2010 NORTHEASTERN NATURALIST 17(Monograph 6):1–78 Analysis of Ecology, Nesting Behavior, and Prey in North American, Central American, and Caribbean Tachysphex (Hymenoptera: Crabronidae) Frank E. Kurczewski* Abstract - An analysis of ecology, nesting behavior, and prey in 40 North American, Central American, and Caribbean species of the pompiliformis, terminatus, brullii, obscuripennis, and julliani species groups is made. Nesting behavior, prey, and ecological components analyzed include number of generations per year, nesting site, aggregation and nest density, pre-existing burrow use, burrow excavation and leveling behavior, presence or absence of tumulus and temporary closure, orientation, hunting, prey capture, malaxation, mutilation, manner of transport, provisioning time, manner of nest entry, final nest closure, nest structure and dimensions, type and stage of prey, prey size, number of prey per fully provisioned cell, placement of prey in cell, wasp’s egg, seasonal trends in nesting, cleptoparasitism, and adaptability. Introduction Tachysphex is a very large, complex, and highly evolved genus in the Tribe Larrini of the Family Crabronidae (Bohart and Menke 1976). This ground-nesting solitary wasp genus contains 445 species worldwide (Pulawski 2009). The genus is well represented on all continents (Bohart and Menke 1976). Eighty-three species inhabit North America, Central America, and the Caribbean Region (Pulawski 1988). The North American species are especially abundant in the western United States and Mexico. Tachysphex species are all black or black with varying amounts of red on the abdomen and legs. The North American, Central American, and Caribbean species are mainly small in size (6–10 mm). They are difficult to classify, as many of them are very similar morphologically (Williams 1914). Some species have two or more generations per year in the Transition, Upper Austral, and Lower Austral faunal zones. Tropical and subtropical species nest almost continually throughout the year in synchrony with alternating wet and dry seasons (Elliott 1996; Genaro and Sánchez 1992; Pulawski 1974; J.A. Genaro, Museo Nacional de Historia Natural, Habana, Cuba, 2009 pers. comm.; W.J. Pulawski, California Academy of Sciences, San Francisco, CA, 2009 pers. comm.). Species of Tachysphex are mainly cursorial and rapid in their movements (Williams 1914), as the genus name implies (tachys = swift, sphex = wasp). The North American, Central American, and Caribbean species excavate mostly short burrows and shallow cells in sandy, gravelly, or, rarely, loamy soils. They stock their cells with incompletely paralyzed grasshoppers (Acrididae), katydids (Tettigoniidae), crickets (Gryllidae), cockroaches (Blattellidae), and mantids (Mantidae) (Krombein 1979, Pulawski 1988). *PO Box 15251, Syracuse, NY 13215; Fkurczewski@ twcny.rr.com. 2 Northeastern Naturalist Vol. 17, Monograph No. 6 Krombein (1979) briefly summarized the ecology, biology, and prey of Tachysphex from the Nearctic and Caribbean Regions in the Catalog of Hymenoptera in America North of Mexico. Kurczewski (1987a) reviewed the nesting behavior of species from the Nearctic Region in the large pompiliformis species group and presented observations on five of the species. Pulawski (1988) summarized life-history information and prey records for some of the species in his revision of North American, Central American, and Caribbean Tachysphex wasps. A number of ecological and behavioral characteristics define the North American, Central American, and Caribbean species of Tachysphex: • They occur mainly in areas of bare or sparsely vegetated sandy or gravelly soil. • They may be gregarious or solitary. • They usually excavate nests from the soil surface, but some species modify and use the abandoned and active burrows of other wasps and bees. • They excavate the burrow and cell with the mandibles and forelegs before searching for prey. • Their nest entrances may or may not be temporarily closed with soil during prey search; if closed, they must be opened and re-closed with each prey item. • Their prey consists mainly of immature orthopteroid insects; different species groups are prey specific at the family or ordinal level. • They usually transport small prey in flight and large prey on the ground held by their antennae and body with the wasp’s mandibles and legs, respectively. • The final nest closure involves the wasp compactly filling the burrow with soil using the mandibles, forelegs, and end of abdomen. • Their burrows are mostly short and diagonal or, rarely, almost vertical, the enlarged oval to elongate-oval cells being rather shallow. • Their nests are single-, bi- or multi-celled with simple branching side burrows; different species and species groups are rather specific in nest type. • The species tend to stock one or a few larger versus several or many smaller prey per cell, usually in a head inward and ventral side upward position. • The wasp’s egg is often laid on the largest individual, but only after the last prey is placed in the cell. • The small sausage-shaped egg is usually affixed to the soft membrane surrounding the base of a prey’s forecoxa. It extends transversely to the opposite side between the bases of the fore- and midlegs in Caelifera (grasshoppers) and Ensifera (katydids, crickets) or longitudinally posteriorad on the thoracic venter in Blattaria (cockroaches). Pulawski (1988) separated the North American, Central American, and Caribbean species of Tachysphex into four species groups based on external morphological characteristics: (1) pompiliformis group, (2) terminatus group, (3) brullii group, and (4) julliani group. For the purpose of this study, I support Bohart and Menke’s (1976) division of the brullii group into the brullii and obscuripennis groups based on major differences in nesting behavior and prey type. 2010 F.E. Kurczewski 3 The large pompiliformis group contains 57 species in North America, Central America, and the Caribbean Region (Pulawski 1988). When viewed collectively, this species group has somewhat diverse nesting behavior. Such behavioral variation agrees with Pulawski’s (1988, 2007) designation of the group as a “heterogeneous assemblage of unassigned species … lacking the diagnostic [morphological] features of [the] other groups.” Elliott and Kurczewski (1985) mentioned this diversity in nesting behavior and suggested separating the pompiliformis group into subgroups in order to clarify the species relationships. Kurczewski (1987a) identified components in the nesting behavior of 20 species in the pompiliformis group. Some species in the pompiliformis group, such as T. pechumani Krombein, are probably distinct enough on the basis of ecology and nesting behavior to be placed in separate subgroups. Tachysphex pechumani differs from other species in the pompiliformis group by its rather slow and jerky movements, constant waving of the mainly orange antennae, elevation of the wings prior to burrow excavation, genusatypical manner of burrow excavation, rudimentary temporary closure of the nest entrance, reluctance to capture band-winged grasshoppers, and unusual method of prey capture and placement prior to nest entry (Kurczewski 2008b, Kurczewski and Elliott 1978). Tachysphex pechumani resembles other species in the pompiliformis group by capturing rather large nymphal and adult spur-throated (Melanoplinae) and slant-faced (Gomphocerinae) acridids, transporting them on the ground, and stocking one grasshopper in a single-celled nest (Kurczewski 2008b, Kurczewski and Elliott 1978). Tachysphex pompiliformis (Panzer), T. montanus (Cresson), T. aethiops (Cresson), T. orestes Pulawski, T. acutus (Patton), and T. punctifrons W. Fox are morphologically similar, ancestral species in the pompiliformis group (Pulawski 1988). They excavate burrows from the ground surface, but some of the species may modify and use the abandoned and active burrows of other wasps and bees (Alcock 1973; Kurczewski 1987a, 1989; Kurczewski and O’Brien 1988; O’Brien 1987). None of the species makes a temporary closure of the nest entrance. Tachysphex pompiliformis in North America and Europe, T. montanus, and T. acutus dig single-celled nests (Adlerz 1904, Alcock 1973, Kurczewski 1989, Kurczewski and O’Brien 1988, Newton 1956, Peckham and Peckham 1905). Tachysphex aethiops, T. acutus, T. punctifrons, and T. pompiliformis in Europe excavate bi- or multi-celled nests (Adlerz 1904; Alcock 1973; Evans 1973; Kurczewski 1987a, 1989; O’Brien 1987). All six species prey on mostly larger nymphal or adult Acrididae that they transport on the ground. Prey of T. punctifrons and T. pompiliformis are often carried to the nest headfirst and dorsal side upward (Alcock 1973, Bonelli 1966, Kurczewski 1987a, Pulawski 1971). Tachysphex punctifrons and T. acutus capture Melanoplus (Melanoplinae; Kurczewski 1987a, 1989, 2000b), while T. aethiops preys on Oedipodinae (band-winged grasshoppers; Alcock 1973; Evans 1970, 1973; O’Brien 1987; M. Buck, Royal Alberta Museum, Edmonton, AB, Canada, 2009 pers. comm.). One, two, or three prey are stocked in fully 4 Northeastern Naturalist Vol. 17, Monograph No. 6 provisioned cells of five of the species in North America (Alcock 1973; Evans 1970, 1973; Kurczewski 1987a, 1989, 2000b; Kurczewski and O’Brien 1988; Newton 1956; O’Brien 1987; Peckham and Peckham 1898, 1905). Tachysphex pauxillus W. Fox, T. hopi Pulawski, and T. semirufus (Cresson), all in the pompiliformis group, nest in level sandy soil, omit a temporary closure of the nest entrance, and capture moderately large nymphal acridids or tettigoniids (T. semirufus). They transport the prey on the ground and place one or two individuals in a single-celled nest (Evans 1970, Kurczewski and Evans 1986, Newton 1956, Pulawski 1988). Tachysphex laevifrons (F. Smith), T. tarsatus (Say), and probably T. williamsi R. Bohart and T. yolo Pulawski, all in the pompiliformis group, nest in level sandy or gravelly soil, omit a temporary closure of the nest entrance, generally capture moderately large or rather large nymphal (though occasionally small nymphal or rarely adult [T. tarsatus]) acridids and transport them in low flights or on the ground according to their size. They stock one or a few prey in a single-celled nest (Alcock and Gamboa 1975; Elliott and Kurczewski 1985; Evans 1970; Kurczewski 1987b, 1991, 1999, 2000b; Peckham and Peckham 1905; Williams 1914). Tachysphex ashmeadii W. Fox was originally placed in the undatus group (Krombein 1979), but Pulawski (1988) moved this species to the pompiliformis group. Similarities in nesting behavior between this species and T. tarsatus include size of prey, manner of transport, absence of temporary closure, use of Gomphocerinae, Oedipodinae, and Melanoplinae as prey, and stocking one or a few individuals in a single-celled nest (Alcock and Gamboa 1975, Elliott and Kurczewski 1985, Kurczewski 1987a, Pulawski 1988, Williams 1914). Tachysphex antennatus W. Fox, T. krombeini Kurczewski, and T. crassiformis Viereck are morphologically similar species in the pompiliformis group (Pulawski 1988), but they differ in aspects of ecology and nesting behavior (Kurczewski 1987a). Tachysphex antennatus nests in gravelly and loamy soils beneath flat stones, omits a temporary closure, captures and stocks several small nymphal Acrididae per cell, carries them in low flights, and makes a one-celled nest (Buck 2004, Kurczewski and Kurczewski 1987). Tachysphex krombeini nests in sand, leaves the nest entrance open when searching for prey, carries small prey in extended flight, and stocks several nymphal acridids and tettigoniids mixed in a single-celled nest (Kurczewski 1971). Tachysphex crassiformis resembles T. laevifrons in nesting behavior, but stocks from one moderately large to six small grasshoppers in a singlecelled nest (Krombein 1963, Kurczewski 1987b, Williams 1914). Tachysphex psammobius Kohl and T. texanus (Cresson) are morphologically distinct species in the pompiliformis group (Pulawski 1988), but they are similar in nesting behavior. Both species nest in sandy or gravelly soils, leave the nest entrance open while hunting for prey, transport small acridids in extended flight, and stock several to many prey in a one–celled nest (Kurczewski 1987a). 