1 2015 NORTHEASTERN NATURALIST 22(Monograph 11):1–88 Analysis of Spider Wasp Host Selection in the Eastern Great Lakes Region (Hymenoptera: Pompilidae) Frank E. Kurczewski1,* and Diane H. Kiernan2 Abstract - Analysis of host selection in 57 spider wasp species and 1 species-complex from the eastern Great Lakes Region revealed new ecological, taxonomic, and size relationships, supplementing and updating information for the family Pompilidae in the Catalog of Hymenoptera in America North of Mexico (Krombein 1979). The spider wasps preyed on large, moderate-size, or somewhat small spiders, according to their own size, and disregarded very small and tiny spiders when stocking a nest cell with a single host spider and wasp’s egg. Large pompilid species preyed mainly on adult, penultimate, and subadult female spiders, and small spider wasp species captured mostly immature/juvenile spiders. Three differentsized spider wasp species inhabiting sandy soils—Anoplius cleora (large), A. apiculatus (medium-size), and Priocnemis cornica (small)—exclusively or frequently captured different sizes, sexes, and stages of the shoreline sand spider Arctosa littoralis (Lycosidae). Anoplius nigritus, A. semicinctus, and A. cylindricus, living in some of the same sandy areas, stung, paralyzed and stocked a large or small (A. cylindricus) burrow-inhabiting wolf spider (Lycosidae), Geolycosa wrighti, in the spider’s own burrow. Caliadurgus fasciatellus, Agenioideus humilis, Episyron biguttatus, E. quinquenotatus, and Poecilopompilus interruptus preyed on different sizes, sexes, and stages of several common orb-weaver species (Araneidae) according to their own size. Two related, strongly polyphagous spider wasp species of the same size and color—Anoplius splendens, a psammophile, and A. marginatus species-complex, a group of 5 species found in sandy, gravelly and loamy areas and difficult to distinguish from one another in female form—provisioned nests with similar sizes, sexes, and stages of many of the same host spider species. Anoplius semirufus, a slightly smaller pompilid species cohabiting sandy areas with species in the A. marginatus species-complex and A. splendens, was rather oligophagous in host selection and provisioned nests mainly with small and moderate-size cursorial-hunting wolf spiders of several genera. Four large spider wasp species from abandoned, overgrown fields and woodland edges—Entypus unifasciatus, Tachypompilus ferrugineus, Anoplius aethiops, and A. atrox—preyed on large, mainly adult female fishing spiders (Pisauridae) and/or large wolf spiders (A. aethiops). Four different-sized, polyphagous, deciduous woodland pompilid species—Priocnemis minorata, P. germana, P. scitula, and Anoplius virginiensis—partitioned different sizes, sexes, and stages of common woodland cursorial-hunting and retreat-dwelling host spiders of several families, especially the hacklemesh- weaver spider Callobius bennetti (Amaurobiidae), according to their own size. Paired and 3-way statistical analyses of host selection in species of Auplopus (mellipes, nigrellus), Anoplius s. str. (imbellis, ithaca), Ammosphex (angularis, michiganensis), Arachnospila (arctus, scelestus), and Aporinellus (completus, medianus, taeniatus) disclosed significant ecological and predatory differences. Species of Pompilidae were associated with specific natural communities at Presque Isle State Park, Erie County, PA; Selkirk Shores State Park, Oswego County, NY; and Southwick Beach State Park, Jefferson County, NY. 1PO Box 15251, Syracuse, NY 13215. 2State University of New York College of Environmental Science and Forestry, Syracuse, NY 13210. *Corresponding author - fkurczewski@ twcny.rr.com. Manuscript Editor: Christopher Heckscher Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 2 Introduction Most spider wasps are strong fliers with long wings and fast runners with long, slender, spiny legs. They are extremely active and rapid in their movements as they run on the ground or vegetation, flicking their wings and antennating the substrate in search of potential prey. A hunting female locates a suitable host spider, pursues it, subdues it with 1 or more venomous stings to its cephalothorax, may feed on hemolymph exuding from the sting puncture wound, excavates a burrow in the ground or constructs a nest above ground, places the paralyzed prey inside, lays an egg on its abdomen, and closes the nest (Evans 1950). The egg hatches in a few days and the wasp larva easily penetrates the spider’s thin abdominal cuticle with its mandibles, consumes the edible portions of the host, grows relatively rapidly over several days, constructs a cocoon, pupates, and emerges as an adult several weeks or a year later (Kurczewski and Edwards 2012). The host spider must be large enough to provide an adequate amount of food for the developing wasp larva. As this study reveals, there is usually a positive size relationship between the spider wasp and its host spider. Adult, penultimate, and subadult female spiders with a sizeable abdomen are the preferred host stages and sex for large spider wasp species. Small spider wasp species develop successfully on small immature/juvenile spiders. Although more than 2500 host records are available for the approximately 300 species of North American spider wasps, there is much to learn about host selection in most pompilid species (Kurczewski 2010, Kurczewski and Edwards 2012). Early published descriptions of pompilid nesting behavior simply provided host mention (Peckham and Peckham 1898, 1905; Rau and Rau 1918), often with a general identification or misidentification of the spider. Even Evans and Yoshimoto’s (1962) landmark study of the ecology and nesting behavior of the spider wasps of the northeastern United States made no attempt to measure the wasps and their prey, except to distinguish some spiders as “large” and some others as “small”. Krombein (1955b, 1956, 1958a) used comparative body-length measurements (in mm) for some North American spider wasps and their host spiders, and Yoshimoto (1954) reported minimal and maximal wet body weight (in mg) for 2 host spiders of Priocnemis minorata Banks. Kurczewski and Kurczewski (1968a, 1968b, 1972) provided comparative size and wet-weight measurements (in mm and mg) for many North American spider wasp species and their host spiders; however, all such measurements were removed from their 1973 host records paper by an editor who deemed them unnecessary and changed them to a single mean weight ratio for each wasp species. The 400+ individual measurements removed from that paper coupled with qualitative and quantitative data from earlier and more recent spider wasp host record papers provided the foundation for the host-selection information presented in this monograph. Little is known about the quantitative comparisons and size relationships of host selection in the North American pompilid species except for Episyron quinquenotatus (Say), a common species inhabiting sandy soils in the Great Lakes Region (Evans and Yoshimoto 1962; Kurczewski 2001; Kurczewski and Kurczewski 1968a, 1968b, 1973; Peckham and Peckham 1898, 1905), and some western North Northeastern Naturalist 3 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 American and Central American spider wasp species included in a host-association study (Wilson and Pitts 2007). In an attempt to fill this void, we report herein on host selection and size comparison in 57 spider wasp species and 1 species-complex from the eastern Great Lakes Region gathered over a 58-year period (1957–2014) by one of us (F.E. Kurczewski; references are cited under individual pompilid species). Thirty-seven of the 57 species and 1 species-complex (63.8%) had sufficient quantitative data on prey selection (>10 host records) to permit reliable statistical testing and analysis by the second author (D.H. Kiernan). Methods Most of the spider wasps collected with host spiders were observed during prey transport or nesting, hand-netted by the first author (F.E. Kurczewski, 1957–2014) or his deceased father, Edmund J. Kurczewski (1961–1986), and freeze-killed. Lyle J. and Eileen Buss and G.B. Edwards furnished comparative body-length data for Tachypompilus ferrugineus and host spiders from northern Florida in order to supplement the few such records for this species from the northeastern US. Additional body-length measurements for this species and Entypus unifasciatus were gathered or extrapolated from images and information presented online (1996–2014). Many individuals furnished host records and, in some cases, wasp and spider body-length measurements for species of Pompilidae from the northeastern US. The pompilid species were identified by the first author or Howard E. Evans (now deceased), except for Anoplius imbellis and Ammosphex angularis, which were determined by James P. Pitts (Utah State University, Logan, UT). The host spiders were identified by Allen R. Brady (1964–1968), G.B. Edwards (1986–2014), Henry S. Fitch (1965–1966), Willis J. Gertsch (1960–1975), Wilton Ivie (1962–1968), Benjamin J. Kaston (1960–1962), Robin E. Leech (1972), Herbert W. Levi (1960–1973), Roy A. Norton (1971–1987), Vincent D. Roth (1964–1981), and Howard K. Wallace (1968–1973). The World Spider Catalog, version 12.5 (Platnick 2012) was extremely useful in enabling us to interpret synonyms and make corrections for the older spider species names. The body length of many host spiders was measured to the nearest mm in the field using a ruler or ocular micrometer (binocular microscope) before the spider was preserved in 70% alcohol. The body length and wet weight of the host spider depended on its age, sex, stage, when it last fed, and whether or not an adult female was gravid. The swollen abdomen of gravid females was easily ruptured by rough handling, so care had to be taken with such individuals. The wasps were measured to the nearest mm using an ocular micrometer (binocular microscope) before they were pinned and identified. Wasps and spiders collected at sunset or in the early evening were placed in individually marked vials, refrigerated, and measured the following morning. In addition to being measured to the nearest mm, many wasps and host spiders were weighed on a Mettler balance or similar scale if available. Wilson and Pitts (2007) measured the length of the spider’s cephalothorax, and this may be the most reliable measurement after the prey is preserved in alcohol because the abdomen almost always shrivels or becomes distorted in preserved specimens. However, such measurement enables only Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 4 a “half-measure” representation of the size in recently captured host spiders. The spider wasps were deposited in the collections of Cornell University, University of Kansas, or Utah State University insect museums, and the spiders were given to the Cornell University Insect Museum, SUNY College of Environmental Science and Forestry Invertebrate Collection, or Florida State Collection of Arthropods. Study Sites The majority of spider wasps with host spiders (1475/2269 records, 65.0%) were collected from Erie County in northwestern Pennsylvania (Fig. 1). Most sandinhabiting and some woodland pompilids with host spiders (1066/2269, 47.0%) were collected from Presque Isle State Park (PI; Fig. 1), an 11-km-long, compound, recurved sandspit on Lake Erie directly north of the city of Erie. Deep alluvial sand beaches characterize its lake shoreline. Behind the beaches, the sand is shaped into active and relict dunes by wave and wind action or leveled into a rather extensive sand plain. Farther inland, more-fertile sandy and loamy soils at the center of the park support a mesophytic deciduous–coniferous forest (Kurczewski 1999). Many Figure 1. Location and abbreviations for eastern Great Lakes Region spider wasp hostselection sites. Not shown: Norwood, Delaware County, PA (NO); Bethany, New Haven County, CT (CT); Weymouth-New Gretna, Atlantic County, NJ (WE); Bear, New Castle County, DE (DE); Belle Haven, Accomack County, VA (BH); Allegan State Game Area, Allegan County (AL), and Grayling, Crawford County (GY), MI; Baldwin City, Lawrence and Lecompton, Douglas County, KS (KS); and Gainesville, Alachua County, and Cedar Keys National Wildlife Refuge, Seahorse Key, Levy County, FL (FL). Northeastern Naturalist 5 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 woodland pompilid species with host spiders (409/2269, 18.0%) were collected from a mesophytic, predominantly Beech–Sugar Maple–Hemlock–Northern Red Oak forest (Fagus grandifolia Ehrh.–Acer saccharum Marshall–Tsuga canadensis (L.) Carrière –Quercus rubra L.) in or adjacent to Wintergreen Gorge Cemetery (WG; Fig. 1), 1.0–1.6 km SE of the city of Erie. This site was bordered by a large, abandoned, mainly overgrown gravel pit at the southern boundary of the cemetery property (Fig. 2; Kurczewski 1999, Kurczewski et al. 1988). Other sites where spider wasps and host spiders were collected included (Fig. 1): PENNSYLVANIA: Mill Creek, 0.5 km S Erie, Erie County, gravelly streambed (MC); Waterford, Erie County, abandoned overgrown field (WA); Meadville, Crawford County, gravelly parking lot (ME); Norwood, Delaware County, residence and backyard (NO); ONTARIO, CANADA: Port Burwell, Regional Municipality of Haldimand-Norfolk, sandy field (PB); Long Point Provincial Park, same municipality, base of sandspit (LP); NEW YORK: Niagara Falls, Niagara County (no habitat description) (NF); Eighteen Mile Creek, Derby, Erie County, rocky streamside (DR); VI Mile Creek, Ithaca, Tompkins County, sand pit and adjacent deciduous woodland (IT); Inlet Valley, Tompkins County, open deciduous woodland (IV); Etna, Tompkins County, cellar foundation and backyard (ET); Groton, Tompkins County, gravel bank and Figure 2. Abandoned overgrown gravel pit bordering south boundary of Wintergreen Gorge Cemetery, Erie County, PA (WG), September 1981. Fine and coarse gravel, loamy sand, silt loam, gravelly and clayey silt loam, and gravelly and shalely hills comprised the soils of this site. Rare and uncommon pompilid species that nested there included Ageniella fulgifrons (Kurczewski and Kurczewski 1987a), Anoplius hispidulus Dreisbach (Kurczewski and Pitts 2011), A. nigerrimus (Kurczewski et al. 1987), and Aporinellus wheeleri (Kurczewski et al. 1988). Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 6 adjacent field (GR); North Lansing, Tompkins County, abandoned overgrown gravelly field (NL); Ludlowville, Tompkins County, backyard patio (LU); 2 km S, 3 km E and 4 km NE Auburn, Cayuga County, active and defunct sand pits and adjacent sandy fields (AU); Tully, Onondaga County, deciduous–coniferous woodland (TU); Marcellus, Onondaga County, abandoned overgrown field and residence (MR); Otisco Lake, Amber, Onondaga County, cabin porch next to deciduous woodland (OL); Onondaga Hill (ON) and Oakwood Cemetery (OC), Syracuse, Onondaga County, abandoned overgrown field and open deciduous woodland; Clark Reservation State Park, Jamesville, Onondaga County, abandoned overgrown field and woodland border (JA), Limestone Creek, Fayetteville, Onondaga County, rocky streambed (FA); 6 km NW Chittenango, Madison County, sandy field (CH); Granby Center, Mallory, Fulton and Selkirk Shores State Park, Oswego County, active and abandoned sand pits and sand blowouts bordered by overgrown fields, and sandy-gravelly beach (GC, ML, FU, SS); Southwick Beach State Park, Jefferson County, sand beach, sand dunes, dry sand plain, and Poison Ivy–Dune Grape– Eastern Cottonwood plant community (Toxicodendron radicans L. Kuntze–Vitis riparia Michx.–Populus deltoides W. Bartram ex Humphry Marshall) (SB); Penny Settlement Road between Port Leyden and Lyonsdale, Lewis County, sandy opening in deciduous woodland (PS); Camp Road, Boonville, Oneida County, sandy road through coniferous–deciduous woodland (BV); New York State Campus at Wanakena, St. Lawrence County, sandy truck trail through coniferous–deciduous woodland (WN); Beekmantown, Clinton County, abandoned overgrown field (BE); Rensselaerville, Albany County, deciduous–coniferous woodland border (RE); Yonkers, Westchester County, edge of deciduous woodland (YO); CONNECTICUT: Bethany, New Haven County, deciduous woodland or elsewhere in Connecticut (CT); NEW JERSEY: Weymouth-New Gretna, Atlantic County, sandy car trails through pine barrens (WE); DELAWARE: Bear, New Castle County, residence (DE); VIRGINIA: Belle Haven, Accomack County, residence and backyard (BH); MICHIGAN: Allegan State Game Area, Fennville, Allegan County, sandy car trail and pine–oak savanna (AL); 1.5 km N Grayling, Crawford County, sandy pine barrens (GY); KANSAS: Baldwin City, Lawrence and Lecompton, Douglas County, edge of deciduous woodland, sandy fields, and sandy-gravelly railroad bed or elsewhere in Kansas (KS); and, FLORIDA: Gainesville, Alachua County, backyard sandbox, and Cedar Keys National Wildlife Refuge, Seahorse Key, Levy County, sandy lawn or elsewhere in (FL). Data from localities in other states are indicated by state abbreviation in parentheses. Explanation of Results The 57 species of spider wasps and 1 species-complex from the eastern Great Lakes Region are arranged in taxonomic and alphabetical order following the family Pompilidae in the Catalog of Hymenoptera in America North of Mexico (Krombein 1979). Number of female wasps and their inclusive collection dates (in parentheses) with associated ecological information and female wasps collected with host spiders are enumerated under Collection dates and Collection dates with prey, respectively. Number of female wasps collected with prey is further enumerated Northeastern Naturalist 7 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 in parentheses under Localities according to locality code abbreviations as listed above in the Study Sites section. Habitat includes a brief description of the study site(s) and, for some species, habitat-related wasp external-morphological information. Burrow excavation apparatus or Cell construction apparatus describes the wasp’s structural body parts used in burrow excavation or cell construction as a further indicator of habitat type and pompilid taxonomic classification. Host families and species is a taxonomic and alphabetically arranged (according to Platnick 2012) list of the families (in CAPS), genera, and species of host spiders for the 57 species and 1 species-complex of Pompilidae, statement of prey-selection diversity or specificity, and, in some species, associated-host observations and relationships. Host sexes and stages include the number of specimens of each category with percentages of the total number of host specimens for the particular wasp species. Host body length (mm) provides ranges, means ± SE, and number of examples of such measurement for each pompilid species and their host spiders. Records from the entire eastern US were used in statistical analyses. Body lengths of wasps and host spiders comprised actual measurements of specimens or measurements based on online images where the wasp and spider were aligned in longitudinal position. Wet weights (mg) and Spider:wasp wet-weight ratios provide additional spider and wasp size comparisons for each pompilid species by wet weight. A summary statement of spider wasp and host spider species comparative body length and/or wet weight, if known, is given. Where regression models are created and plotted with observed data to describe the relationship between variables, P-values for the regression models are reported along with Pearson’s correlation coefficient (r). Two sample t-tests and one-way analysis of variance are used to test for significant differences in lengths and weights (α = 0.05). Amputation of legs indicates the number of specimens of the total number of prey that had 1 or more legs or pedipalps amputated by the wasp(s) at the coxa-trochanter joints. Amputation of all legs is performed in many Ageniellini species to better fit the host spider into the confined space of the cell and to facilitate forward prey transport and, in other pompilid groups, to provide the adult wasp with hemolymph nourishment at the point of amputation (Evans and Yoshimoto 1962). References include all literature citations, personal communications, and personal observations used to define various aspects of host selection in the eastern Great Lakes species of Pompilidae. Results FAMILY POMPILIDAE Subfamily PEPSINAE Tribe Pepsini Priocnessus nebulosus (Dahlbom) Collection dates: (32; 10 July–19 August). There is a single flight period per year in mid-summer in the eastern Great Lakes Region. Collection dates with prey: 31 July 1906 (KS), 19 July 1914 (PA), 10 July 1947 (Washington, DC), 28 August 1949 (KS), 19 August 1954, 5 August 1960, 13 August 1966, 28 July 1987, 14 July 2014. Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 8 Localities: ON (1), IT (2), YO (1), WG (1), DC (1), WV (1), KS (2). Habitat: Deciduous woodland edges and sunlit interior woodland openings, often on sandy or gravelly soil. Burrow excavation apparatus: Mandibles; hind tibiae with strong dorsal serrations for pushing soil upward and backwards out of burrow. Host family and species: AGELENIDAE – Agelenopsis emertoni (Chamberlin & Ivie), A. naevia (Walckenaer), A. pennsylvanica (C.L. Koch), A. potteri (Blackwall), A. utahana (Chamberlin & Ivie), A. sp. All P. nebulosus host records (8/8, 100.0%) are for funnel-web-weaver spiders of the genus Agelenopsis. Host sexes and stages: Adult female, 3 (37.5%); adult male, 1 (12.5%); immature, 4 (50.0%). Both sexes and all stages of spiders were captured by P. nebulosus. Host body length (mm): 9.0, n = 1 (wasp, 11.0, n = 1); wet weights (mg): range = 61–94, mean = 77.50 ± 16.50, n = 2 (wasps, range = 24–63, mean = 43.50 ± 19.50, n = 2); spider:wasp wet-weight ratios: range = 1.49–2.54:1, mean = 2.02:1, n = 2). One wasp was longer than its host spider, but 2 spiders were somewhat heavier than the wasps. Amputation of legs: 0/8. No amputation of the spiders’ legs was observed despite the long legs of the prey and the cumbersome forward method of transport. References: Evans and Yoshimoto 1962; Krombein 1979; Kurczewski 1961b, 1999, 2010, pers. observ.; F.E. Kurczewski and G.B. Edwards, unpubl. data; Kurczewski and Kurczewski 1968a; Kurczewski and Pitts 2011; Townes 1957. Entypus unifasciatus (Say) [Some of the information was gathered under the species name Priocnemioides unifasciatus (Say).] Collection dates: (57; 8 July–17 September). There is a single flight period per year in mid-late summer in the eastern Great Lakes Region. Collection dates with prey: March 1954 (CT), 25 August 1989 (WV), 5 September 2004 (KS), 28 July 2007 (IL), 3 September 2007 (OK), 16 August 2008 (PA), 27 August 2008 (MO), 9 August 2009 (OH), 16 August 2009 (WV), 20 August 2009 (OK), 3 September 2009 (NJ, KS), 8 July 2010 (AR), 25 July 2010 (ON), 6 August 2011 (OH), 12 July 2014 (WV). Localities: ON (1), OH (1), WV (1) plus 16 online records from eastern and midwestern US. Habitat: Abandoned overgrown fields and deciduous woodland edges in areas of fine-grained, non-sandy soils. Burrow excavation apparatus: Mandibles; prominent serrations on dorsal edges of hind tibiae used to push soil upward and backwards during modification of existing burrow or hole in ground. The nests of this species are multi-celled, and the individual cells are made before the spider is captured. Host families and species: LYCOSIDAE – Hogna antelucana (Montgomery), H. aspersa (Hentz), H. spp., Rabidosa ?punctulata (Hentz), R. rabida (Walckenaer); PISAURIDAE – Dolomedes albineus Hentz, D. tenebrosus Hentz, D. sp. This species provisioned its nest cells with wolf spiders (Lycosidae) and fishing spiders (Pisauridae) in the eastern US and lycosids in the western US and South America. Northeastern Naturalist 9 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Rabidosa rabida (10/17 specimens, 58.8%) was a frequently captured host species in the Northeast and Midwest. Rabidosa rabida and E. unifasciatus are common inhabitants of abandoned overgrown fields and woodland edges in these regions. Dolomedes albineus is arboreal in the Southeast and D. tenebrosus occurs frequently in deciduous woodland, away from water, in the Northeast. Host sexes and stages: Adult female, 10 (47.6%); adult or subadult female, 5 (23.8%); subadult female, 5 (23.8%); subadult female or immature, 1 (4.8%). Adult and subadult females (20/21, 95.2%) are the predominant sex and stages of the host spiders. Host body lengths (mm): range = 13–24, mean = 17.50 ± 0.88, n = 13 (wasps, range = 17–21, mean = 18.69 ± 0.33, n = 13) (Fig. 3). Eleven of 13 (84.6%) wasps were as long as their host spiders (r = 0.771, P = 0.002) (Fig. 3). Adult females of Dolomedes were usually longer or as long (range = 20–24 mm, n = 2) as the longest adult or subadult females or immatures of Hogna spp. and Rabidosa rabida (range = 13–21 mm, n = 11). Amputation of legs: 0/21. References: Evans and Yoshimoto 1962; Hurd and Wasbauer 1956; Janvier 1930; Krombein 1979; Kurczewski 1961a, 1999, pers. observ.; Kurczewski and Edwards 2012, unpubl. data; Townes 1957. Priocnemis (Priocnemissus) minorata Banks Collection dates: (282; 15 April–1 July [PI-WG], 21 April–14 June [IT]). There is 1 flight period per year in mid–late spring. Priocnemis minorata was the most noticeable and frequently collected species of 4 rather common deciduous woodland pompilids. Figure 3. Spider body length plotted against wasp body length in Entypus unifasciatus, Tachypompilus ferrugineus, and Anoplius aethiops. Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 10 Collection dates with prey (NY or PA unless indicated otherwise): 9–16 May 1953, 11–14 May 1961, 21 May–17 June 1968, 5 May–22 June 1969, 14–22 May 1970, 15 May–9 June 1971, May–June 1978, 30 April 2004, 17–18 May 2010, 4 June 2010 (WV), 22–27 May 2011, 15 April 2012 (WV), 24 May 2012, 27 April 2013, 13–14 April 2014 (WV), 13–21 April 2014, 21 April 2014 (ND). Localities: WG (98), IT (25), ON (12), PI (2), IV (2); plus records from WV (3), ND (1), and Suffolk County, NY (1). Habitat: Open deciduous woodland. Forewings without dark bands or spots, an adaptation for inhabiting open woodland, and occurrence in spring when there is reduced foliage and ample sunlight. Burrow excavation apparatus: Mandibles; hind tibiae with large protuberant teeth used for pushing soil upward and backwards out of burrow. The nests of this species are multi-celled and the individual cells are made before the spider is captured. Host families and species: DYSDERIDAE - Dysdera crocata C.L. Koch; LYCOSIDAE - Arctosa rubicunda (Keyserling), Gladicosa gulosa (Walckenaer), G. sp., Hogna frondicola (Emerton), Pardosa xerampelina (Keyserling), Schizocosa sp., Trochosa ruricola (De Geer), T. sp. probably ruricola, T. terricola Thorell, Varacosa avara (Keyserling); PISAURIDAE - Dolomedes tenebrosus, Pisaurina mira (Walckenaer); AGELENIDAE - Agelenopsis pennsylvanica; AMAUROBIIDAE - Amaurobius ferox (Walckenaer), Callobius bennetti (Blackwall), Coras juvenilis (Keyserling), C. sp., Wadotes calcaratus (Keyserling), W. hybridus (Emerton), W. sp.; ANYPHAENIDAE - Hibana gracilis (Hentz); LIOCRANIDAE - Agroeca ornata Banks; CLUBIONIDAE - Clubiona canadensis Emerton, C. obesa Hentz, C. spiralis Emerton, C. sp.; PHILODROMIDAE – Thanatus formicinus (Clerck). Priocnemis minorata is strongly polyphagous, preying on 9 families of cursorial-hunting and retreat-dwelling spiders in the northeastern US including 7 families in Erie County, PA (Table 1). Callobius bennetti was the predominant prey (39/98, 39.8%) at 1 site (WG) followed by Clubiona spiralis, Wadotes hybridus, and Coras juvenilis. Wadotes hybridus (15/25, 60.0%) was the most prevalent host spider at another location (IT). Dysdera crocata (Dysderidae) is a rare family, genus, and species for a North American pompilid wasp; the only other D. crocata record is for the Anoplius marginatus species-complex. Host sexes and stages: Adult female, 118 (86.8%); penultimate female, 1 (0.7%); immature female, 3 (2.2%); adult male, 4 (3.0%); immature, 10 (7.4%). Adult females were, by far, the predominant host stage and sex. Host body lengths (mm) (all species): range = 5–14, mean = 9.56 ± 0.18, n = 108 (wasps, range = 8–14, mean = 11.38 ± 0.11, n = 108) (Fig. 4); wet weights (mg): range = 55–404, n = 2. Priocnemis minorata, one of the largest medium-sized spider wasps in deciduous woodland, captured host spiders with much body length variability. Nonetheless, larger females tended to hunt larger spiders (Fig. 4). The majority of wasps (81/108, 75.0%) were longer than their host spiders (Fig. 4). One P. minorata, 12.0 mm long, was unsuccessful in attempting to capture an adult female of Pisaurina mira, 15.5 mm long with a leg span of 45 mm. An immature Northeastern Naturalist 11 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Dolomedes tenebrosus, 9.0 mm long, was captured by another P. minorata, 10.5 mm long, in open deciduous woodland some distance from water. Host body lengths (mm) (Callobius bennetti): range = 6–14, mean = 9.68 ± 0.28, n = 37 (wasps, range = 9–13, mean = 11.24 ± 0.16, n = 37) (Fig. 5). The majority of wasps (32/37, 86.5%) were as long or longer than their host C. bennetti (Fig. 5). The comparative body lengths of P. minorata and all host spiders and P. minorata and C. bennetti showed rather moderate and weak positive linear relationships, respectively (r = 0.401, P < 0.001, n = 108 [Fig. 4]; r = 0.238, P = 0.156, n = 37 [Fig. 5]). Amputation of legs: 12/108 (11.1%) spiders had 1 (7) or 2 (5) legs amputated at the coxa-trochanter joints. An immature long-legged fishing spider, Dolomedes tenebrosus, 9.0 mm long, had the 3rd left and 4th right legs missing but another immature fishing spider, Pisaurina mira, 10.5 mm long with a leg span of 35 mm, showed no evidence of leg amputation (Kurczewski and Kurczewski 1972, Kurczewski et al. 1987). Callobius bennetti, the predominant prey species of P. minorata in Erie Table 1. Families and genera of host spiders preyed on by 4 species of polyphagous woodland pompilids— Priocnemis minorata, Priocnemis germana, Priocnemis scitula, and Anoplius virginiensis—in Erie County, PA. Families of host spiders are arranged in taxonomic order following Platnick (2012) with host genera listed alphabetically. Host-spider family and genus P. minorata P. germana P. scitula A. virginiensis LYCOSIDAE X X Arctosa X Gladicosa X Pardosa X X Trochosa X X Varacosa X PISAURIDAE X Dolomedes X Pisaurina X AGELENIDAE X X X X Agelenopsis X X X X AMAUROBIIDAE X X X X Callobius X X X X Coras X X X Wadotes X X X ANYPHAENIDAE X X Hibana X X LIOCRANIDAE X X Agroeca X X CLUBIONIDAE X X X Clubiona X X X THOMISIDAE X Xysticus X SALTICIDAE X X Maevia X X Naphrys X Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 12 Figure 4. Spider body length (all host species) plotted against wasp body length in Priocnemis minorata, P. germana, P. scitula, and Anoplius virginiensis. Figure 5. Spider body length (Callobius bennetti) plotted against wasp body length in Priocnemis minorata, P. germana, P. scitula, and Anoplius virginiensis. Nine other pompilid species preyed on Callobius bennetti in the eastern Great Lakes Region but in fewer numbers (see text). Northeastern Naturalist 13 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 County, PA (n = 39/98), showed no examples of leg amputation. Priocnemis minorata females may amputate the spiders’ legs to facilitate prey transport or to feed on hemolymph exuding from the point of amputation. This species occasionally visits flowers to obtain nectar. References: E.R. Eaton, 2013 pers. comm.; Evans and Yoshimoto 1962; Krombein 1979; Kurczewski 1962, 1999, pers. observ.; Kurczewski and Edwards 2012, unpubl. data; Kurczewski and Kurczewski 1972; Kurczewski and Pitts 2011; Kurczewski et al. 1987; Townes 1957; R.K. Walton, Concord, MA, 2013 pers. comm; Yoshimoto 1954. Priocnemis (Priocnemis) cornica (Say) Collection dates: (398; 2 May–8 November). There are 2 or, perhaps, 3 generations per year in the northeastern US. Collection dates with prey: 30 September 1957, 26 August 1960, 28 July–4 September 1961, 12 September–11 October 1962, 19 October 1963, 17 September 1965, 13 September–17 October 1967, 4 September–17 October 1968, 20 August– 23 September 1969, 24 August–8 October 1970, 26 July–16 August 1971, July–September 1974, 1986, 2 September 1991, 28 August 2008, 21 September 2009, 15–18 June 2010, 1 June–10 October 2014. Localities: PI (89), WG (19), AU (2), ME (2), GR (3), ON (5); St. Lawrence County, NY (1), Ontario (2). Habitat: Bare sandy and gravelly soils, and bare fine-grained soils of gardens and waste areas. Forewings without bands or spots, an adaptation for inhabiting open, not wooded areas. Burrow excavation apparatus: Mandibles; hind tibiae with low chevron-shaped teeth used for pushing soil upward and backwards out of burrow during modification of existing burrow or hole in the ground. The nests of this species are multi-celled, and the individual cells are made before the spider is captured. Host families and species: LINYPHIIDAE - Hypselistes florens (O.P.-Cambridge); LYCOSIDAE - Allocosa funerea (Hentz), Arctosa littoralis (Hentz), A. sp., Hogna helluo (Walckenaer), Hogna spp., Pardosa distincta (Blackwall), P. ?groenlandica (Thorell), P. milvina (Hentz), P. sp. probably milvina, P. moesta Banks, P. saxatilis (Hentz), P. spp., Pirata arenicola Emerton, P. sedentaria Montgomery, P. sp., Schizocosa avida (Walckenaer), S. crassipalpata Roewer, Trochosa ruricola, T. terricola, Varacosa avara, Lycosidae sp. not Hogna; OXYOPIDAE - Oxyopes salticus Hentz; MITURGIDAE - Cheiracanthium inclusum (Hentz); ANYPHAENIDAE - Hibana gracilis, H. sp.; LIOCRANIDAE - Agroeca sp.; CLUBIONIDAE - Clubiona abbotti L. Koch, C. kastoni Gertsch, C. maritima Emerton, C. sp.; CORINNIDAE – Trachelas tranquillus (Hentz); GNAPHOSIDAE - Drassylus rufulus (Banks), D. sp., Haplodrassus signifer (C.L. Koch), ?Herpyllus sp.; THOMISIDAE - Xysticus sp.; SALTICIDAE - Eris militaris (Hentz), Evarcha hoyi (Peckham & Peckham), Pelegrina proterva (Walckenaer), Habronattus borealis (Banks), H. decorus (Blackwall), H. viridipes (Hentz), H. spp., Salticus scenicus (Clerck), Tutelina harti (Peckham), Zygoballus nervosus (Peckham & Peckham). Priocnemis cornica is strongly polyphagous and has been documented capturing 11 Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 14 families of cursorial-hunting, burrowing and retreat-dwelling spiders in the eastern Great Lakes Region. Hypselistes florens is exceptional prey for P. cornica and the first host record for the family Linyphiidae for a North American pompilid (Kurczewski et al. 1987). The wolf spiders Arctosa littoralis (28/94, 29.8%) and Pardosa milvina (30/94, 31.9%) were major host species at Presque Isle State Park, PA, on dry sand plain behind the beaches and on shrub-dry sand plain and oak-dominant savanna farther inland, respectively. Priocnemis cornica did not capture immatures of Geolycosa wrighti (Emerton), a burrowing wolf spider, despite its abundance on sand beach, sand dunes, and dry sand plain at Presque Isle State Park, PA. Host sexes and stages: Adult female, 18 (14.9%); penultimate female, 3 (2.5%); immature female, 1 (0.8%); adult male, 11 (9.1%); immature male, 11 (9.1%); immature, 77 (63.6%). Non-adult spiders (92/121, 76.0%) were the predominant hosts. Host body lengths (mm) (all species): range = 3–9, mean = 5.13 ± 0.14, n = 94 (wasps, range = 5–9, mean = 6.93 ± 0.08, n = 94) (Fig. 6); host wet weights (mg): range = 7–22, mean = 12.00 ± 2.19, n = 6 (wasps, range = 5–16, mean = 9.50 ± 1.95, n = 6); spider:wasp wet-weight ratios: range = 1.00–1.80:1, mean = 1.36:1, n = 6. Most wasps (77/94, 81.9%) were longer than their host spiders (r = 0.247, P = 0.017) (Fig. 6), although the spiders were usually somewhat heavier than the wasps. Host body lengths (mm) (Pardosa milvina): range = 3.0–6.5, mean = 4.25 ± 0.18, n = 30 (wasps, range = 6.0–8.0, mean = 6.67 ± 0.13, n = 30 (Fig. 6); P. milvina comprised the smallest and lightest (weight) spiders captured by P. cornica (Fig. 6). All wasps (30/30) were longer than their host P. milvina (Fig. 6). Figure 6. Spider body length (Pardosa milvina, Arctosa littoralis, other species) plotted against wasp body length in Priocnemis cornica. Northeastern Naturalist 15 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Host body lengths (mm) (Arctosa littoralis): range = 3.0–9.0, mean = 5.93 ± 0.30, n = 28 (wasps, range = 6.0–9.0, mean = 7.04 ± 0.16, n = 28) (Figs. 6, 7); host wet weight (mg): 12, n = 1 (wasp, 12; n = 1); spider:wasp wet-weight ratios: 1.00:1, n = 1. Most wasps were longer than their host A. littoralis (17/28, 60.7%, r = 0.359, P = 0.061, Figs. 6, 7), but less long than those provisioning with other host spider species (Figs. 6). Although there was no significant difference in P. cornica body length between females that captured A. littoralis (7.04 mm) and those that preyed on other spider species (6.88 mm) (P = 0.403), there was a difference in mean host spider body length, with A. littoralis being significantly longer (5.93 mm) than P. milvina (4.25 mm) or other spider species (5.11 mm) (P < 0.001). Host body lengths (mm) (other species): range = 3.0–7.0, mean = 5.11 ± 0.15, n = 36 (wasps, range = 5.0–8.0, mean = 7.00 ± 0.14, n = 36) (Fig. 6); host wet weights (mg): range = 7–22, mean = 12.75 ± 3.33, n = 4 (wasps, range = 5–16, mean= 10.0 ± 2.68, n = 4). Other spider species were generally intermediate in body length between Arctosa littoralis, the longest overall host, and Pardosa milvina, the shortest overall host of P. cornica (Fig. 6). Nearly all wasps (35/36) were longer than or as long as other host spider species (Fig. 6). Amputation of legs: 2/119 (1.7%) spiders had 1 leg amputated at a coxa-trochanter joint. References: Evans and Yoshimoto 1955, 1962; Krombein 1979; Kurczewski 1961a, 1961b, 1962, 1963, 1981 (as Priocnemis sp., possibly new), 1999, 2010, pers. observ.; Kurczewski and Acciavatti 1990; Kurczewski and Edwards 2012, unpubl. data; Kurczewski and Kurczewski 1968a, 1968b, 1972; Kurczewski and Spofford 1986; Kurczewski et al. 1987; Peckham and Peckham 1898; Rau and Rau 1918; Townes 1957; Wasbauer 1982. Figure 7. Spider body length (Arctosa littoralis) plotted against wasp body length in Priocnemis cornica, Anoplius cleora, and A. apiculatus. Anoplius splendens, A. illinoensis, and A. ithaca also preyed on Arctosa littoralis but in fewer numbers. Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 16 Priocnemis (Priocnemis) germana (Cresson) Collection dates: (174; 1 June–27 September). There may be 2 generations per year in the eastern Great Lakes Region with optimal weather conditions. Priocnemis germana was the second most frequently collected species among 4 rather common deciduous woodland pompilids in Erie County, PA. Collection dates with prey: 24 July–6 September 1962, 25 July–6 September 1968, 29 July–25 August 1969, 9 September 1970, 23 July–9 August 2011. Localities: PI (29), WG (13), ON (2), NH (1). Habitat: Moist deciduous woodland. Forewings with dark bands or spots, an apparent adaptation for inhabiting closed woodland with sunlit openings (see Townes 1957:plate 2). Burrow excavation apparatus: Mandibles; hind tibiae with low, chevron-shaped teeth used for pushing soil upward and backwards out of burrow. Host families and species: AGELENIDAE - Agelenopsis utahana; AMAUROBIIDAE - Callobius bennetti, Coras sp. probably juvenilis, Wadotes calcaratus, W. hybridus, W. sp.; MITURGIDAE – Cheiracanthium mildei L. Koch; ANYPHAENIDAE - Hibana gracilis; CLUBIONIDAE - Clubiona spiralis, C. sp.; SALTICIDAE - Maevia inclemens (Walckenaer). Priocnemis germana is polyphagous, preying on 6 families of cursorial-hunting and retreat-dwelling spiders in the eastern Great Lakes Region and 5 families in Erie County, PA (Table 1). Callobius bennetti was the prevalent host spider at 2 sites in Erie County, PA (PI, WG; 29/44, 65.9%). Host sexes and stages: Adult female, 11 (25.0%); immature female, 9 (20.5%); adult male, 2 (4.5%); immature male, 1 (2.3%); immature, 21 (47.7%). Immature spiders comprised the major host stage (31/44, 70.5%) and female spiders were preferred over male spiders (20/44, 45.5%). Host body lengths (mm) (all species): range = 5.0–12.0, mean = 7.93 ± 0.23, n = 44 (wasps, range = 7.0–11.0, mean = 8.07 ± 0.14, n = 44) (Fig. 4). Host size variability was similar to that of P. minorata with larger wasps often capturing larger spiders and smaller wasps, smaller spiders (Fig. 4). The majority of wasps were the same length or longer than their host spiders (35/44, 79.5%; Fig. 4). Host body lengths (mm) (Callobius bennetti): range = 5.0–10.0, mean = 7.97 ± 0.23, n = 29 (wasps, range = 7.0–11.0, mean = 8.10 ± 0.17, n = 29) (Fig. 5). Most wasps (20/29, 69.0%) were as long or longer than their host C. bennetti (Fig. 5). The comparative body lengths of P. germana and all host spiders and P. germana and C. bennetti showed a moderate positive linear relationship (r = 0.640, P < 0.001, n = 44 [Fig. 4]; r = 0.556, P = 0.002, n = 29 [Fig. 5]). Amputation of legs: 0/44. References: Evans and Yoshimoto 1962; Krombein 1979; Kurczewski 1999, pers. observ.; Kurczewski and Edwards 2012, unpubl. data; Kurczewski and Kurczewski 1968a, 1972; Townes 1957. Priocnemis (Priocnemis) hestia (Banks) Collection dates: (49; 26 May–11 September). There are probably 2 generations per year in the northeastern US. Collection date with prey: 20 July 1954. Northeastern Naturalist 17 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Locality: IT (1). Habitat: Moist deciduous woodland. Forewings with dark bands or spots, an apparent adaptation for inhabiting closed woodland with sunlit openings (see Townes 1957:plate 2). Burrow excavation apparatus: Mandibles; hind tibiae with low, chevron-shaped teeth used for pushing soil upward and backwards out of burrow. Host family and species: LIOCRANIDAE – Agroeca sp. Host sex and stage: Immature, 1 (100.0%). Amputation of legs: 0/1. References: Evans and Yoshimoto 1962, Krombein 1979, Townes 1957. Priocnemis (Priocnemis) notha (Cresson) Collection dates: (14; 2 June–26 October). There are 2 generations per year in the eastern Great Lakes Region. Collection dates with prey: 31 August 1905, 14 September 1958, 21 August 1962, 8 September 1968, 10 October 2014. Localities: AU (1), CH (1), IT (1), ON (1). Habitat: Bare soil of garden, edge of sand pit, and sandy field. There are no dark bands or spots on the forewings, an adaptation for inhabiting open areas. Burrow excavation apparatus: Mandibles; hind tibiae with low, chevron-shaped teeth used for pushing soil upward and backwards out of burrow. Host families and species: LYCOSIDAE – Schizocosa crassipes (Walckenaer), S. sp., Trochosa terricola; MITURGIDAE – Cheiracanthium inclusum or C. mildei; CLUBIONIDAE – Clubiona sp.; SALTICIDAE – Eris militaris. Although there are only a handful of host records, this species is polyphagous in the eastern Great Lakes Region. Host sexes and stages: Adult female, 1 (25.0%); immature male, 1 (25.0%); immature, 2 (50.0%). Immature spiders are probably the preferred host stage. Host body lengths (mm): range = 5.0–8.0, mean = 6.17 ± 0.93, n = 3 (wasps, range = 7.0–8.0, mean = 7.50 ± 0.29, n = 3); host wet weight (mg): 49, n = 1 (wasp, 13, n = 1); spider:wasp wet-weight ratio: 3.77:1. One host spider was longer and much heavier than the wasp; 2 wasps were longer than their host spiders. Amputation of legs: 1/4 (25.0%). References: Evans and Yoshimoto 1962; Krombein 1979; F.E. Kurczewski pers. observ.; Kurczewski and Acciavatti 1990; F.E. Kurczewski and G.B. Edwards, unpubl. data; Kurczewski and Kurczewski 1968a, 1972, 1987b; Townes 1957. Priocnemis (Priocnemis) scitula (Cresson) Collection dates: (91; 10 July–9 October). There is probably only a single generation per year in most years in the eastern Great Lakes Region but 2 generations per year farther southward. Priocnemis scitula was less commonly collected than P. minorata or P. germana in deciduous woodland probably because it was smaller and less frequently seen. Collection dates with prey: 10 August 1954, 26 July–29 August 1968, 22 July– 22 August 1969, 19 July 1970, July–August 1972, 25 July–22 September 2010, 10 July–18 September 2011. Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 18 Localities: PI (14), WG (13), ON (14), IT (1). Habitat: Moist deciduous woodland. Forewings with dark bands or spots, an apparent adaptation for inhabiting closed woodland with sunlit openings (see Townes 1957:plate 2). Burrow excavation apparatus: Mandibles; hind tibiae with low, chevron-shaped teeth used for pushing soil upward and backwards out of burrow. Host families and species: AGELENIDAE - Agelenopsis sp. probably utahana; AMAUROBIIDAE - Callobius bennetti; MITURGIDAE - Cheiracanthium ?mildei; LIOCRANIDAE - Agroeca ornata, Agroeca sp.; CLUBIONIDAE - Clubiona kastoni, C. spiralis, C. sp. probably spiralis, C. sp.; PHILODROMIDAE - Philodromus sp.; THOMISIDAE - Xysticus sp.; SALTICIDAE - Maevia inclemens (Walckenaer), Naphrys pulex (Hentz), Phidippus audax (Hentz); Platycryptus undatus (De Geer), Sitticus fasciger (Simon). Priocnemis scitula is strongly polyphagous, capturing 8 families of cursorial-hunting and retreat-dwelling spiders in the eastern Great Lakes Region and 6 families in Erie County, PA (Table 1). Clubiona was the predominant host genus at 2 localities in Erie County, PA (PI, WG; 19/27, 70.4%). Host sexes and stages: Adult female, 5 (12.2%); immature female, 1 (2.4%); adult male, 8 (19.5%); immature male, 2 (4.9%); immature, 25 (61.0%). Immature spiders comprised the major host stage (28/41, 68.3%). Host body lengths (mm) (all species): range = 4.0–7.0, mean = 5.47 ± 0.12, n = 39 (wasps, range = 6.0–7.0, mean = 6.24 ± 0.08, n = 39) (Fig. 4). The wasps were longer than or the same length as their host spiders in nearly all examples (38/39, 97.4%, Fig. 4). Host body lengths (mm) (Callobius bennetti): range = 4.0–7.0, mean = 5.33 ± 0.88, n = 3 (wasps, range = 6.0–7.0, mean = 6.67 ± 0.33, n = 3) (Fig. 5). All 3 wasps were as long as their host C. bennetti (Fig. 5). The comparative body lengths of P. scitula and all host spiders and P. scitula and C. bennetti showed weak and strong positive linear relationships, respectively (r = 0.179, P = 0.282, n = 38 [Fig. 4]; r = 0.756, P = 0.454, n = 3 [Fig. 5]), although the latter number is based on only 3 data points. Amputation of legs: 3/39 (7.7%) spiders had 1 (2) or 2 (1) legs amputated at the coxa-trochanter joints. References: Evans and Yoshimoto 1962; Krombein 1979; Kurczewski 1999, pers. observ.; Kurczewski and Edwards 2012; Kurczewski and Kurczewski 1972; Kurczewski et al. 1987; Townes 1957. Caliadurgus fasciatellus (Spinola) [Information on this Holarctic species was gathered mainly under the species name Calicurgus hyalinatus Fabricius.] Collection dates: (208; 19 May–1 November). There are at least 2 generations per year in the eastern Great Lakes Region. Collection dates with prey: 4 August 1952 (KS), 3–8 August 1957, 4–17 September 1965, 28 September 1965 (KS), 14 September–2 October 1967, 18 July–29 August 1968, 9 July–23 September 1969, 27 July–22 September 1970, 10 September 1983, 30 June 1984, 7 August 2004, 5–10 July 2011. Northeastern Naturalist 19 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Localities: PI (14), AU (4), ON (3), MA (1), WV (2), KS (2). Habitat: Moist deciduous woodland sunlit openings and woodland edges with sandy, gravelly, or loamy soils. Forewings with dark bands or spots, an apparent adaptation for inhabiting closed woodland with sunlit openings (see Townes 1957:plate 1). Burrow excavation apparatus: Mandibles; hind tibiae with low, chevron-shaped teeth used for pushing soil upward and backwards out of burrow. Caliadurgus fasciatellus uses all 3 pairs of legs and tip of abdomen when removing soil from burrow. Host family and species: ARANEIDAE - Acanthepeira stellata (Walckenaer), Araneus marmoreus Clerck, A. sp., Araniella displicata (Hentz), Cyclosa conica (Pallas), Eustala anastera (Walckenaer), Larinioides patagiatus (Clerck), Neoscona sp., Ocrepeira ectypa (Walckenaer). Caliadurgus fasciatellus preyed on 8 genera of orb-weaver spiders (Araneidae) in the eastern US and 7 genera of Araneidae in Erie County, PA (Table 2). Host sexes and stages: Adult female, 4 (14.8%); immature female, 2 (7.4%); penultimate male, 5 (18.5%); immature male, 4 (14.8%); immature, 12 (44.4%). Most host spiders (23/27, 85.2%) were non-adult. Host body lengths (mm): range = 4.0–7.0, mean = 5.33 ± 0.17, n = 24 (wasps, range = 5.7–9.7, mean = 7.23 ± 0.18, n = 24) (Fig.8); host wet weights (mg): range = 8–48, mean = 23.86 ± 17.29, n = 7 (wasps, range = 8–22; mean = 13.43 ± 5.29; n = 7) (Fig. 9); spider:wasp wet-weight ratios: range = 0.45–6.00:1, mean = 2.29, n = 7. Twenty-two of 24 (91.7%) wasps were longer than their host spiders (Fig. 8), although the host spider weights varied considerably (Fig. 9). The comparative body lengths and wet weights of C. fasciatellus and their host Figure 8. Spider body length plotted against wasp body length in Caliadurgus fasciatellus, Episyron biguttatus, and E. quinquenotatus. Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 20 spiders showed moderate negative and negative linear relationships, respectively (r = -0.156, P = 0.535, n = 18, Fig. 8; r = -0.422, P = 0.345, n = 7, Fig. 9). Such negative relationships may be due to small sample sizes and few atypically larger wasps preying on smaller spiders. Amputation of legs: 3/20 (15.0%) spiders had a pedipalp (1) or leg (1) or 2 legs (1) missing beyond the coxa-trochanter joint. References: Evans and Yoshimoto 1962; Krombein 1958a, 1958c, 1979; Kurczewski 1999, pers. observ.; Kurczewski and Acciavatti 1990; Kurczewski and Edwards 2012; Kurczewski and Kurczewski 1968a, 1968b, 1972, 1987b; Kurczewski and Spofford 1985; Townes 1957. Dipogon (Deuteragenia) calipterus (Say) Collection dates: (11; 28 June–6 September). There may be 2 generations per year in the eastern Great Lakes Region, but more collection records are needed to confirm this aspect of the species’ life history. Collection dates with prey: 6 September 1969, 23 July 2011 (IL). Localities: WG (4), IL (1). Habitat: Moist deciduous woodland. Forewings with dark bands or spots, an apparent adaptation for inhabiting closed woodland with sunlit openings. Burrow excavation apparatus: Species of Dipogon have short legs for a pompilid and nest in existing tubular cavities. They use the mandibles with assistance from legs for removal of wood debris when nesting in plants. Paired tufts of bristles on the mouthparts are used for carrying various organic debris and moist soil to seal off serial cells in the tubular nest chamber with assistance from the pygidium (last dorsal abdominal segment). Figure 9. Spider (wet) body weight plotted against wasp (wet) body weight in Caliadurgus fasciatellus, Episyron biguttatus, and E. quinquenotatus. Northeastern Naturalist 21 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Host families and species: AMAUROBIIDAE – Amaurobius sp., Callobius bennetti; CORINNIDAE – Trachelas tranquillus; GNAPHOSIDAE – Gnaphosa sp.; THOMISIDAE – Misumena sp. This species is polyphagous in the eastern Great Lakes Region. Host sexes and stages: Immature, 4 (100.0%). Immature spiders were the preferred host stage. Host body lengths (mm): range = 4.0–5.0, mean = 4.75 ± 0.25, n = 4 (wasps, range = 5.5–6.0, mean = 5.88 ± 0.13, n = 4). All 4 wasps were longer than their host spiders. Amputation of legs: 0/4. References: E.R. Eaton, 2013 pers. comm.; Krombein 1979; Kurczewski 1999, pers. observ.; Kurczewski and Edwards 2012; Townes 1957; Wasbauer and Powell 1962. Dipogon (Deuteragenia) papago (Banks) Collection dates: (15; 9 June–23 August). There may be 2 generations per year in the eastern Great Lakes Region but more collection records are needed to confirm this. Collection dates with prey: 27 July 1954, 9 June 1957 (CT), 19 July 1967, 14 July 2014 (WV). Localities: WG (1), IT (1), CT (1), WV (1). Habitat: Moist deciduous woodland. Forewings with dark bands or spots, an apparent adaptation for inhabiting closed woodland with sunlit openings (see Townes 1957:plate 2). Burrow excavation apparatus: See Dipogon (Deuteragenia) calipterus. Host families and species: AMAUROBIIDAE – Callobius bennetti; CLUBIONIDAE – Clubiona canadensis Emerton; GNAPHOSIDAE – Haplodrassus hiemalis (Emerton), Sergiolus capulatus (Walckenaer) (= Sergiolus variegatus [Hentz]), Poecilochroa capulata (Walckenaer); SALTICIDAE – Paraphidippus aurantius (Lucas), Phidippus sp. Dipogon papago is polyphagous in the eastern Great Lakes Region. Host sexes and stages: Adult female (4, 100.0%). Adult females were the preferred host stage and sex. Host body length (mm): 10.0, n = 1 (wasp, 9.0, n = 1). One host spider was longer than the wasp. Amputation of legs: 1/4 (25.0%). References: Evans and Yoshimoto 1962; Krombein 1967, 1979; Kurczewski 1999, pers. observ.; F.E. Kurczewski and G.B. Edwards, unpubl. data; Kurczewski and Kurczewski 1968b; Townes 1957. Dipogon (Deuteragenia) pulchripennis (Cresson) Collection dates: (24; 17 June–27 September). There are probably 2 generations per year in the eastern Great Lakes Region with optimal weather conditions. Collection dates with prey: 17 June 1968, 22 June 1970, 14 September 2014. Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 22 Localities: IT (17), PI (1), WG (1), ON (1), Ontario (1). Habitat: Moist deciduous woodland. Forewings with dark bands or spots, an apparent adaption for inhabiting closed woodland with sunlit openings (see Townes 1957:plate 2). Burrow excavation apparatus: See Dipogon (Deuteragenia) calipterus. Host families and species: AMAUROBIIDAE – Callobius bennetti; THOMISIDAE – Xysticus sp.; SALTICIDAE – Phidippus audax, Platycryptus undatus (De Geer). Dipogon pulchripennis is polyphagous in the eastern Great Lakes Region. Host sexes and stages: Adult female, 1 (33.3%); immature, 2 (66.7%). Host body lengths (mm): range = 4.0–9.0, mean = 7.33 ± 1.67, n = 3 (wasps, range = 7.0–9.0, mean = 8.33 ± 0.67, n = 3). All 3 wasps were as long as or longer than their host spiders. Amputation of legs: 1/3 (33.3%). Amputation involved 1st and 2nd right legs of 1 spider. References: Evans and Yoshimoto 1962; Krombein 1979; Kurczewski 1999; F.E. Kurczewski and G.B. Edwards, unpubl. data; Kurczewski and Kurczewski 1972; Rau and Rau 1918. Dipogon (Deuteragenia) sayi Banks Collection dates: (120; 26 May–20 September). There are 2 generations per year in the eastern Great Lakes Region. Collection dates with prey: 25 June 1940 (NM), 6 August 1954, 30 July 1957, 29 August 1968, 7 July 2010 (PA), 19 June 2011. Localities: PI (1), IT (1), ON (1), WV (1), NM (1). Habitat: Moist deciduous woodland. Forewings with dark bands or spots, an apparent adaptation for inhabiting closed woodland with sunlit openings (see Townes 1957:plate 1). Burrow excavation apparatus: See Dipogon (Deuteragenia) calipterus. Host families and species: AGELENIDAE – Agelenopsis utahana; AMAUROBIIDAE – Callobius bennetti, Callobius sp.; GNAPHOSIDAE – Nodocion melanie Levi, Sergiolus capulatus, S. montanus (Emerton); THOMISIDAE –Bassaniana utahensis (Gertsch), B. versicolor (Keyserling), Misumena vatia (Clerck), Misumenoides formosipes (Walckenaer), Tmarus angulatus (Walckener), Xysticus bicuspis Keyserling, X. canadensis Gertsch, X. discursans Keyserling, X. elegans Keyserling, X. ferox (Hentz), X. fraternus Banks, X. funestus Keyserling, X. luctuosus (Blackwall), X. obscurus Collett, X. pellax O.P.-Cambridge, X. punctatus (Keyserling), X. triguttatus Keyserling, X. spp.; SALTICIDAE- Phidippus whitmani Peckham & Peckham. Dipogon sayi is polyphagous in the eastern Great Lakes Region provisioning its nests with 5 families of cursorial-hunting and retreatdwelling spiders including a preponderance of crab spiders (Thomisidae) of the genus Xysticus, particularly X. elegans and X. ferox. Host sexes and stages: Adult, penultimate, and immature female, 242 (97.2%); immature male, 1 (0.4%); immature, 6 (2.4%). Adult, penultimate, and immature females of small crab spider species comprised the predominant sex and stages. Northeastern Naturalist 23 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Host body lengths (mm): range = 5.0–5.5, mean = 5.15 ± 0.12, n = 4 (wasps, range = 6.5–7.5, mean = 6.88 ± 0.24, n = 4). All 4 host spiders were shorter than the wasps. Amputation: 1/5 (20.0%). One wasp bit off its host spider’s leg at a coxa-trochanter joint and fed on hemolymph that exuded from the point of amputation. References: Evans and Yoshimoto 1962; Fye 1965; Krombein 1958a, 1967, 1979; Kurczewski 1999, pers. observ.; Kurczewski and Edwards 2012, unpubl. data; Kurczewski and Kurczewski 1972; Kurczewski and Pitts 2011; Medler and Koerber 1957; K.M. O’Neill 2011, pers. comm. citing H.E. Evans’ unpublished field notes; Peckham and Peckham 1898; Townes 1957. Dipogon (Dipogon) brevis (Cresson) Collection dates: (12; 12 June–19 September). There are probably 2 generations per year in the eastern Great Lakes Region, but more collection records are needed to confirm this aspect of its life history. Collection dates with prey: 3 July 1941, 5 September 1951, 16 August 1954, 10 September 1964, 10 September 1965. Localities: PI (2), IT (1). Habitat: Moist deciduous woodland. The dark bands or spots on the forewings of other species of Dipogon (Deuteragenia) are not as obvious in D. brevis (see Townes 1957:plate 1). Burrow excavation apparatus: See Dipogon (Deuteragenia) calipterus. Host families and species: SALTICIDAE – Evarcha hoyi, Habronattus sp., Phidippus purpuratus Keyserling, Phidippus sp. Dipogon brevis is host specific on species of jumping spiders (Salticidae) based on this small sample size. Host sex and stages: Immature female, 2 (50.0%), immature, 2 (50.0%). Immature spiders are the preferred host stage. Host body lengths (mm): range = 5.0–6.0, mean = 5.50 ± 0.50, n = 2 (wasps, range = 5.0–6.0, mean = 5.50 ± 0.50, n = 2). One wasp was slightly longer than its host spider and 1 spider was slightly longer than another wasp. Amputation of legs: 0/5. References: Evans and Yoshimoto 1962; Krombein 1979; Kurczewski 1999, pes. observ.; Kurczewski and Kurczewski 1968a; Townes 1957. Tribe Ageniellini Phanagenia bombycina (Cresson) Collection dates: (14; 1 June–28 October). There are 2 generations per year in the eastern Great Lakes Region. Collection dates with prey: 28 October 1949 (KS), 6 August 1954, 10 July 1960, 29 June– 16 August 1965, 21 August 1982 (WV), 14 –20 June 2007 (FL), 9 October 2007 (AR), 10 July 2011, 13 July 2014 (WV). Localities: IT (2), ON (1), KS (3), WV (3), FL (1), AR (1). Habitat: Deciduous woodland interior and edges, sand pit, gravel quarry, rock piles, and backyard near building. Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 24 Cell construction apparatus: This species and species of Auplopus make mud cells, usually above ground, in well-concealed places protected from rain. Long, stiff bristles on mentum (mouthpart) and large clypeus of female assist in carrying mud pellets for cell construction. Oval bare pygidium is used as trowel in applying the mud to cell exterior. The first apparent abdominal segment is slender, somewhat constricted sub-basally with concave sides, allowing for considerable movement and flexibility at the juncture with the propodeum (last apparent dorsal thoracic segment) during making of cell. The dorsal edges of hind tibiae are smooth and not used in cell construction. Host families and species: LYCOSIDAE – Alopecosa aculeata (Clerck), Gladicosa gulosa, Rabidosa hentzi (Banks), Schizocosa avida, ?S. sp., Varacosa avara, Lycosidae sp.; AGELENIDAE – Agelenopsis sp.; SALTICIDAE – Maevia inclemens, Platycryptus undatus. Although at first glance seemingly polyphagous in prey selection, 22 of 28 (78.6%) P. bombycina host records are for species of wolf spiders (Lycosidae). Host sexes and stages: Adult female, 5 (45.5%); penultimate female, 2 (18.2%); immature female, 1 (9.1%); adult or penultimate male, 1 (9.1%); immature, 2 (18.2%). Most of the prey records (8/11, 72.7%) are for female spiders. Host body lengths (mm): range = 11.0–16.0, mean = 12.90 ± 0.93, n = 5 (wasps, range = 13.0–13.0, mean = 13.0, n = 5). Two host spiders were slightly longer than the wasps, while 2 wasps were slightly longer than their host spiders. Amputation of legs: All (100.0%) of the host spiders had some or all of the legs cut off at the coxa-trochanter joints to facilitate prey transport and placement of prey in the confines of the mud cells. Some spiders had only 2 front legs remaining and 1 had the first and second pairs left intact. References: Evans and Yoshimoto 1962; Krombein 1979; Kurczewski 1961b, pers. observ.; Kurczewski and Edwards 2012, unpubl. data; Kurczewski and Kurczewski 1968a; Kurczewski et al. 1987; Peckham and Peckham 1898, 1905; Savin 1924; Townes 1957. Auplopus architectus (Say) Collection dates: (58; 26 May–19 October). There are at least 2 generations per year in the eastern Great Lakes Region. Collection dates with prey: 1 June 1960, 12 October 1965 (KS), 16–17 August 1996, 17 June 2007, 30 September–9 October 2011 (VA), 19 October 2013 (VA). Localities: ON (2), IT (1), ME (1), KS (1), MA (1), VA (3). Habitat: Abandoned overgrown fields, meadows, pastures, woodland edges, and active and abandoned gravel pits. Cell construction apparatus: See Phanagenia bombycina. Host families and species: MITURGIDAE - Cheiracanthium inclusum; ANYPHAENIDAE - ?Hibana gracilis; CLUBIONIDAE – Clubiona lutescens Westring, C. sp.; CORINNIDAE – Trachelas tranquillus; THOMISIDAE – Misumenops oblongus (Keyserling); SALTICIDAE – Eris sp., Metaphidippus aeneolus (Curtis), Phidippus audax, P. princeps (Peckham & Peckham), P. whitmani, P. sp. Northeastern Naturalist 25 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 near whitmani, Phidippus sp., Tutelina similis (Banks). Auplopus architectus is polyphagous in the eastern Great Lakes Region provisioning its nests with at least 6 families of cursorial-hunting and retreat-dwelling spiders. There are several host records for species of Phidippus (Salticidae). Host sexes and stages: Adult female, 5 (55.6%); subadult female, 1 (11.1%); adult or subadult male, 1 (11.1%); immature, 2 (22.2%). Most of the host records (6/9, 66.7%) are for female spiders. Host body lengths (mm): 9, n = 1 (wasp, 9, n = 1); host wet weights (mg): range = 59–61, mean = 60.00 ± 1.00, n = 2 (wasps, range = 24.0–24.0, mean = 24.00, n = 2); spider:wasp wet-weight ratios: range = 2.46–2.54:1, mean = 2.50:1, n = 2). One wasp and its host spider were the same length, and 2 other spiders were much heavier than the wasps. Amputation of legs: Except for one example, all of the host spider’s legs were amputated at the coxa-trochanter joints to facilitate prey transport and its placement in the cell. One spider with all legs intact was collected in an early stage of prey transport, before amputation had occurred. References: Evans and Yoshimoto 1962; Krombein 1955b, 1961, 1979; Kurczewski 1961b, 1999, 2010, pers. observ.; Kurczewski and Edwards 2012, unpubl. data; Kurczewski and Kurczewski 1968a, 1987b; Kurczewski and Pitts 2011; Rau and Rau 1918; Townes 1957; Wasbauer 1982. Auplopus caerulescens (Dahlbom) Collection dates: (18; 24 May–5 October). There are 2 generations per year in the eastern Great Lakes Region. Collection dates with prey: summer 1952–1955 (KS), 11 August–7 September 1987, 12 August 1991, 8–27 July 2007, 8 July 2011. Localities: ET (10), ON (5), IT (2), AU (1), KS (2). Habitat: Abandoned overgrown fields, woodland edges, sandy field, cellar foundation, and windows of buildings. Cell construction apparatus: See Phanagenia bombycina. Host families and species: ANYPHAENIDAE – Anyphaena pectorosa L. Koch; CLUBIONIDAE – Clubiona obesa, C. sp.; CORINNIDAE – Trachelas tranquillus; THOMISIDAE – Xysticus sp.; SALTICIDAE – Eris militaris, Habronattus decorus, Maevia sp., Phidippus audax, P. sp., Platycryptus undatus, Sitticus fasciger. Auplopus caerulescens is polyphagous in the eastern Great Lakes Region preying on 5 families of cursorial-hunting and retreat-dwelling spiders including a preponderance of Salticidae. Host sexes and stages: Adult female, 2 (13.3%); penultimate female, 1 (6.7%); adult male, 1 (6.7%); penultimate male, 1 (6.7%); immature, 10 (66.7%). Most (12/15, 80%) host spiders were non-adult. Host body length (mm): 4.5, n = 1 (wasp, 6.5, n = 1); host wet weights (mg): range = 11.5–20.0, mean = 17.30, n = 11 (wasps, range = 6.5–16, mean = 13.50, n = 5); spider:wasp wet-weight ratios: range = 1.14–1.77, mean = 1.30:1, n = 5. Nearly all wasps were longer than their host spiders, but almost all spiders were heavier than the wasps. Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 26 Amputation of legs: All legs of all spiders (15/15, 100.0%) had been amputated at the coxa-trochanter joints. References: Evans and Yoshimoto 1955, 1962; Krombein 1967, 1979; Kurczewski 1989a, 2010, pers. observ.; Kurczewski and Edwards 2012, unpubl. data; Medler 1964; Townes 1957. Auplopus carbonarius (Scopoli) Auplopus carbonarius was introduced into the eastern US (Rockland and Nassau Counties, NY) before 1967 and the species has dispersed or been re-introduced into Ontario, Quebec, Michigan, and Washington since that time. Collection dates: (29; 10 June–26 October). Anoplius carbonarius has 2 generations per year in the eastern Great Lakes Region, including Ontario from which the extreme dates were obtained. Collection dates with prey: 15 July-14 August 1988 (MI), June–August 1999 (MI), 4 August 2007 (Quebec), 12 August 2007 (Ontario), 21 July 2012 (WA). Localities: MI (1), Ontario (2), Quebec (1), WA (1). Habitat: Woodland, especially near damp soil; sand dunes; abandoned overgrown field; and houses, garages, barns, stone walls, and garden s. Cell construction apparatus: See Phanagenia bombycina. Host families and species (North America): LIOCRANIDAE - ?Agroeca sp.; MITURGIDAE: Cheiracanthium mildei, C. ?mildei; SALTICIDAE – Eris militaris. Auplopus carbonarius is strongly polyphagous in Europe provisioning nests with 8–12 families of cursorial-hunting and retreat-dwelling spiders, including a preponderance of Clubionidae, depending on region, and will likely prove to be strongly polyphagous in North America based on the few available host records. Host sex and stages: Adult female, 2 (66.7%); penultimate female, 1 (33.3%). Females of small retreat-dwelling spiders were the predominant hosts. Amputation of legs: In Europe, not all host spiders had all of their legs amputated at the coxa-trochanter joints, possibly because they were collected before amputation had been completed. In North America, 2 wasps were observed in the process of cutting off some of the spider’s legs before continuing prey transport to the nest. References: Buck 2005; Day 1988; Grandi 1954, 1961; Gros and Durand 2013; F.E. Kurczewski pers. observ.; Kurczewski and Edwards 2012, unpubl. data; Kurczewski and O’Brien 1991; Nolfo 1983; Richards and Hamm 1939. Auplopus mellipes (Say) Collection dates: (91; 8 June–16 September). There are sometimes 2 generations per year in the northeastern US contingent on locality and optimal weather conditions. Collection dates with prey: 30 June 1987, 1 June 2008 (VA), 30 June–5 July 2009, 15 July 2010, 25 May 2011 (PA), 19 July–2 September 2011, 8 June–22 July 2012, 5 September 2013. Localities: ON (15), NO (2), BH (1), OL (1), PA (2), VA (1), Ontario (1). Habitat: Moist deciduous woodland sunlit interior and edges, and backyards, often in the vicinity of houses, garages, and outbuildings. Northeastern Naturalist 27 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Cell construction apparatus: See Phanagenia bombycina. Host families and species: PISAURIDAE – Pisaurina mira; MITURGIDAE – Cheiracanthium inclusum, C. mildei, C. sp.; ANYPHAENIDAE – Anyphaena sp., Hibana ?gracilis, H. ?velox (Becker); CLUBIONIDAE – Clubiona obesa, C. sp.; GNAPHOSIDAE – Herpyllus vasifer; PHILODROMIDAE – Philodromus sp.; THOMISIDAE – Misumena vatia, specimen not identified to genus or species; SALTICIDAE – Paraphidippus aurantius, Phidippus arizonensis Peckham & Peckham, P. audax, P. sp., Platycryptus undatus. Auplopus mellipes is strongly polyphagous in the eastern Great Lakes Region provisioning its nests with at least 8 families of cursorial-hunting and retreat-dwelling spiders. Host sexes and stages: Adult female, 9 (40.9%); adult male, 3 (13.6%); subadult female, 1 (4.5%); immature, 9 (40.9%). Adult female and immature spiders (18/22, 86.3%) were the predominant host sex and stage. Host body lengths (mm): range = 5.5–10.3, mean = 8.76 ± 0.27, n = 18 (wasps, range = 8.8–9.5, mean = 9.05 ± 0.06, n = 18) (Fig. 10). Two spiders each weighed 38 mg (wasps were not weighed). Most wasps (12/18, 66.7%) were longer or the same length as their host spiders (Fig. 10). Amputation of legs: All 22 (100.0%) host spiders had most or all their legs and, occasionally, 1 pedipalp amputated at the coxa-trochanter joints to facilitate prey transport and placement in the mud cells. During prey transport, the wasp straddled the paralyzed spider ventral side upward, grasped its spinnerets with her mandibles, and walked or flew forward. Two wasps interrupted cumbersome prey transport to cut off the remaining legs of their spiders. References: Evans and Yoshimoto 1962; Krombein 1952, 1955b, 1967, 1979; Kurczewski 1989a, 1999, pers. observ.; Kurczewski and Edwards 2012, unpubl. Figure 10. Spider body length plotted against wasp body length in Auplopus mellipes and A. nigrellus. Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 28 data; Kurczewski and Spofford 1986; O’Neill and O’Neill 2010; Rau 1922, 1928; Rau and Rau 1918; M.L. Schmidt, Norwood, PA, 2012 and 2013 pers. comm.; Townes 1957. Auplopus nigrellus (Banks) Collection dates: (109; 22 May–22 October). There are at least 2 generations per year in the eastern Great Lakes Region. Collection dates with prey: 6 September 1954, 24 September 1962, 23 June 1964, 26 May 1965 (KS), 23 August 1967. Localities: PI (2), AU (1), IT (1), GR (1), KS (1). Habitat: Moist deciduous woodland sunlit interior and edges. Cell construction apparatus: See Phanagenia bombycina. Host families and species: MITURGIDAE – Cheiracanthium sp.; ANYPHAENIDAE – Anyphaena fraterna (Banks), Hibana gracilis, H. incursa (Chamberlin); CLUBIONIDAE – Clubiona abboti; CORINNIDAE – Trachelas tranquillus, T. sp.; SALTICIDAE – Maevia inclemens, Phidippus spp. Auplopus nigrellus is probably strongly polyphagous in the eastern Great Lakes Region as evidenced by the variety of 5 cursorial-hunting and retreat-dwelling host spider families. Host sexes and stages: Adult female, 2 (20.0%); adult male, 1 (10.0%); immature female, 1 (10.0%); immature, 6 (60.0%). Immature spiders (7/10, 70.0%) were the preferred hosts of this spider wasp species. Host body lengths (mm): range = 5.0–7.0, mean = 6.00 ± 0.41, n = 4 (wasps, range = 6.5–7.0, mean = 6.75 ± 0.14, n = 4) (Fig.10); host wet weights (mg): range = 9–21, mean = 14.33 ± 3.53, n = 3 (wasps, range = 6–8, mean = 7.33 ± 0.67, n = 3); spider:wasp wet-weight ratios: range = 1.13–2.63:1, mean = 1.98:1, n = 3. Three of 4 (75.0%) wasps were longer than their host spiders (Fig. 10), but all spiders outweighed the wasps. Amputation of legs: Five of 8 (62.5%) host spiders had all legs amputated at the coxa-trochanter joints while 3 others (37.5%), in earlier stages of prey transport, had 4 to 7 legs cut off. References: Evans and Yoshimoto 1962; Krombein 1954, 1955b, 1979; Kurczewski 1999, pers. observ.; Kurczewski and Acciavatti 1990; Kurczewski and Edwards 2012; Kurczewski and Kurczewski 1968a, 1968b; Townes 1957; Wasbauer 1982. Ageniella (Leucophrus) fulgifrons (Cresson) Collection dates: (10; 17 July–6 September). There is a single flight period per year in mid-summer in the eastern Great Lakes Region. Collection dates with prey: 24 July 1963, 6 September 1965 (KS), 17 July 1985, 28 July 1986, 28 July 1987. Localities: WG (4), NL (1), KS (2). Habitat: Abandoned overgrown gravel pit, abandoned field with gravelly soil, and meadow. Burrow excavation apparatus: Species of Ageniella do not construct mud cells. Ageniella fulgifrons modifies existing underground (mole) burrows or cavities in soil using the mandibles assisted by hindlegs with row of weak serrations on outer edge of longitudinally concave hind tibia. Northeastern Naturalist 29 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Host family and species: SALTICIDAE – Eris militaris, Phidippus audax, P. sp. Ageniella fulgifrons is host specific on jumping spiders (Salticidae) belonging to 2 genera (Eris, Phidippus). Host sex and stages: Adult female, 3 (42.9%); immature female, 4 (57.1%). Female spiders (7/7, 100.0%) were the preferred host sex of A. fulgifrons. Host body length (mm): range = 11.0–11.0, mean = 11.00, n = 2 (wasps, range = 11.5–13.0, mean = 12.25 ± 0.75, n = 2); host wet weights (mg): range = 55–90, mean = 72.60 ± 7.50, n = 5 (wasps, range = 48–62, mean = 52.67 ± 4.67, n = 3); spider:wasp wet-weight ratios: range = 1.43–1.88:1, mean = 1.62:1, n = 3. Six wasps were longer than their host spiders, but 5 of the spiders outweighed the wasps. Amputation of legs: 7/7 (100.0%). All host spiders had all legs amputated at the coxa-trochanter joints. References: Evans 1959; Evans and Yoshimoto 1962; Krombein 1979; Kurczewski 1989a, 1999, pers. observ.; Kurczewski and Kurczewski 1968a, 1987a, 1987b. Ageniella (Leucophrus) semitincta (Banks) Collection dates: (5; 24–29 July). There is a single flight period per year in mid-summer in the eastern Great Lakes Region, whereas there probably are 2 generations per year in the southeastern US (23 May–8 September). Collection dates with prey: 24 July 1960, 23 June 1965 (KS), 29 July 1965 (KS). Localities: IT (1), KS (2). Habitat: Sandy field, abandoned overgrown sand pit, and sunlit bare red clay bank. Females explored cracks and crevices in soil, and 1 wasp had the top of her head and thorax plastered with dried red mud. Burrow excavation apparatus: Similar to Ageniella fulgifrons. Host family and species: AGELENIDAE – Agelenopsis pennsylvanica, A. potteri, A. spp. Ageniella semitincta is host specific on funnel-web-weaver spiders (Agelenidae) of the genus Agelenopsis. Host sexes and stages: Penultimate male, 1 (20.0%); immature, 4 (80.0%). Nonadult spiders (5/5, 100.0%) were the preferred host stage. Host body lengths (mm): range = 9.5–10.0, mean = 9.83 ± 0.17, n = 3 (wasps, range = 9.0–10.0, mean = 9.33 ± 0.33, n = 3); host wet weights (mg): range = 59–66, mean = 62.50 ± 3.50, n = 2 (wasps, range = 19–22, mean = 20.50 ± 1.50, n = 2; spider:wasp wet-weight ratios: range = 2.68–3.47:1, mean = 3.08:1, n = 2. Two of 3 (66.7%) spiders were longer and weighed much more than the wasps. Amputation of legs: 5/5 (100.0%). All legs of all spiders were amputated at the coxa-trochanter joints. References: Krombein 1979; Kurczewski 1961b, pers. observ.; Kurczewski and Kurczewski 1968a; Townes 1957. Ageniella (Priophanes) agenioides (Fox) Collection dates: (14; 19–27 July). There is a single flight period per year in mid-summer in the eastern Great Lakes Region, but there probably are 2 generations per year in southern Pennsylvania, Maryland, and the Washington, DC area (29 May–23 September). Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 30 Collection dates with prey: 3 October ?1954, 12 October 1964 (KS), 9 July 1965. Localities: KS (2), Hawpeg, NY (locality not found) (1). Habitat: Sandy fields and open areas near upright vegetation. Burrow excavation apparatus: Modifies existing underground burrows or cavities in soil using mandibles assisted by hindlegs with chevron-shaped teeth on outer edge of hind tibiae. Host families and species: THOMISIDAE – Xysticus sp.; SALTICIDAE – Maevia inclemens, Salticidae sp. (unidentified genus). Host spiders include species of crab spiders (Thomisidae) and jumping spiders (Salticidae). Host sexes and stages: Immature female, 1 (33.3%); immature male, 1 (33.3%); immature, 1 (33.3%). All hosts spiders were immature. Host body lengths (mm): range = 5.0–6.5, mean = 5.75 ± 0.75, n = 2 (wasps, range = 6.0–7.0, mean = 6.50 ± 0.50, n = 2; host wet weights (mg): 9, n = 1 (wasp, 9, n = 1); spider:wasp wet-weight ratio: 1.00:1, n = 1. Two wasps were longer than their host spiders, although 1 wasp and spider weighed the same. Amputation of legs: Only 1/3 (33.3%) spiders had all legs amputated at coxatrochanter joints. Another spider had 3 legs and a third spider 2 legs left intact at the time of collection. References: Krombein 1979; Kurczewski 1999, pers. observ.; Kurczewski and Kurczewski 1968a; Townes 1957. Subfamily POMPILINAE Tribe Pompilini Agenioideus (Agenioideus) cinctellus (Spinola) Agenioideus cinctellus was introduced into North America before 1997 and now occurs in southern Ontario in the eastern Great Lakes Region. Collection dates: (14; 17 July–13 September). This species may have only a single generation per year in the eastern Great Lakes Region, as in the related A. humilis (Cresson). Collection date with prey: 13 September 2004. Locality: Rockwood, Wellington County, Ontario (1). Habitat: Moist woodland and cavity in old stone wall; in Europe this species nests in natural cavities and crevices in walls and dead and decayed wood, deserted aculeate burrows, and snail shells. Burrow excavation apparatus: Legs are weakly spinose and tarsal comb is absent. Agenioideus cinctellus is thus restricted to using pre-existing cavities and crevices in various substrates. Host family and species: SALTICIDAE – Phidippus audax. In Europe A. cinctellus provisions nests mainly with species of Salticidae, rarely Thomisidae and Agelenidae. Prey transport involves grasping a front leg with the mandibles and proceeding backwards, often up a vertical substrate, as in the related A. birkmanni (Banks). Host sex and stage: Immature (1/1, 100.0%). Northeastern Naturalist 31 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Amputation of legs: 0/1. References: Buck 2005; Day 1988; Evans 1950; Evans and Yoshimoto 1962; Gros and Durand 2013; F.E. Kurczewski and G.B. Edwards, unpubl. data; Richards and Hamm 1939. Agenioideus (Agenioideus) humilis (Cresson) Collection dates: (15; 5 July–27 August). There is a single annual flight period in mid-summer in the eastern Great Lakes Region. Collection dates with prey: 28 July 2009, 20 July 2010, 5 July–27 August 2013. Localities: NF (1), Putnam County, NY (1), Suffolk County, NY (1), PA (1), MI (1), Quebec (1). Habitat: Existing cavities and crevices in sandy places; debris around cliffs, walls, and buildings; and sunny spots in open deciduous woodland. Bifasciate wings imply a preference for sunny openings in closed woodland and woodland edges. Burrow excavation apparatus: Mandibles; forebasitarsus with 3 long slender comb-spines, the apical spine on basitarsus about as long as 2nd tarsal segment. Host family and species: ARANEIDAE – Acacesia hamata (Hentz), Araneus bispinosus (Keyserling), A. ?diadematus Clerck, A. pegnia (Walckenaer), A. sp., Conepeira excelsa (Banks), Eustala anastera, Larinioides cornutus (Clerck), Zygiella x-notata (Clerck). Agenioideus humilis provisioned its nests with 4 genera and several species of orb-weaver spiders (Araneidae) in the eastern Great Lakes Region. Host sexes and stages: Adult female, 2 (33.3%); subadult or immature female, 2 (33.3%); immature, 2 (33.3%). Female spiders (4/6, 66.7%) were the preferred host sex and immatures (4/6, 66.7%) were the preferred host stage. Host body length (mm): range = 5.6–7.2, mean = 6.40 ± 0.80, n = 2 (wasps, range = 7.7–8.0, mean = 7.85 ± 0.15, n = 2). Both wasps were longer than their host spider. Amputation of legs: 0/6. References: Evans 1950, 1951b; Evans and Yoshimoto 1962; Krombein 1953c, 1979; Kurczewski 1999, pers. observ.; Kurczewski and Edwards 2012, unpubl. data; Rau 1922; Wasbauer 1982. Episyron biguttatus (Fabricius) Collection dates: (167; 26 May–20 September). There are 2 generations per year in the eastern Great Lakes Region. Collection dates with prey: 26 August 1946, 1948, 5 September 1961, 5–6 September 1967, 19 August 1969, June–August 1981, August 1986, 23 July 2005, 25 June 2006, 6 August 2007, 8 July–4 September 2008, 25 June–23 September 2010, 20 August 2011. Localities: WG (8), MR (5), GR (1), Ulster County, NY (1), CT (2), MA (2), NH (1), VT (1), ME (1), Ontario (3). Habitat: Woodland edges and open deciduous woodland on sandy and gravelly soils, exposed gardens, sand and gravel banks and pits, abandoned overgrown field, and gravelly parking lot. Burrow excavation apparatus: Mandibles; forebasitarsus with 3 comb-spines, the apical spine equal in length to 2nd tarsal segment. Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 32 Host family and species: ARANEIDAE – Acanthepeira stellata, Araneus bicentarius McCook, A. diadematus, A. gemmoides Chamberlin & Ivie, A. marmoreus, A. nordmanni (Thorell), A. spp., Argiope aurantia Lucas, A. trifasciata (Fȏrskal), Eriophora ravilla (C. L. Koch), Eustala anastera, Larinioides cornutus, L. patagiatus, Metepeira labyrintha (Hentz), Neoscona benjamina (Walckenaer), N. crucifera (Lucas), N. sp. near sacra (Walckenaer), N. spp. Episyron biguttatus provisioned its nests with 7 genera of orb-weaver spiders (Araneidae) in the eastern Great Lakes Region and in Erie County, PA (Table 2). Host sexes and stages: Adult female, 13 (48.1%); penultimate female, 2 (7.4%); subadult female, 3 (11.1%); adult or subadult female, 2 (7.4%); immature female, 4 (14.8%); adult male, 1 (3.7%); immature, 2 (7.4%). Episyron biguttatus provisioned its nests mainly with female spiders (24/27, 88.9%). Host body lengths (mm): range = 9.5–16.0, mean= 12.65 ± 0.61, n = 10 (wasps, range = 11.0–17.0, mean = 13.70 ± 0.60, n = 10) (Fig.8); host wet weights (mg): range = 126–165, mean = 140.67 ± 12.25, n = 3 (wasps, range = 54–61, mean = 57.67 ± 2.03, n = 3) (Fig. 9); spider:wasp wet-weight ratios: range = 2.07–2.84:1, mean = 2.44:1, n = 3). Nine of 10 wasps (90.0%) were longer than their host spider (Fig. 8), but all spiders were heavier than the wasps (Fig. 9). Amputation of legs: 0/27. References: Evans 1950; Evans and Yoshimoto 1962; Krombein 1953a, 1953b, 1979; Kurczewski 1962, 1999, pers. observ.; Kurczewski and Edwards 2012, unpubl. data; Kurczewski and Kurczewski 1968a, 1968b, 1973; Kurczewski and Pitts 2011; Kurczewski et al. 1987; Peckham and Peckham 1898; Rau 1922. Episyron quinquenotatus (Say) Collection dates: (654; 28 May–30 October). There are 2 or, rarely, 3 generations per year in the eastern Great Lakes Region with optimal weather conditions. Episyron quinquenotatus was, by far, the most abundant pompilid collected with host spiders in the eastern Great Lakes Region. This species often nested in aggregations of 25 or more individuals. Table 2. Host genera of orb-weaver spiders (Araneidae) preyed on by 3 species of pompilids, Caliadurgus fasciatellus, Episyron biguttatus and Episyron quinquenotatus, in Erie County, Pennsylvania. Host araneid genera are arranged in alphabetical order. Host araneid genus Caliadurgus fasciatellus Episyron biguttatus Episyron quinquenotatus Acanthepeira X X X Araneus X X X Araniella X X Argiope X X Cyclosa X X Eustala X X X Larinioides X X X Metepeira X Neoscona X X X Singa X Northeastern Naturalist 33 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Collection dates with prey: 19–26 June 1960, 28 June–5 August 1961, 28 June–12 September 1962, 6 June–12 October 1963, 13 June–16 October 1964, 5 June–27 September 1965, 6 June–25 July 1966, 13 July–15 September 1967, 14 June–16 September 1968, 12 June–22 September 1969, 9 June–6 October 1970, 16 June–28 July 1971, 19 June 1972, 11 June 1993, 10 July 2006 (Ontario). Localities: PI (377), AU (43), SS (11), CH (1), WE (4), AL (1), MR (1), Ontario (1). Habitat: Sandy soils near water courses. Females of this species are numerous on Great Lakes sand beaches, sand dunes, and dry sand plain. Burrow excavation apparatus: Mandibles; forebasitarsus with 4 or 5 slightly flattened, very long comb-spines, the apical spine considerably longer than 2nd tarsal segment. Foretarsus is adapted for excavating in coarse-grained sandy soils. Females raise and hold their wings at a 45° or greater angle when disturbed, possibly an intention movement to fly. Host family and species: ARANEIDAE - Acanthepeira stellata, Araneus cingulatus (Walckenaer), A. diadematus, A. sp. near diadematus, A. gemmoides, A. guttulatus (Walckenaer), A. marmoreus, A. montereyensis (Archer), A. ozarkensis (Archer), A. thaddeus (Hentz), A. sp., Araniella displicata, Argiope trifasciata, Cyclosa conica, Eustala anastera, E. cepina (Walckenaer), E. emertoni (Banks), Larinioides cornutus, L. patagiatus, Metepeira arizonica Chamberlin & Ivie, Neoscona arabesca (Walckenaer), N. crucifera, N. minima F.O.P.-Cambridge, Singa eugeni Levi, Zygiella x-notata. Episyron quinquenotatus provisioned its nests with 9 genera of orb-weaver spiders (Araneidae) in the eastern Great Lakes Region including Presque Isle State Park, PA (Table 2). Larinioides cornutus, L. patagiatus, Araneus diadematus, and Eustala anastera were predominant prey species at Presque Isle State Park, PA (PI), while Araniella displicata and Neoscona arabesca were prevalent host species at Auburn, NY (AU). Host sexes and stages: Adult female, 102 (65.8%); immature female, 3 (1.9%); adult male, 4 (2.6%); immature male, 5 (3.2%); immature, 41 (26.5%). The majority of host spiders were adult female. In a prior study (Kurczewski 2001), 102 of 154 (66.2%) host spiders were adult and 52 (33.8%) were immature. Of 115 sexed prey, 104 (90.4%) were female and only 11 (9.6%) were male. Size of spider was important in host selection. Adult females of the smaller to medium-size and immatures of the larger araneid species comprised commonly provisioned host spiders. Host body lengths (mm): range = 4–10, mean = 6.91 ± 0.11, n = 144 (wasps, range = 7–12, mean = 9.15 ± 0.10, n = 144) (Fig.8); host wet weights (mg): range = 16–106, mean = 44.55±20.05, n = 55 (wasps, range = 11–60, mean = 27.67±12.06, n = 55) (Fig. 9); spider:wasp wet-weight ratios: range = 0.66–5.11:1, mean = 1.64:1, n = 55. Nearly all wasps (136/144, 94.4%) were as long as their host spiders (Fig. 8), but the host spiders often (45/55, 81.8%) weighed as much or more than the wasps (Fig. 9). Large wasps tended to prey on large araneids, and small wasps preyed on small orb-weavers (Fig. 8). The largest E. quinquenotatus females often captured large spiders with little size variation. Many of the host spiders were considerably heavier than the wasps (Fig. 9). There was much variability in host-spider weight for the small to medium-size wasps (Fig. 9). The comparative body lengths Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 34 and wet weights of E. quinquenotatus and their host spiders showed a very weak positive linear relationship (r = 0.130, P = 0.125, n = 141 [Fig. 8]; r = 0.185, P = 0.175, n = 55 [Fig. 9]). At Presque Isle State Park, host spiders averaged slightly larger in June (mean = 7.41 mm) than in July–August (6.57) or September–October (6.66), signifying the use of overwintering adult females. There was a positive correlation between size of host spider and size of wasp reared from such prey at both Presque Isle State Park, PA, and Auburn, NY. At both localities, the largest spiders produced the largest wasps, and the smallest spiders yielded the smallest wasps in the next generation. Amputation of legs: 6/154 (3.9%) had 1 (1), 3 (1), or 4 (2) legs or 2 legs and a pedipalp (2) cut off at the coxa-trochanter joint(s). Three amputations of 3 or 4 legs were on the same side of the spider’s body. References: Evans 1950, 1963, 1970; Evans and Yoshimoto 1955, 1962; Krombein 1958b, 1961, 1979; Kurczewski 1961b, 1962, 1999, 2001, pers. observ.; Kurczewski and Acciavatti 1990; Kurczewski and Edwards 2012, unpubl. data; Kurczewski and Kurczewski 1968a, 1968b, 1973; Peckham and Peckham 1898, 1905; Wasbauer 1982. Poecilopompilus interruptus (Say) Collection dates: (17; 5 July–5 October). There is a single flight period per year in mid-late summer in the eastern Great Lakes Region. Collection dates with prey: 5 July ?1896 (WI), 26 August 1906 (KS), 1948, 8 September 1962, 5 October 1965 (KS). Localities: KS (4), AU (1), plus 20 host records from the eastern and midwestern US for the nominate subspecies. Habitat: Fields with bare sandy and gravelly openings, and edges of sand pits and gravel pits near woodland. Burrow excavation apparatus: Mandibles; forebasitarsus with 4 somewhat short comb-spines, the apical spine up to ¾ length of 2nd tarsal segment. Females raise and hold their wings at a 45° or greater angle when disturbed, possibly an intention movement to fly. Host family and species: ARANEIDAE – Acanthepeira stellata, Araneus trifolium (Hentz), Argiope aurantia, A. trifasciata, Eriophora ravilla, Eustala anastera, Larinioides cornutus, Neoscona benjamina, N. crucifera, N. vertebrata McCook, N. sp. Poecilopompilus interruptus was host specific on 5 genera of orb-weaver spiders (Araneidae) in the eastern Great Lakes Region. Host sexes and stages: Adult female, 21 (72.4%); subadult female, 3 (10.3%); immature female, 3 (10.3%); adult male, 1 (3.4%); immature, 1 (3.4%). Poecilopompilus interruptus preyed mainly on adult female spiders. Host body lengths (mm): range = 13–17, mean = 15.00 ± 2.00, n = 2 (wasps, range = 13.5–14.0, mean = 13.75 ± 0.25, n = 2); host wet weights (mg): 189, n = 1 (wasp, 98, n = 1); spider:wasp wet-weight ratio: 1.93:1, n = 1. One wasp was longer than its host spider, and another host spider was longer than the wasp. The single weighed host spider was somewhat heavier than the wasp. Amputation of legs: 0/29. Northeastern Naturalist 35 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 References: Bugbee 1939; Evans 1950; Evans and Yoshimoto 1955, 1962; Krombein 1953a, 1979; F.E. Kurczewski pers. observ.; Kurczewski and Edwards 2012, unpubl. data; Kurczewski and Kurczewski 1968a, 1987b; Peckham and Peckham 1898; Strandtmann 1953. Tachypompilus ferrugineus (Say) Collection dates: (91; 23 June–3 August [IT]; 16 July–27 September [PI]). This species has a single flight period per year in mid-late summer in the eastern Great Lakes Region. The 23 June record from Ithaca, NY (IT) is exceptionally early for T. ferrugineus. Collection dates with prey: 3 August (no year), 23 July–19 August 1988, 10 August– 5 September 1989, 19 August 1992, 14 July (WV)–18 September 2005 (NJ), 30 August–27 September 2006 (IL), 20 August 2007 (Ohio), 17 July–14 September 2008 (DE), 16 July (PA)–1 August 2009, 20 August 2011 (MD), 31 August 2012, 24 July 2013 (NC), 7 August 2013 (TX), 31 August 2013 (PA). Localities: OC (9), JA (1), DE (1), PA (2), plus 52 online host records from the eastern and midwestern US. Habitat: Rock piles; crevices and openings in stone walls and stone and wooden buildings; and inside and beneath man-made structures in sandy, gravelly, and loamy soils, and artificial composite. Burrow excavation apparatus: Mandibles; raised tubercles above antenna bases; 4 or 5 comb-spines on slender forebasitarsus, the individual spines 3 times as long as tarsal width. The resulting prey-deposition chamber is a shallow concave depression in the soil or powdered substrate material. Host families and species: LYCOSIDAE – Hogna ?antelucana, H. carolinensis (Walckenear), H. helluo, H. lenta (Hentz), H. osceola (Wallace), H. spp., Isohogna timuqua (Wallace), Lycosa sp. near impavida Walckenaer, Rabidosa punctulata, R. rabida, Lycosidae spp.; PISAURIDAE – Dolomedes albineus, D. scriptus Hentz, D. tenebrosus; CTENIDAE – Cupiennius salei (Keyserling), unidentified species. Tachypompilus ferrugineus preys on wolf spiders (Lycosidae) and fishing spiders (Pisauridae) in the eastern US and adds wandering spiders (Ctenidae) in the tropics and subtropics. Hosts in the eastern Great Lakes Region included Hogna spp., Rabidosa rabida, and Dolomedes tenebrosus away from water. Dolomedes albineus, an arboreal fishing spider, was a common host species in the southeastern US. Host sexes and stages: Adult female, 56 (87.5%); penultimate female, 1 (1.6%); subadult or immature female, 5 (7.8%); immature, 2 (3.1%). Adult females were the predominant host stage and sex. Male spiders were not reported as hosts. Host body lengths (mm): range = 17.0–22.5, mean = 19.24 ± 0.46, n = 12 (wasps, range = 14.0–19.0, mean = 17.08 ± 0.48, n = 12) (Fig. 3); host wet weights (mg): range = 179–1167, mean = 492.45 ± 101.133, n = 11 (wasps, range = 78–278, mean = 197.50 ± 44.20, n = 4); spider:wasp wet-weight ratios: range = 2.30–4.71:1, mean = 3.73:1, n = 4. The host spider was longer or as long and weighed more than the wasp in all measured examples (r = 0.412, P = 0.208; n = 11; Fig. 3). Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 36 Amputation: 1/15 (0.7%). Front left leg cut off at coxa-trochanter joint of 1 specimen. References: N. Elhardt, 2013 pers. comm.; Evans 1950, 1951b; Evans and Yoshimoto 1962; Krombein 1979; Kurczewski 1981, 1989b, 1990, 1999, 2010, pers. observ.; Kurczewski and Edwards 2012, unpubl. data; Kurczewski and Kurczewski 1968a, 1973; Rau 1922; Rau and Rau 1918; Strandtmann 1953; Wasbauer 1982; Wilson and Pitts 2007. Anoplius (Lophopompilus) aethiops (Cresson) Collection dates: (84; 27 July–8 November). Adults fly annually in mid-summer– early fall. There is a single generation per year in the eastern Great Lakes Region but 2 or more generations per year farther southward. Collection dates with prey: July 1951, 27 August 1956, 12 September 1958, 12 August 1962, 2 September 1962, 8–10 September 1963, 18 September 1964, 28 August 1966, 5 October 1967, 29 August–20 September 1969, 5–6 September 1972, 29 August 1973, September–October 1978, 8 October 2011, 1 October 2012. Localities: PI (6), MR (4), LU (2), IT (2), OC (1), ON (1), JA (1), BE (1), WA (1), CT (1), MA (1), NJ (2). Habitat: Abandoned overgrown fields and deciduous woodland edges where the soil is fine-grained or gravelly with cavities and crevices. Anoplius aethiops females do not inhabit loose sand because they need a firm substrate with existing holes in which to nest. Burrow excavation apparatus: Mandibles; 3 or, rarely, 4 comb-spines on forebasitarsus, the apical spine ½ the length of 2nd tarsal segment. Foretarsus is adapted for excavating in non-sandy soils, according to the short foretarsal digging comb-spines. Host family and species: LYCOSIDAE - Gladicosa gulosa, Hogna annexa (Chamberlin & Ivie), H. aspersa, H. baltimoriana (Keyserling), H. carolinensis, H. frondicola, H. helluo, H. ?helluo, H. spp., Schizocosa ocreata (Hentz). All host records are for cursorial-hunting and temporary burrowing genera of Lycosidae, especially species of Hogna. Hogna helluo, H. aspersa, and H. frondicola were prevalent host species (16/19, 84.2%) in the eastern Great Lakes Region. Host sexes and stages: Adult female, 17 (81.0%); adult male, 3 (14.3%); immature, 1 (4.8%). Adult females were the predominant host stage and sex. Host body lengths (mm): range = 10–21, mean = 16.79 ± 1.01, n = 14 (wasps, range = 13–19, mean = 16.96 ± 0.60, n = 14) (Fig. 3); host wet weights (mg): range = 124–1209, mean = 620.75 ± 227.28, n = 4 (wasps, range = 75–197, mean = 166.50 ± 30.500, n = 4); spider:wasp wet-weight ratios: range = 1.65–6.14:1, mean = 3.33:1, n = 4. The majority of spiders were longer or the same length (10/14, 71.4%; r = 0.664, P = 0.010, Fig. 3) and heavier than the wasps. Amputation of legs: 2/21 (9.5%) spiders had a hind leg cut off at a coxa-trochanter joint. References: Evans 1951a; Evans and Yoshimoto 1962; Krombein 1979; Kurczewski 1999, pers. observ.; Kurczewski and Acciavatti 1990; Kurczewski and Edwards 2012, unpubl. data; Kurczewski and Kurczewski 1968a, 1968b, 1973; Kurczewski and Pitts 2011; Kurczewski et al. 1987. Northeastern Naturalist 37 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Anoplius (Lophopompilus) atrox (Dahlbom) Collection dates: (24; 18 July–28 September). There is a single flight period per year in mid-summer–early fall in the eastern Great Lakes Region but 2 or more generations per year farther southward. Collection dates with prey: 23 June 2004 (VA), 8 July 2006 (AR), 25 August 2012. Localities: ON (1), AR (1), VA (1). Habitat: Abandoned overgrown fields, meadows, edge of gravel pit, gravelly parking lot, and deciduous woodland edge. Anoplius atrox females usually do not inhabit loose sand because they need a firm substrate with existing burrows, cavities and crevices in which to shape a nest. Burrow excavation apparatus: Mandibles; forebasitarus with comb-spines 1½ times as long as thickness of tarsus; 3 comb-spines on forebasitarsus, the apical one ½ as long as 2nd tarsal segment. This species probably modifies existing holes in soil for use as nests, as indicated by its short foretarsal digging comb-spines. Host families and species: LYCOSIDAE – Hogna sp., Rabidosa rabida; PISAURIDAE – Dolomedes scriptus, D. tenebrosus, D. vittatus Walckenaer, D. sp. Anoplius atrox provisioned its nests with wolf spiders (Lycosidae) and fishing spiders (Pisauridae). Dolomedes tenebrosus frequently inhabits deciduous woodland, away from water. Host sexes and stages: Adult female, 1 (50.0%), immature, 1 (50.0%). Host body length (mm): 18.5, n = 1 (wasp, 20.0, n = 1). Amputation of legs: 0/6. References: Evans 1951a; Evans and Yoshimoto 1962; Krombein 1955a, 1959, 1979; Kurczewski 1999, pers. observ.; Kurczewski and Edwards 2012, unpubl. data; Peckham and Peckham 1900; Rau and Rau 1918. Anoplius (Lophopompilus) carolina (Banks) Collection dates: (17; 27 June–28 August). There is 1 flight period per year in mid-summer in the eastern Great Lakes Region. Collection dates with prey: 28 August 1956, 5 August 1968, 18 July 2008. Localities: IT (8), PI (1), WV (2). Habitat: Sunny openings in moist deciduous woodland. Burrow excavation apparatus: Mandibles; tarsal comb scarcely longer than width of tarsus, forebasitarus with 3 comb-spines, the apical one about 1/3 length of 2nd tarsal segment. Anoplius carolina nests in mammal burrows or openings in bare, loamy soil in deciduous woodland. Host family and species: AMAUROBIIDAE – Callobius bennetti, ?Coras sp., Wadotes hybridus, W. sp. The predominant host species in the eastern Great Lakes and Finger Lakes Regions was Wadotes hybridus (7/9 examples, 77.8%). Host sexes and stages: Adult female, 1 (25.0%); immature female, 1 (25.0%); adult male, 2 (50.0%). Male and female spiders are both selected as hosts. Host body lengths (mm): range = 9–13, mean = 11.67 ± 1.33, n = 3 (wasps, range = 8–12, mean = 10.33 ± 1.20, n = 3). Two host spiders were longer and heavier than the wasps. Amputation of legs: 0/10. Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 38 References: Evans 1951a; Evans and Yoshimoto 1962; Krombein 1958a, 1961, 1979; Kurczewski 1999, pers. observ.; Kurczewski and Edwards unpubl. data; Kurczewski and Kurczewski 1973. Anoplius (Lophopompilus) cleora (Banks) Collection dates: (136; 6 July–31 October). There is 1 or sometimes 2 generations per year in the eastern Great Lakes Region depending on locality and weather conditions. Collection dates with prey: 10 July 1954, 21 August 1962, 22 July–15 October 1964, 21 September 1965, 29 August 1966, 1 August–16 October 1967, 4 September– 14 October 1968, 9 July–10 October 1969, 21 September 1970, August 1990, 27 June 2007 (MA), 19 August–17 September 2014. Localities: PI (43), SS (13), SB (3), CT (1), MA (2). Habitat: Bare sandy soils near water courses, especially sand beaches along the Great Lakes. Burrow excavation apparatus: Mandibles; forebasitarsus with 4 or, rarely, 5 long comb-spines, the apical spine equal to length of 2nd tarsal segment. Foretarsus is adapted for excavating in coarse-grained sandy soils. Host family and species: LYCOSIDAE - Arctosa littoralis, Hogna helluo, Trochosa terricola, Varacosa avara. Nearly all host records (57/59, 96.6%) from the eastern Great Lakes Region (PI, SS, SB) were for the shoreline sand spider Arctosa littoralis. There is no record of Anoplius cleora preying on Geolycosa wrighti, a burrowing wolf spider abundant on the sand beach, sand dunes, and dry sand plain at Presque Isle State Park, PA. Host sexes and stages: Adult female, 13 (30.2%); penultimate female, 2 (4.7%); immature female, 4 (9.3%); adult male, 8 (18.6%); immature male, 5 (11.6%); immature, 11 (25.6%). Adult female and unsexed immature were the predominant host sex and stages from Presque Isle State Park, PA. There was distinct seasonality in the sexes and stages of the host spiders: July–August, 12 immature (85.7%), 2 adult females (14.3%); September–October, 8 immature (27.6%), 8 adult males (27.6%), and 13 adult and penultimate females (44.8%); and, by month: July, 2 immature; August, 2 adult females, 10 immature; September, 8 adult and penultimate females, 5 adult males, 7 immature; October, 5 adult and penultimate females, 3 adult males, 1 immature. Host body lengths (mm): range = 8–18, mean = 12.35 ± 0.35, n = 43 (wasps, range = 12–18, mean = 14.67 ± 0.28, n = 43) (Fig. 7). Most wasps (34/43, 79.1%) were longer than their host spiders (r = 0.054, P = 0.730, n = 43, Fig. 7). As wasp body length increased, wasps 15 mm and longer tended to prey on spiders of many sizes (8–17 mm), not just large spiders (Fig. 7). Mean body length (mm) of host Arctosa littoralis females was significantly longer (13.76 ± 0.44) than that of host A. littoralis males (11.54 ± 0.58) or immatures (10.96 ± 0.43) (P < 0.001) (Fig. 11). Mean body length (mm) of the wasps associated with the different host sexes and stages was not statistically different (P = 0.699): 14.94 ± 0.32, 14.46 ± 0.49, 14.65 ± 0.45, respectively (Fig. 11). Although September and October A. littoralis hosts were noticeably larger than July–August A. littoralis hosts, there was no Northeastern Naturalist 39 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 statistically significant difference (P = 0.139) in mean body length (mm) between any of these months (July–August: 11.67 ± 0.47, range = 9–16, n = 15; September: 12.47 ± 0.57, range = 8–17, n = 19; October: 13.56 ± 0.73, range = 11.0–17.5, n = 9; Fig. 12). There was a statistically significant difference in A. cleora female mean body length (mm) between July–August and October (P = 0.042) but not July–August and September or September and October (July–August: 14.33 ± 0.39, Figure 11. Spider body length (Arctosa littoralis) plotted against wasp body length (Anoplius cleora) according to spider stage and sex. Figure 12. Spider body length (Arctosa littoralis) plotted against Anoplius cleora month of capture. Ranges are shown as vertical lines, and means as widened portion of lines; n = number of observations. Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 40 range = 12–17, n = 15; September: 14.68 ± 0.29, range = 12–17, n = 19; October: 15.89 ± 0.56, range = 14–18, n = 9). Amputation of legs: 0/59. References: Evans 1951a; Evans and Yoshimoto 1955, 1962; Krombein 1953b, 1979; Kurczewski 1999, pers. observ.; Kurczewski and Edwards 2012, unpubl. data; Kurczewski and Kurczewski 1968a, 1968b, 1973, 1987b. Anoplius (Arachnophroctonus) apiculatus (Smith) Collection dates: (262; 8 June–6 November). Anoplius apiculatus has 2 generations per year in the eastern Great Lakes Region. Collection dates with prey: 28 July 1961, 30 August–1 September 1962, 1 July 1963, 3 July–15 October 1964, 19 June–20 October 1965, 29 August 1966, 2 September–31 October 1967, 24 June–17 October 1968, 3 July–10 October 1969, 28 July–29 October 1970, 6–16 August 1971, 17 September 2014. Localities: PI (173), SB (3). Habitat: Bare sandy soils, especially sand beaches along water courses. This species is abundant along the shores of the eastern Great Lakes. Burrow excavation apparatus: Mandibles; forebasitarsus with 3 long, flattened comb spines, each about 2½ times tarsal width, the apical one longer than 2nd tarsal segment. Foretarsus is adapted for excavating in coarse-grained sandy soils. Host family and species: LYCOSIDAE - Arctosa littoralis. All 173 host records from a single locality (PI) were for the shoreline sand spider A. littoralis. There is no record of A. apiculatus capturing Geolycosa wrighti, a burrowing wolf spider abundant on the sand beach, sand dunes, and dry sand plain at Presque Isle State Park, PA. Host sexes and stages: Adult female, 3 (1.7%); immature female, 5 (2.9%); adult male, 2 (1.2%); immature male, 10 (5.8%); immature, 153 (88.4%). The vast majority of host spiders (168/173, 97.1%) were immature. Host body lengths (mm): range = 3–14, mean = 7.30 ± 0.12, n = 167 (wasps, range = 6–15, mean = 9.49 ± 0.12, n = 167) (Fig. 7); host wet weights (mg): range = 11–138, mean = 63.29 ± 16.84, n = 7 (wasps, range = 10–30, mean = 19.43 ± 2.94, n = 7); spider:wasp wet-weight ratios: range = 0.85–5.75:1, mean = 3.26:1, n = 7. The vast majority of wasps were longer (147/167, 88.0%) than the host spiders (r = 0.424, P < 0.001, Fig. 7), but weighed less. Some small spiders (less than 5 mm) were captured, immobilized by stinging, abandoned, and not used as prey, though the wasps fed on their hemolymph. Anoplius apiculatus does not visit flowers for nectar. Amputation of legs: 7/167 (4.2%) spiders had 1 (4), 2 (2) or 4 (1) legs cut off at the coxa-trochanter joints. One female (13 mg) stung her prey (11 mg), amputated its left hindleg, and fed on hemolymph exuding from the point of amputation. In the case of more than 1 leg, such amputations were on the same side of the host spider’s body. Spider size was not a determinant for leg amputation as amputated individuals ranged in length from 3.5 to 9.0 mm. References: Evans 1951a; Evans et al. 1953; Evans and Yoshimoto 1955, 1962; Krombein 1953a, 1979; Kurczewski 1999, pers. observ.; Kurczewski and Edwards, unpubl. data; Kurczewski and Kurczewski 1968a, 1968b, 1973. Northeastern Naturalist 41 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Anoplius (Arachnophroctonus) nigritus (Dahlbom) [Most information on this species was gathered under the name Anoplius (Arachnophroctonus) relativus (Fox).] Collection dates: (53; 15 June–21 August). This species has 1 flight period per year mainly in early–mid-summer in the eastern Great Lakes Region, often coinciding with the appropriate sex, size, and stage of its host spider, but 2 or more generations per year (9 April – 3 December) farther south- and westward. Collection dates with prey: 12 October 1964 (KS), 7 July 1965 (KS), 2 October 1965 (KS), 7 August 1968, 26 June–9 August 1969, 22 June–10 July 1970, 15–16 June 2006 (NJ), 30 June 2006 (IL), 6 August 2010 (VA). Localities: PI (16), WE (1), KS (3), NJ (8), CT (1), IL (1), VA (1). Habitat: Sandy and gravelly soils near water courses, sand beaches, sand dunes, gravel pit, inland sand blowouts, sandy fields, and fine-grained soils of open fields, gardens, and prairies. Burrow excavation