati; 20 August 2007; R. Pfaltzgraff-Carlson. Host: Hogna helluo, juvenile. The wasp grasped the spider by its right pedipalp with her mandibles and pulled it backwards up the side of a house to a hole between two bricks (Pfaltzgraff-Carlson 2007). OK: Cleveland County, Oklahoma City; 7 June 2009; K. Gunter. Host: Hogna sp., adult female. The wasp proceeded backwards up a wall with the spider, grasping it by a pedipalp with her mandibles (Gunter 2009). OK: Rogers County, near Inola; 4 October 2008; no collector. Host: Hogna sp., adult female. The wasp dragged the spider backwards, grasping its left pedipalp with her mandibles (OK Photos 2008). PA: Washington County, Coal Center; 16 July 2009; A. Taracido. Host: Hogna sp., adult or subadult female. This wasp dragged her spider across a driveway, through grasses and leaves, up a house wall, and into a crevice of a basement window. She grasped the spider’s left pedipalp with her mandibles (Taracido 2009). 2012 F.E. Kurczewski and G.B. Edwards 29 TN: Carter County, Elizabethton; 16 July 2006; T. McNeil. Host: Hogna sp., adult or subadult female, 19 mm (wasp, 15 mm). The wasp pulled the spider backwards up a wall, grasping its left pedipalp with the mandibles (McNeil 2006). TN: Unicoi County; 2 September 2010; W. McGigham. Host: Dolomedes tenebrosus, subadult female or juvenile. This wasp pulled the spider up an earthen incline and plant stems, grasping its right pedipalp with her mandibles (Mc- Gigham 2010). TX: Rockwall County, Rockwall; 11 September 2005; no collector; 1754. Host: Lycosidae sp., adult female. “This wasp was moving backwards dragging the spider.” She appeared to grasp the spider’s left foreleg about midway with her mandibles (Blazeblaze 2005). TX: Fort Bend County, E of Needville, Brazos Bend State Park; 7 July 2002; R.M. Dashnau. Host: Dolomedes albineus, adult female. This large spider was observed “hopping/running” on the ground, being pursued closely by a "small red wasp ... 1/2 to 3/4 inches long." The wasp landed on the ground just behind the spider and chased it further, crossing in front, circling, and then running beneath it, apparently in an attempt to sting it in the underside of its cephalothorax. The spider raised its body high on its long legs to avoid being stung. After an unsuccessful attempt, the wasp flew away, perhaps having been bitten defensively by the spider (Dashnau 2002–2004). TX: Saline County, Hickory Ridge near Hemphill; 17 June 2007; Fiskurmanns. Host: Dolomedes albineus, adult or subadult female. The wasp dragged the spider backwards along the bottom of a wall, grasping its left pedipalp with her mandibles (Fiskurmanns 2007). TX: Tarrant County, Fort Worth; 29 August 2010; T. Lewis. Host: Hogna sp., adult female. The wasp dragged her spider backwards, grasping its left pedipalp with her mandibles (Lewis260 2010). WI: Dane County, Devil’s Lake State Park; 1 August 2009; K. St. Denis; at = 23°. Host: Dolomedes tenebrosus, adult female. This wasp dragged her spider backwards, grasping it by a pedipalp with her mandibles (katestdenis 2009). WV: Upshur County, near Ruraldale; 14 July 2005; S. Cresswell. Host: Lycosidae sp. The wasp dragged the paralyzed spider across a garden by the base of its 2nd right leg, then up a wooden shed wall by its left pedipalp, grasping it all the time with the mandibles (Cresswell 2005). A 1.5-minute-long video on You Tube shows a female of T. ferrugineus running after a large adult female of D. albineus on sandy-stony soil. The wasp attempted to grasp the spider's leg with her mandibles, causing it to stop and rear up defensively. The wasp appeared to sting the spider in its ventral cephalothorax at about 15 sec and circled it repeatedly grasping at its legs with the mandibles for 15 sec after which the spider grasped the wasp by wrapping its legs around her. The wasp may have stung the spider again at this time. She escaped the spider's clutches, mounted the spider dorsal side upward in a head-to-tail position, and stung it repeatedly near the bases of its chelicerae, pedipalps, and forecoxae for 30 sec. The legs of the spider collapsed inward and then extended outward. The wasp dismounted, walked a few centimeters away, and cleaned her mouthparts and antennae with her forelegs (Greatnewtini, no date). 30 Southeastern Naturalist Vol. 11, Monograph No. 4 Tachypompilus unicolor unicolor (Banks) AZ: Maricopa County, Phoenix; 3 June 2006; R. Watkins. Host: Olios sp. (Sparassidae), juvenile, 16.0 mm (wasp, 17.0 mm). The spider was dragged backwards by the wasp, dorsal side upward, being grasped by a pedipalp with her mandibles (Watkins 2008). CA: Alameda County, Fremont, Mission Peak Ohlone Trail; 18 July 2010; J.W. Chan. Host: Schizocosa mccooki or S. maxima, adult female. The wasp grasped her prey by its right pedipalp with her mandibles and proceeded backwards on the ground over stones and vegetation (Chan 2010). CA: 2, 14–15, 30 August, 25 September 2010; KEK; 1330–1648; at = 16–31°, st = 33–53°. Males pursued hunting conspecific females on the ground and in flight during the 1st and 2nd weeks in August. Seven females, 15.0–18.5 mm long, searched for prey on the sand surface, beneath decumbent vegetation, and in crevices and holes 20–30 mm in diameter. They ran rapidly across the sand and made quick flights 1-m-long. Hunting wasps tapped the sand surface with their antennae and held their wings upward at a 45° or greater angle, flicking them once or twice per second. They searched for many minutes beneath Conicosia elongata (L.) N.E. Br. (Aizoaceae) (Narrow-leaved Ice Plant) and Caprobrotus chilensis (Molina) N.E. Br. (Chilean Ice Plant) X Caprobrotus edulis (L.) N.E. Br. (Aizoaceae) (Highway Ice Plant) hybrid. The wasps stopped hunting and rested at the onset of coastal marine fog. An old or injured female with frayed wings, 17.0 mm long, was seen “staggering” across the sand 30 August. Other females were observed as late in the season as 25 September 2010 (K.E. Kurczewski, 2010 pers. comm.). CA: 31 July, 8, 15 August, 5, 12 September 2011; KEK; 1111–1446; at = 16– 23°, st = 27–60°. Males searched for females by running through vegetation and Figure 20. Tachypompilus u. unicolor female dragging ?Alopecosa kochi adult male backwards on ground, grasping base of its right pedipalp with her mandibles (R.E. Coville, pers. comm.). 2012 F.E. Kurczewski and G.B. Edwards 31 making short flights. Five females, 15.0–17.5 mm long, searched for prey on and under vegetation, investigated holes in the ground, made short flights, and constantly flicked and raised their wings. An old female was observed hunting more slowly with less wing flicking than usual on 12 September 2011 (1124–1131; at = 18.6°, st = 51.5° (K.E. Kurczewski, 2011 pers. comm.). CA: Santa Barbara County, 25 June 2010, A. Abela. A Stenopogon sp. (Asilidae) was photographed feeding on a T. u. unicolor after capturing and paralyzing it (Abela 2010a). CA: Tulare County, Sequoia National Forest near Ponderosa, 36°06'20'N, 118°31'38"W, 2179 m (elevation), 5 August 2011; KEK. A female, 15.5 mm long, hunted for prey at the edge of a coniferous forest, turning over fallen dried leaves, needles, and pieces of bark with her mandibles and forelegs and entering holes in the ground (K.E. Kurczewski, 2011 pers. comm.). NV: Washoe County, Red Rock Canyon, Red Rock Road N of Halleluja Junction; 8 July 2008; R.E. Coville. Host: ?Alopecosa kochi (Keyserling) (Lycosidae), adult male. The wasp dragged the spider backwards on the ground, grasping the base of its right pedipalp with her mandibles (Fig. 20; Coville 2008; R.E. Coville, Richmond, CA, 2012 pers. comm.). Anoplius (Lophopompilus) aethiops (Cresson) FL: Palm Beach County, Belle Glade; 25 October 1963; C.E. Seiler. Host: Hogna annexa (Chamberlin & Ivie), adult female, 12.0 mm (shriveled) (wasp, 18.5 mm) (J.R. Wiley, 2011 pers. comm.). NY: 8 October 2011; FEK; 1421; at = 23°. Host: Hogna ?helluo, juvenile, 14.5 mm (wasp, 16.5 mm). The wasp held the spider by the base of a hindleg with her mandibles and dragged it backwards on the ground toward a crack in a concrete garage foundation. Anoplius (Lophopompilus) atrox (Dahlbom) AR: Johnson County, Hurricane Wilderness Area; 8 July 2006; H. Raney: Host: Dolomedes sp., juvenile. The wasp grasped the spider by the base of a hindleg with her mandibles and dragged it backwards on the ground (Raney 2006). Anoplius (Lophopompilus) cleora (Banks) MA: Nantucket County, Jetties Beach; 27 June 2007; C. Biegler. Host: Arctosa littoralis (Hentz) (Lycosidae), adult or subadult female. The wasp grasped the base of the spider’s left hindleg with her mandibles and dragged it across the sand surface into an open burrow (Biegler 2007). Anoplius (Notiochares) lepidus atramentarius (Dahlbom) AR: Sevier County, De Queen; 24 July 2010; D. Arbour. Host: Hogna sp., adult or subadult female. This wasp dragged her partly paralyzed spider backwards through a garden (Arbour 2010). 32 Southeastern Naturalist Vol. 11, Monograph No. 4 AR: Washington County, Fayetteville; 28 August, 20 September 2006; E. Trammel. Host: Rabidosa rabida or R. punctulata, adult or subadult female. One female extracted a spider from its burrow, grasping it by a front leg with her mandibles, and stung and paralyzed it. It is unknown whether the wasp used the spider’s burrow as a nest. Another wasp pulled the same species of spider backwards on the ground, grasping it with her mandibles by the base of its left foreleg or pedipalp. She temporarily deposited the spider atop some grass blades before taking it to her burrow (Trammel 2006a, b). FL: Hillsborough County, Environmental Lands Acquisitions Program (ELAP); 22 October 2011; K.H. Allen. Host: Rabidosa punctulata, adult or subadult female (L.S. Atherton, 2011 pers. comm.). GA: Cobb County, Chattahoochee National Recreation Area, Cochran Shoals Unit; 2 July 2009; T. Bell. Host: Rabidosa punctulata, subadult female. The wasp grasped the spider by the base of its left 3rd or 4th leg and dragged it backwards across the sand for 20+ minutes to its burrow (T. Bell 2009). LA: Lafayette Parish, Broussard; 2 June 2010; JWB. Host: Rabidosa rabida, adult male. The wasp dragged the spider backwards on the ground, grasping the base of its right foreleg with her mandibles (Fig. 21; Beck 2010; J.W. Beck, 2011 pers. comm.). Anoplius (Arachnophroctonus) americanus trifasciatus (Palisot de Beauvois) FL: Highlands County, off CR 635 (scrub habitat); 16 April 2011; LSA; “very late in the day.” Host: Arctosa littoralis, juvenile, 13.5 mm (wasp, 12.5 mm; torn forewing). The wasp and prey were photographed prior to the wasp dragging the spider into her burrow (L.S. Atherton, 2011 pers. comm.). Figure 21. Anoplius lepidus atramentarius female dragging Rabidosa rabida adult male backwards on ground, grasping base of its right foreleg with her mandibles (Beck 2010). 