2010 F.E. Kurczewski 5 The terminatus group is a rather small (11 species), homogeneous assemblage of New World species (Pulawski 1988). Tachysphex clarconis Viereck, T. antillarum Pulawski, T. alpestris Rohwer, T. linsleyi R. Bohart, T. terminatus (F. Smith), and T. similis Rohwer nest mainly in level or mildly sloping, barren or sparsely vegetated sandy soils (Kurczewski 2009). Tachysphex apicalis W. Fox nests in sand cliffs and steeply sloping sandbanks (Kurczewski 2009, Kurczewski and Snyder 1968). Under optimal nesting conditions, a finished nest in the terminatus group often contains two to five cells with several or many small grasshoppers usually placed head inward and ventral side upward in each cell (Kurczewski 2009). Except for T. apicalis, nesting behavior in the terminatus group is characterized by several group-specific components. Females spend much time on the sand surface leveling the soil removed from burrow excavation. After completing the burrow, a female temporarily closes her entrance with sand and then makes an orientation flight above the area before flying in search of prey. The mostly small grasshoppers are captured on vegetation, incompletely paralyzed by stinging, and transported to the nest usually in extended flight. The prey individuals are released headfirst in front of the entrance, followed by removal of the sand closure, nest entry, exit, and re-entry with prey. The grasshoppers are usually dragged into the open burrow by an antenna with the wasp’s mandibles (Elliott 1996, Elliott and Kurczewski 1985, Evans 1970, Genaro and Sánchez 1992, Krombein 1964, Kurczewski 2009, Rau and Rau 1918, Strandtmann 1953, Williams 1914). In T. apicalis, leveling the sand and making and removing the temporary sand closure in the entrance are absent due to the limitations of the sand cliffs and steep sloping sandbanks. Provisioning females usually enter the burrow without pausing as they are restricted from releasing the prey on the surface by the mainly vertical situation. The method of final closure in this species is group-atypical in accord with its unusual nesting site (Kurczewski 2009, Kurczewski and Snyder 1968). Only two of the 12 North and Central American species in the brullii group have been observed nesting. Tachysphex belfragei (Cresson) nests in sand, preys on nymphal meadow katydids (Tettigoniidae), transports them on the ground (and probably also in flight), and takes them directly into an open entrance. The nest is relatively deep for a Tachysphex, uncharacteristically almost vertical and single-celled, and contains several prey per cell (Kurczewski 1979). Tachysphex mundus W. Fox, in the brullii group, excavates its nest from the abandoned burrows of other Hymenoptera. Females make an orientation flight above the entrance, hunt and capture small nymphal meadow katydids and, occasionally, tree crickets (Gryllidae), transport them in extended flight, and store several individuals in a single-celled nest (Kurczewski 1979). Tachysphex menkei Pulawski, also in the brullii group, preys on shield-backed katydids (Tettigoniidae; Pulawski 1988). 6 Northeastern Naturalist Vol. 17, Monograph No. 6 Tachysphex inconspicuus (Kirby), T. iridipennis (F. Smith), and T. alayoi Pulawski, all in the obscuripennis group, nest in sandy soils, often behind beaches (Buys 2007, Callan 1993, Genaro 2004, Kurczewski et al., in press, Vesey-FitzGerald 1956). Tachysphex inconspicuus levels the soil removed from burrow excavation, makes a temporary closure of the entrance and orientation flight, and removes the sand fill and enters the nest without releasing the cockroach (Buys 2007; Kurczewski et al., in press). Tachysphex alayoi levels the soil but does not make a temporary closure (Genaro 2004). Prey transport in both species is predominantly in flight, the wasp grasping the prey’s antennae with her mandibles and its body with her legs (Buys 2007; Elliott et al. 1979; Kurczewski et al., in press). Females excavate a rather short and relatively shallow, one- or two-celled nest. They usually provision the cell with a few to several small adult or nymphal cockroaches of the family Blattellidae (Buys 2007; Genaro 2004; Kurczewski et al., in press). There is one reported use of a cricket (Gryllidae) as prey of T. inconspicuus (Buys 2007). The egg is affixed to the base of a forecoxa and extends longitudinally posteriad on the thoracic venter (Genaro 2004; Kurczewski et al., in press). The only record of nesting behavior for North, Central, and South American species in the julliani group is a single, unfinished nest of T. coquilletti Rohwer (Alcock and Gamboa 1975). This species excavates a relatively short arcuate burrow and shallow cell and captures and stocks nymphal Mantidae. The palearctic T. julliani Kohl captures small to moderately large nymphal mantids, carries them in flight or on the ground according to their size, and stocks up to six prey in a fully provisioned cell (Berland 1923; Ferton 1897, 1901). The varied ecology, nesting behavior, and prey of Tachysphex species offer a rewarding opportunity for comparative study (Bohart and Menke 1976). A number of significant papers on the nesting behavior of the nearctic and neotropical Tachysphex have been published since Krombein’s (1979) summary and Pulawski’s (1988) revision. More recent publications describe the ecology, nesting behavior, and prey of many hitherto unstudied North American, South American, and Caribbean species in the pompiliformis, terminatus, and obscuripennis groups, prompting my updated synthesis and analysis of information for this very large and dominant genus of solitary wasps. There has been, to date, little attempt to amalgamate, compare, and analyze the various aspects of the biology and life history of Tachysphex species. This paper analyzes the ecology, nesting behavior, and prey of 40 North American, Central American, and Caribbean species of Tachysphex. The species groups and species are treated and discussed under each section mainly in phylogenetic order following Pulawski (1988). Number of generations per year, nesting site, aggregation and nest density, pre-existing burrow use, burrow excavation and leveling behavior, presence or absence of tumulus and temporary closure, orientation, hunting, prey capture, malaxation, mutilation, manner of transport, provisioning time, manner of nest entry, 2010 F.E. Kurczewski 7 final nest closure, nest structure and dimensions, type and stage of prey, prey size, number of prey per fully provisioned cell, placement of prey in cell, wasp’s egg, seasonal trends in nesting, cleptoparasitism, and adaptability are addressed below. Number of Generations per Year Tachysphex pompiliformis, T. pechumani, and T. laevifrons, all in the pompiliformis group, are univoltine and have obligate diapause in North America (Kurczewski 1971, 1987b, 2008b; Kurczewski and Elliott 1978; Kurczewski and O’Brien 1988; Kurczewski et al. 1970; Pulawski 1988). Tachysphex orestes, T. powelli R. Bohart, T. eldoradensis Rohwer, and T. bohartorum Pulawski, four western montane species in the pompiliformis group (Pulawski 1988), are probably also univoltine and have obligate diapause. The flight season of T. orestes is mainly in June and July (Pulawski 1988). Tachysphex pompiliformis has an abbreviated flight season in cooler northern latitude areas (Alaska, Yukon Territory, Northwest Territories), at higher elevation southward through the western mountains, and from the Great Lakes eastward through the Maritime Provinces (Pulawski 1988). Nesting in this species occurs only in June or July (Alcock 1973; Kurczewski and O’Brien 1988; Newton 1956; Peckham and Peckham 1905; F.E. Kurczewski, pers. observ.). Both T. pechumani and T. laevifrons also have limited flight seasons. Tachysphex pechumani flies mainly in late spring–early summer in the Transition and Upper Austral faunal zones (Kurczewski 2008b, Kurczewski and Elliott 1978, Kurczewski et al. 1970). Adults of the more southern T. laevifrons are active only in spring in the Lower Austral faunal zone (Kurczewski 1971, 1987b, 2000b). Some Tachysphex with facultative diapause have one generation per year at northerly latitudes but two or more generations per year at southerly latitudes. Tachysphex tarsatus is such a species (Fig. 1), and many other North American, Central American, and Caribbean Tachysphex in the pompiliformis group presumably fall in this category (Pulawski 1988). Tachysphex tarsatus has only one generation per year in Wyoming (Evans 1970), but at least two generations a year in Pennsylvania and New York, where it nests from late May through September (Fig. 1; Kurczewski 1991, Kurczewski and Acciavatti 1990). This species is especially common in mid–late summer. Tachysphex tarsatus may have several generations annually in the Carolinas based on its inclusive dates of collection (May–October; Kurczewski 2000b). Tachysphex antennatus is predominantly an early summer species at northern latitudes, although it may be bivoltine in the southern Great Lakes Region (Fig. 1; Buck 2004, Kurczewski and Kurczewski 1987). Most collection records for T. antennatus from northwestern Pennsylvania (81 of 100 specimens, or 81.0%; Fig. 1; Kurczewski and Kurczewski 1987) and from southern Ontario (114 of 130 specimens, or 87.7%; Buck 2004) are for June and July. 8 Northeastern Naturalist Vol. 17, Monograph No. 6 Tachysphex terminatus, in the terminatus group, has at least two generations per year in Pennsylvania and New York (Fig. 1; Kurczewski 2009, Kurczewski and Acciavatti 1990), but it probably has only one generation per year farther northward in Quebec and the Maritime Provinces (F.E. Kurczewski, pers. observ.). Tachysphex terminatus nests from May through October in Kansas (Kurczewski 2009) and Texas (Strandtmann 1953), Figure 1. Number of specimens of Tachysphex antennatus, T. terminatus, and T. tarsatus collected from two locations in Erie County, PA: Presque Isle State Park and 2.4 km southeast of Erie (Kurczewski and Kurczewski 1987; F.E. Kurczewski, pers. observ.). 2010 F.E. Kurczewski 9 producing successive generations at approximately five- or six-week intervals. Tachysphex alpestris and T. clarconis, both in the terminatus group, have only one generation per year in Alaska and Wyoming, respectively (Elliott and Elliott 1973, Evans 1970), but they probably have two or more generations per year in the southwestern US. Species in the terminatus group have quasi-continuous flight periods, including several generations per year, in tropical and subtropical regions (Elliott 1996, Genaro and Sánchez 1992, Kurczewski 2009). Tachysphex apicalis and T. similis, both in the terminatus group, nest from February through November in southern Florida (Kurczewski 2009), and T. similis probably nests during that period in the Bahamas (Elliott 1996). Tachysphex antillarum, another species in the terminatus group, nests throughout the year in Cuba (Genaro and Sánchez 1992). The nesting season of species in the obscuripennis group (T. inconspicuus, T. iridipennis, T. alayoi) fluctuates throughout the year in the tropics in connection with alternating wet and dry periods (W.J. Pulawski, 2009 pers. comm.). In the Bahamas, T. alayoi may nest throughout much of the year, but is especially active during the warmer months (Elliott et al. 1979). In the Caribbean Region, T. alayoi nests from March through December and possibly through the entire year (J.A. Genaro, 2009 pers. comm.; W.J. Pulawski, 2010 pers. comm.). In Central and South America, T. inconspicuus and T. iridipennis probably nest throughout the year (S. Brady, Smithsonian Institution, Washington, DC, 2009 pers. comm.), especially during the dry season (March–April) when rainfall is less and solar radiation is at its peak (Buys 2007; Kurczewski et al., in press; Williams 1941). Nesting Site Tachysphex tarsatus and many other species in the pompiliformis group nest mainly in level sandy or gravelly soil with sparse to moderate amount of vegetation (Fig. 2). Tachysphex antennatus uniquely nests beneath flat stones in gravelly and loamy soil (Fig. 3; Kurczewski and Kurczewski 1987), often in “hard-packed soil” (Buck 2004). Tachysphex acutus nests in sandy or gravelly loam, usually in or at the edges of fields (Fig. 4; Kurczewski 1989). The psammophilous T. pechumani inhabits oak/pine-dominant barrens, savanna, and open woodland (Fig. 5; Kurczewski 2008a, b; Kurczewski and Elliott 1978; Kurczewski et al. 1970). Historically, this species occupied ancestral pine-barrens and oak savanna prior to their disturbance and destruction (Kurczewski 1998, 2000a). Species in the terminatus group (T. clarconis, T. antillarum, T. alpestris, T. linsleyi, T. terminatus, T. similis) nest mostly in level or mildly sloping, barren or sparsely vegetated sandy soil (Fig. 6; Elliott 1996, Elliott and Kurczewski 1985, Evans 1970, Genaro and Sánchez 1992, Krombein 1964, Kurczewski 2009). Tachysphex apicalis, an ecologically atypical species in the group, usually inhabits sand cliffs and steeply sloping sandbanks (Fig. 7; Kurczewski 2009, Kurczewski and Snyder 1968). 