apparatus: Mandibles; forebasitarsus with 3 rather long comb-spines, the spines 2¼ times tarsal thickness. Foretarsus is adapted for excavating in sandy, gravelly, or loamy soils. Host families and species: LYCOSIDAE – Arctosa littoralis, Geolycosa domifex (Hancock), G. missouriensis (Banks), G. ?turricola (Treat), G. wrighti, G. spp., Hogna ?antelucana, H. lenta, Hogna spp., Rabidosa rabida, Schizocosa avida, S. sp.; AGELENIDAE - Agelenopsis naevia, A. pennsylvanica. The burrowing lycosids G. domifex, G. wrighti and G. ?turricola were exclusive prey at Uxbridge, Ontario; Presque Isle State Park, PA (16/16 host records, 100.0%); and Vineland, Cumberland County, NJ (8/8, 100.0%), respectively. The wasps modified and used the spiders’ burrows for nests at all 3 localities. Elsewhere in the eastern US, A. nigritus females were variable in nesting behavior, excavating their own burrows from the ground surface (Evans 1951a, Kurczewski and Edwards 2012, Kurczewski and Kurczewski 1968a) or, when preying on Geolycosa spp., modifying and using the spiders’ burrows as nests (Evans and Yoshimoto 1962; F.E. Kurczewski, pers. observ.; Kurczewski and Kurczewski 1973; McQueen 1978). Although there are 2 such host records from north-central Texas (Kurczewski 1975), A. nigritus did not prey on Arctosa littoralis, a rather large sometimes burrowing wolf spider abundant on the sand beach, sand dunes, and dry sand plain at Presque Isle State Park, PA. The hunting behavior of A. nigritus females at this location must, therefore, differ from those of A. apiculatus and A. cleora that capture A. littoralis exclusively or nearly so. Cursorial-hunting and temporary burrowing lycosids and funnel-webweaver spiders (Agelenopsis spp.) comprised hosts of A. nigritus in other sections of the eastern and midwestern US (Evans and Yoshimoto 1962, Kurczewski and Edwards 2012, Kurczewski and Kurczewski 1968a). Host sexes and stages: Adult female, 13 (52.0%); penultimate female, 5 (20.0%); subadult female, 3 (12.0%); penultimate male, 1 (4.0%); immature, 3 (12.0%). Nearly all (21/25, 84.0%) host spiders were female. Host spiders consisted almost exclusively of adult, penultimate, and subadult females of G. wrighti at Presque Isle State Park (13/16; 81.3%) and G. ?domifex at Vineland, NJ (8/8; Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 42 100.0%). All prey (100.0%, n > 100) at Uxbridge, ON, were adult reproductive (75.0%) or non-reproductive (25.0%) females of G. domifex (McQueen 1978). In Ontario, nearly all adult female spiders (99.0%) in the aggregation were preyed on, but they were captured only after they had reproduced, thus having a minimal effect on population numbers (McQueen 1978). Geolycosa males were rarely preyed on (1 record) probably because they were not of suitable host size and biomass to enable successful development of the wasp larva. At Presque Isle State Park, PA, and Southwick Beach State Park, NY, males of G. wrighti were exposed on the sand surface searching for and courting adult females in September–October, well after the wasps had disappeared for the year. This timing may be another reason why they escaped capture by the wasps. Host body lengths (mm) (all species): range = 9–18, mean = 15.14 ± 0.48, n = 22 (wasps, range = 12–17, mean = 15.09 ± 0.29, n = 22; host wet weights (mg): 170–501, mean = 292.17 ± 51.01, n = 6 (wasps, range = 56–224, mean = 114.67 ± 25.76, n = 6); spider:wasp wet-weight ratios: 1.70–4.73:1, mean = 2.55:1, n = 6. Half of the host spiders (10/20, 50.0%) were longer than the wasps (r = 0.625, P = 0.002). Host body lengths (mm) (Geolycosa wrighti): range = 9–18, mean = 14.83 ± 0.64, n = 15 (wasps, range = 13.0–16.5, mean = 15.07 ± 0.32, n = 15 (Fig. 13); host wet weight (mg): 208, n = 1 (wasp, 74, n = 1); spider:wasp wet-weight ratio: 2.81:1, n = 1. Ten of 15 (66.7%) host G. wrighti at Presque Isle State Park, PA were as long as the wasp (Fig. 13) and all spiders weighed much more than the wasps. Most of the G. domifex reproductive females captured by A. nigritus at Uxbridge, ON, weighed 350–600 mg (McQueen 1978). A rather large A. nigritus (142 mg) was unsuccessful in attempting to capture a Geolycosa sp. (601 mg, spider: wasp wet-weight ratio, 4.23:1) inside its burrow entrance in southern Florida (Kurczewski and Kurczewski 1968b). Figure 13. Spider body length (Geolycosa wrighti) plotted against wasp body length in Anoplius nigritus, A. cylindricus, A. marginatus species-complex, and A. splendens. Northeastern Naturalist 43 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Amputation of legs: 0/22. References: Evans 1951a; Evans and Yoshimoto 1955, 1962; Krombein 1979; Kurczewski 1975, 1999, pers. observ.; Kurczewski and Edwards unpubl. data; Kurczewski and Kurczewski 1968a, 1968b, 1973, 1987b; Kurczewski and Pitts 2011; McQueen 1978. Anoplius (Arachnophroctonus) semicinctus (Dahlbom) [Most information on A. semicinctus was gathered under the name Anoplius (Arachnophroctonus) marginalis (Banks).] Collection dates: (73; 22 June–5 September). There is a single flight period per year, usually in mid-summer in the eastern Great Lakes Region, coinciding with appropriate host spider sex, size, and stage. Collection dates with prey: 16 July 1952, 7 July 1953, 28 June 1959, 1961, 18 April 1963, 9 April 1971 (FL), April 1973 (FL), 11–25 July 2004 (MA), 5 September 2005 (MA). Localities: PI (1), SB (1), MA (3), KS (4), CO (15), FL (3). Habitat: Sand beaches, sand dunes, sand plain, abandoned overgrown fields with bare openings and friable soil, and cornfields. Burrow excavation apparatus: Mandibles; forebasitarsus usually with 4 combspines, the spines up to 2 times as long as tarsal width. Foretarsus is adapted for excavating in coarse-grained sandy or gravelly soils. Host family and species: LYCOSIDAE – Arctosa littoralis, Geolycosa missouriensis, G. pikei (Marx), G. rafaelana (Chamberlin), G. wrighti, G. spp., Hogna baltimoriana, H. carolinensis, H. frondicola, H. helluo, H. sp. near ceratiola Gertsch, H. sp. near lenta, H. sp., Lycosa sp. near impavida, Rabidosa rabida, Schizocosa avida, S. sp. Anoplius semicinctus preyed exclusively on adult or subadult females of Geolycosa wrighti (7 records) or G. rafaelana (8) in Colorado (Gwynne 1979). The wasps used the spiders’ burrows for nests and excavated side burrows midway down, each terminating in a round or ovoid cell to hold the paralyzed prey. A Phidippus princeps (Salticidae) was paralyzed and dragged backwards on the ground, but not taken into the wasp’s burrow (R.K. Walton, 2012 pers. comm.). A Habronattus coronatus (Hentz) (Salticidae), captured by a small female of this species, was abandoned on the ground and not placed in a nest (Rau and Rau 1918). Host sexes and stages: Adult, penultimate, subadult, and immature female, 14 (70.0%); adult and subadult male, 2 (10.0%); immature, 4 (20.0%). Adult, penultimate, subadult, and immature females were the preferred host stages and sex. Host body lengths (mm): range = 14.0–18.5, mean = 16.38 ± 1.11, n = 4 (wasps, range = 12.5–16.5, mean = 14.88 ± 0.85, n = 4); host wet weights (mg): range = 401–501, mean = 451.0 ± 50.0, n = 2 (wasps, range = 99–154, mean = 126.5 ± 27.5, n = 2); spider:wasp wet-weight ratios: range = 3.25–4.05:1, mean = 3.65:1, n =2. All 4 host spiders were as long as and considerably heavier than the wasps. Amputation of legs: 0/20. References: Evans 1951a; Evans and Yoshimoto 1955, 1962; Gwynne 1979; Krombein 1953a, 1953b, 1958b, 1964, 1979; Kurczewski 1962, 1981, 1999, pers. Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 44 observ.; Kurczewski and Edwards 2012; Kurczewski and Kurczewski 1973; Kurczewski et al. 1987; Rau and Rau 1918. Anoplius (Arachnophroctonus) semirufus (Cresson) Collection dates: (338; 22 May–14 October). There are 2 generations per year in the eastern Great Lakes Region. Collection dates with prey: 26 July 1961, 1 August–23 September 1962, 17 July–16 September 1964, 16 May 1965 (KS), 5 June–19 September 1965, 6 June–6 August 1966, 23 August–14 September 1967, 24 June–16 September 1968, 25 June–20 September 1969, 19 July–6 October 1970, 5 July–2 September 1971, June–September 1973, 1974, 1975, 1981, 1982. Localities: PI (120), AU (7), IT (3), WG (2), MC (1), CH (1), PS (1), KS (1). Habitat: Bare areas of sandy soils near or just inside open woodland, sand plain, sandy soils near water courses, sandy firetrail, inland sand blowouts, and sand and gravel pits. Burrow excavation apparatus: Mandibles; forebasitarsus with 3 long, slender comb-spines up to 3 times width of tarsus. Foretarsus is adapted for excavating in coarse-grained sandy soils. Host families and species: LYCOSIDAE - Gladicosa gulosa, Hogna helluo species group, Pardosa distincta, P. milvina, P. sp. probably milvina, P. aff. modica (Blackwall), P. moesta, P. xerampelina, P. sp. near floridana Banks, P. sp., Pirata insularis Emerton, P. sp., Rabidosa punctulata, R. rabida, Schizocosa avida, S. bilineata (Emerton), S. crassipalpata, S. crassipes, S. ocreata, S. saltatrix (Hentz), S. sp., Trochosa abdita (Gertsch), T. ruricola, T. terricola, Varacosa avara, V. parthenus (Chamberlin); AGELENIDAE - Agelenopsis pennsylvanica, A. sp.; AMAUROBIIDAE - Callobius sp. probably bennetti. Anoplius semirufus usually captured cursorial-hunting lycosids (120/144, 83.3%), occasionally funnel-webweaver agelenids (23/144, 16.0%), and, rarely, hacklemesh-weaver amaurobiids (1/144, 0.7%) at Presque Isle State Park, PA. Varacosa avara (33/120, 27.5%), Agelenopsis sp. probably pennsylvanica (23/120, 19.2%), Schizocosa saltatrix (13/120, 10.8%), and S. crassipes (12/120, 10.0%) were the most captured host spiders. Anoplius semirufus did not provision its nests with Arctosa littoralis or Geolycosa wrighti, 2 wolf spiders abundant on the sand beach, sand dunes, and dry sand plain at Presque Isle State Park, PA., implying that it either did not hunt in these habitats or did not search in burrows for its hosts. Host sexes and stages: Adult female, 59 (43.4%); penultimate female, 1 (0.7%); immature female, 10 (7.4%); adult male, 4 (2.9%); penultimate male, 3 (2.2%); immature male, 6 (4.4%); immature, 53 (39.0%). Adult females of smaller species and unsexed immatures of larger species were predominant (112/136, 82.4%) stages and sex of the host spiders. Host body lengths (mm): range = 5–10, mean = 7.80 ± 0.13, n = 114 (wasps, range = 5–11, mean = 8.30 ± 0.11, n = 114) (Fig. 14); host wet weights (mg): range = 10–97, mean = 43.68 ± 5.67, n = 19 (wasps, range = 6–46, mean = 19.08 ± 2.52, n = 19) (Fig. 15); spider:wasp wet-weight ratios: range = 0.91–3.77:1, mean = 2.29:1, n = 19. One-half of the host spiders were longer or the same length (57/114, Northeastern Naturalist 45 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 50.0%) as the wasps (r = 0.436, P < 0.001) (Fig. 14), and most of the prey (17/19, 89.5%) weighed more than the wasps (Fig. 15). Amputation of legs: 13/125 (10.4%) spiders had 1 (11) or 2 (2) legs amputated at the coxa-trochanter joints. In the case of 2 amputated legs, both were cut off the same side of the spider. Figure 14. Spider body length plotted against wasp body length in Anoplius semirufus and A. marginatus species-complex. Figure 15. Spider (wet) body weight plotted against wasp (wet) body weight in Anoplius semirufus, A. marginatus species-complex, and A. splendens. Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 46 References: Evans 1951a; Evans and Yoshimoto 1955, 1962; Krombein 1953a, 1953b, 1958b, 1959, 1979; Kurczewski 1962, 1999, pers. observ.; Kurczewski and Acciavatti 1990; Kurczewski and Edwards 2012; Kurczewski and Kurczewski 1968a, 1968b, 1973; Kurczewski and Pitts 2011; Kurczewski et al. 1987. Anoplius (Pompilinus) cylindricus (Cresson) Collection dates: (75, 13 June–17 September, East Hartford, CT; 30 June–11 September [PI]). There are probably 2 generations per year in the northeastern US. Collection dates with prey: 29 August 1964, 6–11 September 1967, 30 June 1970, 10 March–2 May 1973 (FL). Localities: PI (6), KS (1), FL (4). Habitat: Sand dunes, sand plain, sand ridge, and inland sand blowout. This species is strongly psammophilous throughout its range. Burrow excavation apparatus: Mandibles; forebasitarsus with 3 or 4 stout and rather lengthy comb-spines, often 2 times as long as tarsal thickness. The foretarsi are adapted for excavating in coarse-grained sandy soils. Host family and species: LYCOSIDAE – Geolycosa hubbelli Wallace, G. micanopy Wallace, G. rafaelana, G. wrighti, G. spp. Anoplius cylindricus is host specific on burrowing wolf spiders (Lycosidae) of the genus Geolycosa. All wasps (7/7, 100.0%) extended the spider’s burrow into a short side chamber and crude oval cell in which they stocked the paralyzed prey. Host sexes and stages: Immature, 11 (100.0%). All host spiders were immature. Host body lengths (mm): 5.0–8.5, mean = 7.50 ± 0.46, n = 7 (wasps, range = 6.0–9.0, mean = 8.14 ± 0.40, n = 7 (Fig. 13); host wet weights (mg): range = 22– 35, mean = 28.40 ± 2.91, n = 7 (wasps, range = 7–26, mean = 17.40 ± 3.44, n = 7); spider:wasp wet-weight ratios: range = 1.22–5.00:1, mean = 1.63:1, n = 7. Five of 7 wasps (71.4%) were as long as or longer than their host spiders (Fig. 13), but all 7 spiders weighed more than the wasps. Amputation of legs: 0/11. References: Evans 1951a; Evans and Yoshimoto 1962; Gwynne 1979; Krombein 1979; Kurczewski 1981, 1999, pers. observ.; Kurczewski and Kurczewski 1968a, 1968b, 1973. Anoplius (Pompilinus) insolens (Banks) Collection dates: (26; 19 June–4 August [IT], 20 June–24 September [WG]). There are probably 2 generations per year in the eastern Great Lakes Region. Collection dates with prey: mid-June–September 1985. Localities: WG (1), CT (1). Habitat: Sandy and gravelly soils near vegetation. Burrow excavation apparatus: Mandibles; forebasitarsus with 3 comb-spines, the spines short and not or only slightly longer than tarsal width. Host families and species: LYCOSIDAE – Pardosa milvina; PHILODROMIDAE – Tibellus gertschi Chamberlin & Ivie; THOMISIDAE – Xysticus sp.; SALTICIDAE – Maevia inclemens. Anoplius insolens is probably strongly polyphagous in the eastern Great Lakes Region based on these 4 host records. Northeastern Naturalist 47 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Host sexes and stages: Adult female (3/3, 100.0%). Adult females of small species were the preferred hosts. Amputation of legs: 0/2. References: Evans 1951a, Evans and Yoshimoto 1962, Krombein 1979, Kurczewski 1999, Kurczewski and Edwards 2012, Kurczewski et al. 1987, Wasbauer 1982. Anoplius (Pompilinus) marginatus (Say) species-complex This species-complex may consist of 5 species identifiable only in the male sex by their distinctive genitalia: A. bequaerti (Dreisbach), A. marginatus (Say), A. rectangularis (Dreisbach), A. stenotus (Banks), and A. townesi Evans. Collection dates: (389; 24 May–1 November). There are at least 2 generations per year in the eastern Great Lakes Region. Collection dates with prey: 24 August 1960, 19–31 August 1961, 20–30 August 1962, 4 August 1963, 30 June–18 July 1964, 18 July–4 October 1967, 13 June–30 August 1968, 13 June–13 August 1969, 7 June–10 August 1970, 16–23 June 1971, June–September 1972, 1973, 1975, 1978, 1980, 1981, 1982, 1985, 1986, 13 September 1991, 9 August 1992, 2 July 2010, 6 July–9 October 2011. Localities: PI (29), WG (28), AU (44), IT (25), ON (14), GY (1), GR (1), CH (3), SS (3), PB (1), LP (1), WE (2), AL (1), Ontario (1). Habitat: Bare ground in sandy and gravelly soils, loamy soils of gardens and waste areas, and friable soil at woodland edges. Such variation in habitat and soil type is related to 5 species being in A. marginatus species-complex. Burrow excavation apparatus: Mandibles; forebasitarsus with 3 comb-spines of moderate length, not or scarcely longer than thickness of tarsus. Species in this species-complex often modify existing burrows in soil as nests. Host families and species: DYSDERIDAE - Dysdera crocata; ARANEIDAE - Argiope aurantia; LYCOSIDAE - Arctosa emertoni Gertsch, Arctosa sp., Geolycosa domifex, G. fatifera (Hentz), G. wrighti, Pardosa distincta, P. saxatilis, P. xerampelina, P. sp., Pirata insularis, Schizocosa avida, S. crassipes, S. saltatrix, S. spp., Trochosa ruricola, T. terricola, T. sp., Varacosa avara; AGELENIDAE - Agelenopsis naevia, A. pennsylvanica, A. sp.; AMAUROBIIDAE - Callobius bennetti, Amaurobiidae sp.; ANYPHAENIDAE - Anyphaena sp.; CORINNIDAE - Castianeira descripta (Hentz), C. longipalpa (Hentz), C. sp.; CLUBIONIDAE - Clubiona kastoni, C. mixta Emerton, C. sp.; GNAPHOSIDAE - Callilepis imbecilla (Keyserling), Drassodes neglectus (Keyserling), Drassyllus niger (Banks), Gnaphosa muscorum (L. Koch), G. sp., Haplodrassus signifer, Herpyllus vasifer; PHILODROMIDAE - Philodromus infuscatus Keyserling, P. keyserlingi Marx, Thanatus formicinus, T. sp., Tibellus duttoni (Hentz), T. oblongus (Walckenaer); THOMISIDAE - Misumena vatia, Xysticus auctificus Keyserling, X. bicuspis, X. elegans, X. ferox, X. funestus, X. gulosus, X. sp.; SALTICIDAE - Evarcha hoyi, Habronattus borealis, H. decorus, H. viridipes, H. sp., Maevia inclemens, Metaphidippus sp., Phidippus audax, P. clarus Keyserling, P. purpuratus Keyserling, P. rimator (Walckenaer), P. whitmani. The Anoplius marginatus species-complex, consisting of perhaps 5 species unidentifiable in the female sex, is strongly polyphagous preying on at least 12 host Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 48 families of orb-weaving, cursorial-hunting, burrowing, and retreat-dwelling spiders in the eastern Great Lakes Region (Table 3). Dysderidae is an unusual host family for a pompilid, Priocnemis minorata being the only other Nearctic spider wasp to utilize such a host species. Varacosa avara and Trochosa terricola (Lycosidae) and Thanatus formicinus (Philodromidae) were commonly provisioned host species. There is a highly unusual record of “A. marginatus” capturing a harvestman, Odiellus pictus (Wood) (Opiliones: Phalangiidae) (Evans 1948). Host sexes and stages: Adult female, 56 (36.6%); penultimate female, 1 (0.7%); immature female, 20 (13.1%); adult male, 5 (3.3%); penultimate male, 1 (0.7%); immature male, 8 (5.2%); immature, 62 (40.5%). Adult female and unsexed immature spiders constituted 118/153 (77.1%) of the host records. Host body lengths (mm): range = 5–11, mean = 7.78 ± 0.19, n = 76 (wasps, range = 7–12, mean = 9.13 ± 0.14, n = 76) (Figs. 14, 16); host wet weights (mg): range = 13–131, mean = 51.00 ± 4.06, n = 53 (wasps, range = 10–49, mean = 26.54 ± 1.33, n = 53) (Fig. 15); spider:wasp wet-weight ratios: range = 0.35–6.77:1, mean = 1.91:1, n = 53. The wasps were mainly longer than their host spiders (57/76, 75.0%; r = 0.429, P < 0.001; Figs. 14, 16), but most of the spiders (47/53, 88.7%) weighed more than the wasps (Fig. 15). Larger wasps often provisioned their nests with larger spiders (Figs. 14–16). Wasps with prey 4.23–5.17 times as heavy as themselves had considerable difficulty in prey transport. Wasps with prey 5.53–6.77 times as heavy as themselves were unable to transport their spiders and abandoned them because of the relatively large sizes and heavy weights. Amputation of legs: 7/114 (6.1%) spiders had 1 (5) or 3 (2) legs amputated at the coxa-trochanter joints. References: Evans 1948, 1951a, 1951b; Evans and Yoshimoto 1962; Krombein 1953a, 1959, 1979; Kurczewski 1962, 1999, 2010, pers. observ.; Kurczewski and Figure 16. Spider body length plotted against wasp body length in Anoplius marginatus species-complex and A. splendens. Northeastern Naturalist 49 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Table 3. Families and genera of host spiders preyed on by 2 closely related, strongly polyphagous pompilid taxa—Anoplius marginatus species-complex and Anoplius splendens—in Erie County, PA. Families of host spiders are arranged in taxonomic order following Platnick (2012) with host genera listed alphabetically. Anoplius marginatus Host spider family and genus species-complex Anoplius splendens DYSDERIDAE X Dysdera X ARANEIDAE X X Argiope X Larinioides X LYCOSIDAE X X Arctosa X X Geolycosa X X Hogna X Pardosa X X Pirata X Schizocosa X X Trochosa X X Varacosa X X PISAURIDAE X Pisaurina X AGELENIDAE X X Agelenopsis X X AMAUROBIIDAE X X Callobius X X ANYPHAENIDAE X Anyphaena X CORINNIDAE X X Castianeira X X CLUBIONIDAE X Clubiona X GNAPHOSIDAE X X Callilepis X Drassodes X X Drassylus X Gnaphosa X Haplodrassus X Herpyllus X PHILODROMIDAE X X Philodromus X Thanatus X X Tibellus X X THOMISIDAE X X Misumena X Xysticus X X SALTICIDAE X X Evarcha X Habronattus X X Maevia X Metaphidippus X Phidippus X X Platycryptus X Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 50 Acciavatti 1990; Kurczewski and Edwards 2012, unpubl. data; Kurczewski and Kurczewski 1968a, 1968b, 1973; Kurczewski and Pitts 2011; Kurczewski et al. 1987; Peckham and Peckham 1898; Rau and Rau 1918; Wasbauer 1982. Anoplius (Pompilinus) splendens (Dreisbach) Collection dates: (114; 6 June–5 October). This species often has 2 generations per year in the eastern Great Lakes Region depending on locality and optimal weather conditions. Collection dates with prey: 7 July 1962, 29 June 1964, 9 June–27 September 1965, 12 October 1965 (KS), 23 July 1966, 1 August–5 October 1967, 1 August–14 September 1968, 25 June–22 September 1969, 20 June–18 September 1970, 3 July–17 August 1971, June–September 1972, 1976, 1978, 9 August 1992, 13 June 1993, 19 August 2004. Localities: PI (77), AL (1), GY (1), MA (1), KS (1). Habitat: Sandy soils near water courses, sand dunes, sand beaches, sand plain, and inland sand blowouts. Burrow excavation apparatus: Mandibles; forebasitarsus with 3 comb-spines of moderate length, each about as long as width of tarsus. Anoplius splendens uncharacteristically does not have the long comb-spines typical of other psammophilous Anoplius species that nest in coarse-grained sandy soils. Host families and species: ARANEIDAE - Larinioides patagiatus; LYCOSIDAE - Arctosa littoralis, Geolycosa wrighti, Hogna frondicola, H. sp., Pardosa distincta, P. milvina, P. sp. near floridana, P. sp., Schizocosa avida, S. crassipes, S. ocreata, S. saltatrix, S. sp., Trochosa terricola, T. sp., Varacosa avara; PISAURIDAE - Pisaurina mira; AGELENIDAE - Agelenopsis pennsylvanica, A. sp.; AMAUROBIIDAE - Callobius bennetti, C. sp.; CORINNIDAE - Castianeira longipalpa, C. sp.; GNAPHOSIDAE - Drassodes auriculoides Barrows, D. neglectus; PHILODROMIDAE - Thanatus formicinus, T. sp. probably formicinus, Tibellus oblongus; THOMISIDAE - Xysticus elegans, X. ferox, X. sp. probably ferox, X. funestus, X. pellax (= X. ontariensis Emerton); X. sp.; SALTICIDAE - Habronattus agilis, H. coronatus, H. viridipes, H. sp., Phidippus audax, P. cardinalis Keyserling, P. clarus, P. ?rimator, Platycryptus undatus. This species is strongly polyphagous preying on at least 10 families of orb-weaving, cursorial-hunting, burrowing and retreat-dwelling spiders in the eastern Great Lakes Region (Table 3). Varacosa avara (Lycosidae) and Thanatus formicinus (Philodromidae) were commonly used host species. Host sexes and stages: Adult female, 29 (36.3%); penultimate female, 1 (1.3%); immature female, 8 (10.0%); adult male, 3 (3.8%); immature male, 8 (10.0%); immature, 31 (38.8%). Adult female and unsexed immature spiders accounted for 60/80 (75.0%) of the host records. Host body lengths (mm): range = 5–13, mean = 7.59 ± 0.20, n = 70 (wasps, range = 7–12, mean = 9.57 ± 0.15, n = 70) (Fig. 16); host wet weights (mg): range = 21–127, mean = 44.67 ± 16.89, n = 6 (wasps, range = 12–36, mean = 21.17 ± 3.42, n = 6) (Fig. 15); spider:wasp wet-weight ratios: range = 1.05–3.53:1, mean = 2.11:1, n = 6. The wasps were often longer than their host spiders (54/70, 77.1%; Northeastern Naturalist 51 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 r = 0.367, P = 0.002; Fig. 16), but all spiders (6/6, 100.0%) weighed more than the wasps (Fig. 15). Larger wasps tended to provision their nests with larger spiders (Figs. 15, 16) including the shoreline sand spider Arctosa littoralis and the burrowing wolf spider Geolycosa wrighti. Amputation of legs: 3/79 (3.8%) spiders had 1 leg cut off at a coxa-trochanter joint. References: Evans 1951a; Evans and Yoshimoto 1962; Krombein 1959, 1979; Kurczewski 1999, 2010, pers. observ.; Kurczewski and Edwards unpubl. data; Kurczewski and Kurczewski 1968a, 1968b, 1973; Kurczewski et al. 1987; Wasbauer 1982; Wasbauer and Powell 1962. Anoplius (Pompilinus) subcylindricus (Banks) Collection dates: (31, 1 June–10 October). There are 2 generations per year in the eastern Great Lakes Region. Collection dates with prey: 6 June–20 August 1954, 19 September–14 October 1964, June–September 1982, 1984, 1985. Localities: KS (4), WG (3), IT (3). Habitat: Bare sandy and gravelly soils near vegetation. Burrow excavation apparatus: Mandibles; forebasitarsus with 3 short combspines, the spines about as long as thickness of tarsus. Host families and species: GNAPHOSIDAE – Micaria sp.; PHILODROMIDAE – Philodromus sp.; THOMISIDAE – Misumenops asperatus (Hentz), Xysticus banksi Bryant, X. discursans, X. ferox, X. funestus, X. gulosus, X. triguttatus. Anoplius subcylindricus is rather oligophagous on crab spiders (Thomisidae), particularly species of Xysticus, but rarely captures Philodromidae and Gnaphosidae. Host sexes and stages: Adult female, 2 (20.0%); immature female; 2 (20.0%), adult male, 1 (10.0%); immature male, 1 (10.0%); immature, 4 (40.0%). Immature (7/10, 70.0%) was the predominant host stage. Host body lengths (mm): range = 5.0–6.5, mean = 5.50 ± 0.35, n = 4 (wasps, range = 7.0–9.0, mean = 7.75 ± 0.48, n = 4; host wet weights (mg): range = 20–48, mean = 29.50 ± 6.40, n = 4 (wasps, range = 11–27, mean = 17.75 ± 3.82, n = 4); spider:wasp wet-weight ratios: range = 0.81–4.00:1, mean = 2.09:1, n = 4. All 4 wasps were longer than their host spiders, but 2 of 4 (50.0%) spiders were heavier than the wasps. Amputation of legs: 0/10. References: Evans 1951a, 1951b; Evans and Yoshimoto 1962; Krombein 1953a, 1979; Kurczewski 1999, pers. observ.; Kurczewski and Acciavatti 1990; Kurczewski and Kurczewski 1968a; Kurczewski et al. 1987. Anoplius (Pompilinus) tenebrosus (Cresson) Collection dates: (101; 24 April–25 September). Adults emerge in July in upstate New York and mate shortly thereafter. Males die after a few weeks; females overwinter in the fall in deep burrows dug in sandy soil and nest during the first warm days of spring of the following year. There is thus a single generation of females that overlaps 2 years. Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 52 Collection dates with prey: 3 June 1968, 29 May 1970, 22 May–19 June 1971, May–June 1972, May–June 1973, 20 May 1977, 20 May–11 June 1978, 24 April–2 May 2005 (MA), 30 April 2009, 18 June 2011, 7 May 2014. Localities: BV (11), WN (5), ML (5), FU (1), PS (1), SS (1), CH (1), MA (5), Quebec (3). Habitat: Sandy soils in the vicinity of woodland; sand pits; and sandy fields, roads, and trails. Burrow excavation apparatus: Mandibles; forebasitarsus with 3 comb-spines not or scarcely longer than tarsal width. Host families and species: LYCOSIDAE – Alopecosa kochi (Keyserling), Arctosa sp., Gladicosa gulosa, Hogna baltimoriana, H. frondicola, Pardosa moesta, Schizocosa avida, S. crassipalpata, S. ocreata, S. saltatrix, S. sp. probably avida, Trochosa ruricola, T. terricola, T. sp., Varacosa avara; AGELENIDAE – Tegenaria domestica (Clerck); AMAUROBIIDAE – Callobius bennetti, Wadotes hybridus, W. sp.; CORINNIDAE – Agroeca ornata; GNAPHOSIDAE – Drassylus sp., Haplodrassus signifer, Zelotes subterraneous (C. L. Koch), Z. sp.; PHILODROMIDAE – Thanatus formicinus; THOMISIDAE – Misumena vatia, Ozyptila distans Dondale & Redner, Xysticus ampullatus Turnbull, Dondale & Redner, X. canadensis, X. elegans, X. ferox, X. gulosus, X. spp.; SALTICIDAE – Habronattus sp., Naphrys pulex, Sitticus sp. Anoplius tenebrosus provisioned its nests with 8 families of cursorial-hunting, burrowing, and retreat-dwelling spiders in the eastern Great Lakes Region; wolf spiders (Lycosidae) comprised the majority (55/80, 68.8%) of the host records. Host sexes and stages: Adult female, 25 (32.5%); immature female, 38 (49.4%); adult male, 5 (6.5%); immature male, 1 (1.3%); immature, 8 (10.4%). Adult and immature females (63/77, 81.8%) were the preferred stages and sex. Host body lengths (mm): range = 6.0–9.0, mean = 7.00 ± 1.00, n = 3 (wasps, range = 8.5–9.5, mean = 9.00 ± 0.29, n = 3); host wet weights (mg): range = 16–48, mean = 30.33 ± 9.39, n = 3 (wasps, range = 25–29, mean = 26.667 ± 1.20, n = 3). Two of 3 (66.7%) wasps were longer than their host spider but 2 of 3 spiders were heavier than the wasps. Alm and Kurczewski (1984) reported the following host wet-weight (mg) data: range = 16–210, mean = 90.41, n = 94 (wasps, range = 11–72, mean = 39.73, n = 94); spider:wasp wet-weight ratios: 0.64–3.92:1, mean = 2.28:1, n = 94. Amputation of legs: 2/11 (18.2%). References: Alm and Kurczewski 1984; Evans 1951a, 1970; Evans and Yoshimoto 1962; Krombein 1979; F.E. Kurczewski, pers. observ.; Kurczewski and Edwards 2012, unpubl. data; Kurczewski and Kurczewski 1973; Kurczewski et al. 1987; Wasbauer and Powell 1962. Anoplius (Anoplius) depressipes Banks Collection dates: (18; 8 June–12 August). There is probably only a single generation per year in the eastern Great Lakes Region but 2 or more generations per year (February–September) in Florida. Collection dates with prey: July 1954, 18 July 1957, 23 June–1 July 1962. Localities: PI (3), SS (1), TX (3), OK (1), LA (1), FL (2). Northeastern Naturalist 53 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Habitat: Shores of ponds, swamps, lagoons, and slow-moving streams. Females nest in existing cavities in decaying wood or soil on the shore and return to the same cavities for successive cells. The wasps are highly adept at walking on the water surface, diving below the surface in search of and pursuit of fishing spiders, and towing their host spider across the water surface to an existing burrow on the shore. Burrow excavation apparatus: Mandibles; forelegs with small comb-spines; tarsal segments are flattened or slightly concave and ridged with very small spines or setae. Such a structural mechanism is evidently an adaptation for a semi-aquatic existence. Host family and species: PISAURIDAE – Dolomedes scriptus, D. striatus Giebel, D. tenebrosus, D. triton sexpunctatus Hentz, Pisaurina mira. All host records from the eastern Great Lakes Region (13/13, 100.0%) are for fishing spiders of the genus Dolomedes (Pisauridae), including semi-aquatic and quasiterrestrial species. Host sexes and stages: Adult female, 11 (84.6%); subadult female, 1 (7.7%); subadult male, 1 (7.7%). Adult female spiders were the preferred host stage and sex. Host body lengths (mm): range = 21.0–23.0, mean = 22.00 ± 0.58, n = 3 (wasps, range = 17.0–19.0, mean = 18.33 ± 0.67, n = 3). All 3 host spiders were longer and much heavier than the wasps. Amputation of legs: 0/11. There was no evidence of leg amputation despite the long legs of the host spiders. The method of prey transport—towing the spider across the water surface using the wings while grasping a foreleg—probably influenced the lack of amputation. References: Evans 1949, 1951a; Evans and Yoshimoto 1962; Krombein 1979; Kurczewski 1999, pers. observ.; Kurczewski and Edwards 2012, unpubl. data; Kurczewski and Kurczewski 1968a, 1987b; Roble 1985. Anoplius (Anoplius) illinoensis (Robertson) Collection dates: (104; 13 June–26 September). There may be 2 generations per year in the eastern Great Lakes Region with optimal weather conditions, although this species is noticeable only in late summer. Collection dates with prey: 26 August 1951, 23 September 1956, 22 June 1969, August 1975, August–September 1979, August–September 1981, September 1984. Localities: WG (5), PI (2), IT (1), WE (1) CT (1). Habitat: Abandoned overgrown fields, meadows, shrub and tree savannas, gravel pits, and gravelly parking lots. Burrow excavation apparatus: Mandibles; forebasitarsus somewhat strongly spined compared to other species of Anoplius s. str., the spines being mostly as long as tarsal width. Host family and species: LYCOSIDAE – Arctosa littoralis, Hogna helluo, H. sp., Schizocosa avida, S. ocreata, Trochosa terricola. All host records (10/10, 100.0%) were for species of wolf spiders (Lycosidae). Host sexes and stages: Adult female, 4 (40.0%); penultimate female, 1 (10.0%); immature female, 1 (10.0%); adult male, 2 (20.0%); immature, 2 (20.0%). The majority of host records (6/10, 60.0%) were for female spiders. Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 54 Host body lengths (mm): range = 9–15, mean = 12.00 ± 3.00, n = 2 (wasps, range = 12–13, mean = 12.50 ± 0.50, n = 2); host wet weights (mg): range = 98–286, mean = 192.00 ± 94.00, n = 2 (wasps, range = 54–89, mean = 71.50 ± 17.50, n = 2); spider:wasp wet-weight ratios: range = 1.81:1–3.21:1, mean = 2.51:1, n = 1. One of 2 (50.0%) host spiders was longer than the wasp and both spiders weighed significantly more than wasps. Amputation of legs: 0/10. References: Evans 1951a; Evans and Yoshimoto 1962; Krombein 1979; Kurczewski 1999, pers. observ.; Kurczewski and Acciavatti 1990; Kurczewski and Kurczewski 1973; Kurczewski et al. 1987. Anoplius (Anoplius) imbellis Banks Collection dates: (37; 9 June–26 October). There are 2 generations per year in late spring and late summer in the eastern Great Lakes Region. Collection dates with prey: 14 October 1968, August–September 1982, August– September 1985, 24–25 September 2009, 15 June 2010, 17 August–6 October 2011, 25–26 October 2012, 1–2 October 2013. Localities: PI (4), WG (4), ON (21). Habitat: Gravelly and loamy, often damp soils near edges of overgrown fields and deciduous woodland; and gravel pits, sometimes near water. Burrow excavation apparatus: Mandibles; forebasitarsus with 3 short combspines, those of upper row half as long as width of tarsus and those of lower row as long as tarsal width. Foretarsus is adapted for excavating in fine-grained soils such as silt loam. Host families and species: LYCOSIDAE – Arctosa sp., Pardosa distincta, P. distincta species group, P. milvina, P. ramulosa McCook, P. sp., Schizocosa mccooki (Montgomery), S. sp., Trochosa ruricola, T. sp. probably ruricola, T. terricola, Varacosa avara; PISAURIDAE - Pisaurina mira; AMAUROBIIDAE - Amaurobius ferox; THOMISIDAE – Xysticus sp. Anoplius imbellis preys mainly on wolf spiders (Lycosidae) (28/31 host records, 90.3%) but rarely captures fishing spiders (Pisauridae) and hacklemesh-weaver spiders (Amaurobiidae) in the eastern Great Lakes Region and crab spiders (Thomisidae) in the western US. Most host records were for species of Pardosa and Trochosa. Host sexes and stages: Adult female, 8 (25.8%), immature female, 2 (6.5%), adult male, 4 (12.9%), penultimate male, 2 (6.5%), immature male, 2 (6.5%), immature, 13 (41.9%). Anoplius imbellis captured both sexes of host spiders about equally (10 females, 8 males), but immature spiders were the predominant host stage (19/31, 61.3%). Host body lengths (mm): range = 5.2–11.0, mean = 8.66 ± 0.35, n = 23 (wasps range = 6.5–11.0, mean = 9.22 ± 0.32, n = 23) (Fig. 17). The wasps were mainly as long as or the same length as their host spiders (23/26, 88.5%) (Fig. 17). Amputation of legs: 0/23 (0.00%) including a fishing spider, Pisaurina mira, with all legs intact and a leg span of 39.5 mm. References: Evans 1951a; Evans and Yoshimoto 1962; Krombein 1979; Kurczewski 1999, 2010 (reported as A. (A.) nigerrimus [Scopoli]), pers. observ.; Northeastern Naturalist 55 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Kurczewski and Acciavatti 1990; Kurczewski and Edwards 2012, unpubl. data; Kurczewski and Kurczewski 1968a, 1973; Kurczewski and Pitts 2011; Kurczewski et al. 1987; Wasbauer 1957, 1982; Wasbauer and Powell 1962; Wilson and Pitts 2007. Anoplius (Anoplius) ithaca (Banks) Collection dates: (51; 24 May–10 October). Anoplius ithaca has 2 generations per year in the eastern Great Lakes Region. Collection dates with prey: 28 August 1955, 30 July–19 August 1961, 9 September 1966, 25 September–10 October 1968, 4–10 September 1969, June 1984. Localities: PI (9), IT (2), WG (1), DR (1), MA (1). Habitat: Rocky shorelines and stream sides, sandy beaches with pebbles and stones, and water courses with dried streambeds. Burrow excavation apparatus: Mandibles; forebasitarsus with some spines of both upper and lower rows as long as tarsal width. Tarsal claws large, bifid, with inner claw of each pair distinctly longer than outer claw and about as long as last tarsal segment. Such a structural mechanism is evidently an adaptation for running across round stones and pebbles. Host families and species: LYCOSIDAE – Arctosa littoralis, Hogna sp., Pardosa groenlandica, P. lapidicina Emerton, P. lowriei Kronestedt, P. milvina, P. sp. probably milvina, P. steva Lowrie and Gertsch, P. sp.; GNAPHOSIDAE – Drassylus depressus (Emerton). Anoplius ithaca often captured small lycosids belonging to the genus Pardosa, especially near water courses. Host sexes and stages: Adult female, 10 (37.0%); penultimate female, 1 (3.7%); adult male, 1 (3.7%); immature female, 1 (3.7%); immature, 14 (51.9%). Non-adult spiders were the predominant A. ithaca prey (16/27, 59.3%), and female spiders were much preferred over males where the 2 sexes occurred together. Host body lengths (mm): range = 4.5–9.0, mean = 6.51 ± 0.48, n = 10 (wasps, range = 7.0–9.0, mean = 7.97 ± 0.21, n = 10) (Fig. 17). Although the wasps were Figure 17. Spider body length plotted against wasp body length in Anoplius imbellis and A. ithaca. Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 56 approximately the same size, Arctosa littoralis prey (range = 7.0–9.0 mm, mean = 7.80, n = 5 [wasps, range = 7.0–8.0, mean = 7.80, n = 5]) were larger than Pardosa milvina prey (range = 4.5–5.6 mm, mean= 5.22, n = 5 [wasps, range = 7.0–9.0, mean = 8.14, n = 5]). Only 2 of 5 (40.0%) wasps were longer than their A. littoralis prey, whereas all 5 wasps were much longer than their P. milvina prey (Fig. 17). Amputation of legs: 1/10 (10.0%). Amputation of a single leg in A. ithaca probably enabled feeding on the exuding hemolymph at the point of amputation as this species has never been observed obtaining nectar from flowers. References: Cooper 1953; Evans 1948, 1951a; Evans and Yoshimoto 1955, 1962; Krombein 1956, 1979; Kurczewski 1962, 1999, pers. observ.; Kurczewski and Kurczewski 1968a, 1973; Kurczewski et al. 1987; Ricards 1969; Wasbauer 1957, 1982. Anoplius (Anoplius) nigerrimus (Scopoli) Anoplius nigerrimus is a common Palearctic species that was not recognized as a distinct species in North America until the mid-20th century (Evans 1951a). Collection dates: (24; June, 1 July–9 August). There are too few records to ascertain the number of generations per year in the eastern Great Lakes Region. This species has 2 or more generations per year in southern Europe, based on early and late collection records. Collection date with prey: June 1981. Locality: WG (1). Habitat: Overgrown gravel pit bordering deciduous woodland, and sand barren near open woodland. Burrow excavation apparatus: Mandibles; forebasitarsus with short combspines, the spines usually shorter than width of tarsus. Host family and species: LYCOSIDAE – Trochosa terricola. In Europe, this wasp species preys mainly on Lycosidae, including T. terricola, but also captures Pisauridae and Gnaphosidae. Host sex and stage: Immature female, 1 (100.0%). Amputation of legs: 0/1. References: Evans 1951a; Evans and Yoshimoto 1962; Krombein 1979; Kurczewski 1999, 2010 (re-identified as A. imbellis); Kurczewski and Kurczewski 1987b; Kurczewski et al. 1987; Richards and Hamm 1939. Anoplius (Anoplius) ventralis (Banks) Collection dates: (61; 19 June–6 November). There may be 2 generations per year in the eastern Great Lakes Region with optimal weather conditions, although this species is noticeable only in late summer. Collection dates with prey: 13 September 1965, 2 September 1966, 16 August–4 October 1967, September 1978. Localities: AU (2), PI (1), WG (1), KS (1), Ontario (1). Habitat: Edge of abandoned overgrown field, crevice at bottom of sand cliff, open deciduous woodland, gravel pit, and border of sandy field. Burrow excavation apparatus: Mandibles; forebasitarsus with spines of moderate length, often as long as or slightly longer than width of tarsus. Northeastern Naturalist 57 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Host families and species: LYCOSIDAE – Hogna frondicola, H. helluo, Schizocosa saltatrix, Trochosa terricola; AGELENIDAE – Agelenopsis naevia. Host records include cursorial-hunting spiders of the family Lycosidae and, less frequently, funnel web-weaver spiders of the family Agelenidae. Host sexes and stages: Adult female, 4 (66.7%); immature female, 1 (16.7%); immature, 1 (16.7%). Host records demonstrate a preference for females of moderate- size wolf spider species (Lycosidae). Host body lengths (mm): range = 9–12, mean = 11.25 ± 0.75, n = 4 (wasps, range = 11–13, mean = 12.50 ± 0.50, n = 4); host wet weights (mg): range = 63–169, mean = 123.00 ± 31.39, n = 3 (wasps, range = 67–102, mean = 82.00 ± 10.41, n = 3); spider:wasp wet-weight ratios: range = 0.82–2.04:1, mean = 1.51:1, n = 3. All 4 (100.0%) wasps were longer than their host spiders, but 2 of 3 (67.7%) spiders outweighed the wasps. Amputation of legs: 1/6 (16.7%). One spider had its 1st and 3rd left legs cut off at coxa-trochanter joints. References: Evans 1951a; Evans and Yoshimoto 1962; Krombein 1979; Kurczewski 1999, pers. observ.; Kurczewski and Acciavatti 1990; Kurczewski and Kurczewski 1968a, 1968b; Kurczewski et al. 1987; Powell 1958; Wasbauer and Powell 1962. Anoplius (Anoplius) virginiensis (Cresson) Collection dates: (120; 24 May–3 October). Anoplius virginiensis has 2 generations per year in the eastern Great Lakes Region. Collection dates with prey: 18 August–3 October 1966, 6 June–6 September 1968, 6–25 August 1969, 31 August–17 September 2009. Localities: PI (16), WG (5), IT (5), TU (1), ON (4). Habitat: Moist deciduous woodland with sunlit openings, rotting stumps, and fallen timber. Burrow excavation apparatus: Mandibles; forebasitarsus with 3 comb-spines, those of upper row very short and those of lower row as long as tarsal width. Foretarsus is adapted for excavating in soft and rotting wood materials. Host families and species: LYCOSIDAE - Pardosa lapidicina, Trochosa terricola; AGELENIDAE - Agelenopsis pennsylvanica, A. utahana, A. sp. probably utahana, A. sp.; AMAUROBIIDAE - Callobius bennetti, Coras juvenilis, Wadotes calcaratus, W. hybridus. Anoplius virginiensis is moderately polyphagous, preying on 3 cursorial-hunting and retreat-dwelling families of spiders in the eastern Great Lakes Region (Table 1). Agelenopsis spp. (12/21, 57.1%) and Callobius bennetti (5/21, 23.8%) were predominant host spider species at 1 locality (PI). Three of 5 host spiders (60.0%) from another site (IT) were Agelenopsis spp. Host sexes and stages: Adult female, 19 (44.2%); immature female, 4 (9.3%); adult male, 6 (14.0%); penultimate male, 1 (2.3%), immature male, 4 (9.3%); immature, 9 (20.9%). More adult female spiders were captured by the wasps than any other sex or stage. Host body lengths (mm) (all species): range = 7–14, mean = 9.46 ± 0.30, n = 26 (wasps, range = 8–14, mean = 10.12 ± 0.22, n = 26) (Fig. 4); host wet weight (mg): 55, n = 1 (wasp, 26, n = 1); spider:wasp wet-weight ratio: 2.12:1, n = 1. There was Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 58 less variability in host spider size in A. virginiensis than in Priocnemis minorata and P. germana, 2 other commonly seen deciduous woodland pompilids, probably the result of smaller sample size (Fig. 4). Most A. virginiensis females were longer or the same size (22/26, 84.6%) as their host spiders but weighed less (Fig. 4). Host body lengths (mm) (Callobius bennetti): range = 9–11, mean = 9.60 ± 0.40, n = 5 (wasps, range = 9–11, mean = 10.00 ± 0.32, n = 5) (Fig. 5). All wasps (5/5, 100.0%) were the same length or longer than their host C. bennetti (Fig. 5). The comparative body lengths of A. virginiensis and all host spiders and A. virginiensis and Callobius bennetti showed moderate and strong positive linear relationships, respectively (r = 0.573, P = 0.002, n = 26 [Fig. 4]; r = 0.751, P = 0.111, n = 5 [Fig. 5]), although the latter is based on only 5 data points. Amputation of legs: 0/31. References: Evans 1951a; Evans and Yoshimoto 1962; Krombein 1979; Kurczewski 1999, 2010, pers. observ.; Kurczewski and Kurczewski 1968a, 1973. Ammosphex angularis (Banks) Collection dates: (12; 21 June–5 September). There may be 2 generations per year in the eastern Great Lakes Region, but few collection reco rds to confirm this. Collection dates with prey: 11 August 1968, 21 June 1969, 2 September 1971. Localities: CH (1), PS (1), WE (1), CA (18), WA (1), WY (1). Habitat: Sand dunes, bare sand at woodland border, sandy open woodland, and sandy field. Burrow excavation apparatus: Mandibles; forebasitarsus with comb-spines longer than width of tarsus; apical spine on forebasitarsus half the length of 2nd foretarsal segment. Host families and species: LYCOSIDAE – Schizocosa sp.; ANYPHAENIDAE – unidentified genus and species; CLUBIONIDAE – Clubiona sp.; CORINNIDAE – Castianeira thalia Reiskind, C. sp.; GNAPHOSIDAE – Callilepis altitudonis Chamberlin, C. imbecilla, ?Litophyllus sp., Sergiolus ocellatus (Walckenaer) species group; PHILODROMIDAE – Thanatus formicinus; THOMISIDAE – Xysticus knowltoni Gertsch, X. spp; SALTICIDAE – Habronattus spp.. Ammosphex angularis is strongly polyphagous throughout North America, provisioning with 8 families of cursorial-hunting and retreat-dwelling spiders. Host sexes and stages: Adult female, 10 (47.6%); penultimate female, 2 (9.5%); adult male, 3 (14.3%); immature, 6 (28.6%). Adult female and immature spiders were the predominant (16/21, 76.2%) sex and stages. Host body lengths (mm): range = 4.5–8.0, mean = 6.09 ± 0.27, n = 18 (wasps, range = 6.0–8.5, mean = 7.20 ± 0.23, n = 18) (Fig.18); host wet weights (mg): range = 13–34, mean = 23.50 ± 10.50, n = 2 (wasps, range = 9–17, mean = 13.0 ± 4.00, n = 2); spider:wasp wet-weight ratios: range = 1.44–2.00:1, mean = 1.72, n = 2). All 18 (100.0%) wasps were as long as or longer than their host spiders (Fig. 18). Amputation of legs: 1/18 (5.6%). References: Alcock 1973; Evans 1951b, 1959, 1963, 1970; Evans and Yoshimoto 1962; Krombein 1979; Kurczewski 1999, pers. observ.; Kurczewski and Edwards 2012; Kurczewski and Kurczewski 1973; Wasbauer and Powell 1962. Northeastern Naturalist 59 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Ammosphex michiganensis (Dreisbach) Collection dates: (19; 25 May–9 October). There are 2 generations per year in the eastern Great Lakes Region. Collection dates with prey: 25 May 1963, 9 October 1966, 5 September 1967, 5 August 1968, 30 August 1969. Localities: MR (3), AU (2), GR (1), CH (1). Habitat: Edges of gravel bank and sand pit, border of overgrown field, sandy field, and open woodland with loamy soil. Burrow excavation apparatus: Mandibles; foretarsal comb-spines as long as width of tarsus, the apical spine of forebasitarsus half as long as 2nd foretarsal segment. Host family and species: THOMISIDAE – Xysticus conctator Thorell, X. ferox, X. funestus, X. transversatus (Walckenaer), X. spp. Ammosphex michiganensis is oligophagous on crab spiders (Thomisidae) of the genus Xysticus. Host sexes and stages: Penultimate female, 2 (28.6%); immature female, 3 (42.9%); immature, 2 (28.6%). Ammosphex michiganensis mainly selected penultimate and immature females (5/7, 71.4%) of small thomisid species. Host body lengths (mm): range = 5.0–8.0, mean = 6.00 ± 0.55, n = 5 (wasps, range = 6.5–10.0, mean = 8.90 ± 0.64, n = 5) (Fig. 18); host wet weights (mg): range = 22–76, mean = 42.25 ± 12.76, n = 4 (wasps, range = 26–40, mean = 32.50 ± 3.78, n = 4); spider:wasp wet-weight ratios: range = 0.85–2.00:1, mean = 1.23:1, n = 4. All 5 wasps were longer than their host spiders (Fig. 18), although 2 of 4 host spiders were heavier than the wasps. Amputation of legs: 0/7. References: Evans 1951b; Evans and Yoshimoto 1962; Krombein 1979; F.E. Kurczewski, pers. observ; Kurczewski and Acciavatti 1990; F.E. Kurczewski and Figure 18. Spider body length plotted against wasp body length in Ammosphex angularis and A. michiganensis. Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 60 G.B. Edwards, unpubl. data; Kurczewski and Kurczewski 1968a, 1968b, 1973; Kurczewski and Snyder 1964. Arachnospila arctus (Cresson) Collection dates: (85; 22 May–5 October). There are 2 generations per year in the eastern Great Lakes Region. Collection dates with prey: 14 September 1967, 3–5 October 1967, 7 September 1988, 9–17 October 2008. Localities: PI (4), ON (3), AU (1), OC (1). Habitat: Sloping or flat open woodland or parkland, sunlit openings in deciduous woodland, gravelly soil at woodland border, backyard patio, and silt loam soil near house foundation. Burrow excavation apparatus: Mandibles; forebasitarsus with 3 or 4 combspines, the spines up to 3 times as long as foretarsal width. The structure of the foretarsal comb of this species indicates that it should be much more psammophilous than it demonstrates. Host families and species: LYCOSIDAE – Arctosa rubicunda, A. sp., Schizocosa crassipes, S. saltatrix; AGELENIDAE – Cybaeus sp., Hololena sp., Novalena pina Chamberlin & Ivie; AMAUROBIIDAE – Amaurobius ferox, Callobius bennetti; CLUBIONIDAE – Clubiona sp.; GNAPHOSIDAE – Gnaphosa muscorum, Orodrassus coloradensis (Emerton); SALTICIDAE – Phidippus johnsoni (Peckham & Peckham). Arachnospila arctus is polyphagous throughout North America, capturing cursorial-hunting and retreat-dwelling spiders. Host sexes and stages: Adult female, 7 (46.7%); subadult female; 1 (6.7%), immature female, 3 (20.0%); adult male, 2 (13.3%); immature, 2 (13.3%). Female spiders (11/15, 73.3%) were the preferred host sex. Host body lengths (mm): range = 8.0–11.5, mean = 9.06 ± 0.39, n = 8 (wasps, range = 10.0–12.0, mean = 10.75 ± 0.31, n = 8) (Fig. 19); host wet weights (mg): range = 41–81, mean = 53.25 ± 9.33, n = 4 (wasps, range = 28, mean = 28.00, n = 4); spider:wasp wet-weight ratios: range = 1.46–2.89:1, mean = 1.90:1, n = 4. All but 1 wasp (7/8, 87.5%) were longer than their host spider (Fig. 19), but all wasps were lighter in weight than their spiders. Amputation of legs: 0/13. References: Evans 1951b; Evans and Yoshimoto 1962; Krombein 1979; Kurczewski 1999, 2010, pers. observ.; Kurczewski and Edwards 2012; Kurczewski and Kurczewski 1968b, 1973; Wasbauer 1982. Arachnospila scelestus (Cresson) Collection dates: (71, 10 June–6 October). Arachnospila scelestus has 2 generations per year in the eastern Great Lakes Region. Collection dates with prey: 24 July 1965, 14 September–3 October 1967, 21 September 1968, August–September 1969, June–September 1972, June–September 1974, 13 June 1993. Localities: PI (6), SS (4), WE (2), AL (1). Habitat: Relict and degraded sand dunes, sand plain, shrub savanna, inland sand blowout, bare sand at deciduous woodland borders, and sandy openings in deciduous–coniferous woodland. Northeastern Naturalist 61 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Burrow excavation apparatus: Mandibles; forebasitarsus usually with 4 welldeveloped comb-spines, the spines up to 3 times as long as foretarsal width. Foretarsus is adapted for excavating in coarse-grained sandy soils. Host families and species: LYCOSIDAE – Geolycosa rafaelana, Gladicosa gulosa, Hogna frondicola, Hogna sp., Schizocosa avida, S. crassipalpata, S. pacifica (Banks), Varacosa avara, Lycosidae sp.; PISAURIDAE – Dolomedes sp.; SALTICIDAE – Phidippus sp. Arachnospila scelestus is rather oligophagous predominantly capturing relatively large wolf spiders (Lycosidae) but rarely provisioning with fishing spiders (Pisauridae) and jumping spiders (Salticideae). Schizocosa avida and Gladicosa gulosa were prevalent host spider species in the eastern Great Lakes Region. Host sexes and stages: Adult female, 8 (40.0%); subadult or immature female, 5 (25.0%); adult male, 3 (15.0%); immature male, 2 (10.0%); immature, 2 (10.0%). Female was the predominant sex of A. scelestus host spiders (13/20, 65.0%). Host body lengths (mm): range = 8.0–17.5, mean = 13.46 ± 0.84, n = 13 (wasps, range = 11.0–14.0, mean = 12.62 ± 0.25, n = 13) (Fig. 19); host wet weights (mg): range = 49–379, mean = 205.40 ± 57.64, n = 5; (wasps, range = 35–83, mean = 50.40 ± 8.65, n = 5); spider:wasp wet-weight ratios: range = 1.11:1–9.97:1, mean = 4.39:1, n = 5. The spider was longer or the same length as the wasp in 9/13 (69.2%) examples (Fig. 19) and heavier than the wasp in all 13 examples. Host spider size was positively correlated with wasp size (Fig. 19). Based on 212 specimens, Evans (1951b) gives 13.5 mm, compared to 12.62 mm from our study, as the mean body length of females of A. scelestus, which would compare more closely to the mean body length of the host spiders. Figure 19. Spider body length plotted against wasp body length in Arachnospila arctus and A. scelestus. Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 62 Amputation of legs: 2/11 (18.2%). Both spiders had a single leg amputated at the coxa-trochanter joint. References: Evans 1951b, 1970; Evans and Yoshimoto 1962; Gwynne 1979; Krombein 1979; Kurczewski 1999, 2010, pers. observ.; Kurczewski and Kurczewski 1968a, 1968b, 1973; Kurczewski et al. 1987; Peckham and Peckham 1898, 1905; Rau and Rau 1918; Wasbauer 1982. Aporinellus completus Banks Collection dates: (49; 13 May–27 September). Aporinellus completus has at least 2 generations per year in the eastern Great Lakes Region. Collection dates with prey: 30 July–25 August 1896 or 1897 (A. ?completus) (WI), 21 August–14 September 1967, 4 September 1968, 10–23 September 1969, 27–31 July 1970, 17 August 1971, August–September 1986. Localities: PI (12), WG (2), WE (2), GC (1). Habitat: Bare sandy and gravelly soils at deciduous woodland edge, relict and degraded sand dunes, and inland sand blowout. Burrow excavation apparatus: Mandibles; forebasitarsus with long and slender comb-spines, and the apical spine as long as 2nd tarsal segment. The foretarsus is adapted for excavating in coarse-grained sandy or gravelly soils. Host family and species: SALTICIDAE – Eris militaris, Evarcha hoyi, Habrocestum pulex (Hentz), Habronattus borealis, H. sp. probably borealis, H. viridipes, H. spp., Maevia inclemens, Phidippus clarus, Salticus scenicus, Sitticus floricola palustris (Peckham & Peckham). Aporinellus completus provisioned its nests with jumping spiders (Salticidae) in the eastern Great Lakes Region. Peckham and Peckham’s (1898) observations on A. fasciatus (Smith) (= A. medianus Banks) probably pertain to A. completus based on prey type (Maevia inclemens, Phidippus sp., Salticus sp.) (Salticidae) (Evans 1951b; F.E. Kurczewski, pers. observ.). Host sexes and stages: Adult female, 6 (30.0%), immature female, 1 (5.0%), adult male, 2 (10.0%), immature male, 4 (20.0%), immature, 7 (35.0%). Immature jumping spiders (12/20, 60.0%) were the predominant hosts of A. completus. Host body lengths (mm): range = 4.5–6.0, mean = 5.40 ± 0.14, n = 15 (wasps, range = 5.0–7.0, mean = 6.23 ± 0.14, n = 15) (Fig. 20); host wet weights (mg): range = 16–20, mean = 18.33 ± 1.20, n = 3 (wasps, range = 6–9, mean = 7.33 ± 0.88, n = 3); spider:wasp wet-weight ratios: range = 2.22–3.17:1, mean = 2.56:1, n = 3. The wasps were longer or the same length as their host spiders in all examples except 1 (14/15, 93.3%, r = -0.224, P = 0.423) (Fig. 20), but all spiders weighed 2 or 3 times more than the wasps. Amputation of legs: 0/18. References: Evans 1951b; Evans and Yoshimoto 1962; Krombein 1979; Kurczewski 1999, pers. observ.; Kurczewski and Edwards 2012; Kurczewski and Kurczewski 1968b, 1973; Kurczewski and Pitts 2011; Kurczewski et al. 1987; Wasbauer 1982. Aporinellus medianus Banks Collection dates: (9; 24 May–3 October). There are 2 generations per year in the eastern Great Lakes Region. Northeastern Naturalist 63 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Collection dates with prey: 15 August 1926, 19–29 September 1964, 13 October 1965, 3 October 1966. Localities: KS (3), AU (1); additional host records from the upper Midwest, Southeast, and CA. Habitat: Sand dunes, sandy garden, sand pit bordering woodland, and sandy field. Burrow excavation apparatus: Mandibles; foretarsal comb of long slender spines; forebasitarsus with 3 comb-spines, and the apical spine is equal in length to 2nd foretarsal segment. Host families and species: OXYOPIDAE – Oxyopes salticus; MITURGIDAE – Cheiracanthium inclusum; PHILODROMIDAE – Tibellus duttoni, T. oblongus; THOMISIDAE – Xysticus sp. near gulosus. Aporinellus medianus is polyphagous, preying on 4 families of cursorial-hunting and retreat-dwelling spiders. If the host records for Maevia inclemens, Phidippus sp., and Salticus sp. actually pertain to A. completus (Evans 1951b), instead of A. fasciatus (= A. medianus) (Peckham and Peckham 1898), then this strengthens the case for A. medianus being polyphagous and unlike other species in the genus insofar as exclusive host selection on Salticidae and/or Thomisidae. Host sexes and stages: Adult female, 7 (63.6%); immature male, 1 (9.1%); immature, 3 (27.3%). Adult females of small spider species are the preferred host stage and sex. Host body lengths (mm): range = 4.0–12.0, mean = 7.42 ± 1.19, n = 6 (wasps, range = 5.0–9.5, mean = 7.25 ± 0.59, n = 6) (Fig. 20); host wet weights (mg): range = 12–22, mean = 16.67 ± 2.91, n = 3 (wasps, range = 7–11, mean = 8.67 ± 1.20, n = 3); spider:wasp wet-weight ratios: range = 1.50–2.29:1, mean = 1.93:1, n = 3. Figure 20. Spider body length plotted against wasp body length in Aporinellus completus, A. medianus, and A. taeniatus. Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 64 Four of 6 (66.7%) host spiders were as long as the wasps (r = 0.778, P = 0.069; Fig. 20), and all spiders weighed more than the wasps. Amputation of legs: 0/11. References: Evans 1951b; Evans and Yoshimoto 1962; Krombein 1959, 1979; F.E. Kurczewski, pers. observ.; Kurczewski and Acciavatti 1990; Kurczewski and Edwards 2012; Kurczewski and Kurczewski 1968a; Peckham and Peckham 1898; Powell 1985. Aporinellus taeniatus (Kohl) Collection dates: (21, 28 June–22 September [PI], 13 June–19 September [CT]). There are probably 2 generations per year in the northeastern US, but not enough host records to confirm this aspect of its life history . Collection dates with prey: 7 August 1968, 7 July 1969, 1–12 September 1976, 23 August 1978. Localities: PI (6), WE (2), additional host records from central coastal CA from 2010–2012 (28). Habitat: Sand dunes, sand plain, and sandy car trail in pine barrens. Burrow excavation apparatus: Mandibles; forebasitarsus with 2 or 3 combspines, and the apical one is approximately equal to length of 2nd tarsal segment. Foretarsal digging comb is adapted for coarse-grained sandy soils. Host family and species: SALTICIDAE – Habronattus calcaratus (Banks), H. agilis, H. borealis, H. viridipes, H. sp. Aporinellus taeniatus is oligophagous on jumping spiders (Salticidae) of the genus Habronattus. Host sexes and stages: Adult female, 18 (52.9%); immature female, 3 (8.8%); adult male, 5 (14.7%); penultimate male 2 (5.9%); immature, 6 (17.6%). Female spiders (21/34, 61.8%) were the preferred host sex and adult spiders (23/34, 67.6%), the preferred host stage. Host body lengths (mm): range = 4.0–7.0, mean = 5.97 ± 0.17, n = 29 (wasps, range = 5.8–7.5, mean = 6.60 ± 0.09, n = 29) (Fig. 20); host wet weights (mg): range = 9–24, mean = 15.33 ± 4.49, n = 3 (wasps, range = 8–13, mean = 10.33 ± 1.45, n = 3); spider:wasp wet-weight ratios: range = 1.00–2.40:1, mean = 1.51:1, n = 3. Twenty-seven of 29 (93.1%) wasps were the same length or longer than their host spider (r = 0.527, P = 0.003) (Fig. 20), but all spiders were heavier than or as heavy as the wasps. Amputation of legs: 0/34. References: Evans 1951b; Evans and Yoshimoto 1962; Krombein 1964, 1979; Kurczewski 1999, pers. observ.; Kurczewski and Edwards 2012; Kurczewski and Kurczewski 1973, 1987b; Kurczewski et al. 1988. Aporinellus wheeleri Bequaert Collection dates: (205; 22 May–2 October). There are 2 generations per year in the eastern Great Lakes Region. Collection dates with prey: 21 July–2 October 1979, 9 July–27 September 1980, 15 June–14 September 1981, 1 June–14 September 1982, 6 July–6 September 1983, 11 June–19 September 1984, 19 June–20 September 1985 (WG); 4–19 September 1985, 26 May–27 July 1986, 28–31 July 1987 (AU). Northeastern Naturalist 65 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Localities: WG (194), AU (6). Habitat: Gravelly and shalely soils in abandoned overgrown gravel pit (Fig. 2), edge of gravel quarry, and gravelly roadside deposits. Burrow excavation apparatus: Mandibles; forebasitarsus with 2 short combspines and an additional moderate length spine just below basal comb-spine; apical basitarsal comb-spine slightly more than half as long as 2nd basitarsal segment and not adapted for coarse-grained sandy soils. Host families and species: THOMISIDAE – Xysticus ferox, X. sp.; SALTICIDAE – Eris sp., Habronattus borealis, H. decorus, Metaphidippus protervus, M. sp., Tutelina elegans. Habronattus borealis (161/200, 80.5%) and H. decorus (27/200, 13.5%) accounted for the vast majority of host records. Xysticus ferox and X. sp. (Thomisidae, 1 record each) were extremely rare host species. Host spiders were larger in late summer (adult and subadult females) and late spring (mainly overwintering adults) and smaller in mid-summer (mostly immatures). Host sexes and stages: Adult female, 27 (13.5%); immature female, 94 (47.0%); adult male, 14 (7.0%); immature male, 48 (24.0%); immature, 17 (8.5%). Immature spiders (159/200, 79.5%) were the predominant host stage and female spiders (121/200, 60.5%), the predominant host sex. Host wet weights (mg) (Salticidae): range = 22–34, mean = 27.00, n = 5 (wasps, range = 8–11, mean = 8.80, n = 5); spider:wasp wet-weight ratios: range = 2.75– 3.09:1, mean = 3.07:1, n = 5; host wet weights (mg) (Thomisidae): 12, n = 1 (wasp, 7, n = 1); spider:wasp wet-weight ratio: 1.71:1, n = 1. Only the 5 largest salticids and their wasps and 1 thomisid and its wasp were weighed. Amputation of legs: 0/200. References: Evans 1951b; Kurczewski 1999, pers. observ.; Kurczewski and Acciavatti 1990; Kurczewski and Kurczewski 1987b; Kurczewski et al. 1988. Discussion Species in the tribe Pepsini were more numerous in eastern Great Lakes Region deciduous woodland (10 species) than in abandoned overgrown fields and woodland edges (2) or open sandy, gravelly, and loamy areas (2). There were more species of Ageniellini in abandoned overgrown fields and woodland edges (6 species) than in deciduous woodland (2) or open sandy, gravelly, and loamy areas (1). Species in the tribe Pompilini were predominant in open sandy, gravelly, and loamy areas (20 species), not uncommon in abandoned overgrown fields and woodland edges (11), but scarce in deciduous woodland (3). The preponderance of Pepsini in deciduous woodland, Ageniellini in abandoned overgrown fields and woodland edges, and Pompilini in open sandy, gravelly, and loamy areas is connected with female morphological adaptations related to specific habitat and nesting requirements, narrowness (oligophagy) or broadness (polyphagy) in host-spider selection, and microclimate, length of flight season, and number of generations per year. For example, most North American Pompilini (85%) have a rake or comb on the foretarsus for excavating a burrow in bare, friable soil (Evans 1950, 1951a, 1951b). Species of Pepsini and Ageniellini do not use a foretarsal comb or rake for burrow Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 66 excavation but, instead, have hind tibial (Pepsini) or mouthpart and metasomal structural modifications (Dipogon, Ageniellini) (Townes 1957). Small pompilid species (mean body length < 8 mm) were numerous in open sandy, gravelly, and loamy areas (8 species), and deciduous woodland (6), and less so in abandoned overgrown fields and woodland edges (3) in the eastern Great Lakes Region. The number of medium-size pompilid species (8–13 mm) was similar in open sandy, gravelly, and loamy areas (9 species), abandoned overgrown fields and woodland edges (11), and deciduous woodland (9). Large spider wasp species (>13 mm) were found in open sandy, gravelly, and loamy areas (6 species) and abandoned overgrown fields and woodland edges (5), but not in deciduous woodland (0). Some large species, such as Anoplius aethiops, entered deciduous woodland to hunt for prey (Gladicosa gulosa, Hogna frondicola, H. helluo) but nested in more open sunlit areas. In addition to being limited by a scarcity of large host spider species in deciduous woodland, large spider wasps require a high amount of radiant energy in order to maintain optimal body temperature. Many large species such as Entypus unifasciatus, Tachypompilus ferrugineus, Anoplius aethiops, A. atrox, A. cleora, A. nigritus, A. semicinctus, and A. depressipes have heavily infuscate or deeply fuliginous, often violaceous wings that function in absorbing radiant energy in an open, often sunny environment. Large pompilids usually preyed on large spiders, medium-size spider wasps on medium-size spiders, and small pompilids on small spiders in an almost straightline relationship (Fig. 21; Kurczewski and Kurczewski 1968a, 1968b, 1972, 1973). Body-length measurements for host Araneidae (orb-weaver spiders) or Thomisidae (crab spiders) were usually less than those for other spider families of the same weight because of their compact body shape and rotund, as opposed to ovoid or ovoid-elongate, abdomen. Large pompilids preyed mainly on adult spiders, and small spider wasps sought out mostly immature spiders (Kurczewski and Kurczewski 1968a, 1968b, 1972, 1973). The largest spider wasps captured a high percentage of adult, penultimate, and subadult female spiders, whereas the smallest pompilids usually provisioned with a large proportion of immature spiders that hadn’t reached sexual maturity (Fig. 22). Anoplius cleora, a large species, was an exception to this rule (Fig. 22). This species preyed nearly exclusively on Arctosa littoralis, a rather large wolf spider found on sandy shorelines along the Great Lakes. Anoplius cleora captured a large proportion of slightly smaller, unsexed immature spiders in July–August and a preponderance of larger adult female and male spiders in September–October (Figs. 11, 12). Conversely, some small pompilids such as Anoplius ithaca, Ammosphex angularis, and Aporinellus taeniatus captured an inordinately high percentage of adult and subadult female spiders of families and genera that contain small species (Fig. 22). Large spider wasp species were mostly univoltine with only a single generation per year in the eastern Great Lakes Region (9/11 species, 81.8%). Episyron biguttatus and Arachnospila scelestus, 2 of the smaller large pompilids, usually had 2 generations per year under optimal weather patterns. Anoplius cleora, a larger spider wasp species, sometimes had a partial second generation if the first Northeastern Naturalist 67 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 generation emerged in early summer and atypically warm, summer-like conditions persisted through early fall (Evans 1951a, Kurczewski 1999). Large spider wasps have considerably more biomass than small or medium-size pompilids, usually take longer to develop underground from egg to adult stage, tend to emerge in mid–late summer, and provision their nests mainly with large adult, penultimate, or subadult female spiders that reach maturity in late summer. Small and medium-size pompilid species were mainly bivoltine with 2 generations per year in the eastern Great Lakes Region (36/46 species, 78.3%), especially in deciduous woodland (12/15, 80.0%). Small spider wasps probably develop more rapidly from egg to adulthood than large pompilids because of their considerably less biomass. Many small species typically flew intermittently from late spring to early fall and had an inclusive flight season of approximately 4 successive months with optimal weather patterns. Priocnemis minorata (mean body length = 11.4 Figure 21. Host spider mean body length plotted against pompilid species mean body length. Regression line is based on observed data (≥10 host records per species): ● designates cursorial-hunting, burrowing, and retreat-dwelling spider species; o designates orb-weaver spider species. Abbreviations are as follows: Eu = Entypus unifasciatus; Tf = Tachypompilus ferrugineus; Aa = Anoplius aethiops; An = Anoplius nigritus; Ac = Anoplius cleora; Eb = Episyron biguttatus; Ar = Arachnospila scelestus; Pm = Priocnemis minorata; Av = Anoplius virginiensis; Al = Anoplius splendens; Ap = Anoplius apiculatus; Ab = Anoplius imbellis; Eq = Episyron quinquenotatus; Am = Anoplius marginatus species-complex; Au = Auplopus mellipes; As = Anoplius semirufus; Pg = Priocnemis germana; At = Anoplius ithaca; Cf = Caliadurgus fasciatellus; Ag = Ammosphex angularis; Pc = Priocnemis cornica; Ae = Aporinellus taeniatus; Ps = Priocnemis scitula; and Ao = Aporinellus completus. Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 68 mm) and Anoplius carolina (10.3 mm), 2 of the larger medium-size pompilid species in deciduous woodland, were both univoltine with abbreviated flight seasons: P. minorata in mid–late spring, and A. carolina in mid-summer (Evans and Yoshimoto 1962, Kurczewski 1999, Yoshimoto 1954). Small and medium-size pompilid species tended to be more polyphagous (26/46 species, 56.5%) and less oligophagous (20/46, 43.5%), whereas large spider wasps were unequivocally oligophagous (11/11, 100.0%) in the eastern Great Lakes Region. Some of the small and medium-size pompilid species were strongly polyphagous: Priocnemis minorata (9 host families of spiders), P. cornica (11), P. germana (6), P. scitula (8), Auplopus carbonarius (8–12 in Europe), Anoplius marginatus species-complex (12), A. splendens (10), A. tenebrosus (8), and Ammosphex angularis (8). The largest spider wasps in the eastern Great Lakes Region were invariably oligophagous: Entypus unifasciatus (2 similar host families of spiders), Episyron biguttatus (1), Poecilopompilus interruptus (1), Tachypompilus ferrugineus (2), Anoplius aethiops (1), A. atrox (2), A. cleora (1), A. nigritus (2), A. semicinctus (1), and A. depressipes (1). Such oligophagy is undoubtedly connected with the correlative sizes of the spider wasp species and their host spider species. Bivoltine spider wasps tended to be polyphagous (23/38 species, 60.5%), preying on several unrelated families of spiders, often a combination of various Figure 22. Percent adult, penultimate, and subadult female spiders plotted against pompilid species mean body length. Regression line is based on ≥10 host records per species and was created using computed means and species percent, not total observed data. Illustrated is a strong linear relationship between sex (female) and stage (adult) of spider and increased wasp species body length. ● designates cursorial-hunting, burrowing, and retreat-dwelling spider species; o designates orb-weaver spider species. Pompilid species abbreviations are the same as in Figure 21 plus Pi = Poecilopompilus interruptus; Ax = Anoplius tenebrosus; Ad = A. depressipes; Ai = A. illinoensis; Ay = Arachnospila arctus; and Aw = Aporinellus wheeleri. Northeastern Naturalist 69 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 cursorial-hunting and retreat-dwelling species, whereas univoltine pompilid species were primarily oligophagous (16/19, 84.2%), capturing species belonging to 1 or 2 similar spider families in the eastern Great Lakes Region. Bivoltine pompilid species largely have access to a wide variety of types and stages of spiders throughout late spring, summer, and early fall, while univoltine spider wasps are mainly restricted to hunting fixed stages and sexes of specific families, genera, and species of spiders at certain times of year, especially late summer. Deciduous woodland pompilids were overwhelmingly polyphagous (13/15 species, 86.7%), except for A. carolina, which captured only species of Amaurobiidae. This characteristic of woodland pompilids is not surprising given that there are more families, genera, and species of spiders in deciduous woodland than in other habitats in the eastern Great Lakes Region. Spiders prefer dark and shaded conditions with high humidity (Kaston 1948, Kurczewski and Edwards 2012). Dead leaves and other plant litter on the damp forest floor, decomposing logs and stumps, loose bark, and foliage, and branches and twigs of countless trees and shrubs provide a haven for a multitude of spider types. Spider wasp species from open sandy, gravelly, and loamy areas, abandoned overgrown fields, and woodland edges were mainly oligophagous (28/42, 66.7%) in the eastern Great Lakes Region. Many spider species that inhabit open areas are crespuscular or nocturnal to avoid the solar-generated hot and desiccative ground conditions. Nocturnal activity enables many spider species to avoid predation by strictly diurnal spider wasp species, but crepuscular spider activity often does not provide such protection as many pompilids hunt up to sunset. Anoplius bengtssoni (Regan), the largest pompilid species in the eastern Great Lakes Region (mean body length = 22.0 mm), is crepuscular or nocturnal with enlarged ocelli (simple eyes) for enhanced nighttime vision (Kurczewski 1999). In fact, the holotype female of this species was collected at night inside a lighted house (Evans 1951a)! A psammophilous species with 4 long comb-spines on the forebasitarsus, A. bengtssoni was observed at Presque Isle State Park, PA, in mid–late summer behind the beaches at sunset. This species has never been observed nesting or collected with prey. Two rather large lycosids common at night on/in the sand along the shores of the eastern Great Lakes are the shoreline sand spider Arctosa littoralis and the burrowing wolf spider Geolycosa wrighti. Both species represent potential prey for A. bengtssoni, a species in the predominantly wolf-spider-hunting subgenus Lophopompilus. A third prey option for A. bengtssoni at the edge of the deciduous–coniferous woodland at Presque Isle State Park is the large, often terrestrial fishing spider (Pisauridae) Dolomedes tenebrosus. The spider wasp and host spider species of the eastern Great Lakes Region were categorized ecologically on the basis of their occurrence in specific natural communities. At Presque Isle State Park, PA, from which approximately half of the host records were obtained, spider wasp species and host spider species inhabited a natural-community gradient characterized by increasing amounts of vegetation (Fig. 23) ranging from sand beach through sand dunes (Fig. 24, 25), dry sand plain (when lake level is average or low) (Figs. 26), shrub–dry sand plain savanna, and Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 70 Figure 23. Species are listed taxonomically from top to bottom according to their arrangement in the Family Pompilidae in the Catalog of Hymenoptera in America North of Mexico (Krombein 1979), and their inclusive natural communities at Presque Isle State Park, Erie County, PA (PI) are ordered by increasing amounts of vegetation from sand beach (left) to deciduous–coniferous woodland (right). Episyron quinquenotatus also nested along the Lake Erie shore in a narrow sandy and rocky beach and ln degraded sand dunes 1.6–6.4 km west of the park base where there was little human interference. *Natural community names are modified from Bissell and Bier (1987) and Olivero et al. (2002). Northeastern Naturalist 71 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 oak-dominant savanna (Fig. 27) to mesophytic deciduous–coniferous woodland (Fig. 28). Arctosa littoralis and Geolycosa wrighti were 2 rather large, common wolf spider species of the sand beach, sand dunes, and, less so, dry sand plain at this locality (Figs. 29, 30). They were, more or less, confined to these natural communities because they construct silk-lined cylindrical burrows downward into damp sand (Truman 1942). Farther inland at slightly higher elevation the sand is not moist enough for burrow maintenance and spider burrows tend to cave in rather than hold their tubular form. In the reported records, Arctosa littoralis left its burrow, usually under dimly lit conditions, to wander on the sand surface where it became prey for several pompilid species or it hid beneath driftwood and flat stones during the daytime where it was discovered and pursued by the hunting spider wasps. Anoplius apiculatus (173/173 host records, 100.0%), A. cleora (57/59, 96.6%), A. ithaca (5/10, 50.0%), A. splendens (3/77, 3.9%), A. semicinctus (1/4, 25.0%), A. illinoensis (2/2, 100.0%), and Priocnemis cornica (28/89, 31.5%) all provisioned nests at Presque Isle State Park with different sizes and stages of A. littoralis according to their own size (Fig. 7; Figure 24. Eastern Lake Ontario sand beach and sand dunes, the latter with Ammophila breviligulata Fernald (Beach-grass) and Populus deltoides Marshall (Cottonwood), Southwick Beach State Park, Jefferson County, NY (SB) (A.M. Olivero, New York Natural Heritage Program, Albany, NY, 2013 pers. comm.). Species of Pompilidae that occupied these natural communities included Episyron quinquenotatus, Anoplius cleora, A. apiculatus, A. nigritus, A. semicinctus, A. cylindricus, A. ithaca, and Aporinellus taeniatus. Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 72 Figure 26. Dry sand plain, Presque Isle State Park, PA (PI), with various grasses and forbs, especially Rubus hispidus L. (Swamp-dewberry), July 1968. Species that nested in this natural community included the same species as in Figure 25 plus Priocnemis cornica, Anoplius marginatus species-complex, A. splendens, and Arachnospila scelestus. Figure 25. Relict sand dunes, Southwick Beach State Park, Jefferson County, NY (SB), May 2014 (Mark DeFilippo, Syracuse, NY, 2014 pers. comm.). Species that nested in this habitat later in the year included Episyron quinquenotatus, Anoplius cleora, A. apiculatus, A. nigritus, A. semicinctus, A. cylindricus, and Aporinellus taeniatus. Northeastern Naturalist 73 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Figure 27. Quercus rubra L. (Northern Red Oak) and Pteridium aquilinum (L.) Kuhn (Bracken Fern) savanna and successional northern hardwoods interface, Selkirk Shores State Park, Oswego County, NY (SS), June 1971. Species of Pompilidae that occupied these plant communities included Priocnessus nebulosus, Priocnemis cornica, Caliadurgus fasciatellus, Auplopus mellipes, A. nigrellus, Agenioideus humilis, Episyron biguttatus, E. quinquenotatus, Tachypompilus ferrugineus, Anoplius aethiops, A. atrox, A. semirufus, A. marginatus species-complex, A. splendens, A. subcylindricus, A. depressipes, A. illinoensis, A. imbellis, A. ventralis, Arachnospila scelestus, and Aporinellus completus. Figure 28. Mesophytic deciduous–coniferous woodland, typical of that found along Lake Erie, Grand River Terraces, Morgan Township, Ashtabula County, OH (J. Bissell and R. Boronka, Cleveland Museum of Natural History, Cleveland, OH, 2012 pers. comm.). Likely pompilid species found here include Priocnemis minorata, P. germana, P. scitula, Caliadurgus fasciatellus, Dipogon papago, D. pulchripennis, D. sayi, D. brevis, Auplopus mellipes, A. nigrellus, Agenioideus humilis, Anoplius carolina, A. depressipes, A. ventralis, A. virginiensis, and Arachnospila arctus. Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 74 Figure 29. Arctosa littoralis, adult or subadult female, on driftwood, Wilson Lake shore, Russell County, KS (E.R. Eaton, Colorado Springs, CO, 2013 pers. comm.). Anoplius apiculatus, A. cleora, A. ithaca, and Priocnemis cornica were major predators of this spider species at Presque Isle State Park, PA (PI). Figure 30. Geolycosa wrighti, adult male, on relict sand dune near Wisconsin River, east of Spring Green, Iowa County, WI (L. Bartholomay, Iowa State University, Ames, IA, and P.J. DeVries, University of New Orleans, New Orleans, LA, 2013 pers. comm.). Anoplius nigritus and A. cylindricus were major predators on adult females and immatures of this spider species, respectively, at Presque Isle State Park, PA (PI). F.E. Kurczewski, pers. observ.; Kurczewski and Kurczewski 1968a, 1968b, 1972, 1973; Kurczewski et al. 1987). Geolycosa wrighti occurred in the same natural communities at Presque Isle State Park as A. littoralis, and adult, penultimate, and subadult females or, rarely, penultimate males and large unsexed juveniles of this species were preyed on by Anoplius nigritus (16/16 host records, 100.0%) and A. semicinctus (7/7, Northeastern Naturalist 75 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 100.0% in Colorado; Gwynne 1979). Small, unsexed, immature G. wrighti were preyed upon by A. cylindricus (6/6, 100.0%), A. marginatus species-complex (3/29, 10.3%) and A. splendens (1/77, 1.3%) (Fig. 13). Host G. wrighti captured by A. nigritus and A. semicinctus were more mature and significantly longer and heavier than host G. wrighti preyed on by the other species of Anoplius (P < 0.001). Anoplius nigritus, A. semicinctus, and A. cylindricus captured and paralyzed G. wrighti inside or near their burrow entrance and then modified and used the spider’s burrow as a nest (Evans and Yoshimoto 1962; Gwynne 1979; Krombein 1953a, 1953b, 1958b; Kurczewski 1981, pers. observ.; Kurczewski and Kurczewski 1968b, 1973; McQueen 1978). Anoplius marginatus speciescomplex and A. splendens females transported captured and paralyzed G. wrighti elsewhere and stocked them in separate nests dug by the wasps (F.E. Kurczewski, pers. observ.). Arachnospila scelestus practiced the same nesting behavior in Colorado when preying on Geolycosa rafaelana (8/8, 100.0%; Gwynne 1979). Deciduous–coniferous woodland was at the opposite end of the natural community vegetation-gradient at Presque Isle State Park from sand beach and sand dunes. The spider fauna here was characterized by a number of typical cursorialhunting and retreat-dwelling sylvestral species: Arctosa rubicunda and Gladicosa gulosa (Lycosidae); Agelenopsis utahana (Agelenidae); Wadotes hybridus, W. calcaratus, Coras juvenilis, and Callobius bennetti (Amaurobiidae); Hibana gracilis (Anyphaenidae); Clubiona spiralis (Clubionidae); Agroeca ornata (Liocranidae); and Maevia inclemens (Salticidae) (Table 1). Various sizes, stages, and sexes of these spider species comprised the predominant hosts of 4 common polyphagous woodland pompilid species (listed in order of decreasing wasp mean body length): Priocnemis minorata (11.4 mm), Anoplius virginiensis (10.1), P. germana (8.1), and P. scitula (6.2). Selection of prey spiders was size-related—larger spiders being captured by the larger pompilid species, and smaller spiders, often of the same species but different stages, by the smaller spider wasp species (spider mean body length): P. minorata (9.6 mm), A. virginiensis (9.5), P. germana (7.9), and P. scitula (5.5) (Figs. 4, 5). Priocnemis minorata and Anoplius virginiensis prey included host-spider families with larger species (Lycosidae, Pisauridae, Agelenidae), whereas Priocnemis germana and P. scitula prey incorporated host-spider families with smaller species (Thomisidae, Salticidae) (Table 1). Priocnemis minorata and Anoplius virginiensis captured a low percentage of immature spiders (9.6%, 39.5%, respectively), whereas Priocnemis germana and P. scitula preyed on a high percentage of immature spiders (70.5%, 68.3%, respectively). The percentage of adult female host spiders better illustrates the size relationship between the 4 deciduous woodland spider wasp species and their hosts: Priocnemis minorata (86.8%), Anoplius virginiensis (44.2%), P. germana (25.0%), and P. scitula (12.2%). The remainder of the spider wasp fauna at Presque Isle State Park for which host spider records are available occurred in 1 or more natural communities ranging from sand beach, sand dunes, and dry sand plain to the edge of or just inside deciduous–coniferous woodland (Fig. 23). Caliadurgus fasciatellus, Episyron biguttatus, E. quinquenotatus, Anoplius semirufus, Arachnospila scelestus, Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 76 Aporinellus completus, and A. taeniatus nested in bare or sparsely vegetated sandy soil within this natural-community continuum (Fig. 23). Anoplius aethiops, A. atrox, A. subcylindricus, A. illinoensis, A. imbellis, A. ventralis, and Arachnospila arctus nested near or at the deciduous–coniferous woodland edge where the soil is more fertile, loamy, and somewhat compact. Anoplius imbellis preferred damp loamy or loamy-gravelly soils, whereas A. aethiops, A. atrox, A. illinoensis, and A. ventralis inhabited drier, fine-grained loamy soils with occasional cavities and depressions. Agenioideus humilis and Tachypompilus ferrugineus frequently nested around man-made structures such as rock piles, stone fences, walls, and buildings. Anoplius ithaca was restricted to stony beaches or sand beaches with flat stones and pebbles. Anoplius depressipes is semi-aquatic and nested along lagoons, ponds, and swamps in the interior of the park. Priocnessus nebulosus, species of Dipogon and Auplopus, Anoplius carolina, and Arachnospila arctus inhabited sunlit openings in the deciduous–coniferous woodland. A number of spider species from Presque Isle State Park and elsewhere in the eastern Great Lakes Region served as hosts for multiple pompilid species: LYCOSIDAE: Hogna helluo (Priocnemis cornica, Tachypompilus ferrugineus, Anoplius aethiops, A. cleora, A. semicinctus, A. illinoensis, A. ventralis); Pardosa distincta (Priocnemis cornica, Anoplius semirufus, A. marginatus species-complex, A. splendens, A. imbellis); P. milvina (Priocnemis cornica, Anoplius semirufus, A. insolens, A. splendens, A. imbellis, A. ithaca); Rabidosa rabida (Entypus unifasciatus, Tachypompilus ferrugineus, Anoplius atrox, A. nigritus, A. semicinctus, A. semirufus); Schizocosa avida (Priocnemis cornica, Phanagenia bombycina, Anoplius nigritus, A. semicinctus, A. semirufus, A. marginatus species-complex, A. splendens, A. tenebrosus, A. illinoensis, Arachnospila scelestus); S. crassipes (Priocnemis notha, Anoplius semirufus, A. marginatus species-complex, A. splendens, Arachnospila arctus); Trochosa ruricola (Priocnemis minorata, Anoplius semirufus, A. marginatus species-complex, A. tenebrosus, A. imbellis); T. terricola (Priocnemis minorata, P. cornica, P. notha, Anoplius cleora, A. semirufus, A. marginatus species-complex, A. splendens, A. tenebrosus, A. illinoensis, A. imbellis, A. nigerrimus, A. ventralis, A. virginiensis, Arachnospila arctus); Varacosa avara (Priocnemis minorata, P. cornica, Phanagenia bombycina, Anoplius cleora, A. semirufus, A. marginatus species-complex, A. splendens, A. imbellis, Arachnospila arctus, A. scelestus); PISAURIDAE: Dolomedes tenebrosus (Entypus unifasciatus, Priocnemis minorata, Tachypompilus ferrugineus, Anoplius atrox, A. depressipes); AGELENIDAE: Agelenopsis pennsylvanica (Priocnessus nebulosus, Priocnemis minorata, Ageniella semitincta, Anoplius nigritus, A. semirufus, A. marginatus species-complex, A. splendens, A. virginiensis); AMAUROBIIDAE: Callobius bennetti (Priocnemis minorata, P. germana, P. scitula, Dipogon calipterus, D. pulchripennis, D. sayi, Anoplius carolina, A. semirufus, A. marginatus species-complex, A. splendens, A. tenebrosus, A. virginiensis, Arachnospila arctus); and SALTICIDAE: Habronattus viridipes (Priocnemis cornica, Anoplius marginatus species-complex, A. splendens, Aporinellus completus, A. taeniatus); Maevia inclemens (Priocnemis scitula, Phanagenia bombycina, Auplopus Northeastern Naturalist 77 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 nigrellus, Ageniella agenioides, Anoplius insolens, A. marginatus species-complex, A. splendens) . Caliadurgus fasciatellus, Agenioideus humilis, Episyron biguttatus, E. quinquenotatus, and Poecilopompilus interruptus were the only species in the eastern Great Lakes Region known to provision their nests exclusively with orb-weaver spiders (Araneidae; Evans 1950; Evans and Yoshimoto 1962; Kurczewski 2001; Kurczewski and Kurczewski 1968a, 1968b, 1972, 1973; Kurczewski and Spofford 1985; Kurczewski et al. 1987). These 5 species are non-specific insofar as the genus and species of host orb-weaver, C. fasciatellus provisioning with 8, A. humilis with 4, E. biguttatus with 7, E. quinquenotatus with 9, and P. interruptus with 5 genera of Araneidae in the eastern Great Lakes Region (Table 2), often sharing the same spider species, albeit different host sizes and stages. Caliadurgus fasciatellus (mean body length = 7.2 mm) captured a preponderance (85.2%) of non-adult spiders commensurate with its small size, whereas the larger E. biguttatus (13.7 mm) and E. quinquenotatus (9.2 mm) preyed on a high percentage (88.9%, 67.7%, respectively) of larger female spiders. Nearly all C. fasciatellus (22/24, 91.7%), E. quinquenotatus (136/144, 94.4%), and E. biguttatus (9/10, 90.0%) females were as long in body length as their rotund host spiders (Fig. 8). The host orb-weaver spiders of the 3 pompilid species were significantly different in mean body length (mm): C. fasciatellus at 5.3 mm; E. biguttatus at 12.7 mm; and E. quinquenotatus at 6.9 mm (P < 0.001). The mean wet body weight of E. quinquenotatus females (27.7 mg) was more than double that of C. fasciatellus females (13.4), while that of E. biguttatus females (57.7) was more than double that of E. quinquenotatus females (P < 0.001). The mean wet host-spider weight of E. quinquenotatus (44.6 mg) was nearly double that of C. fasciatellus (23.9) (P = 0.00956), while that of E. biguttatus (140.7) was about thrice that of E. quinquenotatus. Fewer than half the host spiders of C. fasciatellus (42.9%) outweighed the wasps (Fig. 9). In E. quinquenotatus, nearly all spiders (45/55, 81.8%) weighed as much or more than the wasps, and all 3 prey of E. biguttatus (100.0%) weighed considerably more than the wasps (Fig. 9). Caliadurgus fasciatellus and E. quinquenotatus readily fly with their host spider if it is lighter in weight, the same weight, or slightly heavier than the wasp and may transport it forward instead of dragging it backwards on the ground as in many other pompilid species (Evans and Yoshimoto 1962, Kurczewski 2001, Kurczewski and Spofford 1985). This manner of prey handling allows these 2 pompilid species to nest farther from their source of host spiders than E. biguttatus. Episyron biguttatus, on the other hand, has never been observed to fly with its relatively heavy prey, the wasp has always been reported dragging it backwards on the ground (Kurczewski and Edwards 2012; F.E. Kurczewski, pers. observ.). Five species of Auplopus inhabited the abandoned overgrown fields, woodland edges, and deciduous–coniferous woodlands in the eastern Great Lakes Region. Females of all 5 species build mud nests using the stiff bristles of the mentum (mouthpart) and large clypeus for carrying individual mud pellets and the smooth pygidium (last abdominal tergite) as a trowel to manipulate the mud during cell conNortheastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 78 struction (Evans and Yoshimoto 1962, Townes 1957). Species of Auplopus amputate the host spider’s legs and, sometimes 1 or both pedipalps, at the coxa-trochanter joints to facilitate prey transport and allow the paralyzed prey to fit inside the barrelor ovoid-shaped mud cell (Kurczewski and Edwards 2012). Auplopus mellipes is a medium-size (mean body length = 9.1 mm) species that occurs in moist deciduous woodland, especially at woodland edges, where it often nests on or near man-made structures. Auplopus nigrellus is a small (6.8 mm) species that is largely restricted to moist deciduous woodland where it nests on or near the ground (Townes 1957). All species of eastern Great Lakes Auplopus are seemingly strongly polyphagous, capturing a variety of cursorial-hunting and retreat-dwelling spiders, predominantly in moist deciduous woodland, with the larger A. mellipes taking significantly larger host spiders than the smaller A. nigrellus (P < 0.001; Fig. 10). Four of the largest pompilid species in the eastern Great Lakes Region—Entypus unifasciatus, Tachypompilus ferrugineus, Anoplius aethiops, and A. atrox—sometimes occurred together in abandoned overgrown fields or at deciduous woodland edges in mid–late summer (Evans and Yoshimoto 1962, Kurczewski 1999, Kurczewski and Pitts 2011). All 4 species provisioned their nests with large, predominantly adult female wolf spiders (Lycosidae) and, except A. aethiops, fishing spiders (Pisauridae), showing little difference between wasp species in spider mean body length (P = 0.131; Fig. 3). Entypus unifasciatus (18.7 mm) and T. ferrugineus (17.1) and T. ferrugineus and A. aethiops (17.0) females were not significantly different but E. unifasciatus and A. aethops were slightly different in wasp mean body length (P = 0.035; Fig. 3). Except for A. aethiops, these pompilid species preyed on adult females of Rabidosa rabida (16–20 mm long), a common wolf spider in abandoned overgrown fields (Kaston 1948; Kurczewski 1989b, 1990, 2010; Kurczewski and Edwards 2012). Entypus unifasciatus and T. ferrugineus were photographed fighting over such a paralyzed host spider (Figs. 31–34; Scott 2007). Anoplius semirufus inhabited shrub–dry sand plain savanna and oak-dominant savanna at Presque Isle State Park, often nesting in bare sand bordering deciduous– coniferous woodland. Females were significantly smaller (mean body length = 8.3 mm) than females of A. splendens (9.6) or A. marginatus species-complex (9.1) (P < 0.001; Fig. 14). Anoplius marginatus species-complex females were significantly heavier than A. semirufus females (P = 0.016; Fig. 15). Host selection in A. semirufus incorporated mainly adult females of smaller species and immatures of larger species of 8 genera of wolf spiders (Lycosidae; 120/144 specimens, 83.3%). Anoplius splendens, on the other hand, provisioned its nests with 10 spider families, and A. marginatus species-complex preyed on 12 spider families and a harvestman in the eastern US (Krombein 1979, Kurczewski 2010, Kurczewski et al. 1987). Mean host-spider body length in A. semirufus was 7.8 mm (range = 5–10 mm). There was no significant difference in host-spider body length between A. semirufus and A. marginatus species-complex (P = 0.924). There was no significant difference in host-spider body weight between A. semirufus, A. marginatus species-complex, and A. splendens (P = 0.594). Anoplius semirufus females frequently captured wolf spiders that were longer or the same length (57/114, 50.0%; Northeastern Naturalist 79 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Figure 31. An Entypus unifasciatus female has captured and paralyzed by stinging a Rabidosa rabida, adult female, and is pulling it backwards, dorsal side upward, up the vertical side of a railroad tie grasping it with her mandibles by the end of its 2nd right leg (Scott 2007; S. Scott, Fairfield, IL, 2012 pers. comm.). Figure 32. The Entypus unifasciatus female has temporarily released the Rabidosa rabida adult female and is reconnoitering ahead, presumably searching for a direct path to her preexisting burrow. Another spider wasp, a Tachypompilus ferrugineus female, has entered the area and is inspecting the paralyzed prey before attempting to steal the spider and transport it to a secure hiding place (Scott 2007; S. Scott pers. comm. 2012). Northeastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 80 Figure 34. The victorious Entypus unifasciatus female resumes transporting the paralyzed Rabidosa rabida adult female, dorsal side upward, backwards toward her preexisting burrow, grasping it with her mandibles by a pedipalp (Scott 2007; S. Scott, 2012 pers. comm.). Figure 33. The Entypus unifasciatus female has returned to her paralyzed spider, sees the Tachypompilus ferrugineus female attempting to steal this prey, and violently attacks the intruding wasp, using her mandibles, legs, and stinger (Scott 2007; S. Scott, 2012 pers. comm.). Northeastern Naturalist 81 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Fig. 14) and weighed more (17/19, 89.5%; spider:wasp mean wet-weight ratio = 2.29:1; Fig. 15) than themselves. Anoplius splendens (16/70, 22.9%; 2.11:1) and A. marginatus species-complex (19/76, 25.0%; 1.91:1) captured a variety of shorter and lighter spiders relative to their own body length and weight than A. semirufus (Figs. 14–16). Anoplius splendens is closely related to species of the A. marginatus speciescomplex and had the same range in body length (7–12 mm), although A. splendens females were significantly longer than A. marginatus species-complex females (P < 0.017; Fig. 16). The host spiders of the 2 species were similar in body length: A. splendens mean = 7.6 mm (range = 5–13 mm) and A. marginatus speciescomplex mean = 7.8 mm (range = 5–11 mm) (P = 0.493; Fig. 16). Both taxa were common inhabitants of dry sand plain, shrub–dry sand plain savanna, and oak-dominant savanna at Presque Isle State Park, but A. splendens was more psammophilous than species of the A. marginatus species-complex and sometimes entered the sand dunes in order to capture prey. Nine (69.2%) families, 15 (44.1%) genera, and 20 (31.7%) species comprised common host spiders of the 2 wasp taxa in Erie County, PA (Table 3; Kurczewski 1968a, 1968b, 1973; Kurczewski et al. 1987). The 2 taxa were strikingly similar in their percentage of host adult female and unsexed immature spiders combined: A. marginatus species-complex, 77.1%; and A. splendens, 75.0%. Unsexed immature spiders are considered as female behaviorally and ecologically because they resemble this sex in general appearance, do not distinguish themselves as males in size or behavior, and inhabit the same niches as the females (Gertsch 1949). Anoplius marginatus species-complex and A. splendens were also very similar in the percentage of wasps being longer in body length than their host spiders: 75.0% and 77.1%, respectively. Anoplius imbellis is not psammophilous like many congeners at Presque Isle State Park. Instead this species is often found on damp loamy soils that are sometimes near water. Females of A. imbellis excavated burrows from the soil surface or they used existing depressions, as in the related A. nigerrimus (Scopoli), a Holarctic species (Richards and Hamm 1939). Anoplius imbellis was similar to A. semirufus in prey selection, predominantly capturing moderate-size wolf spiders (Lycosidae) and, rarely, other similar-size cursorial-hunting and retreat-dwelling spiders (Kurczewski and Edwards 2012). Anoplius imbellis (mean body length = 9.2 mm) and its host spiders (8.7 mm) were, on average, both larger than A. semirufus (8.3 mm) and its prey (7.8 mm). Anoplius imbellis captured about equal numbers of male (8) and female (10) spiders, and most wasps were longer (23/26, 88.5%) than their host spiders (Kurczewski and Edwards 2012). In contrast A. semirufus captured mainly female spiders (70) with few male spiders (13), and, including immature hosts, spiders were longer than or as long as the wasps in half the examples (57/114). There was a strong correlation between wasp size and host-spider size in A. imbellis (r = 0.875) but not in the related A. ithaca (r = -0.300) (Fig. 17). Anoplius imbellis mainly captured medium-size Trochosa terricola or T. ruricola (Lycosidae), whereas A. ithaca (mean = 8.0 mm) preyed on equivalent-size (7.8 mm) immatures of Arctosa littoralis but also much smaller (5.2 mm) Pardosa distincta, both Lycosidae, in equal numbers. Anoplius imNortheastern Naturalist F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 82 bellis females and their host spiders were significantly longer than A. ithaca females and their host spiders (P < 0.001). The genus Ammosphex contained both polyphagous (A. angularis) and oligophagous (A. michiganensis) species in the eastern Great Lakes Region. Ammosphex angularis was predaceous on no less than 8 families of cursorial-hunting and retreatdwelling spiders throughout its transcontinental range (Kurczewski and Edwards 2012), whereas A. michiganensis captured only species in the crab spider genus Xysticus (Thomisidae) (Krombein 1979). There was a rather strong correlation between wasp size and host-spider size in A. angularis (r = 0.767) but not in A. michiganensis (r = 0.285) (Fig. 18). In A. angularis, the wasps were as long as their host spiders in all (18) examples, whereas in A. michiganensis, the wasps were considerably longer than their shorter and rotund crab spiders (5/5, 100.0%; Fig. 18). Species of Arachnospila were, likewise, polyphagous (A. arctus) or somewhat oligophagous (A. scelestus). Arachnospila arctus captured 6 families of cursorialhunting or retreat dwelling spiders, albeit with a predilection for wolf spiders (Lycosidae; Kurczewski and Edwards 2012), while A. scelestus preyed almost entirely on rather large lycosids and, very rarely, fishing spiders (Pisauridae) and jumping spiders (Salticidae) (Krombein 1979, Kurczewski 2010, Kurczewski et al. 1987). Arachnospila scelestus females and their host spiders were significantly longer than A. arctus and their host spiders (P < 0.001). There was a moderate correlation between wasp size and host-spider size in A. scelestus (r = 0.558) but not in A. arctus (r = -0.054) (Fig. 19). In A. arctus, the wasp was longer than its host spider in nearly all examples (7/8, 87.5%; Fig. 19), whereas, conversely, the spider was longer or the same length as A. scelestus in most examples (9/13, 69.2%; Fig. 19). The genus Aporinellus contains some of the smallest (range in mean body length = 6.2–7.3 mm) pompilid species in the eastern Great Lakes Region. Aporinellus taeniatus was the most psammophilous of the 4 species in the region, occurring mainly on sand dunes and dry sand plain at Presque Isle State Park, PA, and Southwick Beach State Park, NY (Kurczewski et al. 1988). Aporinellus medianus nested in sand dunes and sand plain in central coastal California (Kurczewski and Edwards 2012) and in sandy fields, gardens, roadways, and trails in the northeastern US (Evans 1951b, Kurczewski and Kurczewski 1968a). Evans (1951b) suspected that Peckham and Peckham’s (1898) observations on A. fasciatus (= A. medianus) pertain to A. completus based on the exclusive use of salticid prey, and we support his assumption. Aporinellus completus was found commonly in sandy or gravelly areas at the edge of deciduous woodland in the eastern Great Lakes Region (Kurczewski 1999). Aporinellus wheeleri was restricted to gravelly-shalely areas and, unlike the other 3 species, has a foretarsal digging rake with forebasitarsal comb-spines only half as long as in the other species, commensurate with its non-psammophilous habitat (Evans 1951b, Kurczewski et al. 1988). Aporinellus medianus, the largest of the 4 species (mean = 7.3 mm), was polyphagous in provisioning nests with Oxyopidae, Miturgidae, Philodromidae, and Thomisidae (Krombein 1979, Kurczewski and Edwards 2012). Peckham and Peckham’s (1898) salticid host records for this species must be excluded if their observations pertain instead to A. completus (Evans 1951b, Northeastern Naturalist 83 F.E. Kurczewski and D.H. Kiernan 2015 Vol. 22, Monograph 11 Kurczewski and Edwards 2012). Furthermore, unlike A. medianus females, which often provisioned with spiders as large as themselves, A. completus, A. taeniatus, and A. wheeleri usually captured spiders smaller than themselves (Fig. 20). In regards to host sex and stage, A. completus, the smallest of the 4 species (6.2 mm), preyed mainly on unsexed immature spiders, A. medianus and A. taeniatus (6.6 mm) captured mainly adult female spiders of small species, and A. wheeleri (6.8 mm; Evans 1951b) provisioned mostly with immature female spiders (Kurczewski et al. 1988). There was a rather strong correlation between wasp size and host spider size in A. medianus (r = 0.778), moderate correlation in A. taeniatus (r = 0.527), and slightly negative correlation in A. completus (r = -0.224) (Fig. 20). Acknowledgments We are indebted to Edmund J. Kurczewski, father of the first author, who collected the majority of spider wasps and host spiders from Erie County, PA. We also thank Robert Acciavatti, Steven Alm, John Brennan, Matthias Buck, George Byers, Eric Eaton, Noah Elhardt, Nancy Elliott, Henry Fitch, Grant Gaumer, Wilton Ivie, Benjamin Kaston, Keith Kurczewski, George Matuza, Richard Miller, Verne Pechuman, David Peckham, Tom Pucci, Vince Roth, Margery Spofford, and Richard Walton for pompilid prey records from the eastern Great Lakes Region and elsewhere. Lyle and Eileen Buss and G.B. Edwards provided color photographs, species identifications, and spider wasp and host spider bodylength measurements for Tachypompilus ferrugineus from northern Florida. James R. Wiley confirmed the identity of T. ferrugineus. Front and back cover photographs were kindly provided by Mark DeFilippo, Steve Scott, John Smith, and Naomi Smith. Mark DeFilippo also kindly provided the Figure 25 photograph. Steve Scott allowed us to use his images of Entypus unifasciatus provisioning with Rabidosa rabida and interacting with Tachypompilus ferrugineus. Marie Schmidt sent us photographs and measurements of Auplopus mellipes with prey. Eric Eaton permitted us to use his image of Arctosa littoralis, and Peter DeVries (photographer) and Lyric Bartholomay (collector) provided the image of Geolycosa wrighti. Samuel Marshall confirmed the identity of G. wrighti. Howard E. Evans identified several species of Pompilidae. James Pitts determined Anoplius imbellis and Ammosphex angularis. We acknowledge the host-spider identifications made by the multitude of arachnologists listed under Methods. James Bissell and Renee Boronka assisted with the classification of natural communities from Presque Isle State Park, PA. Sandra Bonanno, Julie Lundgren, and Adele Olivero aided with the classification of Selkirk Shores and Southwick Beach State Parks, NY, natural communities. John Lerg provided the facilities of the Allegan State Game Area, near Fennville, MI. Joe Stoll finalized Figure 1 and reconfigured the front and back cover photographs. Field studies by the first author (F.E. Kurczewski) were aided by a series of grants from Sigma Xi-RESA Grants-in-Aid of Research, Cornell University Faculty Research Fund, National Science Foundation Postdoctoral Fellowship and Undergraduate Research Participation Programs, National Institutes of Health Postdoctoral Fellowship, State University of New York-RESA Grants-in-Aid of Research, Michigan Chapter of the Nature Conservancy, Michigan Department of Natural Resources, and New York State United University Professions. 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