2012 F.E. Kurczewski and G.B. Edwards 33 FL: Pinellas County, Fort De Soto Park; 4 July 2008; LSA. Host: Hogna ?carolinensis (Walckenaer), juvenile. This wasp dragged her spider backwards, grasping the base of its right hindleg with her mandibles (Fig. 22; L.S. Atherton, 2011 pers. comm.). Anoplius (Arachnophroctonus) insignis bahamas Krombein BAHAMAS: San Salvador Island, East Beach; 29 January 1982; NBE. Host: Arctosa sp., adult female, 13.0 mm (wasp, 12.0 mm). “Prey of Anoplius insignis bahamas” on label (N.B. Elliott, 2010 pers. comm.). Anoplius (Arachnophroctonus) nigritus (Dahlbom) CA: 4, 9 April 2011; KEK. Host: Hogna grandis (Banks), adult male, 15.0, 15.0 mm (wasps, 16.0, 16.4 mm) (K.E. Kurczewski, 2011 pers. comm.). TX: Hidalgo County, 7.2 km SW Mission, National Butterfly Center, NBC Gardens; 3 December 2010; LFC; 1600; at = 28°. Host: Hogna sp., probably antelucana (Montgomery). This wasp nested in silty clay loam. The female dragged the spider backwards toward an open burrow fronted by a crescent-shaped tumulus, grasping it with her mandibles by the base of its left foreleg or pedipalp (Fig. 23). Using her mandibles, she repositioned the spider so its abdomen faced the opening before entering (Fig. 24). She backed into the circular entrance, pulling the spider inside by its spinnerets with her mandibles (Fig. 25). The spider slowly disappeared from view as it was pulled inside (Fig. 26; L.F. Cooper, Haines City, FL, 2010 pers. comm.). Figure 22. Anoplius americanus trifasciatus female dragging Hogna ?carolinensis juvenile backwards on ground, grasping base of its right hindleg with her mandibles (L.S. Atherton, pers. comm.). 34 Southeastern Naturalist Vol. 11, Monograph No. 4 VA: Alexandria County, Jackson Miles Abbott National Wetlands Reserve/ Fort Belvoir; 6 August 2010; M. Moran. Host: Rabidosa rabida, adult or subadult female. This wasp, with exceedingly frayed wings, fed on fluids exuding from a sting puncture on the spider’s cephalothorax (Moran 2010). Figure 23. Anoplius nigritus female grasping Hogna sp., probably antelucana, by base of its left pedipalp or foreleg with her mandibles and dragging it backwards toward open burrow (L.F. Cooper, pers. comm.). Figure 24. Anoplius nigritus female repositioning Hogna sp., probably antelucana, near entrance with her mandibles in order to pull it inside by spinnerets (L.F. Cooper, pers. comm.). 2012 F.E. Kurczewski and G.B. Edwards 35 Anoplius (Arachnophroctonus) semicinctus (Dahlbom) MA: Middlesex County, Acton, Sudbury; 11, 25 July 2004, 5 September 2005; R.K. Walton. Hosts: Hogna sp., adult or subadult female; Schizocosa sp., adult female; Phidippus princeps (Peckham & Peckham) (Salticidae), penultimate female. Following burrow excavation, one wasp re-stung her spider, which was then presumably cleptoparasitized by an unknown miltogrammine fly (Sarcophagidae) that alit on the spider’s abdomen and larviposited. She dragged the spider Figure 26. Hogna sp., probably antelucana, disappearing from view as it is pulled down burrow by Anoplius nigritus female (L.F. Cooper, pers. comm.). Figure 25. Hogna sp., probably antelucana, being pulled down burrow by its spinnerets by Anoplius nigritus female using her mandibles (L.F. Cooper, pers. comm.). 36 Southeastern Naturalist Vol. 11, Monograph No. 4 with fly larva on its abdomen into her burrow face forward by a front leg with the mandibles rather than the typical rear-end entry during which the spinnerets are grasped (Walton 2004). Another wasp was pursued at her entrance by a cleptoparasitic pompilid of the genus Evagetes (Walton 2005; R.K. Walton, Concord, MA, 2012 pers. comm.). Anoplius (Arachnophroctonus) semirufus (Cresson) FL: Okeechobee County, Kissimmee Prairie Preserve State Park; 13–16 March 2011; LSA. Hosts: Trochosa abdita (Gertsch), adult female, 8.8 mm (wasp, 8.5 mm), with 4th right leg amputated at coxa-trochanter joint; Varacosa parthenus (Chamberlin), juvenile (4 examples); Hogna helluo species group, juvenile. Five of the six wasps cached their spider above ground on vegetation prior to burrow excavation and elevated their wings during the process (Fig. 27; L.S. Atherton, 2011 pers. comm.). Anoplius (Pompilinus) insolens (Banks) CA: Sierra County, San Francisco State University Sierra Nevada Field Campus, along Route 49, 1.6 km NE of Bassetts; 9 July 2008; R.E. Coville. Host: Xysticus sp., adult female (gravid). The wasp dragged the spider backwards on the ground, grasping the base of its 2nd right leg with her mandibles (Fig. 28; R.E. Coville, 2012 pers.comm.). Figure 27. Anoplius semirufus female inspecting Varicosa parthenus juvenile before pulling it into open burrow by its spinnerets. Females typically hold their wings in an elevated position during inspection of the spider (L.S. Atherton, pers. comm.). 2012 F.E. Kurczewski and G.B. Edwards 37 Anoplius (Pompilinus) marginatus (Say) complex* AR: Washington County, Fayetteville; 14 October 2006; E. Trammel. Host: Phidippus audax, subadult, 8.0 mm (wasp, 12.0 mm). The wasp examined the recently stung spider with her antennae and mouthparts (Trammel 2006c). FL: Liberty County, Torreya State Park; 25 May 2007; LSA. Host: Phidippus whitmani Peckham & Peckham, adult female. The wasp dragged the spider backwards, grasping it with her mandibles by the base of its left hindleg (Fig. 29; L.S. Atherton, 2011 pers. comm.). NY: 6, 28 July, 27 September, 8–9 October 2011; FEK; at = 25–31°. Hosts: Schizocosa saltatrix (Hentz), adult female, 7.0 mm (wasp, 8.5 mm); Trochosa ruricola, adult female, 9.5 mm (wasp, 10.5 mm); Amaurobiidae sp., penultimate male, 7.5 mm (wasp, 9.0 mm); Thanatus formicinus (Clerck) (Philodromidae), adult female, 8.0 mm (wasp, 9.0 mm); Phidippus audax, juvenile, 9.0, 10.0, 9.5 mm (wasps, 10.5, 11.0, 11.0 mm). *This complex consists of four or five species presently identifiable only in the male sex by their distinctive genitalia. Anoplius (Pompilinus) tenebrosus (Cresson) MA: Middlesex County, Concord; 24 April, 2 May 2005; D. Walton. Hosts: Arctosa sp., adult female; Xysticus sp., penultimate female; Thanatus formicinus, adult female. The wasps nested in bare sandy soil in April–May, fed on haemolymph exuding from the sting puncture on the spider’s cephalothorax, groomed themselves beside their paralyzed prey, and pulled the spiders into open burrows by their spinnerets with the mandibles (Walton 2005; R.K. Walton, 2012 pers. comm.). Figure 28. Anoplius insolens female dragging a Xysticus sp. female backwards on ground, grasping the base of its 2nd right leg with her mandibles (R.E. Coville, pers. comm.). 38 Southeastern Naturalist Vol. 11, Monograph No. 4 Anoplius (Anoplius) depressipes Banks FL: Collier County, Naples; 30 March 2009; R. Shackleton. Host: Dolomedes triton (Walckenaer), adult female. This wasp captured her spider on the surface of a pond. She towed the spider across the water surface for 6–7 m, grasping it by the end of its left foreleg with her mandibles and using her wings, before disappearing in dense grasses at the water’s edge (Shackleton 2009a, b). FL: Okaloosa County, Blackwater River State Park; 22 February 2008; LSA: Host: Dolomedes triton, subadult male. The wasp dragged her spider backwards on a sandy beach, released it dorsal side upward on the sand surface, and examined it with her antennae (Fig. 30). She then grasped the spider by the end of its foreleg with her mandibles and, using her wings, towed it across the water surface to the other side of a channel where she apparently had a preexisting burrow (Fig. 31; L.S. Atherton, 2011 pers. comm.). LA: Terrebonne Parish, Theriot; 6 August 2009; R. Madere. Host: Dolomedes triton, adult female (large). The wasp pulled her spider backwards through weeds near a pond, grasping it with her mandibles by the base of its left foreleg and holding it rather upright (Madere 2009). TX: Travis County, Austin; 10 April 2006; V. Engel. Host: Dolomedes triton, adult female, ?subadult female. The larger of two wasps pulled the D. triton adult Figure 29. Anoplius marginatus complex female pulling Phidippus whitmani adult female backwards across dried leaf, grasping the base of its left hindleg with her mandibles (L.S. Atherton, pers. comm.). 2012 F.E. Kurczewski and G.B. Edwards 39 female backwards by its pedipalp with the mandibles, examined it with her antennae, and obtained nourishment from the sting puncture on the underside of its cephalothorax with her mouthparts. She interrupted transporting her prey to rest, defecate, and clean herself with her hindlegs (Engel 2006). Figure 30. Anoplius depressipes female examining Dolomedes triton subadult male with her antennae on sand surface after stinging and subduing it (L.S. Atherton, pers. comm.). Figure 31. Anoplius depressipes female towing Dolomedes triton subadult male across water surface toward preexisting burrow, grasping the end of its foreleg with her mandibles and using her wings (L.S. Atherton, pers. comm.). 40 Southeastern Naturalist Vol. 11, Monograph No. 4 Anoplius (Anoplius) imbellis Banks NY: 15 June 2010; FEK; 1305; at = 26°, gt = 48°. One female, 11.0 mm long, searched for spiders in gravelly loam on and under stones. On 14 July 2010, perhaps the same female, now with heavily frayed wings, 11.0 mm long, was observed lengthening a preexisting burrow beneath a shrub in the shade. She spent 50 min (1223–1313) removing soil alternately with her forelegs, midlegs, and hindlegs, eventually forming a crescent-shaped tumulus 47 mm wide and 40 mm long in front of the opening. She backed from the burrow and began running on the surrounding soil, searching on and under stones and decumbent vegetation (at = 26–29°, gt = 33–43°). She returned and entered her burrow at 1326, exited, entered at 1327, exited, returned at 1339 without entering, walked away, returned at 1351 at the onset of light rain, entered the burrow, and plugged the opening with damp soil from below. The sun reappeared at 1440, but the female remained inside (1441–1815). The next day (15 July 2010), she exited this burrow at 1235 and excavated a new burrow beneath a dried Asclepias syriaca L. (Asclepiadaceae) (Common Milkweed) leaf in the shade under the same shrub, 23 cm N of her prior excavation (at = 28°, gt = 38°). At 1251, she exited the new burrow headfirst, turned around, entered it, and stayed inside, possibly because of the high temperatures (1252–1840; at = 29–32°, gt = 50–57°). Two recently emerged females with non-frayed wings, 9.0 and 10.0 mm long, respectively, and probably representing a 2nd generation, were observed hunting on sparsely vegetated gravelly loam 9 and 11 August 2010 at 1250–1310 and 1239–1256 (at = 30–31°, 27–28°; gt = 47–48°, 38–43°), respectively. Females with frayed wings, 9.0 and 10.0 mm long, respectively, and perhaps the same wasps, searched for prey beneath stones, in crevices and holes, and around exposed tree roots on 27 August 2010 at 1425–1455 (at = 22–23°, gt = 43–46°) and 7 and 20 September 2010 at 1648 and 1336–1358 (at = 29°, 19°; gt = 42–49°, 33–38°). Such wasps periodically made 30-cm- to1-m-long flights. NY: 17, 29–31 August, 2–3, 13, 18 September 2011; FEK; 1038–1528; at = 18–28°. Hosts: Pardosa sp., juvenile, 6.5 mm (wasp, 7.0 mm); Trochosa ruricola, adult female, 11.0, 10.0 mm (wasps, 11.0, 9.5 