10 Northeastern Naturalist Vol. 17, Monograph No. 6 Tachysphex mundus, in the brullii group, nests in gravelly, sandy, silty, or clayey soil as a consequence of renovating and using the pre-existing burrows of other wasps and bees (Kurczewski 1979). Tachysphex belfragei, in Figure 2. Man-made opening in sandy field, 2 km west of Sennett, Cayuga County, NY. Tachysphex terminatus nested in bare sand areas, and T. tarsatus nested in more vegetated, peripheral areas (Kurczewski 1991, Kurczewski and Snyder 1968). Figure 3. Man-made gravel pit, 2.4 km southeast of Erie, Erie County, PA. Tachysphex antennatus nested beneath flat stones in gravelly loam (Kurczewski and Kurczewski 1987). 2010 F.E. Kurczewski 11 this group, probably also renovates and uses pre-existing burrows, but has been found nesting only in sandy soil (Kurczewski 1979). Figure 4. Field growing on gravelly sandy loam, 3 km east of Auburn, Cayuga County, NY. Tachysphex acutus nested in field, and T. tarsatus nested in bare areas along a man-made path (Kurczewski 1989). 12 Northeastern Naturalist Vol. 17, Monograph No. 6 Tachysphex inconspicuus, T. iridipennis, and T. alayoi, all in the obscuripennis group, nest in sandy soil near beaches or inland in man-made areas in the tropics and subtropics (Buys 2007; Callan 1993; Elliott et al. 1979; Figure 5. Man-made vehicle path in pine-barrens, 1 km north of Weymouth, Atlantic County, NJ. Tachysphex pechumani and T. terminatus nested in bare sand (Kurczewski and Elliott 1978). Figure 6. Man-made sandbank, 3 km east of Auburn, Cayuga County, NY in which Tachysphex terminatus and T. similis (rarely) nested. 2010 F.E. Kurczewski 13 Genaro 2004; Kurczewski et al., in press; Vesey-FitzGerald 1956; W.J. Pulawski, 2009 pers. comm.). Tachysphex coquilletti, one of two North, Central, and South American species in the julliani group, nests in arid regions of the southwestern United States in “desert scrubland” (Alcock and Gamboa 1975; F.E. Kurczewski, pers. observ.). Aggregation and Nest Density Species in the terminatus group excavate multi-celled nests with the result that emerging adults are immediately in close contact, thereby ensuring a certain amount of aggregation (Kurczewski 2009). A limited amount of bare or sparsely vegetated sandy soil further enhances the gregarious capacity of the site (Evans 1974). Tachysphex antillarum, T. terminatus, T. similis, and T. apicalis, all in the terminatus group, often form dense aggregations numbering 50–75 or, sometimes, more than 100 females nesting simultaneously at a site (Genaro and Sánchez 1992, Kurczewski 2009, Strandtmann 1953). A “colony” of T. terminatus nesting beneath a farm implement shed in Texas numbered “hundreds of nests,” including 10 cells in one square foot of soil (Strandtmann 1953). An aggregation of T. terminatus in a sloping sandbank in central New York contained upwards of 150 nests in an area 9.7 x 1.8 m (F.E. Kurczewski, pers. observ.). Nests were so dense in the center of the aggregation that five pairs of females used common entrances, with one Figure 7. Western side of man-made drainage ditch near Lakeport, Glades County, fl. Tachysphex apicalis nested in top third of vertical sand cliff, and T. similis nested in level loose sand of field atop cliff (Kurczewski 2009, Kurczewski and Snyder 1968). 14 Northeastern Naturalist Vol. 17, Monograph No. 6 wasp excavating her burrow off the side of another wasp’s entrance. Thirty cells of T. terminatus at another central New York site were found in a square meter of sandy soil (F.E. Kurczewski, pers. observ.). Ninety-five nests of T. apicalis were excavated in a section of vertical sand cliff in southern Florida. Nineteen of the nests were located in an area 55 cm long. Entrances at the center of the aggregation were only 1.7–4.0 cm apart (Kurczewski 2009). Tachysphex inconspicuus, in the obscuripennis group, nests in the tropics in bare sandy soil in large aggregations of more than 100 females (Buys 2007). Tachysphex pechumani, in the pompiliformis group, forms smaller aggregations in bare sand in the central Great Lakes Region, almost never exceeding 50 nesting females (Kurczewski 2008b, Moan and Tramer 2008). In addition to multi-celled nests and limited space enhancing the density of nesting aggregations, periodic grasshopper outbreaks can increase the size and density of Tachysphex aggregations, even in members of the pompiliformis group that make well-spaced, single-celled nests. In Idaho, an outbreak of grasshoppers resulted in 22 prey (cells?) of T. pompiliformis, T. montanus, and several other species in the pompiliformis group being unearthed in a square foot of soil (Newton 1956). Nine pairs of 96 T. tarsatus nests at a site in central New York, where grasshoppers were abundant during an unusually dry year, had entrances as close together as 2.4–12.7 cm (Kurczewski 1991). However, overall, only three of 23 aggregations of T. tarsatus examined during a 30-year-long period had entrances closer together than 15 cm (Kurczewski 1991). Tachysphex females tend to localize their nesting and excavate several consecutive burrows in a small area. In T. pechumani, nests in close proximity usually denote the work of a single wasp (Kurczewski 2008b). Females excavate and complete successive burrows in one area before moving a short distance to make additional consecutive nests in another area. Tachysphex acutus often excavates a second burrow close to or from the filled entrance of a previously finished nest; this behavior is perhaps an early stage in the evolution of bi-celled nests (Kurczewski 1989). Tachysphex terminatus females build a “chain of nests” one “right after another,” an inch apart (Rau and Rau 1918). Such a series of nests in close proximity eliminates the need for an accompanying sequence of orientation flights (Rau and Rau 1918). Pre-existing Burrow Use The majority of Tachysphex excavate burrows beginning from the ground surface. However, some ancestral North American species in the pompiliformis group renovate and use pre-existing burrows of other insects (Pulawski 1988). Tachysphex aethiops generally renovates and uses for nesting the abandoned burrows of other wasps and bees (Alcock 1973, O’Brien 1987), but, Evans (1970) found this species excavating burrows from the ground surface. Some nests of T. punctifrons in sandy soil contain 2010 F.E. Kurczewski 15 long empty spurs implying the takeover and renovation of pre-existing burrows (Kurczewski 1987a). Tachysphex acutus often appropriates the filled entrance of a recently completed conspecific burrow providing easy penetration into the soil, instead of moving elsewhere to begin a new nest from the ground surface (Kurczewski 1989). Tachysphex mundus, in the brullii group, excavates burrows below ground level from the walls of abandoned burrows of other ground-nesting wasps and bees (Kurczewski 1979). The genus-atypical, nearly vertical, relatively deep nest of T. belfragei implies the use of a pre-existing [bee] burrow (Kurczewski 1979). Tachysphex terminatus and T. apicalis, both in the terminatus group, rarely use pre-existing conspecific burrows to start nests. Tachysphex terminatus, when nesting in dense aggregations, may excavate a burrow from an active entrance of another female, the two wasps sharing an enlarged common entrance and temporary closure (Kurczewski 2009). Needless to say, much intraspecific interaction takes place when the two females, each with prey, arrive at the entrance at the same time. Tachysphex apicalis may complete a burrow started and abandoned by another female in the cliff face or steep slope (Kurczewski 2009). Burrow Excavation and Leveling Behavior Many Tachysphex, during excavation, remove soil from the burrow in a common manner. As the mandibles loosen the soil, the forelegs, working in unison, rake it backward beneath the synchronously lifting abdomen. During this behavior, the foretarsi are fishhooked medially to expose the largest number of rake spines (Fig. 8). Even T. mundus, when renovating abandoned bee burrows beneath the surface, uses her mandibles to loosen soil and her forelegs in unison in typical Tachysphex manner to remove the loosened soil from her modified burrow (Kurczewski 1979). Except for T. pechumani, species in the pompiliformis group make little attempt to level the soil removed from burrow excavation. The soil simply accumulates in front of the entrance as an enlarging convex mound or tumulus with each soil-removal episode (Fig. 9). The distance to which T. tarsatus removes soil from its burrow during excavation decreases from start to finish (Kurczewski 1991), while that of T. acutus remains about the same as the tumulus increases in size (Kurczewski 1989). Unlike other species in the pompiliformis group, T. pechumani rarely backs onto the surface when removing loosened sand from its burrow. When she does, the female almost never proceeds backward from her entrance beyond her body length. Instead, remaining below ground level and out of sight, she periodically flings an arc-shaped “spray” of sand backward from inside the burrow. This practice results in a well-distributed, very shallow, dark-colored (charcoal-laced), fan-shaped sand deposit on the lightercolored sand surface (Fig. 10; Kurczewski 2008b). Tachysphex pechumani burrows of equivalent length dug at higher sand surface temperatures take 16 Northeastern Naturalist Vol. 17, Monograph No. 6 less time to excavate than burrows dug at lower sand surface temperatures (Kurczewski 2008b). In the terminatus group, early stages of burrow excavation are similar to those in the pompiliformis group, with the wasp loosening soil with her Figure 9. Tachysphex tarsatus female resting on convex-shaped tumulus following excavation of burrow. Figure 8. Foretarsus of Tachysphex apicalis female fishhooked medially in raking position to show lateral row of long, slender, close-set, flexible rake-spines (Kurczewski 1964). 2010 F.E. Kurczewski 17 mandibles and then raking it backward from the burrow with her forelegs in unison beneath her synchronously lifting abdomen (Kurczewski 2009). However, females gradually proceed backward farther with the sand and, about midway through the excavation, except for T. apicalis, begin extensively leveling the surface in front of the entrance by distributing the sand with the forelegs in various directions. This lengthy behavior removes most traces of the tumulus in preparation for a subsequent temporary closure of the entrance, after which the entire area is more or less indistinguishable from the surrounding sand surface (Elliott 1996, Elliott and Kurczewski 1985, Genaro and Sánchez 1992, Krombein 1964, Kurczewski 2009). Some females of T. terminatus and T. similis spend twice as long or longer leveling the sand surface in front of their entrance as they do excavating below the surface inside their burrow (Kurczewski 2009). Tachysphex apicalis nests in sand cliffs and steep slopes. The soil flung backward with the forelegs during excavation usually falls out of the wasp’s reach. There is no loose soil available near the entrance with which to temporarily close the opening (Kurczewski 2009). Tachysphex inconspicuus, in the obscuripennis group, resembles most species in the terminatus group in leveling the sand removed from burrow excavation in preparation for making a temporary closure of the nest entrance (Buys 2007; Kurczewski et al., in press). Temporary Closure of the Entrance Temporarily closing the nest entrance with soil during the search for prey characterizes the terminatus group except T. apicalis (Kurczewski 2009), Figure 10. Shallow, evenly distributed, fan-shaped, dark-colored tumulus (left) and depressed, sand-filled entrance (white arrow) of Tachysphex pechumani. 