mm), adult male, 10.0 mm (wasp, 11.0); Trochosa sp., probably ruricola, penultimate male, 8.0 mm (wasp, 11.0); Pisaurina mira, juvenile, 10.0 mm, 39.5 mm leg span (wasp, 11.0); Amaurobius ferox, penultimate male, 11.0 mm (wasp, 11.0 mm), juvenile, 9.5, 9.0 mm (wasps, 10.0, 9.0 mm). Wasp-hunting movements increased noticeably in the presence of a spider. Instead of walking on the ground and periodically flicking her wings, such a female ran rapidly in a circuitous manner with raised wings and made short hopping flights that sometimes extended to straight-line flights of 50–75 cm. One female transported her small (8.0 mm) penultimate male prey rapidly up a vertical 2 x 6 back-deck board, grasping the spider by the base of its hindleg with her mandibles. Another female paused during prey transport to feed on haemolymph exuding from a coxa-trochanter joint. A female with slightly frayed wings, 9.0 mm long, was followed for three days (NY; 25–26, 28 September 2011) through successive burrow excavation, 2012 F.E. Kurczewski and G.B. Edwards 41 provisioning, and final closure of a five-celled nest located at the base of a moss clump in damp gravelly loam. (September 27 was cool and rainy and there was no wasp activity). She walked very slowly on 25 September, flicking her wings, while searching for a place to excavate her burrow (1129–1141; at = 21°, gt = 31°). She sampled the ground in many places before remaining in one place and digging. She backed from her entrance removing damp soil with her forelegs, midlegs, and hindlegs 57 times, eventually forming a broad crescent-shaped tumulus, 27 mm long, 28 mm wide, and 9 mm high. Her movements increased in rapidity as the soil temperature rose. She appeared less frequently in her entrance as she dug deeper, especially when molding a cell at the end of the burrow. She eventually enlarged the entrance to 6.0 mm. Twice, during excavation, a Lasius niger (L.) (Formicidae) (Black Garden Ant) worker entered the burrow and was promptly seized with the wasp’s mandibles, walked several centimeters away, and released. The female completed her burrow in 79 minutes, stood inside the entrance and cleaned her antennae with the forelegs, walked around the opening several times, and flew straight into nearby vegetation to search for prey. She excavated four side burrows and cells off the main burrow following placement of a paralyzed spider in the 1st cell, ovipositing, and spending 50, 46, 69 (4th cell), and 34 minutes for subsequent excavations. Unlike the 57 times when she removed soil from her entrance during excavation of the main (1st) burrow, she was seen only rarely in the entrance for a side-burrow excavation except for that leading to the 4th cell. The wasp evidently filled a burrow leading to a previous cell with soil from excavation of the new side burrow and cell belowground and out of sight. This wasp captured one spider on 25 September and two spiders each on 26 and 28 September. She returned to her nest with a paralyzed wolf spider in 84, 45, 34, 47 and 48 minutes, respectively, dragging the prey rapidly backwards on the ground while grasping the base of a 3rd or 4th leg with her mandibles. The spiders were transported a distance of 1.3–1.8 m in 4–5 minutes following a circuitous route, but only in 2–3 minutes following a direct route. Twice, she temporarily lost her spider enroute to her nest and searched frantically for it in flight and by running rapidly on the ground. After its recovery, she released a paralyzed prey on the tumulus, dorsal side upward or on its side, and pulled it down the burrow by its spinnerets with her mandibles after turning around inside. With the largest (adult female) lycosid, she spent 7 minutes enlarging the diameter of the main burrow and entrance in order to accommodate its girth. As direct sunlight waned and air and ground temperature decreased in late afternoon or early evening (1645–1821), she returned without prey, entered her burrow and empty cell, and closed herself inside, pushing damp soil into the entrance from below (at = 23–25°, gt = 24–28°). Final closure of the nest occurred after the largest (last) spider was taken inside cell 5, and a worker L. niger interfered with the provisioning process (28 September; 1330–1342; at = 27°, gt = 36°; see below). The wasp exited headfirst and, raking soil alternately backwards with her forelegs, backed down the burrow. She hammered this soil in place at the end of the burrow with the tip of her 42 Southeastern Naturalist Vol. 11, Monograph No. 4 abdomen. This behavior was repeated for about 5 minutes of the 12-minute-long closure. The wasp then began smoothing over the tumulus using her foretarsi and mandibles. She completed the closing process by placing tiny leaves and twigs, pebbles, and small clods of soil from a nearby solitary bee tumulus atop the area of the entrance. She searched for a place to excavate a new burrow for 10 minutes in the same area, finally selecting the base of a mossy ledge of damp loam in the shade (1352; at = 26°, gt = 30°). She closed herself inside this new excavation from below with damp soil at the onset of rain. The main burrow, 5.0 mm wide, entered the ground at a 75–80° angle with the surface and went, more or less, straight downward to the 1st cell at a depth of 51 mm (Fig. 32). The 2nd and 3rd cells were excavated off short side burrows at depths of 49 and 46 mm, respectively. The 4th cell was located at a depth of 38 mm off a side burrow branching from the main burrow near the entrance, and a 5th cell was located just above it at a depth of 35 mm (Fig. 32). The cells were roughly 6 mm wide and high and 10–12 mm long. A paralyzed spider was placed in each cell in a face outward and dorsal side upward position. It exhibited movements of the appendages and mouthparts. The spiders were identified as Trochosa ruricola, 1 adult female, 10 mm, and Trochosa sp., probably ruricola, 4 juveniles (8.0, 8.0, 8.0, 8.5 mm). The adult female T. ruricola found in the shallowest cell was the last spider taken into the nest by the wasp (Fig. 32). The wasp’s eggs, ca. 2.0 mm long, were placed on the side of the spider’s abdomen, about midway from the base. Figure 32. Completed five-celled nest of Anoplius imbellis with cells numbered in order of completion. Tumulus is stippled and shown before being leveled by the wasp during final nest closure. Fill inside burrows is not shown. Nest is dr awn to scale. 2012 F.E. Kurczewski and G.B. Edwards 43 Lasius niger workers twice stole prey spiders from A. imbellus cachement sites. An L. niger worker entered a nest as an adult female T. ruricola was being pulled down the burrow by the wasp. The wasp was pulling the spider downward by its spinnerets from below, and the ant was pulling the spider upward by its front leg from above! One A. imbellus female, 9.0 mm long, was captured by a male Machimus ?sadyates (Walker) (Diptera: Asilidae), 15.5 mm long, on 31 August 2011 (1524; at = 27°). A small female A. imbellis, 6.5 mm long, slowly searched for a place to excavate a burrow on 6 October 2011. She spent two successive nights underground at air temperatures as low as 2–3°. She examined areas of damp loamy soil between pebbles, tapping the soil with her antennae and flicking her wings sporadically as she walked zigzag from place to place (1221–1307; at = 16– 17°, gt = 23–28°). She started burrow excavation at 1308 in a depression at the base of a mossy ledge (at = 17°, gt = 26°) and finished at 1409 (at = 17°, gt = 29°), a total of 61 minutes. She then began hunting on the surrounding soil and pebbles and under nearby vegetation (1410–1453; at = 17–18°, gt = 27–29°). From 6–9 October, this female excavated, provisioned, and completed a threecelled nest with cells at depths of 31, 34, and 37 mm below the ground surface. She stocked the cells with and oviposited on Pardosa milvina adult females, 5.5 and 5.2 mm long, and a Trochosa sp., probably ruricola, juvenile, 7.0 mm long, with its right hindleg cut off at the coxa-trochanter joint. Ammosphex angularis angularis (Banks) CA: 3–4, 14, 18, 25 April 2011, 24 March, 2, 7, 8–9, 18 April 2012; KEK; 1113–1528; at = 15–21°, st = 37–48°. Hosts: Clubiona sp. (Clubionidae), juvenile, 5.1 mm; Castianeira thalia Reiskind (Corinnidae), adult male, 6.5 mm; Castianeira sp., penultimate female, 7.5 mm; ?Litophyllus sp. (Gnaphosidae), penultimate female, 6.0 mm; Sergiolus ocellatus (Walckenaer) species group (Gnaphosidae), juvenile, 5.0 mm; Thanatus formicinus, adult female, 8.0, 8.0, 6.0, 6.0 mm, adult male, 5.0, 5.0 mm; Xysticus knowltoni Gertsch, adult female, 7.0, 6.0 mm, X. sp., juvenile, 4.5 mm (wasps: 7.8, 8.1, 8.3, 6.0, 6.0, 8.4, 8.5, 7.5, 7.5, 6.5, 7.0, 7.0, 6.0, 6.0 mm) (K.E. Kurczewski, 2011–2012 pers. comm.). CA: Santa Barbara County, Vandenberg Air Force Base; 2 September 2010; A. Abela. Host: Anyphaenidae or Clubionidae sp., ?adult female, 7.1 mm (wasp, 8.5 mm). The wasp alternately grasped the spider by the base of its left or right hindleg with her mandibles and dragged it backwards across the sand, stopping occasionally to inspect it and change the leg grasp (Fig. 33; Abela 2010b; A.J. Abela, 2011 pers. comm.). Hunting females were observed from mid-March through late November at Montana de Oro State Park and early December 2009 at nearby Los Osos (K.E. Kurczewski, 2009–2012 pers. comm.). Females searched for host spiders by running through grasses and on and under other low vegetation, periodically flicking their wings, antennating the substrate, and making short quick flights. Once captured, the paralyzed spiders were transported backwards, the wasp grasping her prey by the base of its leg, often a hindleg, with the mandibles. The 44 Southeastern Naturalist Vol. 11, Monograph No. 4 spider was placed above ground, usually on a plant leaf or in an axile. Three prey were cached 39, 53, and 91 cm, respectively, from the wasp’s burrow entrance, 15 to 45 mm above ground level. Six burrows were excavated: 2 in bare sand, 1 each beneath clumps of grass and overhanging dead twigs, 1 underneath a Frangula californica (Eschsch.) A. Gray (Rhamnaceae) (California Coffee-berry) plant, and 1 in a sloping sand cliff off the side of an unidentified solitary bee burrow entrance. The wasps excavated their burrows, using the forelegs alternately. They periodically removed sand from their burrows by walking backwards, depositing the sand atop an enlarging tumulus in front of the entrance. The intervals between consecutive sand removals usually increased in duration as the burrow lengthened. When finished with the excavation, each wasp emerged from her entrance headfirst, walked around it one or more times, sometimes reentered the burrow, reexited, and walked or flew to her cached prey. She dragged the spider backwards to her nest as described above, released it abdomen toward the opening, entered the burrow, exited headfirst, grasped the spider by its spinnerets with her mandibles, and pulled it inside. Five entrances ranged from 4.0 to 6.5 mm in diameter—larger entrances being associated with larger wasps and smaller entrances with smaller wasps. Two burrows entered the sand at a 45° angle and went diagonally downward; three other burrows were more vertical in their descent. The five burrows ranged in length from 34 to 54 mm—longer burrows being associated with Figure 33. Ammosphex a. angularis female dragging an anyphaenid or clubionid ?adult female backwards on sand, grasping the base of its left hindleg with her mandibles (Abela 2010b). 2012 F.E. Kurczewski and G.B. Edwards 45 larger wasps and shorter burrows with smaller wasps. Four tumuli associated with these burrows were 41 x 42, 50 x 37, 34 x 32, and 28 x 29 mm long and wide, respectively. In each case, the paralyzed spider was placed in a rounded cell at the end of the burrow abdomen in, face out, and dorsal side upward. The wasp’s egg, 1.5 mm long, was affixed vertically to the side of the spider’s abdomen near its base. Ammosphex anomalus anomalus (Dreisbach) CA: 6 June, 18 July 2010; KEK; 1106–1525; at = 17–31°, st = 27–60°. Hosts: Lycosidae sp., not Hogna, adult female, 10.0 mm (wasp, 9.5 mm); Herpyllus hesperolus Chamberlin (Gnaphosidae), adult female, 9.5 mm (wasp, 9.5 mm), 3rd left leg missing beyond coxa-trochanter joint. One female captured her prey on the sand surface. A 2nd wasp emerged from beneath an unidentified succulent dragging her paralyzed spider backwards, grasping it by a leg with the mandibles. She carried her prey up vegetation and released it atop a plant, 11 cm above the sand surface, prior to burrow excavation. She dug her burrow in the shade in sand beneath overhanging vegetation. The entrance was 5 mm in diameter. After finishing her burrow she retrieved her spider and, grasping it with her mandibles by a leg, flew to the ground. She spent nearly 5 minutes transporting the spider backwards toward her open nest entrance (K.E. Kurczewski, 2010 pers. comm.). Ammosphex solonus solonus (Banks) CA: 4–6 April 2011, 24 March, 2, 15, 27 April 2012; KEK; 1345–1547; at = 19°. Hosts: Alopecosa kochi, adult female, 8.0, 9.0, 10.0, 10.0, 9.0, 10.5 mm (wasps, 8.9, 9.5, 9.3, 9.5, 9.2, 9.5 mm); Pardosa californica Keyserling, adult male, 7.5 mm (wasp, 8.5 mm); ?Schizocosa sp., juvenile, 8.5 mm (wasp, 9.0 mm); Zelotes cayucos Platnick & Shadab (Gnaphosidae), adult female, 9.5, 7.5, 8.5 mm (wasps, 9.5, 8.3, 8.7 mm), one spider had its left hindleg missing beyond coxa-trochanter joint (K.E. Kurczewski, 2011–2012 pers. comm.). CA: San Luis Obispo County, Los Osos, El Moro Elfin Forest; 23 March 2012; FEK; 1220–1341; at = 19°. Host: Alopecosa kochi, adult male, 8.5 mm (wasp, 9.0 mm). The paralyzed spider was dragged backwards on the sand, being held in an almost perpendicular position by the base of a hindleg with the wasp’s mandibles. The wasp cached the spider above ground in Artemisia californica (Lessing) (Asteraceae) (California Sagebrush) and then spent 81 minutes (1220– 1341) excavating a burrow in sand beneath dried plant litter. She interrupted her excavation seven times to fly or walk to examine the spider . Arachnospila arctus (Cresson) CA: San Luis Obispo County, Los Osos; 18 July 2011; KEK; 1528; at = 21°. Host: Phidippus johnsoni, subadult female, 9.5 mm (wasp, 12.0 mm). This wasp searched for prey spiders atop and beneath dried Eucalyptus leaves (K.E. Kurczewski, 2011 pers. comm.). 46 Southeastern Naturalist Vol. 11, Monograph No. 4 Arachnospila fumipennis eureka (Banks) CA: 3–4 April 2011; KEK; 1351–1547; at = 19–21°. Hosts: Pardosa californica, adult female, left hindleg cut off at coxa-trochanter joint, 9.5 mm (wasp, 10.0 mm); Alopecosa kochi, adult female, right hindleg amputated at coxa-trochanter joint, 10.0 mm (wasp, 10.5 mm) (K.E. Kurczewski, 2011 pers. comm.). CA: San Luis Obispo County, Los Osos, El Moro Elfin Forest; 4–6 April 2011; FEK; 1138–1248; at = 18–19°. Hosts: Pardosa tuoba Chamberlin, adult female, 7.5, 7.5, 8.5 mm (Lycosidae) (wasps, 9.0, 9.5, 9.5 mm); P. californica, adult male, 7.5, 7.5 mm (wasps, 8.5, 8.5 mm); Thanatus formicinus, adult male, 5.0 (wasp, 10.0 mm). Thanatus formicinus may have been used only for adult feeding because of its small size. Aporinellus completus Banks CA: Santa Barbara County, sandy coastal back dune habitat; 28 June 2011; A. Abela. Host: Habronattus californicus (Banks), ?adult female (Salticidae), 5.0 mm long (wasp, 6.5 mm; frayed wings). This wasp transported the spider backwards across the sand, grasping its face near the eyes with her mandibles (Fig. 34; A.J. Abela, 2011 pers. comm.). ID: Fremont County, St. Anthony Sand Dunes; July 1977; NBE. Host: Habronattus sp., juvenile, 5.0 mm long (wasp, 6.5 mm) (N.B. Elliott, 2010 pers. comm.). Figure 34. Aporinellus completus female pulling Habronatus californicus ?adult female backwards on sand, grasping its face near the eyes with her mandibles. This is an old wasp as indicated by her tattered forewings (A. Abela, pers. comm.). 2012 F.E. Kurczewski and G.B. Edwards 47 Aporinellus medianus Banks CA: 22 May 2011, 27 April 2012; KEK; 1152–1303; at = 15–16°, st = 38–39°. Hosts: Tibellus oblongus (Walckenaer) (Philodromidae), adult female (gravid), 9.5, 12.0 mm (wasps, 7.5, 9.5 mm); Cheiracanthium inclusum, adult female 7.5 mm (wasp, 7.5 mm). The female from 22 May 2011 had tattered wings (K.E. Kurczewski, 2011–2012 pers. comm.). Aporinellus taeniatus taeniatus (Kohl) CA: 3–6 April, 22, 30 May, 6, 20, 27 June, 10 July, 15 August, 5 September 2011, 2–3, 7–8, 15 April, 28 May 2012; KEK; 1023–1605; at = 14–23°, st = 27– 60°. Hosts: Habronattus californicus, adult female, 6.5, 7.0, 6.5, 6.5 mm (wasps, 6.5, 7.0, 6.5, 6.8 mm), adult male, 5.5, 5.0 mm (wasps, 6.5, 6.0 mm), penultimate male, 5.5, 5.0 mm (wasps, 6.0, 6.0 mm); H. peckhami (Banks), adult female, 7.0, 7.0, 7.0, 6.0, 7.0, 6.5, 7.0, 6.5 mm (wasps, 7.5, 7.0, 7.0, 6.5, 7.0, 7.0, 7.0, 6.8 mm), adult male, 5.5, 5.5 mm (wasps, 6.5, 6.5 mm); Habronattus sp., juvenile female, 5.0 mm (wasp, 5.8 mm) (K.E. Kurczewski, 2011–2012 pers. comm.). CA: San Luis Obispo County, Los Osos, El Moro Elfin Forest; 4–5 April 2011; 29–30 March 2012; FEK; 1110–1635; at = 17–19°; sst = 36–44°. Hosts: H. californicus, adult male, 4.5 mm (wasp, 5.8 mm); H. peckhami, adult female, 7.0, 6.5, 7.0 mm (wasps, 6.7, 6.8, 6.8 mm); Habronattus sp., juvenile, 6.0, 5.0 mm (wasps, 6.5, 5.8 mm). All burrows were excavated in bare sand in the open or underneath overhanging twigs or plants. Burrow excavation involved the wasp loosening the sand with her mandibles and then using the tarsal comb of her forelegs alternately to rake it backwards beneath her synchronously raising abdomen. As the female dug deeper, damp sand accumulated in a crescent-shaped tumulus in front of and partly around her entrance. After finishing her burrow, the wasp walked around her entrance up to three times and then flew or walked to retrieve her paralyzed cached spider. Four burrow excavations took 26, 28, 36, and 62 minutes, respectively. Paralyzed prey were cached up to 0.9 m from the wasp’s burrow entrance. The spiders were placed dorsal or ventral side upward on decumbent or upright plants, usually on a leaf, at heights ranging from 15 to 85 mm above the sand surface. Prey transport involved retrieving the paralyzed spider from its cache atop a plant, examining it with the antennae, grasping it dorsal side upward by the face with the mandibles, and flying to the ground. Transport then usually included walking rather straight backwards interspersed with turning around and making short low flights forward, in either case maintaining the grasp of the spider’s face with the mandibles. The two largest (heaviest) salticids were pushed forward (abdomen first) across the sand instead of being carried forward in flight. The spider’s abdomen moved from side to side as the wasp’s hindlegs kicked sand backwards while pushing the relatively heavy prey. Placed in a vial, neither wasp relinquished her grasp of the spider’s face as she continued to push the spider forward. Upon close inspection, the female grasped the spider with her mandibles near the anterior lateral eyes or between the anterior and posterior lateral eyes. At her entrance, the wasp positioned the spider with its spinnerets toward the 48 Southeastern Naturalist Vol. 11, Monograph No. 4 opening and, grasping them with her mandibles, pulled the prey down the open burrow after turning around inside. Three burrow entrances were 5–6 mm in diameter. The burrows ranged in length from 38–44 mm, the longest burrow being associated with the 62 minutelong burrow excavation. The tumuli associated with these burrows were 36–45 mm wide and 32–48 mm long prior to final nest closure. Hedychridium coruscum and Argochrysis toralis Kimsey, both Chrysididae, lurked on the sand surface and low vegetation near A. taeniatus females excavating burrows, but they did not enter an excavation. Males of the asilid Machimus occidentalis (Hine) trailed two female A. taeniatus carrying paralyzed spiders to their burrows. The robber flies did not attempt to capture the wasps possibly because of their small sizes and the fact that they remained on the ground and did not attempt to fly. Aporinellus yucatanensis (Cameron) CA: 14 April 2011; KEK; 1323–1518; at = 20–21°, st = 40–41°. Host: Xysticus nevadensis (Keyserling), adult male, 4.5, 4.5, 4.5 mm (wasps, 6.5, 6.5, 7.0 mm) (K.E. Kurczewski, 2011 pers. comm.). Discussion Cursorial-hunting, retreat-dwelling, orb-weaving, and burrowing spiders comprise typical hosts for the approximately 300 North American pompilid species (Evans and Wiley 2001; Evans and Yoshimoto 1962; Krombein 1979; Kurczewski and Kurczewski 1968a, 1968b, 1972, 1973; Wasbauer and Kimsey 1985). Many spider wasps, especially those living in woodlands with more than a single generation per year, include several unrelated families of cursorial-hunting and retreat-dwelling spiders among their hosts (Table 1; Evans and Yoshimoto 1962, Kurczewski 1999, Richards and Hamm 1939). These wasp species are poor “spider taxonomists” but good “spider ecologists” (Evans and Yoshimoto 1962). Pompilids with “narrow ecological ranges” are more restricted in prey selection than species with “broad ecological ranges” (Krombein 1953). Some species of Priocnemis, Dipogon, and Auplopus (Pepsinae), and Sericopompilus, Anoplius (Pompilinus), Ammosphex, and Arachnospila (Pompilinae) are rather unselective of their host spider families (Table 1). Such polyphagous prey selection may, in part, be related to their broad habitat preferences, at least in the Pepsinae. Most such species nest in closed and open woodlands, brush and shrub edges, and overgrown fields where spider families, genera, and species are numerous (Evans and Yoshimoto 1962, Kurczewski 1999). Many species of oligophagous pompilids, especially in the Pompilinae, nest in open sandy and gravelly habitats where spider families, genera, and species are fewer in number (Evans and Yoshimoto 1962, Kurczewski 1999, Wasbauer and Kimsey 1985). There is a positive size relationship between spider wasp and host spider (Kurczewski and Kurczewski 1968a, 1968b, 1972, 1973). Larger pompilids 2012 F.E. Kurczewski and G.B. Edwards 49 Table 1. Species of Pompilidae and their host families of spiders.1, 2 Spider families Cy Dy Th Li Ne Ar Ly Pi Ox Ct Ag Ha Am Mi An Cl Gn Sp Ph To Sa FAMILY POMPILIDAE Subfamily PEPSINAE Tribe Pepsini Pes ● Enf ● Enu ● ● Prm ● ● ● ● ● ● ● Prc ● ● ● ● ● ● ● ● ● Pr g ● ● ● ● ● ● Prs ● ● ● ● ● ● ● ● Caf ● Dic ● ● ● ● Dis ● ● ● ● ● Tribe Ageniellini Phb ● ● ● Aua ● ● ● ● ● Aub ● ● Auc ● ● ● ● Aur3 ● ● ● ● ● ● ● ● ● ● ● Aum ● ● ● ● ● ● ● Aun ● ● ● ● ● Aga ● ● ● Agp ● Agc ● ● ● Age ● ● ● ● Ags ● ● Subfamily POMPILINAE Tribe Aporini Aph ● Tribe Pompilini Agh ● Sea ● ● ● ● ● ● ● Sen ● ● Epb ● Epc ● Epo ● Epq ● Eps ● Poa ● ● Poi ● Taf ● ● ● Tau ● ● Ana ● Anx ● ● Anc ● Anl ● Anr ● ● ● Ani ● Ann ● ● Ane ● ● 50 Southeastern Naturalist Vol. 11, Monograph No. 4 Table 1, continued. Spider families Cy Dy Th Li Ne Ar Ly Pi Ox Ct Ag Ha Am Mi An Cl Gn Sp Ph To Sa Ans ● ● ● Ano ● ● ● ● Anm ● ● ● ● ● ● ● ● ● ● ● Ant ● ● ● ● ● ● ● ● And ● Anb ● ● ● ● Amg ● ? ● ● ● ● ● Ama ● ● ● Ams ● ● Ara ● ● ● ● ● ● Arf4 ● ? Apc ● Apm ● ● ● ● ● ● Apt ● Apy ● 1Spider families arranged taxonomically according to Platnick (2012). Family abbreviations: Cy = Cyrtaucheniidae-Theraphosidae, Dy = Dysderidae-Segestriidae, Th = Theridiidae, Li = Linyphiidae, Ne = Nephilidae, Ar = Araneidae, Ly = Lycosidae, Pi = Pisauridae, Ox = Oxyopidae, Ct = Ctenidae-Zoropsidae, Ag = Agelenidae, Ha = Hahniidae, Am = Amaurobiidae, Mi = Miturgidae- Tengellidae, An = Anyphaenidae-Liocranidae, Cl = Clubionidae-Corinnidae, Gn = Gnaphosidae, Sp = Sparassidae, Ph = Philodromidae, To = Thomisidae, Sa = Salticidae. 