18 Northeastern Naturalist Vol. 17, Monograph No. 6 T. inconspicuus in the obscuripennis group (Buys 2007; Kurczewski et al., in press), and, rudimentarily so, T. pechumani in the pompiliformis group (Kurczewski 2008b, Kurczewski and Elliott 1978). This behavior involves the wasp flinging sand backward into the opening with her forelegs beneath her synchronously lifting abdomen in an attempt to fill the entrance. Temporary closure of the nest entrance is probably restricted to species nesting in friable, often mainly barren, coarse-textured sandy or gravelly soils. Tachysphex terminatus spends about twice as long as T. similis for temporary closure of the entrance and removal of the sand fill at the same sand surface temperature. In both species, temporary closure and its removal are made increasingly faster in a straight-line relationship as sand surface temperature rises from 35 to 60 °C. During closure and removal of the sand fill, T. terminatus makes hovering flights into cooler strata of air when surface temperature reaches 55 °C. When surface temperature reaches 59 °C, such flights gradually give way to an aborted temporary closure. Tachysphex similis, on the other hand, does not typically make hovering flights at high sand surface temperatures but, instead, makes increasingly looser temporary closures and leaves the entrance progressively more open above 57 °C. A partly open nest to enable the wasp with prey rapid entry to avoid the hot sand is the end result in both species at very high sand surface temperatures (>58 °C; Kurczewski 2009). The temporary closure of T. pechumani is often loose and incomplete, the entrance frequently remaining partly open and depressed as much as 5–8 mm (Kurczewski 2008b). Even when temporarily fully closed, a 1–3 mm-deep depression identifies the site of an entrance in this species (Fig. 10; Kurczewski and Elliott 1978). Orientation After excavating a burrow and, in some species, making a temporary closure of the nest entrance, females familiarize themselves with their surroundings to facilitate future returns to the nest with and without prey. In the pompiliformis group, T. pechumani makes hovering and turning flights above the entrance during and after temporary closure, although it is not known whether such flights are for orientation or in response to increased sand surface temperature. Females leave the area by walking in enlarging circles or figure eights at lower sand temperatures or making low, circular flights of increasingly larger radii at higher sand temperatures (Kurczewski 2008b). Tachysphex tarsatus orients to its nest entrance by running around the opening on the ground or making low circular flights of increasingly larger radii. Orientation on the ground may include one or more entries into the open burrow, occasionally interspersed with episodes removing additional sand from the opening (Kurczewski 1991). Females of T. acutus, nesting in dense vegetation, run around the entrance interspersed with removing 2010 F.E. Kurczewski 19 additional soil from the burrow before making a low rapid flight to the hunting area (Kurczewski 1989). Other species in the pompiliformis group that prey on larger grasshoppers and transport them to the nest on the ground or in low flights, such as T. laevifrons and T. crassiformis, exhibit similar orientation behavior prior to hunting (Kurczewski 1987b). Females of T. antennatus make circular or figure eight hovering flights above the entrance prior to hunting, capture smaller grasshoppers, and transport them in low flights to the nest. The orientation flights gradually become higher and wider before the wasp departs in a long flight to search for prey (Kurczewski and Kurczewski 1987). Species in the terminatus group make circular orientation flights above the area of the entrance after excavating a burrow and, except for T. apicalis, making a temporary closure of the nest entrance (Elliott 1996, Elliott and Kurczewski 1985, Kurczewski 2009). Females of T. clarconis, T. terminatus, T. similis, and T. apicalis fly in circles of increasingly larger radii, gradually increasing in height, before flying off to hunt for prey (Elliott 1996, Elliott and Kurczewski 1985, Kurczewski 2009). Tachysphex mundus, in the brullii group, makes hovering orientation flights above the area of its entrance, always facing the opening, before flying off in search of prey (Kurczewski 1979). Tachysphex inconspicuus, in the obscuripennis group, makes orientation flights above the area of the entrance after making its initial temporary closure of the nest entrance (Buys 2007; Kurczewski et al., in press). Tachysphex coquilletti, in the julliani group, orients to its nest by flying slowly a few centimeters above the ground in an outward spiral, its abdomen “waggling” strongly until it darts off to hunt for prey (Alcock and Gamboa 1975). Hunting Tachysphex pechumani hunts for prey in vegetation bordering the sand where it nests, usually no farther than 7–10 m from the entrance. Females walk or run on the ground or low vegetation in a zigzag manner, passing by suitable-sized individuals of Dissosteira carolina (L.) (Carolina Grasshopper), Spharagemon collare (Scudder), and S. bolli Scudder, all band-winged grasshoppers of the subfamily Oedipodinae (Kurczewski 2008b, Moan and Tramer 2008). Some hunting females make hovering flights 20–30 cm above the ground, interspersed with slow winding flights among upright plants. If unsuccessful in capturing prey, females make short, quick flights to new hunting areas or they return periodically on the ground to their nests. At the nest, a female examines her partly sand-filled entrance or she removes the sand closure, enters, exits, re-closes the entrance, and returns to the hunting area on the ground (Kurczewski 2008b). Some other species in the pompiliformis group, as exemplified by T. tarsatus, T. laevifrons, T. crassiformis, T. ashmeadii, T. pompiliformis, and T. acutus, hunt randomly in any direction from the nest but usually 20 Northeastern Naturalist Vol. 17, Monograph No. 6 no farther than 5–10 m away. They run on the ground or low vegetation, pivoting and turning, and constantly tap their antennae on the substrate interspersed with short, low, quick flights. In the spring, females of T. tarsatus hunt smaller grasshoppers farther from their nests than during mid– late summer, when prey capture of larger individuals may occur only 1–5 m from the entrances. If unsuccessful in capturing prey, the wasps fly quickly to a new hunting area or they return periodically to their nests. They enter, exit, sometimes remove loose sand from the burrow, often re-orient, and return to the hunting area on the ground or in flight (Alcock 1973; Elliott and Kurczewski 1985; Kurczewski 1987b, 1989, 1991; Kurczewski and O’Brien 1988). Tachysphex antennatus searches for prey 2–10 m from its nest. Females hunt in low, rapid flights and by running on the ground and low vegetation (Kurczewski and Kurczewski 1987). Species in the terminatus group fly to the nearest source of vegetation to hunt for prey, sometimes a distance of 15–60 m from the barren sand in which they nest. They can hunt at such distances from their nests because the grasshoppers they capture are small in size and can be transported relatively quickly in flight. Hunting females hover in flight next to upright plants, sometimes land and walk on the stems and leaves, and tap their antennae constantly. Rarely, where there are no upright plants, T. similis and T. apicalis search for smaller nymphal grasshoppers on low vegetation near the ground (Kurczewski 2009). If unsuccessful in capturing prey, females in the terminatus group, except T. apicalis, fly to other vegetation or they return periodically to their nests, land, and examine the temporary sand fill or remove the closure, enter, exit, refill the entrance, and fly back to the hunting area. Females of T. apicalis unsuccessful in prey capture fly back to their nests and then return to hunt in flight without entering the burrow or they enter, exit, and fly back to the hunting area (Kurczewski 2009). Tachysphex mundus, in the brullii group, searches for prey by making short, quick flights and running rapidly through grasses at ground level as well as flying into higher and denser vegetation where it disappears from view (Kurczewski 1979). Females of T. inconspicuus, after making the initial temporary closure or placing a prey in the nest and re-closing the entrance with soil, fly almost straight upward into the tropical wet forest canopy to hunt for additional blattellid cockroaches (Kurczewski et al., in press). Prey Capture Not all attempted captures of Acrididae are successful. The grasshoppers exhibit a repertoire of anti-predator strategies that enable them to prevent or elude capture (Steiner 1981a). The easiest method of prevention or escape is for the grasshopper to remain perfectly still or leap away. Some adult band-winged grasshoppers attempt to confound prey capture by rapidly unfolding their wings and displaying their bright colors (Steiner 2010 F.E. Kurczewski 21 1981a). This “startle display” is effective in discouraging prey capture by T. ashmeadii (Williams 1914). Hunting females in the pompiliformis group, including T. laevifrons, T. tarsatus, and T. crassiformis, are sometimes discouraged by larger grasshoppers lifting their hindlegs in a threatening manner or repelled by a well-directed kick from the grasshopper’s hindlegs (Kurczewski 1987b, 1991; Steiner 1981a; Williams 1914). Tachysphex tarsatus has been observed rubbing the end of its abdomen in the soil in an attempt to remove dark semifluid, an apparent repellent, regurgitated by its prospective acridid prey (Fig. 11; Kurczewski 1991, Steiner 1981a). Some stung grasshoppers are apparently unaffected by the wasp’s venom, immediately right themselves, and walk or leap away (Kurczewski 1987b, 1991). The different sizes of the wasps and potential prey and appropriate versus inappropriate prey type determine successful or unsuccessful prey capture. Tachysphex laevifrons, T. crassiformis, and T. tarsatus approach grasshoppers much larger than themselves and obviously beyond prey size range, touch them with their antennae, or alight on their dorsum, but quickly fly off without attempting capture (Kurczewski 1987b, 1991). Tachysphex pechumani preys on moderately large grasshoppers belonging to the subfamilies Gomphocerinae and Melanoplinae, but passes by appropriate-sized individuals of Oedipodinae without attempting capture (see above, Kurczewski 2008b). Prey capture is often difficult to observe and study in the field, especially among dense vegetation. One sting is sometimes sufficient to subdue, at Figure 11. Tachysphex tarsatus female inserting sting near base of right midcoxa of nymphal Dissosteira carolina. The grasshopper has regurgitated dark, viscous semifluid defensively against the wasp’s attack (Kurczewski 1991). 22 Northeastern Naturalist Vol. 17, Monograph No. 6 least temporarily small grasshoppers, but four to eight stings may be necessary to immobilize large prey of T. tarsatus (Kurczewski 1991). Tachysphex laevifrons and T. crassiformis may deliver up to five stings in order to subdue larger grasshoppers (Kurczewski 1987b). It is not known whether this number represents a partial or complete stinging sequence. Tachysphex pompiliformis prey are paralyzed by four consecutive, well-placed stings in the ventral thorax and throat under artificial conditions in the laboratory, beginning with a sting in the intersegmental membrane surrounding the base of the grasshopper’s hindleg (Steiner 1981b). Females in the pompiliformis group always sting their prey on the ground, the grasshopper being ventral or dorsal side upward or on its side (Fig. 11). Maintaining her body at about a 90° angle to the longitudinal axis of the prey and clinging to its dorsum, the wasp bends her abdomen underneath and inserts the sting into the underside of the grasshopper’s thorax (Kurczewski 1987b, 1991, 2008b; Williams 1914). The first two stings are usually applied to the hind and forecoxal corium (soft intersegmental membrane) surrounding the bases of the hind- and foreleg (Steiner 1981b). Although the sequence of sting placement in T. pechumani is similar to that of other species in the pompiliformis group, prey capture in this species is unique. Females approach the grasshopper on the ground face to face and circle it clockwise or counter-clockwise, alternately waving their mainly orange antennae. Wasps may circle the grasshopper up to two times in one direction and once in the opposite direction before pouncing on it from the side and stinging it as described above (Kurczewski 2008b). Prey capture in the terminatus group is less demonstrative than in the pompiliformis group because of the smaller sizes of the prey. Once the grasshopper is located on vegetation, the wasp’s slow, winding, and hovering flight transforms into a lightning-fast forward flight to prevent the grasshopper from leaping away. The small nymphal grasshopper is pulled from the plant with the wasp’s legs and mandibles and flown to the ground, where it is stung up to three times in the intersegmental membranes surrounding the leg bases (Kurczewski 2009). Whether or not this sequence of stings represents the full complement is unknown. Prey are stung into a state of partial paralysis with rhythmic movements of the antennae, mouthparts (especially palpi), legs, and abdominal segments. The legs and antennae move less frequently than the mouthparts. The age and condition of the wasp may play a role in the degree and duration of the paralysis. Although most grasshoppers never recover from the effect of the venom, occasionally some prey are able to resume normal activity almost immediately upon being released (Kurczewski 1991). Malaxation Malaxation (Ferton 1901) involves the wasp kneading the soft intersegmental membrane surrounding the prey’s forecoxa or forecoxae with 2010 F.E. Kurczewski 23 the mandibles as the grasshopper lies on its back or side (Fig. 12). Some T. tarsatus females even knead the coxa, trochanter, femur, or tarsus of the foreleg on the side of the grasshopper being malaxated (Kurczewski 1991). Alternatively, the wasp may be imbibing hemolymph exuding from a sting puncture wound in the forecoxal intersegmental membrane (Kurczewski 1991, Williams 1914). Malaxation is observed in most species in the pompiliformis and terminatus groups and is probably universal in the genus (Kurczewski 1987a, 1987b, 1991, 2008b, 2009; Kurczewski and Elliott 1978; Kurczewski and O’Brien 1988; Williams 1914). This behavior is believed to be preparatory to oviposition, since the egg is always affixed to an area at the base of a grasshopper’s foreleg (Newton 1956). Softening this area might facilitate attaching the egg securely. However, in T. tarsatus and other species in the pompiliformis group, half the females attach their egg to the side opposite that which was malaxated, and several wasps malaxate one acridid and then lay their egg on another grasshopper in the cell (Kurczewski 1991). Furthermore, females may malaxate both the left and right forecoxal coria of the grasshopper during prey transport. Grasshoppers extensively malaxated are sometimes abandoned, suggesting that they suffice to feed the adult wasp but do not furnish a satisfactory source of food for the developing wasp larva (Kurczewski 1987b, 1991). Malaxation usually takes place on the ground after the stinging of the grasshopper, often at some distance from the nest. Malaxation of larger prey may occur several times during transport to the nest and, rarely, near the nest entrance (Kurczewski 1987b, 1991, 2008b, 2009). Malaxation may also take place within the cell after the prey has been placed inside. Figure 12. Tachysphex tarsatus female malaxating right forecoxal corium of recently stung, nymphal Dissosteira carolina. 