2Pompilid species arranged taxonomically according to Krombein (1979). Species abbreviations: Pes = Pepsis sp., Enf = Entypus fulvicornis, Enu = Entypus unifasciatus, Prm = Priocnemis minorata, Prc = Priocnemis cornica, Prg = Priocnemis germana, Prs = Priocnemis scitula, Caf = Caliadurgus fasciatellus, Dic = Dipogon calipterus, Dis = Dipogon sayi, Phb = Phanagenia bombycina, Aua = Auplopus architectus, Aub = Auplopus bellus, Auc = Auplopus caerulescens, Aur = Auplopus carbonarius, Aum = Auplopus mellipes, Aun = Auplopus nigrellus, Aga = Ageniella arcuata, Agp = Ageniella accepta, Agc = Ageniella coronata, Age = Ageniella euphorbiae, Ags = Ageniella salti, Aph = Aporus hirsutus, Agh = Agenioideus humilis, Sea = Sericopompilus apicalis, Sen = Sericopompilus neotropicalis, Epb = Episyron biguttatus, Epc = Episyron conterminus, Epo = Episyron oregon, Epq = Episyron quinquenotatus, Eps = Episyron snowi, Poa = Poecilopompilus algidus, Poi = Poecilopompilus interruptus, Taf = Tachypompilus ferrugineus, Tau = Tachypompilus unicolor, Ana = Anoplius aethiops, Anx = Anoplius atrox, Anc = Anoplius cleora, Anl = Anoplius lepidus, Anr = Anoplius americanus, Ani = Anoplius insignis, Ann = Anoplius nigritus, Ane = Anoplius semicinctus, Ans = Anoplius semirufus, Ano = Anoplius insolens, Anm = Anoplius marginatus complex, Ant = Anoplius tenebrosus, And = Anoplius depressipes, Anb = Anoplius imbellis, Amg = Ammosphex angularis, Ama = Ammosphex anomalus, Ams = Ammosphex solonus, Ara = Arachnospila arctus, Arf = Arachnospila fumipennis, Apc = Aporinellus completus, Apm = Aporinellus medianus, Apt = Aporinellus taeniatus, Apy = Aporinellus yucatanensis. 3North American and European records (Grandi 1961, Richards and Hamm 1939, this study). 4North American and European records (Grandi 1961, Krombein 1979, Wasbauer 1982, this study). prey on larger spiders, especially families containing larger species of spiders, and smaller spider wasps capture smaller spiders, particularly families containing smaller species of spiders. Larger species of Pompilidae tend to be more oligophagous than the smaller spider wasps, which tend to be more polyphagous. Thus, species of Pepsis provision their nests with tarantulas; Entypus, Lophopompilus, and Arachnophroctonus spp. capture wolf spiders 2012 F.E. Kurczewski and G.B. Edwards 51 (Lycosidae), fishing spiders (Pisauridae), and wandering spiders (Ctenidae); and Episyron and Poecilopompilus spp. prey on larger orb-weaving spiders (Araneidae, Nephilidae) (Krombein 1979). Smaller spider wasps in the genera (and subgenera) Priocnemis, Dipogon, and Auplopus (Pepsinae) and Pompilinus and Ammosphex (Pompilinae) capture a preponderance of smaller spiders belonging to the Anyphaenidae, Clubionidae, Gnaphosidae, Philodromidae, Thomisidae, Salticidae, and related families—all hunting spiders that do not build a snare to catch their prey (Table 1; Krombein 1979). The majority of Pompilidae in this study can be arbitrarily separated into species that provision their nests with orb-weaving spiders of the families Araneidae and/or Nephilidae (Caliadurgus fasciatellus, Agenioideus humilis, Episyron biguttatus, E. conterminus, E. oregon, E. quinquenotatus, E. snowi, Poecilopompilus algidus, P. interruptus), species that stock their nests mainly with cursorialhunting wolf spiders (Lycosidae) and related families with similar habits such as Ctenidae and Pisauridae (Entypus fulvicornis, E. unifasciatus, Ageniella accepta, Tachypompilus ferrugineus, T. unicolor, Anoplius aethiops, A. atrox, A. cleora, A. lepidus, A. americanus, A. insignis, A. nigritus, A. semicinctus, A. semirufus, A. depressipes, A. imbellis, Ammosphex solonus, Arachnospila fumipennis), and species that prey on both cursorial-hunting and retreat-dwelling spiders (Priocnemis minorata, P. cornica, P. germana, P. scitula, Dipogon calipterus, D. sayi, Phanagenia bombycina, Auplopus architectus, A. bellus, A. caerulescens, A. carbonarius, A. mellipes, A. nigrellus, Ageniella arcuata, A. coronata, A. euphorbiae, Sericopompilus apicalis, Anoplius insolens, A. marginatus complex, A. tenebrosus, Ammosphex angularis, A. anomalus, Arachnospila arctus) (Table 1). Pepsis sp. provisions its nests with large tarantulas (Theraphosidae). Ageniella salti stocks Anyphaenidae and Clubionidae in its nests. Sericopompilus neotropicalis provisions its nests with Latrodectus mactans (Theridiidae) (Southern Black Widow Spider) and Thomisidae. Aporinellus completus and A. taeniatus are host-specific on jumping spiders (Salticidae), A. yucatanensis on crab spiders (Thomisidae), and A. medianus on jumping spiders, crab spiders, running crab spiders (Philodromidae), and other spiders (Philodromidae). Aporus hirsutus paralyzes trapdoor spiders (Cyrtaucheniidae), often on the sand surface, and stocks them in their own burrows (Krombein 1979). Many species in the tribe Ageniellini regularly amputate all of the spider’s legs at the coxa-trochanter joints, and some of them build mud cells (Phanagenia, Auplopus, and some Ageniella in North America) (Evans and Yoshimoto 1962, Krombein 1979). Evagetes padrinus is cleptoparasitic on Episyron quinquenotatus (Kurczewski and Kurczewski 1968a). Species of Pepsis, the large tarantula hawks of the New World, are predaceous on spiders of the family Theraphosidae, often species of Bothriocyrtum, Aphonopelma, and Mygale. The females prepare a burrow before hunting for prey, or they use the tarantula’s burrow as a nest (Krombein 1979, this study). Entypus fulvicornis, like its North American congeners, provisions its nests with large, often female, wolf spiders (Lycosidae). Schizocosa avida is a previously reported prey for this wasp species (Krombein 1979). Entypus 52 Southeastern Naturalist Vol. 11, Monograph No. 4 unifasciatus preys frequently on adult females of Rabidosa rabida (Evans and Yoshimoto 1962; Krombein 1979; this study). Additional hosts of this wideranging spider wasp include large adult females of other Lycosidae (Evans and Yoshimoto 1962, Townes 1957, Wasbauer and Powell 1962, this study). Dolomedes is a new host genus and Pisauridae a new host family for E. unifasciatus. Hogna is a new host genus for E. fulvicornis. Dysderidae is a new host family for P. minorata previously reported to capture Lycosidae, Pisauridae, Agelenidae, Amaurobiidae, Anyphaenidae, Liocranidae, and Clubionidae (Kurczewski 1999, this study). Priocnemis cornica is highly polyphagous, preying on nine families of cursorial- hunting and retreat-dwelling spiders (Krombein 1979; Kurczewski 1999, 2010; Kurczewski et al. 1987; this study). This species nests in preexisting burrows in the ground (Evans and Yoshimoto 1962) and sometimes uses the spider’s own burrow as a nest (Kurczewski 1981). Miturgidae is a new host family for Priocnemis germana, which previously had been reported to provision its nests with Agelenidae, Amaurobiidae, Anyphaenidae, Clubionidae, and Salticidae (Krombein 1979, Kurczewski 1999). The salticids Platycryptus undatus and Sitticus fasciger are new host species and Miturgidae and Philodromidae are new host families for Priocnemis scitula, a polyphagous species that also preys on Agelenidae, Amaurobiidae, Liocranidae, Clubionidae, Thomisidae, and Salticidae (Krombein 1979, Kurczewski 1999, Kurczewski et al. 1987, this study). In North America and Europe, Caliadurgus fasciatellus (as Calicurgus hyalinatus (Fabricius)) provisions its nests with orb-weaving Araneidae belonging to several genera (Krombein 1979, Richards and Hamm 1939, this study). This species demonstrates the unusual behavior of suspending its paralyzed spider from vegetation using the prey’s own silken threads (Kurczewski and Spofford 1985). Dipogon sayi nests mainly in preexisting cavities in wood in deciduous woodlands and stocks linear series of cells with cursorial-hunting and retreat-dwelling spiders belonging to the families Agelenidae, Amaurobiidae, Gnaphosidae, Thomisidae, and Salticidae (Krombein 1979, this study). The genus Xysticus (Thomisidae) is the predominant prey of this species in the eastern United States (Krombein 1979; K.M. O’Neill, Montana State University, 2011 pers. comm., citing H.E. Evans’ unpublished information). Dipogon calipterus has been reared from cells located in the pith and hollow stems of plants and provisions its nests with spiders of the families Amaurobiidae, Corinnidae, Gnaphosidae, and Thomisidae (Krombein 1979, Townes 1957). Most of the prey records for Phanagenia bombycina are for species of Lycosidae, but, very occasionally this species has been known to capture Agelenidae or Salticidae (Krombein 1979, Kurczewski et al. 1987; this study; N. Fensler, Findlay, OH, 2011 pers. comm.). As with many Ageniellini, P. bombycina amputates the host’s legs at the coxa-trochanter joints and makes mud cells (Evans and Yoshimoto 1962, Peckham and Peckham 1905). Prey transport in this species is variable. The wasp proceeds forward straddling the spider dorsal side upward, 2012 F.E. Kurczewski and G.B. Edwards 53 grasping it by a chelicera or a pedipalp (Evans and Yoshimoto 1962, this study), or ventral side upward by its spinnerets (Peckham and Peckham 1905), in either case using the mandibles. Auplopus architectus provisions its nests with cursorial-hunting and retreatdwelling spiders belonging to the families Miturgidae, Clubionidae, Corinnidae, Gnaphosidae, Thomisidae, and Salticidae, stocking a preponderance of jumping spiders (Salticidae) in its cells (Krombein 1979; Kurczewski 1999, 2010; Wasbauer 1982; this study). This species transports its paralyzed prey, with all legs amputated at the coxa-trochanter joints, forward either ventral side upward by grasping its spinnerets or dorsal side upward by holding a pedipalp, in both cases using the mandibles. Host records for Auplopus bellus comprise adult females of Trachelas (Corinnidae) (Genaro 1993) and juveniles of Hibana (Anyphaenidae) (this study). Trachelas prey show incomplete paralysis and legs amputated at the coxa-trochanter joints (J.A. Genaro, Museo Nacional de Historia Natural, Habana, Cuba, 2010 pers. comm.). This species makes mud cells. Auplopus caerulescens provisions its nests with small cursorial-hunting and retreat-dwelling spiders belonging to the families Anyphaenidae, Clubionidae, Corinnidae, Thomisidae, and Salticidae (Krombein 1979; Kurczewski 1989a, 2010; this study). It too amputates the legs of its prey (Kurczewski 1989a). Auplopus carbonarius is a Palearctic species recently introduced into the northeastern United States (Nolfo 1983, Kurczewski and O’Brien 1991). In the US, this species preys on an introduced European species of Miturgidae (this study). In Europe, A. carbonarius (as Pseudagenia carbonaria (Scopoli)) provisions its nests with no less than 12 families of spiders, including a majority of Clubionidae (Grandi 1961, Richards and Hamm 1939). Rau and Rau (1918) depicted mud nests of “Pseudagenia (= Auplopus) mellipes” Banks inside an abandoned Sceliphron caementarium (Drury) (Sphecidae) mud nest, under loose bark of a fallen tree, and inside an old oakapple. Evans and Yoshimoto (1962) found spheroid mud cells of A. mellipes side by side in an abandoned insect burrow in a wood stump. Krombein (1952) located mud cells of A. mellipes side by side on the underside of an exposed, decaying tree root. O’Neill and O’Neill (2010) noted A. mellipes nesting inside a 9.1-mm-wide trap-nest, the largest diameter available to the wasps. Platycryptus undatus is a previously reported host of A. mellipes (Krombein 1952, 1979). Kaston (1948) noted a mud dauber’s nest (Sphecidae) provisioned exclusively with this spider species. Anyphaenidae and Miturgidae are new host families for A. mellipes (this study) previously reported to provision its nests with Pisauridae, Clubionidae, Gnaphosidae, Philodromidae, and Salticidae (Kurczewski 1989a, 1999). Auplopus nigrellus preys on spiders belonging to six families: Anyphaenidae, Corinnidae, Miturgidae, Clubionidae, Thomisidae, and Salticidae (Krombein 1979, Wasbauer 1982). As in other species in this genus, the wasp usually amputates all of the spider’s legs at the coxa-trochanter joints and builds mud nests. 54 Southeastern Naturalist Vol. 11, Monograph No. 4 Ageniella arcuata provisions its nests with Oxyopidae, Thomisidae, and Salticidae (Krombein 1979, Kurczewski and Kurczewski 1987, this study). Evans and Yoshimoto (1962) observed an Ageniella arcuata female transporting forward an Oxyopes salticus (Oxyopidae) female, with all legs amputated at the coxa-trochanter joints, climbing tall vegetation and making short flights, straddling the spider and grasping it with her mandibles by the “anterior end”. Edward Trammel (posted as Diogmites 2009) photographed a female of this pompilid straddling a female of the same spider species and walking forward, carrying it up vegetation and making short flights, but grasping it at the opposite end (spinnerets) with her mandibles (this study). Members of the Ageniella accepta species-group, which includes A. accepta, A. blaisdelli (Fox), and A. conflicta Banks, are evidently host specific on species of Lycosidae according to reliable published reports (Krombein 1979, Wilson and Pitts 2007, this study). Waichert et al. (2011) provided data that members of this species-group are synonymous, but these data are not conclusive, and further study is necessary before making taxonomic changes here. Following amputation of its legs, the paralyzed spider is carried forward, often dorsal side upward and face first, being held with the mandibles by a chelicera or pedipalp (Evans and Yoshimoto 1962, Kurczewski and Kurczewski 1968a, this study). The wasp’s wings are usually held flat on the dorsum and do not assist in the forward transport if the spider is not relatively large in size (Kurczewski and Kurczewski 1968a). Ageniella coronata and A. salti amputate the legs of the spider at the coxatrochanter joints and walk forward with the prey. Lycosidae is a new host family for A. coronata, which previously had been associated with the prey families Tengellidae and Clubionidae (Krombein 1979, this study). Prey records for A. salti, which amputates the spider’s legs, include Anyphaenidae and Clubionidae (Krombein 1979, this study). Ageniella euphorbiae has been collected with cursorial- hunting and retreat-dwelling spiders of the families Lycosidae, Agelenidae, Hahniidae, and Anyphaenidae (Krombein 1979, this study). None of the spiders captured by A. euphorbiae had their legs amputated at the coxa-trochanter joints (Kurczewski and Kurczewski 1968a). Some species of Ageniella transport prey spiders to their nests partly in flight (Evans and Yoshimoto 1962), as observed for A. euphorbiae and A. arcuata. Aporus hirsutus inhabits active and relict sand dunes in the western US and paralyzes, stocks, and oviposits on trapdoor spiders (Cyrtaucheniidae) in their own burrows. All host records for A. hirsutus are for the genus Aptostichus (Krombein 1979, this study). Species of Evagetes are cleptoparasites on other species of Pompilidae (Evans and Yoshimoto 1962). Evagetes padrinus is a known cleptoparasite of Episyron quinquenotatus (Kurczewski and Kurczewski 1968a), as surmised in this study. Numerous prey records for Agenioideus humilis indicate specificity for orb-weaving spiders (Araneidae). Reported prey genera include Acacesia, Araneus, Araniella, Conepeira, and Neosconella (Krombein 1979, Wasbauer 1982, this study). 2012 F.E. Kurczewski and G.B. Edwards 55 Sericopompilus apicalis stocks Araneidae, Lycosidae, Oxyopidae, Anyphaenidae, Philodromidae, Thomisidae, and Salticidae in its nests (Krombein 1979, this study). Sericopompilus neotropicalis preying on an adult female of Latrodectus mactans (Theridiidae) is confirmed from photographs presented by Elhardt (2010) (this study). The only previous host record for this pompilid is Misumenops sp. (Thomisidae) (Wasbauer 1982). The only other pompilid known to capture and provision its nests with black widow spiders is Tastiotenia festiva Evans (Evans 1961). Episyron biguttatus is host specific on orb-weaving spiders (Araneidae) of the genera Acanthepeira, Araneus, Argiope, Eriophora, Eustala, Metepeira, and Neoscona (Krombein 1979, Kurczewski et al. 1987, this study). Episyron conterminus is family-specific on orb-weaving Araneidae belonging to the genera Acacesia, Araneus, Argiope, Eriophora, Eustala, Gea, Larinia, Neoscona, and Neosconella (Krombein 1979, Kurczewski 1981, Kurczewski and Spofford 1986, this study). The few prey records for E. oregon include only the araneid genus Araneus (this study). Episyron quinquenotatus provisions its nests with Araneidae of the genera Allepeira, Araneus, Araniella, Argiope, Conepeira, Eustala, Larinioides, Metepeira, Neoscona, and Singa (Krombein 1979, Kurczewski 2001, Wasbauer 1982, this study). Episyron snowi stocks its nest cells with orb-weaving Araneidae of the genera Cyclosa and Neoscona (Krombein 1979, Kurczewski et al. 1987, this study). Poecilopompilus algidus provisions its nests with large female orb-weaving spiders of the families Araneidae and Nephilidae (Krombein 1979, Kurczewski 1981, Martins 1991, this study). Neoscona, Argiope, and Araneus are new North American host genera for P. algidus (this study). In Brazil, P. algidus stocked the nephilid Nephila clavipes in nests in July but switched to large araneids later in the year (Martins 1991). Poecilopompilus interruptus also provisions its nests with large Araneidae belonging to the genera Acanthepeira, Araneus, Argiope, Epeira, Eriophora, and Neoscona (Krombein 1979, this study). Eriophora ravilla is a new host genus and species for P. interruptus (this study). Levi (1970) noted that H.K. Wallace (pers. observ.) observed P. interruptus attempting to capture Eriophora ravilla both in its orb web and on the ground (L.S. Atherton, 2011 pers. comm.). Tachypompilus ferrugineus preys on large, often adult female Lycosidae, Pisauridae, and Ctenidae, capturing a preponderance of wolf spiders and fishing spiders (Krombein 1979; Kurczewski 1981, 1989b, 2010; Wasbauer 1982; Wilson and Pitts 2007; this study). Geolycosa (?) is a new host genus for T. ferrugineus (this study). The You Tube video reported herein is the first detailed observation of prey capture in this pompilid. Tachypompilus unicolor captures large adult and subadult lycosids and sparassids (Kurczewski 2010, this study). Live-caught T. ferrugineus females were significantly larger than handreared T. ferrugineus females from the same region. Most, if not all, of the hand-reared spiders with affixed wasp eggs or larvae or completed cocoons were removed by sifting sand through a screen followed by placing the devel56 Southeastern Naturalist Vol. 11, Monograph No. 4 oping wasps with spiders in canisters with sand. Such an operation may have partly dislodged or at least disturbed the immature wasps, resulting in smaller individuals. Furthermore, the environment of the canisters may have provided suboptimal conditions for normal wasp development. Anoplius aethiops is host specific on large wolf spiders (Lycosidae) belonging to the genera Gladicosa, Hogna, and Schizocosa (Krombein 1979, Kurczewski et al. 1987, this study). Hogna helluo is a frequent prey of this species, occurring in the same habitats (Krombein 1979; Kurczewski and Kurczewski 1968a, 1973, this study). Anoplius aethiops females nest in preexisting crevices and holes in the ground (Evans and Yoshimoto 1962, Kurczewski et al. 1987, this study). Anoplius atrox is less selective than A. aethiops, capturing large fishing spiders (Pisauridae) as well as large wolf spiders (Krombein 1979, Kurczewski 1999, this study). Anoplius cleora is almost species specific in provisioning its nests with various stages and both sexes of the sand spider Arctosa littoralis (Lycosidae) (Evans and Yoshimoto 1962; Kurczewski and Kurczewski 1968a, 1968b, 1973). Only three of numerous host records for this wasp are for lycosids other than A. littoralis (Evans and Yoshimoto 1962; Krombein 1979; Kurczewski and Kurczewski 1968a, 1968b, 1973). Anoplius lepidus captures and stocks its nests with large Lycosidae of several genera and species. Rabidosa rabida is a previously reported, common host of A. lepidus (Krombein 1979, this study). Most prey records for Anoplius americanus are for species of Lycosidae (Krombein 1979, this study); however, there are single records each for Oxyopidae and Philodromidae (Krombein 1979). The only known hosts of Anoplius insignis are adult females of Pardosa and Arctosa (Lycosidae) (Genaro 1993, this study). Prey transport in this species involves the wasp grasping the spider with her mandibles and dragging it backwards by a hindleg (J.A. Genaro, 2010 pers. comm.). Anoplius nigritus, as A. relativus (Fox), is reported to prey on larger spiders belonging to the families Agelenidae and Lycosidae (Krombein 1979, Kurczewski 1999, this study). Females excavate their own burrows from the ground surface (Evans 1951a, Evans and Yoshimoto 1962, this study) or use the spider’s burrow as a nest (Kurczewski and Kurczewski 1973, McQueen 1978). Anoplius semicinctus also preys predominantly on larger Lycosidae, although there are two host records for Salticidae, one perhaps being an adult feeding record (Pellenes coronatus (Hentz)), and the other, Phidippus princeps (Peckham & Peckham), without predatory details (Krombein 1979, Rau and Rau 1918, this study). Anoplius semirufus provisions its nests with moderate-sized juvenile and adult Lycosidae of the genera Hogna, Pardosa, Pirata, Rabidosa, Schizocosa, Trochosa, and Varacosa, and, rarely, Agelenopsis (Agelenidae) and Amaurobius (Amaurobiidae) (Krombein 1979, Kurczewski et al. 1987, this study). This spider wasp frequently caches its prey spider aboveground on vegetation during burrow excavation and elevates and holds its wings above the dorsum when provisioning its nest (Evans and Yoshimoto 1962, this study). 