24 Northeastern Naturalist Vol. 17, Monograph No. 6 Mutilation Some Tachysphex prey are minus appendages or parts thereof when removed from the cell. The terminal antennal or tarsal segments of one or both hindlegs are occasionally missing. Indented or compressed antennal segments indicate where the prey was held with the wasp’s mandibles during transport. Among species in the pompiliformis group that prey on relatively large grasshoppers (T. acutus, T. punctifrons, T. pechumani, T. tarsatus), prey missing one or both hindlegs occurs at a frequency of 4.9–21.0% (Kurczewski 1987a, 1989, 1991, 2008b; F.E. Kurczewski, pers. observ.). Some larger prey (>40 mg) of T. terminatus, in the terminatus group, have one hindleg missing (Kurczewski 2009). Tachysphex alayoi and T. inconspicuus, both in the obscuripennis group, sometimes amputate or mutilate terminal antennal or tarsal segments of their cockroach prey, possibly during prey capture or transport (Genaro 2004; Kurczewski et al., in press). Prey Transport Some ancestral species in the pompiliformis group, such as T. punctifrons and T. pompiliformis, transport prey on the ground head forward and dorsal side upward (Alcock 1973, Kurczewski 1987a). These and other species in this group, such as T. pompiliformis, T. pechumani, T. tarsatus, T. laevifrons, T. ashmeadii, and T. crassiformis, transport prey on the ground head forward and ventral side upward, but sometimes carry it dorsal side upward, if only for a short distance (Bonelli 1966; Kurczewski 1987b, 1991, 2008b; Kurczewski and Elliott 1978; Kurczewski and O’Brien 1988; Pulawski 1971; Williams 1914). Species in the terminatus group hold the grasshopper in flight head forward and ventral side upward, rarely dorsal side upward or on the side (Elliott 1996, Elliott and Kurczewski 1985, Genaro and Sánchez 1992, Kurczewski 2009, Strandtmann 1953). Tachysphex mundus, in the brullii group, carries its tettigoniid or gryllid prey in flight head forward and dorsal side upward (Kurczewski 1979). Tachysphex belfragei, in this group, also captures Tettigoniidae and transports its prey on the ground (and probably also in flight), perhaps in the same manner (Kurczewski 1979). Method of prey transport is a function of prey size and weight. Wasps stay entirely on the ground if the prey is relatively large and heavy (Fig. 13), alternate short, low flights and ground transport with moderate-sized individuals, or make one or more extended flights if the prey is small and light (Fig. 14). Species in the pompiliformis group carry prey weighing about twice their own weight or less partly or wholly in flight (Kurczewski 1987a, 1987b, 1991), but heavier prey are transported entirely on the ground (Kurczewski 1987a, 1987b, 1989, 1991). An exception is T. pechumani, which was observed transporting one relatively small prey (prey-to-wasp ratio = 1.58:1) entirely on the ground to the nest (Kurczewski 2008b). In this species, where use of relatively large prey is commonplace, only one of >250 observations 2010 F.E. Kurczewski 25 of prey transport (prey-to-wasp weight ratio = 1.96:1) involved short, low flights (Kurczewski and Elliott 1978). Larger and heavier prey in the pompiliformis group are straddled more anteriorly, held with the mandibles farther out on the antennae, and grasped with the hindlegs farther forward on the prey’s thorax (Fig. 13; Kurczewski 1987b, 1989, 1991). The wasp’s first and second pair of legs are used only for walking and running (Williams 1914). The wings beat continually to assist the forward thrust with larger grasshoppers. The largest prey-to-wasp weight ratios recorded for species in this group are: T. tarsatus (8.96:1; Kurczewski 1991), T. acutus (7.78:1; Kurczewski 1989), T. ashmeadii (7.73:1; F.E. Kurczewski, pers. observ.), T. pechumani (7.71:1; Kurczewski 2008b), T. crassiformis (6.75:1; Kurczewski 1987b), and T. laevifrons (5.81:1; Kurczewski 1987b). The same prey transport rules for relative sizes and weights of wasps and their prey and manner of carriage apply also to the terminatus group; however, nearly all observations of prey transport in this group are for flight rather than ground transport. In flight, the wasp grasps the bases of the prey’s antennae with the mandibles and holds the prey’s body with the legs (Fig. 14). Ground transport in T. terminatus and T. similis is very rare (prey-to-wasp weight ratio of >2.1–2.3:1) and has not been observed in T. clarconis, T. antillarum, or T. apicalis (Elliott 1996, Elliott and Kurczewski 1985, Genaro and Sánchez 1992, Krombein and Evans 1955, Kurczewski 2009). Figure 13. Tachysphex tarsatus female transporting nymphal Melanoplus sp. forward on ground, grasping its antennae with her mandibles and its body with her hindlegs. Her antenna tips are tapping the ground surface (R. Jacksy, Toledo Area Metroparks, Toledo, OH, 2001 pers. comm.). 26 Northeastern Naturalist Vol. 17, Monograph No. 6 There is no difference in manner of flight transport and prey carriage between Acrididae and Tettigoniidae in T. terminatus (Kurczewski 1966a) or T. krombeini in the pompiliformis group (Kurczewski 1971). Tachysphex semirufus, also in the pompiliformis group, preys on Tettigoniidae, but practices ground transport, grasping the Mormon Cricket by the base of its antennae with the mandibles and probably also holding its body with the hindlegs (Kurczewski and Evans 1986). Species in the obscuripennis group prey on small adult and nymphal cockroaches and carry them in flight or, rarely, if larger, on the ground. Tachysphex inconspicuus and T. alayoi transport cockroaches in flight, grasping their prey's antennae with their mandibles and body with their legs (Buys 2007; Elliott et al. 1979; Genaro 2004; Kurczewski et al., in press). The ratio of weight of prey to wasp in T. alayoi flight transport is 1.98–2.76:1 (Elliott et al. 1979). Tachysphex inconspicuus prey are mostly only slightly longer in body length than the wasps, but sometimes weigh two to three times as much (Kurczewski et al., in press). Tachysphex iridipennis has been observed carrying its prey on the ground (W.J. Pulawski, 2009 pers. comm.). Nothing is known about prey carriage in T. cockerellae Rohwer or T. coquilletti, the two North American mantid-hunting species in the julliani group. The palearctic T. julliani captures small and moderately large mantids and carries them in flight or on the ground according to their size (Berland 1923, Ferton 1901). Figure 14. Tachysphex apicalis female flying toward her nest entrance, grasping the antennae of a nymphal Achurum carinatum with her mandibles and holding its body with her legs (Kurczewski 2009, Kurczewski and Snyder 1968). 2010 F.E. Kurczewski 27 Provisioning Time Species that fly with smaller prey generally spend less time between consecutive returns to the nest with prey individuals than species that transport larger prey on the ground. Tachysphex terminatus and T. similis in the terminatus group average about 11 min between flights with consecutive smaller prey items in late spring–early summer, but 33 min with slightly larger, consecutive prey individuals in late summer (Elliott 1996, Kurczewski 2009). Tachysphex antennatus, in the pompiliformis group, averages significantly less time (12 min) between flights with prey in June than between flights with prey in July (24 min) (Kurczewski and Kurczewski 1987). In T. antennatus, there is no correlation between prey size (weight) and duration spent between consecutive provisioning trips (Kurczewski and Kurczewski 1987). Tachysphex krombeini and T. psammobius, also in the pompiliformis group, fly with small prey and average about 13 and 6 min, respectively, between successive prey items (Kurczewski 1971, 1987a). Tachysphex mundus, in the brullii group, flies with small prey and averages only about 7 min between consective prey items (Kurczewski 1979). Tachysphex inconspicuus, in the obscuripennis group, flies with small prey and averages about 23 min between consecutive returns to the nest with paralyzed cockroaches (Kurczewski et al., in press). Many species in the pompiliformis group capture larger individuals, practice ground transport, and take, on average, much longer to obtain prey: T. ashmeadii (106 min), T. acutus (58), T. tarsatus (29), T. punctifrons (49), T. laevifrons (69), and T. pechumani (32) (Elliott and Kurczewski 1985; Kurczewski 1987a, 1987b, 1989, 1991, 2008b; F.E. Kurczewski, pers. observ.). Tachysphex crassiformis practices both flight and ground tranport of prey with smaller and larger individuals, respectively, and averages about 20 min between consecutive returns with prey (Kurczewski 1987b). Nest Entry Manner of nest entry with prey is usually related to presence or absence of temporary closure and prey size. In the pompiliformis group, except T. pechumani, smaller prey are usually carried directly into the open nest without being released (Kurczewski 1971, 1987a, 1987b, 1991; Kurczewski and Kurczewski 1987). It is not known whether prey taken into the entrance in this manner are carried all the way to the cell or laid down just inside the burrow by the wasp, out of sight, and, seconds later, dragged to the cell after the female turns around in the nest. Species in the pompiliformis group with larger prey release it outside or within the open entrance, enter the nest, turn around inside, walk up the burrow, grasp it by an antenna with the mandibles and, drag it back down the burrow. The sequence of placement of prey in the entrance, ranging from larger to smaller individuals, respectively, is: (1) head outside entrance, 28 Northeastern Naturalist Vol. 17, Monograph No. 6 (2) head inside entrance, (3) head and thorax inside entrance (Fig. 15, 16), and (4) head, thorax, and abdomen inside entrance (Kurczewski 1987b, 1989, 1991; Kurczewski and Kurczewski 1987). Figure 15. Tachysphex crassiformis female releasing moderate-sized, nymphal Psinidia fenestralis in open entrance (Kurczewski 1987b). Figure 16. Abdomen, midlegs, and hindlegs of moderate-sized, nymphal Psinidia fenestralis protruding from Tachysphex crassiformis burrow after being released in the open entrance of the nest. Wasp is inside the nest (Kurczewski 1987b). 2010 F.E. Kurczewski 29 Species in the terminatus group, except T. apicalis, and T. pechumani in the pompiliformis group make a temporary closure of the entrance and must release the prey on the ground and remove the soil fill from the opening before entering the nest (Elliott 1996; Elliott and Kurczewski 1985; Genaro and Sánchez 1992; Krombein 1964; Kurczewski 2008b, 2009; Kurczewski and Elliott 1978). In the terminatus group, the mostly small grasshoppers are released headfirst directly in front of the entrance followed by removal of the closure, nest entry, exit, and re-entry with prey. The placement of prey headfirst in front of the entrance more or less in a straight line facilitates the wasp pulling the grasshopper into the nest by an antenna (Fig. 17). Tachysphex apicalis, on the other hand, is restricted to carrying its prey directly into the open entrance due to the limitations of the sand cliff or steep sandbank (Fig. 18). In T. pechumani, the female releases the relatively large grasshopper with its head near the loosely filled entrance. However, the longitudinal axis of the prey’s body may be oriented variously with respect to the direction of the burrow, sometimes at a right angle, or even in a diagonal position above the entrance (Fig. 19). Tachysphex pechumani females spend less time to remove the temporary sand fill from the entrance at higher sand surface temperatures and more time to remove the sand fill at lower sand surface temperatures (Kurczewski 2008b). Tachysphex inconspicuus, in the obscuripennis group, is genus-atypical in that it retains its grasp of the prey’s antennae with the mandibles and body to the side with a hindleg as it removes the temporary closure from the entrance. Females enter the nest headfirst without releasing the cockroach (Fig. 20; Buys 2007; Kurczewski et al., in press). Figure 17. Tachysphex terminatus female removing sand fill from the nest entrance, using her forelegs in unison, after releasing a Melanoplus sp. headfirst on the sand surface in front of the burrow (Kurczewski 2009). 