2012 F.E. Kurczewski and G.B. Edwards 57 The Anoplius marginatus complex consists of four or five species recognizable only in the male sex by their genitalia: marginatus, rectangularis (Dreisbach), stenotus (Banks), bequaerti (Dreisbach), and townesi Evans (Evans 1951a). Species in the A. marginatus complex are strongly polyphagous in prey selection as exemplified by “A. marginatus” preying on 11 families of cursorial-hunting and retreat-dwelling spiders plus Araneidae and Phalangiidae (Opiliones) (Krombein 1979; Kurczewski 1999, 2010; this study). Anoplius splendens (Dreisbach), a related species, is known to capture 10 families of cursorial-hunting, retreatdwelling, and orb-weaving spiders (Krombein 1979, Kurczewski 1999). Anoplius tenebrosus is an atypical pompilid species that overwinters as an adult in a deep underground burrow, emerges during the first warm days of spring, and completes its nesting cycle before summer (Alm and Kurczewski 1984). A single female is thus active for two successive years. This species is highly polyphagous, preying on many species in the families Lycosidae, Agelenidae, Amaurobiidae, Liocranidae, Gnaphosidae, Philodromidae, Thomisidae, and Salticidae (Table 1; Alm and Kurczewski 1984, Krombein 1979). Anoplius insolens is evidently another polyphagous species in the subgenus Pompilinus, having been collected four times with a spider, each a different host family: Lycosidae, Philodromidae, Thomisidae, and Salticidae (Krombein 1979, Wasbauer 1982, this study). Anoplius depressipes is host specific on large, semi-aquatic fishing spiders (Pisauridae) of the genus Dolomedes (Krombein 1979, this study). This spider wasp is ecologically restricted to the shores of ponds and slow-moving streams (Evans and Yoshimoto 1962). Anoplius depressipes females are highly adept at walking on the water surface, diving below the surface in search and pursuit of fishing spiders, and towing their paralyzed host spider across the water surface to a preexisting burrow on the shore (Evans and Yoshimoto 1962, Roble 1985, this study). Anoplius imbellis preys almost entirely on wolf spiders (Lycosidae), nesting often in gravelly soil, sometimes near water (Evans and Yoshimoto 1962, Krombein 1979, Kurczewski 2010 [as A. (A.) nigerrimus], Wasbauer 1982, Wasbauer and Powell 1962, this study). Wilson and Pitts (2007) noted this species atypically capturing Xysticus sp. (Thomisidae), and we report herein the atypical capture of Pisaurina mira (Pisauridae) and Amaurobius ferox (Amaurobiidae). The discovery of five- and three-celled nests provides previously unknown nesting information for this species. Several species of Anoplius s. str., mainly those that nest in wood or plant material, build multicelled nests, while those that nest in soil sometimes excavate cells in close proximity within the same cavity (Evans and Yoshimoto 1962, Richards and Hamm 1939). Such conservation of space, excavation time, and energy is probably tied to the absence of a foretarsal digging rake in this subgenus. Host records for Ammosphex angularis (Lycosidae, ?Anyphaenidae, Clubionidae, Corinnidae, Gnaphosidae, Philodromidae, Thomisidae, Salticidae) (Krombein 1979, Kurczewski 1999, this study) and A. anomalus (Lycosidae, 58 Southeastern Naturalist Vol. 11, Monograph No. 4 Gnaphosidae, Thomisidae) (Krombein 1979, this study) indicate a polyphagous propensity for cursorial-hunting and retreat-dwelling spiders. All host records for A. solonus are for species of Lycosidae (Krombein 1979) and, now, Gnaphosidae (this study). Arachnospila arctus preys on several families of cursorial-hunting and retreat-dwelling spiders: Lycosidae, Agelenidae, Amaurobiidae, Clubionidae, Gnaphosidae, and Salticidae (Krombein 1979, Kurczewski 2010, Wasbauer and Kimsey 1985, this study). All North American and European prey records for the holarctic Arachnospila fumipennis (as Pompilus fumipennis Zetterstedt in Europe) are for wolf spiders (Lycosidae) (Grandi 1961, Krombein 1979, Wasbauer 1982, this study). Our record of a prey philodromid may represent its use only for adult feeding. All Aporinellus completus host records are for species of Salticidae, often species of Habronattus (Krombein 1979, Kurczewski et al. 1987, Wasbauer 1982, this study). Aporinellus taeniatus preys exclusively on Salticidae, A. yucatanensis on Thomisidae, and A. medianus on Oxyopidae, Miturgidae, Anyphaenidae, Philodromidae, Thomisidae, and Salticidae (Krombein 1979, Kurczewski et al. 1988, this study). Aporinellus taeniatus typically moves its prey by walking backward interspersed with short flights forward. An unusual method of prey transport during which the wasp grasps the face of the salticid with the mandibles, dorsal side upward, and pushes it forward on the ground is probably related to the spider’s size and weight, with larger and heavier prey being moved in this manner and smaller and lighter individuals being carried forward in short flights interspersed with walking backwards (this study). Kurczewski et al. (1988) described A. wheeleri Bequaert (= A. taeniatus wheeleri) transporting Pellenes (= Habronattus?) species (Salticidae) dorsal side upward, grasping the spider’s face with the mandibles and walking backwards interspersed with turning around and making short flights forward. Aporinellus completus has been photographed exhibiting this same method of prey transport (A.J. Abela, 2011 pers. comm.). Krombein (1959) (as A. fasciatus (Smith)) and Powell (1985) both noted A. medianus grasping a philodromid by its “mouthparts” and dragging it backwards. Evans (1959) observed A. basalis Banks (as A. bridwelli Evans) pulling an oxyopid backwards up a sand bank, grasping the spider near its pedicel with the mandibles. Acknowledgments James P. Pitts, Utah State University, kindly agreed to serve as Guest Editor for this monograph. In addition, he identified several species from California and provided opinions on various other species identifications. Keith E. Kurczewski, Los Osos, CA, and Nancy B. Elliott, Siena College, sent us pompilids hand-collected with prey spiders. James R. Wiley located additional specimens of Pompilidae with prey in the Florida State Collection of Arthropods. Lyle J. and Eileen Buss, University of Florida, Gainesville, observed, photographed (Figs. 17–18), and collected Tachypompilus ferrugineus and prey from their backyard and children’s sandbox. Lyn S. Atherton, Tierra Verde, FL (Figs. 13, 15–16, 22, 27, 29–31), James W. Beck, Broussard, LA (Figs. 2–4, 21), Linda F. Cooper, Haines City, FL (Figs. 23–26), Rollin E. Coville, Richmond, CA (Figs. 20, 28), Wayne Hughes, Wolfskin District, GA (Fig. 19), Edward Trammel, 2012 F.E. Kurczewski and G.B. Edwards 59 Fayetteville, AR (Fig. 10), and Troy R. Zurovec, San Benito, TX (Fig. 1) allowed us to use their spider wasp-host spider images and sent us descriptions of wasp-spider interactions. Carol Davis, Taylorsville, UT, sent us a group of seven mud cells of Auplopus architectus metallicus affixed together (Fig. 5). Joseph W. Stoll, Syracuse University, photographed Figure 8, re-touched and image-scanned other photographs, enhanced Figure 32, and helped compose the cover photographs. Lynette Schimmer, Puyallup, WA, photographed Figs. 6 and 7; Marie L. Schmidt, Norwood, PA, photographed Figure 9; Noah Elhardt, South San Francisco, CA, Figure 11; Sasha Azevedo, Charleston, SC, Figure 12; and Alice J. Abela, Santa Maria, CA, Figures 14, 33, and 34. Ken H. Allen, Temple Terrace, FL sent us a photograph of Anoplius lepidus atramentarius provisioning its nest with Rabidosa punctulata. Kevin O’Neill, Montana State University, forwarded Howard E. Evans’ unpublished notes on Dipogon sayi, and allowed F.E. Kurczewski to review his trap-nesting manuscript before its 2010 publication. Dick Walton, Concord, MA, described his observations on nesting in Anoplius semicinctus and A. tenebrosus. Nick Fensler, Findlay, OH, forwarded a prey record of Agelenopsis (Agelenidae) for Phanagenia bombycina. J.C. Cokendolpher, Lubbock, TX, sent us two prey records for Auplopus nigrellus. Lynn Kimsey, University of California-Davis, identified potential cleptoparasitic cuckoo wasps (Chrysididae), Eric Fisher, El Dorado Hills, CA, predatory robber flies (Asilidae), and David J. Keil, California Polytechnic State University, California plant species. David Bixler, Boreal Studies Research Labs, Tower, MN; Allen Brady, Hope College; Tom Prentice and Rick Vetter, University of California-Riverside; and Margaret A. Hodge, Louisiana School for Math, Science, and the Arts, helped identify some species of Lycosidae. Kelly Kissane, Blinn College, determined Dolomedes albineus and D. tenebrosus (Pisauridae). Wayne Maddison, University of British Columbia, and Marshall Hedin, San Diego State University, named Habronattus californicus and H. peckhami (Salticidae); Rick C. West, Sooke, BC, Canada, named Aphonopelma anax (Theraphosidae); and Darrell Ubick, California Academy of Sciences, identified some unusual species of Gnaphosidae. Julio A. Genaro, Museo Nacional de Historia Natural, Habana, Cuba, sent us his observations on some Caribbean pompilids. David Headrick, California Polytechnic State University, furnished collecting equipment. We acknowledge the use of online information presented by Algidus, David Arbour, Andrew Bateman, M. M. Beckey, Erin K. Bell, Ted Bell, Tom Bentley, Craig Biegler, Blazeblaze, Mike Boone, Margarethe Brummermann, Bug Guide, Bugman, Mark C. Cassino, Alice Cavette, Carmen Champagne, Jason W. Chan, Stephen Cresswell, T. S. Crider, Julia J. Cross, danielj, Richard M. Dashnau, Carol Davis, Dawn419, Mike Deep, Diogmites (Edward Trammel), Drakken, Eric R. Eaton, Charley Eiseman, El Cobrador, Lee Elliott, Victor Engel, Kevin Firth, Fiskurmanns, John Frisch, Futureman1, Stan Gilliam, Greatnewtini, Joe A. Grzybowski (JAGpix), Kevin Gunter, Kenneth D. 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Available online at http://www.alexanderwild.com/Insects/Bees-Wasps-and-Sawflies/4230200_rM6tm 2/8/1397346070_7cgddBC#!i=1397346070&k=7cgddBC&lb=1&s=A. Accessed 11 January 2012. Wilson, J.S., and J.P. Pitts. 2007. New host associations for New World spider wasps (Hymenoptera: Pompilidae). Journal of the Kansas Entomological Society 80:223–228. W00dy4012. 2010. Two-fer: Black wasp? and ? - Auplopus mellipes. Available online at http://bugguide.net/node/view/398807/bgimage. Accessed 9 March 2011. Yuschock, J.M. 2010. Wasp - Dipogon sayi. Available online at http://bugguide.net/node/ view/591755. Accessed 4 January 2012. Zxgirl. 2008. A tragic demise. Available online at http://www. flickr.com/photos/zxgirl/ 2547378922/. Accessed 20 July 2011. 68 Southeastern Naturalist Vol. 11, Monograph No. 4 Appendix 1. Host spider index for species of Pompilidae included in this study.* CYRTAUCHENIIDAE Aptostichus atomarius Simon: Aporus (Plectraporus) hirsutus (Banks). THERAPHOSIDAE Aphonopelma anax (Chamberlin): Pepsis sp. DYSDERIDAE Dysdera crocata C.L. Koch: Priocnemis (Priocnemissus) minorata Banks. THERIDIIDAE Latrodectus mactans (Fabricius): Sericopompilus neotropicalis (Cameron). NEPHILIDAE Nephila clavipes (L.): Poecilopompilus a. algidus (Smith). ARANEIDAE Acanthepeira ?stellata (Walckenaer): Poecilopompilus i. interruptus (Say). Araneidae sp.: Episyron quinquenotatus hurdi Evans. Araneus bicentarius McCook: Episyron b. biguttatus (Fabricius). Araneus diadematus Clerck: Episyron b. biguttatus. Araneus ?diadematus: Agenioideus (Agenioideus) humilis (Cresson). Araneus marmoreus Clerck: Episyron b. biguttatus. Araneus montereyensis (Archer): Episyron oregon Evans, Episyron quinquenotatus hurdi. Araneus nordmanni (Thorell): Episyron b. biguttatus. Araneus sp.: Caliadurgus fasciatellus alienatus (Smith), Agenioideus (Agenioideus) humilis, Episyron b. biguttatus, ?Episyron snowi (Viereck), Poecilopompilus algidus coquilletti (Provancher). Argiope aurantia Lucas: Poecilopompilus a. algidus. Argiope trifasciata (Forskal): Poecilopompilus algidus coquilletti. Cyclosa conica (Pallas): Caliadurgus fasciatellus alienatus. Eriophora ravilla (C.L. Koch): Episyron b. biguttatus, Poecilopompilus a. algidus, Poecilopompilus i. interruptus. Eustala anastera (Walckenaer): Episyron conterminus cressoni (Dewitz). Gea heptagon (Hentz): Episyron conterminus cressoni. Larinioides cornutus (Clerck): Episyron q. quinquenotatus. Larinioides ?cornutus: Episyron quinquenotatus hurdi. Larinioides ?patagiatus (Clerck): Episyron quinquenotatus hurdi. Neoscona crucifera (Lucas): Episyron b. biguttatus, Poecilopompilus a. algidus, Poecilopompilus i. interruptus. Neoscona ?oaxacensis (Keyserling): Poecilopompilus algidus willistoni (Patton). Neoscona sp.: Caliadurgus fasciatellus alienatus, Episyron b. biguttatus, ?Episyron snowi. LYCOSIDAE Alopecosa kochi (Keyserling): Ammosphex s. solonus (Banks), Arachnospila fumipennis eureka (Banks). ?Alopecosa kochi: Tachypompilus u. unicolor (Say). 2012 F.E. Kurczewski and G.B. Edwards 69 Arctosa littoralis (Hentz): Anoplius (Lophopompilus) cleora (Banks), Anoplius (Arachnophroctonus) americanus trifasciatus (Palisot de Beauvois). Arctosa sp.: Anoplius (Arachnophroctonus) insignis bahamas Krombein, Anoplius (Pompilinus) tenebrosus (Cresson). ?Geolycosa sp.: Tachypompilus f. ferrugineus (Say). Hogna annexa (Chamberlin & Ivie): Anoplius (Lophopompilus) aethiops (Cresson). Hogna ?carolinensis (Walckenaer): Anoplius (Arachnophroctonus) americanus trifasciatus. Hogna grandis (Banks): Anoplius (Arachnophroctonus) nigritus (Dahlbom). Hogna helluo (Walckenaer): Tachypompilus f. ferrugineus. Hogna ?helluo: Anoplius (Lophopompilus) aethiops. Hogna helluo species group: Anoplius (Arachnophroctonus) semirufus (Cresson). Hogna lenta (Hentz): Tachypompilus f. ferrugineus. Hogna sp., probably antelucana (Montgomery): Anoplius (Arachnophroctonus) nigritus. Hogna sp.: Entypus fulvicornis (Cresson) Entypus u. unifasciatus (Say), Priocnemis (Priocnemis) cornica (Say), Tachypompilus f. ferrugineus, Anoplius (Notiochares) lepidus atramentarius (Dahlbom), Anoplius (Arachnophroctonus) semicinctus (Dahlbom). ?Hogna sp.: Entypus u. unifasciatus. Isohogna timuqua (Wallace): Tachypompilus f. ferrugineus. Lycosidae sp., not Hogna: Priocnemis (Priocnemis) cornica, Ammosphex a. anomalus (Dreisbach). Lycosidae sp.: Ageniella (Ageniella) coronata Banks, Tachypompilus f. ferrugineus. Pardosa californica Keyserling: Ammosphex s. solonus, Arachnospila fumipennis eureka. Pardosa milvina (Hentz): Priocnemis (Priocnemis) cornica, Anoplius (Anoplius) imbellis Banks. Pardosa tuoba Chamberlin: Arachnospila fumipennis eureka. Pardosa sp.: Anoplius (Anoplius) imbellis. Rabidosa hentzi (Banks): Phanagenia bombycina (Cresson). Rabidosa punctulata (Hentz): Ageniella (Ageniella) accepta (Cresson), Tachypompilus f. ferrugineus, Anoplius (Notiochares) lepidus atramentarius. Rabidosa ?punctulata: Anoplius (Notiochares) lepidus atramentarius. Rabidosa rabida (Walckenaer): Entypus u. unifasciatus, Tachypompilus f. ferrugineus, Anoplius (Notiochares) lepidus atramentarius, Anoplius (Arachnophroctonus) nigritus. Rabidosa ?rabida: Anoplius (Notiochares) lepidus atramentarius. Rabidosa sp.: Entypus u. unifasciatus. Schizocosa avida (Walckenaer): Entypus fulvicornis (Cresson). Schizocosa ?maxima Dondale & Redner: Entypus unifasciatus californicus (Townes), Tachypompilus u. unicolor. Schizocosa ?mccooki Montgomery: Entypus unifasciatus californicus, Tachypompilus u. unicolor. Schizocosa saltatrix (Hentz): Anoplius (Pompilinus) marginatus (Say) complex. Schizocosa sp.: Anoplius (Arachnophroctonus) semicinctus. ?Schizocosa sp.: Phanagenia bombycina, Ammosphex s. solonus. Trochosa abdita (Gertsch): Anoplius (Arachnophroctonus) semirufus. Trochosa ruricola (De Geer): Priocnemis (Priocnemissus) minorata, Anoplius (Pompilinus) marginatus complex, Anoplius (Anoplius) imbellis. Trochosa sp., probably ruricola: Priocnemis (Priocnemissus) minorata, Anoplius (Anoplius) imbellis. Varacosa parthenus (Chamberlin): Anoplius (Arachnophroctonus) semirufus. 70 Southeastern Naturalist Vol. 11, Monograph No. 4 PISAURIDAE Dolomedes albineus Hentz: Entypus u. unifasciatus, Tachypompilus f. ferrugineus. Dolomedes tenebrosus Hentz: Entypus u. unifasciatus, Tachypompilus f. ferrugineus. Dolomedes triton (Walckenaer): Anoplius (Anoplius) depressipes Banks. Dolomedes sp.: Anoplius (Lophopompilus) atrox (Dahlbom). Pisaurina mira (Walckenaer): Priocnemis (Priocnemissus) minorata, Anoplius (Anoplius) imbellis. OXYOPIDAE Oxyopes salticus Hentz: Ageniella (Priophanes) arcuata (Banks). AGELENIDAE Agelenopsis pennsylvanica (C.L. Koch): Priocnemis (Priocnemissus) minorata. HAHNIIDAE Calymmaria lora (Chamberlin & Ivie): Ageniella (Ageniella) euphorbiae (Viereck). AMAUROBIIDAE Amaurobiidae sp.: Anoplius (Pompilinus) marginatus complex. Amaurobius ferox (Walckenaer): Priocnemis (Priocnemissus) minorata, Anoplius (Anoplius) imbellis. ?Coras sp.: Priocnemis (Priocnemissus) minorata. ?Wadotes sp.: Priocnemis (Priocnemissus) minorata. MITURGIDAE Cheiracanthium inclusum (Hentz): Auplopus architectus metallicus (Banks), Aporinellus medianus Banks. Cheiracanthium mildei C.L. Koch: Priocnemis (Priocnemis) germana (Cresson), Auplopus carbonarius (Scopoli), Auplopus mellipes variitarsatus (Dalla Torre). Cheiracanthium ?mildei: Priocnemis (Priocnemis) scitula relicta Banks, Auplopus carbonarius. ANYPHAENIDAE Anyphaena sp.: Auplopus mellipes variitarsatus. ?Anyphaenidae sp.: Ammosphex a. angularis (Banks). Hibana ?gracilis (Hentz): Auplopus m. mellipes (Say), Auplopus mellipes variitarsatus. Hibana incursa (Chamberlin): Auplopus nigrellus (Banks). Hibana ?incursa: Ageniella (Ageniella) euphorbiae. Hibana velox (Becker): Ageniella (Ameragenia) salti (Banks). Hibana sp.: Auplopus bellus (Cresson). CLUBIONIDAE Clubiona sp.: Ammosphex a. angularis. ?Clubionidae sp.: Ammosphex a. angularis. CORINNIDAE Castianeira thalia Reiskind: Ammosphex a. angularis. Castianeira sp.: Ammosphex a. angularis. Trachelas tranquillus (Hentz): Dipogon (Deuteragenia) c. calipterus (Say). 2012 F.E. Kurczewski and G.B. Edwards 71 GNAPHOSIDAE Herpyllus hesperolus Chamberlin: Ammosphex a. anomalus. ?Litophyllus sp.: Ammosphex a. angularis. Sergiolus ocellatus (Walckenaer) species group: Ammosphex a. angularis. Zelotes cayucos Platnick & Shadab: Ammosphex s. solonus. SPARASSIDAE Olios sp.: Tachypompilus u. unicolor. PHILODROMIDAE Philodromus sp.: Priocnemis (Priocnemis) scitula relicta. Thanatus formicinus (Clerck): Anoplius (Pompilinus) marginatus complex, Anoplius (Pompilinus) tenebrosus, Ammosphex a. angularis, ?Arachnospila fumipennis eureka. Tibellus oblongus (Walckenaer): Aporinellus medianus. THOMISIDAE Bassaniana versicolor (Keyserling): Dipogon (Deuteragenia) s. sayi Banks. Xysticus knowltoni Gertsch: Ammosphex a. angularis. Xysticus nevadensis (Keyserling): Aporinellus yucatanensis (Cameron). Xysticus sp.: Dipogon (Deuteragenia) s. sayi, Anoplius (Pompilinus) insolens (Banks), Anoplius (Pompilinus) tenebrosus. SALTICIDAE Eris militaris (Hentz): Auplopus carbonarius. Eris sp.: Auplopus a. architectus (Say). Habronattus californicus (Banks): Aporinellus completus Banks, Aporinellus t. taeniatus (Kohl). Habronattus peckhami (Banks): Aporinellus t. taeniatus. Habronattus sp.: Aporinellus completus, Aporinellus t. taeniatus. Naphrys pulex (Hentz): Priocnemis (Priocnemis) scitula relicta. Paraphidippus aurantius (Lucas): Auplopus m. mellipes. Phidippus audax (Hentz): Priocnemis (Priocnemis) scitula relicta, Anoplius (Pompilinus) marginatus complex. Phidippus johnsoni Peckham & Peckham: Arachnospila arctus (Cresson). Phidippus sp. probably johnsoni: Sericopompilus apicalis (Say). Phidippus princeps (Peckham & Peckham): Anoplius (Arachnophroctonus) semicinctus. Phidippus whitmani Peckham & Peckham: Anoplius (Pompilinus) marginatus complex. Phidippus sp.: Sericopompilus apicalis. Platycryptus undatus (De Geer): Priocnemis (Priocnemis) scitula relicta, Phanagenia bombycina, Auplopus mellipes variitarsatus. Sitticus fasciger (Simon): Priocnemis (Priocnemis) scitula relicta, Auplopus caerulescens subcorticalis (Walsh). *Families of spiders and species of Pompilidae are listed taxonomically following Platnick (2012) and Krombein (1979), respectively, with species of spiders arranged alphabetically. 72 Southeastern Naturalist Vol. 11, Monograph No. 4 The Southeastern Naturalist . . . ♦ A quarterly peer-reviewed and edited interdisciplinary natural history science journal with a regional focus on the southeastern United States(ISSN 1528-7092 [print], ISSN 1938-5412 [online]). ♦ Featuring research articles, notes, and research summaries on terrestrial, fresh-water, and marine organisms, and their habitats. 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