30 Northeastern Naturalist Vol. 17, Monograph No. 6 Final Closure of the Nest Final closure of the nest is similar in species that occupy level ground. After placing the full complement of prey in a cell and laying an egg on a Figure 18. Tachysphex apicalis female flying directly into her open nest entrance, holding a small nymphal Melanoplus sp. underneath (Kurczewski 2009). Figure 19. Tachysphex pechumani female has released a Melanoplus sp. to left of entrance at right angle to direction of burrow. The end of the wasp’s abdomen can be seen as she enters the nest. 2010 F.E. Kurczewski 31 single individual, the wasp compactly fills the burrow and entrance with soil. She breaks down the walls of the burrow with the mandibles and rakes the soil backward into the burrow beneath the synchronously lifting abdomen, using the forelegs in unison. The foretarsi are fishhooked medially to expose the largest number of rake spines (Fig. 21). The wasp rapidly tamps the soil compactly in place at the end of the burrow with the vibrating bent end of the abdomen. The last abdominal tergite, or pygidial plate, is triangular, mildly convex, mainly smooth, carinate marginally, and well adapted for this purpose (Fig. 22). The rapid tamping behavior and mostly smooth, triangular pygidium are highly diagnostic behavioral and morphological characteristics for most members of this genus. After the burrow is filled flush with the surface, the wasp moves her abdominal apex from side to side in trowel-like fashion smoothing over the fill (Elliott 1996; Kurczewski 1987b, 1989, 1991, 2008b, 2009; Kurczewski and Elliott 1978; Kurczewski and Evans 1986; Kurczewski and Kurczewski 1987; Kurczewski and O’Brien 1988; Peckham and Peckham 1898; Williams 1914). Tachysphex inconspicuous, in the obscuripennis group, fills its burrow in the same manner as other species (Buys 2007; Kurczewski et al., in press). Some species in the pompiliformis and terminatus groups make hovering flights toward the end of the closure. It is not known whether such flights serve for orientation for future nest site location or are in response to high sand surface temperatures (Evans 1973; Kurczewski 1987b, 1989, Figure 20. Tachysphex inconspicuus female grasping a Chorisoneura sp. antennae with her mandibles and its body with one of her hindlegs, and, holding the prey to one side, removes the temporary closure from the entrance with her forelegs (Kurczewski et al., in press). 32 Northeastern Naturalist Vol. 17, Monograph No. 6 Figure 22. Shiny, trowel-shaped, mildly convex pygidium (last abdominal tergum) of a Tachysphex pechumani female used for tamping sand in the burrow when bent downward (Kurczewski et al. 1970). Figure 21. Tachysphex tarsatus female making a final nest closure. Her foretarsi are fishhooked medially, exposing lateral rake-spines, as she rakes sand backward into the burrow (Kurczewski 1991). 2010 F.E. Kurczewski 33 1991, 2008b, 2009; Kurczewski and Elliott 1978; Kurczewski and Kurczewski 1987). Tachysphex mundus spends most of the time filling its burrow beneath the surface and out of sight. Near the surface, females use the same behavior for filling their burrow as described above (Kurczewski 1979). Tachysphex semirufus sometimes leaves a depressed entrance after its burrow is filled (Kurczewski and Evans 1986). Tachysphex antennatus and T. tarsatus, when nesting in gravelly soil, use pebbles as part of the fill, especially in the upper portion of the burrow and entrance (Kurczewski 1991, Kurczewski and Kurczewski 1987). Tachysphex acutus, T. pechumani, and T. inconspicuus may dig shallow pits in the ground near the entrance with their mandibles and forelegs in order to obtain additional soil to supplement the fill (Kurczewski 1989, 2008b; Kurczewski et al., in press). Tachysphex pechumani uses only the proximal portion of its shallow tumulus as fill, leaving the distal portion on the surface as darker-colored residue (Fig. 23). Some females of T. acutus, T. tarsatus, and T. pechumani place tiny twigs, pieces of leaves, grass blades, pebbles, dried florets, and moss fragments atop the fill for apparent concealment (Kurczewski 1989, 1991, 2008b; Kurczewski and Elliott 1978). Some T. tarsatus females near the end of the nesting season oviposit on a grasshopper in the cell, but make no attempt to fill the burrow with soil. Other T. tarsatus females, late in the season, fill the burrow but do not oviposit on the grasshopper, perhaps having exhausted their egg supply (Kurczewski 1991). Tachysphex apicalis and, rarely, T. tarsatus, T. similis, and T. terminatus demonstrate a different type of final closure when nesting in sand cliffs and Figure 23. Completed nest of Tachysphex pechumani showing distal portion (right) of scattered tumulus remaining on the sand surface following final closure. The proximal portion of the tumulus (left, white outline) was used as sand fill. 34 Northeastern Naturalist Vol. 17, Monograph No. 6 steep slopes. They close the nest by pulling down sand from the sides of the burrow and entrance with the mandibles and fling the loosened soil backward with the forelegs in unison. They compactly pack the soil at the end of the burrow with the bent end of the vibrating abdomen. During the final stages of such a closure, females stand upright on hindlegs inside the enlarged entrance, grasp the sides of the cavity with the midlegs, and loosen sand with the mandibles (Fig. 24). Such finished nests are never filled flush, but are highly visible as a concave depression on the rather smooth surface (Kurczewski 1991, 2009). Nest Structure and Dimensions Nest structure and dimensions in Tachysphex species are related to whether or not the species uses a pre-existing burrow, soil texture, slope and moisture level, size of female, and wasp generation/time of year. Nests begun from pre-existing burrows tend to be longer and, sometimes, deeper than nests started from the soil surface (Kurczewski 1979, 1987a). Nests in sand are usually longer and deeper than nests excavated in loam, sandy loam, loamy sand, or loamy fine sand (Buys 2007; Kurczewski 1989, 1991, 2008b, 2009). Larger wasps tend to make longer and deeper nests than smaller females (Kurczewski 2009). First generation nests are generally longer and deeper than second and later generation nests in many species (Kurczewski 1987b, 1991, 2009); however, nests of T. antennatus are slightly longer and deeper in July than in June (Kurczewski and Kurczewski 1987). In the Bahamas, June cells of T. similis are deeper than April cells of this species (Elliott 1996). Figure 24. Tachysphex apicalis female standing upright in an enlarged entrance to pull down sand for final nest closure with her mandibles. 2010 F.E. Kurczewski 35 Most species in the pompiliformis group excavate a short, oblique burrow that terminates in a single cell (Fig. 25; Alcock 1973, Elliott and Kurczewski 1985; Evans 1970; Kurczewski 1987a, 1987b, 1991, 2008b; Kurczewski and Elliott 1978; Kurczewski and Evans 1986; Kurczewski and Kurczewski 1987; Newton 1956; Williams 1914). Tachysphex aethiops and T. punctifrons modify pre-existing burrows into moderately long, diagonal main burrows leading to side burrows that may end in five or six cells (Fig. 26; Kurczewski 1987a, O’Brien 1987). Tachysphex acutus frequently excavates two-celled nests with burrows that often bifurcate from a Figure 25. Exposed, one-celled Tachysphex pechumani nest (side view) showing a paralyzed, nymphal Melanoplus sp. positioned head inward and ventral side upward (Kurczewski 2008b). Figure 26. Unfinished, six-celled Tachysphex punctifrons nest (side view) showing placement of side burrows and cells. The lateral splaying of burrows is not shown in this two-dimensional figure. Cells are lettered in the order in which they were completed. All cells contained an egg (e). Sand-filled burrows and tumulus are stippled (Kurczewski 1987a). 36 Northeastern Naturalist Vol. 17, Monograph No. 6 common entrance, a possible rudimentary stage in multi-celled nest development (Fig. 27; Kurczewski 1989). Species in the terminatus group typically excavate a short, rather shallow two- to five-celled nest under optimal nesting conditions (Fig. 28; Elliott 1996, Elliott and Kurczewski 1985, Evans 1970, Genaro and Sánchez 1992, Krombein 1964, Kurczewski 2009). Three-celled nests are common in T. alpestris, T. terminatus, T. similis, and T. apicalis (Evans Figure 27. Burrows, cells, and spurs of one- and two-celled Tachysphex acutus nests (top view). Cells are numbered in the order in which they were made. The scale refers to all nests (Kurczewski 1989). 2010 F.E. Kurczewski 37 Figure 28. Representative configurations of Tachysphex terminatus (top), T. similis (middle), and T. apicalis (bottom) bi- and multi-celled nests. Cells are numbered in the order in which they were made. The scale refers to all nests (Kurczewski 2009). 38 Northeastern Naturalist Vol. 17, Monograph No. 6 1970, Kurczewski 2009). Some nests in the terminatus group are final closed with only a single, fully provisioned cell (Elliott 1996, Elliott and Kurczewski 1985, Evans 1970, Krombein 1964, Kurczewski 2009). Many one-celled nests are closed prematurely at the onset of inclement weather or dusk or during scarcity of prey from drought or other unfavorable enironmental factors (Kurczewski 2009). Nests of species in the brullii group have varying dimensions when excavated in different soil types from the pre-existing burrows of other Hymenoptera (Kurczewski 1979). The portions of nests excavated by T. mundus in non-sandy soil are relatively short and shallow, but one nest of T. belfragei in loose sand was nearly vertical and rather deep (10 cm; Kurczewski 1979). Species in the obscuripennis group excavate rather short, oblique burrows and shallow one- or two-celled nests in sandy soil (Buys 2007; Genaro 2004; Kurczewski et al., in press). One nest of T. coquilletti, in the julliani group, was arcuate, moderately long (12 cm), and rather shallow (4 cm) (Alcock and Gamboa 1975). Prey Type North American, Central American, and Caribbean Tachysphex stock their cells with paralyzed orthopteroid insects, mainly Acrididae, but also Gryllidae, Tettigoniidae, Blattellidae, and Mantidae (Appendix 1; Krombein 1979, Pulawski 1988). Most species in the pompiliformis group prey on nymphal and, rarely, adult Acrididae. Tachysphex ashmeadii, T. pompiliformis, and T. tarsatus prey on as many as three or four subfamilies of Acrididae (Elliott and Kurczewski 1985; Kurczewski 1991; Pulawski 1988; Williams 1914; F.E. Kurczewski, pers. observ.). Tachysphex semirufus and T. tipai Pulawski capture nymphal Mormon Crickets and katydids, respectively (Kurczewski and Evans 1986; Pulawski 1988, 2007). Tachysphex krombeini stocks nymphal acridids and tettigoniids mixed in its cells (Kurczewski 1971). A female of T. apricus Pulawski is pinned with a Parabacillus hesperus Hebard (Phasmatodea: Heteronemiidae) more than four times its length (Elliott and Kurczewski 1985). Pulawski (1988) questions the authenticity of this record. There is specificity at the prey subfamily level among some acrididhunters in the pompiliformis group. Tachysphex aethiops evidently captures only Oedipodinae (Alcock 1973; Evans 1970, 1973; O’Brien 1987). Tachysphex crassiformis preys predominantly on Oedipodinae and, very rarely, Gomphocerinae (Krombein 1963, Kurczewski 1987b, Williams 1914). Tachysphex acutus and T. punctifrons stock their cells with nymphal and adult (T. punctifrons) Melanoplus (Melanoplinae; Kurczewski 1987a, 1989, 2000b). In the northeastern United States, T. tarsatus females inhabiting sandy fields or the edges of sandpits and sandblows prey predominantly on species of Melanoplus, often M. femurrubrum (DeGeer), M. sanguinipes (Fabricius), or M. bivittatus (Say). Females nesting in or near gravel pits 2010 F.E. Kurczewski 39 capture Dissosteira carolina, while those nesting in sand near open water prey on Trimerotropis maritima (Harris) or Psinidia fenestralis (Serville), all Oedipodinae (Kurczewski 1991, 1999). Species in the terminatus group prey mainly on nymphal Acrididae. A few females of T. terminatus stocked several cells with a single nymphal bush katydid, Scudderia, probably furcata Brunner von Wattenwyl (Tettigoniidae), in addition to a few or several nymphal Melanoplus bivittatus during one late spring at a single locality (Kurczewski 1966a, Pulawski 1988). Tachysphex apicalis, the largest species in the group, very rarely stocks its cells with adult male and nymphal acridids of smaller species (Kurczewski 2009). Very large sample sizes for T. terminatus and T. similis reveal equivalent percentages of Gomphocerinae, Oedipodinae, and Cyrtacanthacridinae + Melanoplinae as prey. Tachysphex apicalis stocks its cells with the same subfamilies of prey but in different proportion. Members of three subfamilies of Acrididae and three or four genera and species of prey may be put in a single cell of these species (Kurczewski 2009). Females in the terminatus group are evidently unselective of their prey species and capture whatever grasshoppers of suitable (small) size are available and abundant near the nesting site (A.B. Gurney, United States National Museum, Washington, DC, 1963 pers. comm.). Species in the brullii group stock their cells with nymphal Tettigoniidae (katydids) and, rarely, nymphal Gryllidae (tree crickets; Elliott and Kurczewski 1985, Kurczewski 1979). Species in the obscuripennis group provision their cells with small adult and nymphal blattellid cockroaches (Buys 2007; Callan 1993; Elliott et al. 1979; Genaro 2004; Kurczewski et al., in press; Pulawski 1988; Vesey-FitzGerald 1956). A single paralyzed gryllid was collected, along with the usual blattellid prey, by a female of T. inconspicuus (Buys 2007). There is a preponderance of adult female prey among the cell contents of T. inconspicuus (Kurczewski et al., in press). Tachysphex cockerellae and T. coquilletti, both in the julliani group, prey upon nymphal Litaneutria minor (Scudder) (Minor Ground Mantid) (Mantidae; Alcock and Gamboa 1975, Elliott and Kurczewski 1985). Litaneutria minor is a ground-dwelling species in dry grasslands, but also inhabits low vegetation such as sagebrush and antelope-brush (Cannings 1987). Prey Size and Number per Cell Because of their mainly small size, species of Tachysphex capture and stock their cells mostly with small, relatively small, or moderate-sized nymphal prey. There is a correlation between prey size and number of prey per fully provisioned cell in members of this genus. In the pompiliformis group, T. pechumani captures relatively large nymphal and adult grasshoppers and stores only a single prey per cell in spring and early summer (Kurczewski 2008b, Kurczewski and Elliott 1978). Tachysphex aethiops, T. montanus, T. acutus, T. ashmeadii, T. tarsatus, T. pompiliformis, and T. punctifrons also prey on relatively large nymphal and adult acridids, especially in mid–late 40 Northeastern Naturalist Vol. 17, Monograph No. 6 summer, and store one individual per cell (Alcock 1973; Evans 1970, 1973; Kurczewski 1987a, 1989, 1991; Kurczewski and O’Brien 1988; Newton 1956; O’Brien 1987; Williams 1914). However, most of these species stock two, three, or four smaller acridids per cell in spring or early summer (Alcock 1973; Alcock and Gamboa 1975; Bonelli 1966; Elliott and Kurczewski 1985; Kurczewski 1987a, 1991, 1999; Newton 1956; Williams 1914). Several other species in the pompiliformis group prey on small acridids or acridids and tettigoniids mixed (T. krombeini) and typically stock several or many individuals per fully provisioned cell: T. antennatus (3–9; Kurczewski and Kurczewski 1987), T. krombeini (7; Kurczewski 1971), T. psammobius (11; Kurczewski 1987a), and T. texanus (4; Kurczewski 1987a). Species in the terminatus group usually stock several or many small prey per fully provisioned cell (Kurczewski 2009). Maximal numbers per fully provisioned cell for species in this group: T. clarconis, 6; T. antillarum, 11; T. alpestris, 13; T. linsleyi, 7; T. terminatus, 14; T. similis, 15; and T. apicalis, 12 (Kurczewski 2009). Sometimes, there are so many prey that some of them are aligned in the burrow in front of the cell because they all cannot fit inside (Kurczewski 2009). Fully provisioned cells in the terminatus group with only one or two prey are often closed prematurely at the onset of inclement weather or dusk. One or two prey per cell is also associated with scarcity of grasshoppers due to drought or other unpredictable environmental occurrences (Kurczewski 2009). In the brullii group, from three to six small tettigoniids or tettigoniids and gryllids mixed are stocked in fully provisioned cells of T. mundus (Kurczewski 1979). Four meadow katydids were stocked in one fully provisioned cell of T. belfragei (Kurczewski 1979). From one to five small cockroaches are stocked in fully provisioned cells of species in the obscuripennis group (Buys 2007; Genaro 2004; Kurczewski et al., in press). There is no record of the number of prey mantids per cell or their sizes in T. cockerellae or T. coquilletti in the julliani group (Pulawski 1988). Ferton (1897, 1901) recorded as many as six small mantids per cell in the palearctic T. julliani (Pulawski 2007). Prey Placement in Cell Prey are usually placed in the cell in the manner in which they are taken into the nest. Most prey in the pompiliformis, terminatus, and obscuripennis groups are placed in the cell in a head inward and ventral side upward position (Fig. 25; Alcock 1973; Buys 2007; Elliott 1996; Elliott and Kurczewski 1985; Genaro and Sánchez 1992; Kurczewski 1987b, 1989, 1991, 2000b, 2008b, 2009; Kurczewski and Elliott 1978; Kurczewski and Kurczewski 1987; Kurczewski and O’Brien 1988; Kurczewski et al., in press; O’Brien 1987; Peckham and Peckham 1898, 1905; Williams 1914). Cells that contain the largest number of prey, as in the terminatus group, often have more individuals placed in other positions. Prey in atypical positions may be 2010 F.E. Kurczewski 41 accidentally moved to those positions by wasps maneuvering to oviposit on another prey in the cell (Kurczewski 2009). Females of T. apicalis oviposit by getting on their backs beneath the pedestal prey, curling the abdomen around the grasshopper’s thorax until the apex reaches its venter, and affixing an egg to a forecoxal corium (F.E. Kurczewski, pers. observ.). Tachysphex punctifrons and, rarely, T. pompiliformis, both in the pompiliformis group, and T. mundus in the brullii group, transport prey to their nests on the ground and in flight, respectively, head forward and dorsal side upward. Yet, T. punctifrons and T. pompiliformis place prey in their cells head inward and ventral side upward, and T. mundus places prey head inward and ventral or dorsal side upward (Kurczewski 1979, 1987a). Wasp’s Egg Eggs of Tachysphex species are often laid on the largest and heaviest prey on the bottom or at the back end of the cell. This individual is usually positioned head inward and ventral side upward. Eggs of T. inconspicuus, in the obscuripennis group, are laid mainly on adult female cockroaches, the larger and heavier sex (Kurczewski et al., in press). The egg affixation site is the soft intersegmental membrane surrounding the base of a forecoxa. In acridid- and tettigoniid-hunting species, the wasp’s egg extends to the opposite side of the affixation site, transversely across the ventral thorax between the bases of the fore- and midlegs (Fig. 29; Elliott 1996; Evans 1970, 1973; Genaro and Sánchez 1992; Krombein 1964; Kurczewski 1971, 1979, 1987a, 1987b, 1989, 1991, 2008b, 2009; Kurczewski and Elliott 1978; Kurczewski and Kurczewski 1987; Newton 1956; O’Brien 1987; Williams 1914). Figure 29. Tachysphex antennatus egg affixed to a nymphal Melanoplus sp. in a position typical of species that prey on Acrididae (Kurczewski and Kurczewski 1987). 42 Northeastern Naturalist Vol. 17, Monograph No. 6 In species in the obscuripennis group, the wasp’s egg is also attached to a forecoxal corium, but extends longitudinally posteriad on the thoracic venter (Fig. 30; Genaro 2004; Kurczewski et al., in press). Tachysphex tarsatus very rarely attaches its egg to the cervix (throat) of extremely large grasshoppers, possibly because of difficulty in getting into proper egg-laying position (Kurczewski 1991). Attachment of the egg to the left or right forecoxal corium occurs with about equal frequency in large samples (Kurczewski 1991, 2008b, 2009). In some multi-celled nests, a female places her eggs on one side or the other of prey in most or all of the cells. Other wasps alternate between left and right sides. Egg size is a function of wasp size in this genus (Kurczewski 1987a, 1991). Seasonal Trends in Nesting Behavior Seasonal trends in nesting behavior are seen in T. tarsatus, T. crassiformis, and T. antennatus in the pompiliformis group and T. terminatus and T. similis in the terminatus group: 1) First generation/spring T. terminatus females in Kansas and New York and T. similis females in Kansas spend significantly less time obtaining consecutive prey items than second and later generation/mid–late summer females in these regions (Kurczewski 2009). The amount of time spent between successive arrivals at the nest with prey in T. antennatus is less than half as long with small grasshoppers in spring–early summer than with slightly larger grasshoppers in mid-summer (Kurczewski and Kurczewski Figure. 30. Contents of a two-celled nest of Tachysphex inconspicuus with two and four prey, respectively, showing wasp eggs (white arrows) affixed to the forecoxal coria of 1st and 3rd cockroaches from left and extending longitudinally posteriad on the thoracic venters. Riatia fulgida (Saussure) is on the far right; the other five cockroaches are Chorisoneura spp. Note missing antenna segments on some of the prey. (Kurczewski et al., in press). 2010 F.E. Kurczewski 43 1987). Based on random net sweeps through vegetation near the nesting sites, there are more grasshoppers of suitable (small) size and prey type (genus and species) available to these species earlier in the nesting season (Kurczewski 2009; Kurczewski and Kurczewski 1987; A.B. Gurney, 1963 pers. comm.; F.E. Kurczewski, pers. observ.). In T. tarsatus in New York, on the other hand, females spend less time between captures of Melanoplus femurrubrum in late summer than captures of Dissosteria carolina, Melanoplus sanguinipes, or M. bivittatus in late spring–early summer. Suitable-sized (moderately large) M. femurrubrum are more abundant in mid–late summer than the other species in late spring–early summer at some New York sites (Kurczewski 1991; F.E. Kurczewski, pers. observ.). 2) At nearly all localities, longer burrows and deeper cells characterize first generation nests, and shorter burrows and shallower cells are typical of second and later generation nests. Spring nests of T. crassiformis, T. tarsatus, T. terminatus, and T. similis are often longer and deeper than mid–late summer nests at the same locations (Kurczewski 1987b, 1991, 2009). Nests of T. antennatus, on the other hand, are slightly longer and deeper in July than in June (Kurczewski and Kurczewski 1987). Nests of T. similis in the Bahamas are deeper in June than in April (Elliott 1996). 3) Species of prey Acrididae are different in first and later generation cells of Tachysphex at most localities. In New York, Dissosteira carolina and Melanoplus bivittatus are predominant prey species of T. tarsatus in late spring–early summer, M. sanguinipes in early–mid-summer, and M. femurrubrum in mid–late summer (Kurczewski 1991). Species of prey Acrididae from first and later generation cells of T. terminatus and T. similis, except in Florida, are usually different and tied to prey availability near the nesting site. It is difficult to discern prey differences for the different generations of T. similis and T. apicalis in Florida as both wasps and prey are available almost continually throughout the year (Kurczewski 2009). In the terminatus group, there is a gradual decrease in the availability of potential prey species of appropriate (small) size through the nesting season in temperate regions (A.B. Gurney 1963, pers. comm.). 4) Mean number of prey per fully provisioned cell is reduced from first to later wasp generations. A gradual decrease in the mean number of prey per cell accompanies an increase in mean prey size from late spring to late summer in T. tarsatus (Kurczewski 1991). More grasshoppers are stocked per cell earlier and fewer grasshoppers per cell later in the nesting season in T. similis and T. terminatus (Kurczewski 2009). The mean number of prey per fully provisioned cell in T. terminatus is approximately halved from first to second generation in central New York (Kurczewski 2009). 5) Prey individuals from spring–early summer cells are often smaller and weigh less than prey individuals from mid–late summer cells. This trend of increasing mean body length and mean wet weight of prey individuals occurs throughout the nesting season in T. tarsatus, T. crassiformis, and 44 Northeastern Naturalist Vol. 17, Monograph No. 6 T. antennatus (Kurczewski 1987b, 1991; Kurczewski and Kurczewski 1987). Tachysphex terminatus and T. similis stock smaller and lighter grasshoppers per cell earlier and larger and heavier grasshoppers later in the nesting season (Kurczewski 2009). 6) There is a noticeable reduction in prey biomass from first to later generation cells in T. terminatus and T. similis at many sites, as evidenced by a decrease in mean aggregate prey wet weight per cell despite larger and heavier individuals being stocked in the cells (Kurczewski 2009). 7) A decrease in biomass per cell in T. terminatus and T. similis from spring to late summer results in first generation wasps of the following year being slightly smaller and lighter in weight than second or later generation wasps (Elliott and Kurczewski 1975; Kurczewski 2009; F.E. Kurczewski, pers. observ.). 8) An increase in biomass per cell during the nesting season in T. tarsatus from wasps gradually stocking larger individuals results in the reared wasps increasing in size and weight, except for first generation females of the following year (Kurczewski 1991). 9) First generation females of T. terminatus and T. similis in Kansas and New York complete more cells per day than second and later generation wasps (Kurczewski 2009). Females of T. tarsatus, on the other hand, stock and complete more cells per day later in the summer than in late spring–early summer at the same sites, possibly because they gradually use fewer and larger prey individuals per cell. In addition, their predominant prey at that time of year, Melanoplus femurrubrum, is exceedingly common (Kurczewski 1991; F.E. Kurczewski, pers. observ.). Such variation among the species is related to different prey abundance and availability of appropriate-sized individuals at the various times of year (A.B. Gurney, 1963 pers. comm.). Cleptoparasitism Species of Tachysphex are afflicted with different cleptoparasites that diminish their reproductive success in the various aggregations. Common cleptoparasites include hymenopterans belonging to the families Mutillidae (Callan 1942, 1993; Ferguson 1962; Moan and Tramer 2008; Williams 1914) and Chrysididae (Elliott and Kurczewski 1985; Kurczewski 1967, 2008b) and dipterans of the families Bombyliidae (Kurczewski and Harris 1968) and Sarcophagidae (Alcock 1973; Buys 2007; Elliott 1996; Evans 1970, 1973; Genaro and Sánchez 1992; Kurczewski 1989, 1991, 2008b; Moan and Tramer 2008; Newton 1956; O’Brien 1987; Spofford et al. 1986; Spofford and Kurczewski 1990, 1992; Williams 1914). Sarcophagid flies in the tribe Miltogrammini are common inhabitants at Tachysphex nesting sites (Alcock 1973; Bohart and Menke 1976; Buys 2007; Elliott 1996; Evans 1970; Genaro and Sánchez 1992; Kurczewski 1989, 1991, 2008b; Moan and Tramer 2008; Newton 1956; O’Brien 1987; Pulawski 1971, 1988; Spofford et al. 1986; Spofford and Kurczewski 1990, 2010 F.E. Kurczewski 45 1992). They utilize the wasp’s eggs, larvae, and prey for rearing their own young (Allen 1926, Evans 1966, Evans and West Eberhard 1970, Matthews and Matthews 1978, Newcomer 1930, Williams 1914). Species in the terminatus group are especially heavily cleptoparasitized by miltogrammine flies. Overall miltogrammine cleptoparasitism in a 7-year study of more than 200 nests of T. terminatus was 42.5% (Spofford and Kurczewski 1992), with the frequency of cleptoparasitism at one locality in central New York attaining 57.9% (Spofford et al. 1986). The frequency of cleptoparasitism from miltogrammine flies in a large aggregation of T. antillarum in Cuba was 53.6% (Genaro and Sánchez 1992). In contrast, the cleptoparasitic frequency from miltogrammine flies in the more solitary T. tarsatus was only 23.8% over the 7-year period (Spofford and Kurczewski 1992), and 23.3% for this species during one year at another site in central New York (Kurczewski 1991). Stalking and lurking miltogrammines (Hilarella hilarella (Zetterstedt), Taxigramma heteroneura (Meigen), Sphenometopa tergata Meigen) and open-hole searching miltogrammines (Metopia spp.) are cleptoparasitic on Tachysphex species that nest among or near vegetation and transport large prey on the ground. These fly species focus their attacks on species in the pompiliformis group such as T. pompiliformis, T. montanus, T. aethiops, T. acutus, T. tarsatus, T. antennatus, and T. pechumani (Alcock 1973; Evans 1970; Kurczewski 1989, 1991, 2008b; Kurczewski and Kurczewski 1987; Newton 1956; O’Brien 1987; Spofford and Kurczewski 1990, Williams 1914). Such miltogrammines lurk near excavating wasps, perch on stones or plants beside their nests, trail females transporting prey, or ride to wasp entrances on the prey’s abdomen or legs. They larviposit on the prey or in the open entrances before leaving. Open-hole searching miltogrammines larviposit in the open wasp burrows, probably after ascertaining that the nest is being actively provisioned. Closed-hole searching miltogrammines (Phrosinella spp.) cleptoparasitize Tachysphex species that nest in barren or sparsely vegetated sand and make a temporary closure of the nest entrance (Spofford et al. 1986). Phrosinella aurifacies Downes digs into and larviposits in the temporarily closed entrances of species in the terminatus group such as T. terminatus and T. similis (Spofford and Kurczewski 1990; Spofford et al. 1986; F.E. Kurczewski, pers. observ.). The maggot burrows through the sand closure down to the cell where it attacks the cell contents (Spofford and Kurczewski 1990, Spofford et al. 1986). Ubiquitous satellite-flies of the genus Sphixapata and Amobia floridensis (Townsend) appear to be less selective of their hosts and cleptoparasitize a wide range of ground-nesting and other solitary wasps (Evans 1970), including Tachysphex species in the pompiliformis, terminatus, and obscuripennis groups (Buys 2007, Elliott 1996, Genaro and Sánchez 1992, Spofford and Kurczewski 1990). Sphixapata vigilans (Allen) and S. rubriventris (Macquart) maintain surveillance perches near wasp entrances during burrow 46 Northeastern Naturalist Vol. 17, Monograph No. 6 excavation and hunting forays and larviposit on the prey when the wasp is preoccupied with provisioning activities (Genaro and Sánchez 1992, Kurczewski 2008b, Spofford et al. 1986). Sphixapata trilineata (Wulp) is attracted to the wasp’s provisioning flight and larviposits on the prey in midair or on the ground, often in the presence of the wasp (Genaro and Sánchez 1992, McCorquodale 1986, Spofford et al. 1986). Species of Tachysphex have evolved different methods of behavior to counter the cleptoparasitic attacks of the miltogrammine flies. Tachysphex inconspicuus excavates false burrows in the soil around the main burrow to distract the flies from larvipositing in the nest (Buys 2007). The miltogrammine Amobia floridensis was observed entering such an “accessory burrow” of T. inconspicuus in Brazil (Buys 2007). Tachysphex tarsatus, T. pechumani, T. terminatus, and T. similis exhibit freeze-stops and face-offs to dissuade the miltogrammine flies from larvipositing. The wasps remain motionless for long periods of time, facing the fly, and their success in preventing or reducing cleptoparasitism is dependent on the number of wasps and flies in the immediate area. Other wasps flying nearby often serve as distractions for the flies, allowing the wasp with prey to continue on to her nest unobstructed (Kurczewski 2008b, Spofford and Kurczewski 1992). Provisioning females in the pompiliformis group (T. pompiliformis, T. pechumani, T. tarsatus) often climb vegetation and remain motionless, clinging to the plant, to escape the pursuit of miltogrammine flies (Alcock 1973; Kurczewski 1991, 2008b; Spofford and Kurczewski 1992). The fly sometimes trails the wasp and prey onto the plant stem and remains fixed on the pair (Fig. 31; Alcock 1973; F.E. Kurczewski, pers. observ.). Species in the terminatus group with small prey attempt to elude the satellite-flies by making rapid, winding and circuitous flights through vegetation, away from the nesting site (Elliott 1996, Spofford and Kurczewski 1992). The wasps often modify their flight patterns or do not enter their nests in the presence of the flies (Elliott 1996, Genaro and Sánchez 1992, McCorquodale 1986). Satellite-flies of the genus Sphixapata are particularly adept at following the wasps in flight; hence, the name satellite-fly is quite appropriate. The flies have large eyes and enlarged central eye facets that enable them to effectively pursue the wasps as though tethered to the provisioning wasps by an invisible thread (Allen 1926; W.H. Downes, Jr., Michigan State University, East Lansing, MI, 1983 pers. comm.). Tachysphex tarsatus and T. pechumani attempt to elude pursuing miltogrammine flies by transporting their prey on the ground into shaded areas or through dense vegetation, but the flies are usually able to follow the slow-moving pair (Kurczewski 1991; Williams 1914; F.E. Kurczewski, pers. observ.). Tachysphex terminatus, T. similis, T. pechumani, and T. tarsatus females fly at and chase the flies from the area in an attempt to rid themselves of their constant pursuit (Elliott 1996, Kurczewski 1991, 2008b, Spofford and 2010 F.E. Kurczewski 47 Kurczewski 1992). Such chases are usually ineffectual, with the flies returning almost immediately to the provisioning wasp. Species in the pompiliformis and terminatus groups may abandon a cell infested with maggots or leave a prey on the ground after it has been larviposited upon by miltogrammine flies (Kurczewski 1991, 2008b; Spofford and Kurczewski 1992). Cell and prey abandonment is common in T. tarsatus, with one-third of its nests/prey under attack being abandoned rather than ovipositing and wasting a viable egg or final closing and exerting considerable energy in the process (Spofford and Kurczewski 1992). Tachysphex terminatus and T. antillarum search for maggots on the prey with their antennae and mouthparts and lift the grasshopper’s legs with their mandibles (Genaro and Sánchez 1992, Spofford et al. 1986). Tachysphex terminatus cleans the underside of the prey’s head, cervix, thorax, and abdomen with its mouthparts and then cleans the dorsal surface of the same areas, after turning the grasshopper over. Tachysphex terminatus removes maggots from the prey if aware of satellite-fly larvipositional contact. Wasps often destroy the maggots by cutting them in half with their mandibles (Fig. 32). In one study (Spofford et al. 1986), only 1 of 35 (2.9%) prey of T. terminatus contained maggots after being cleaned by the wasps. Adaptability Species of Tachysphex alter their nesting behavior in order to facilitate certain situations and compensate for suboptimal conditions or atypical Figure 31. Tachysphex pompiliformis female resting on an upright plant stem, holding an acridid prey head forward and dorsal side upward with her mandibles and hindlegs after being pursued by a miltogrammine fly. The fly moved from the hindleg of the grasshopper to the plant stem opposite the wasp (Alcock 1973). 48 Northeastern Naturalist Vol. 17, Monograph No. 6 circumstances. Tachysphex pechumani is univoltine in the central Great Lakes Region, occurring predictably every year in late spring and early summer (Kurczewski 2008b). However, some females with perfect wings, complete foretarsal digging rakes, and clean integument emerge annually in early August in northern Lower Michigan. There are no males associated with these wasps and the females do not mate or nest. Whether or not these late emergents are simply first generation stragglers or a very small partial second generation remains to be determined. Tachysphex pechumani, a highly psammophilous species that typically occurs in pine-barrens and oak savanna, has the capability to nest in finer-grained soils following man-made disturbance of their usual habitat (Kurczewski 2008b). Tachysphex similis and T. terminatus are able to nest in small, man-made sand piles dumped alongside roads or on asphalt or concrete parking lots following destruction of their natural habitat (Kurczewski 1966b; F.E. Kurczewski, pers. observ.). In dense nesting aggregations, T. terminatus females share entrances or, they sometimes move and occupy strongly sloping soils at the periphery of the aggregation (Kurczewski 2009). Rarely, this species, T. similis, T. pompiliformis, and T. tarsatus nest in nearly vertical or vertical sand cliffs rather than level soil (Kurczewski 1991, 2009; Pulawski 1971). Conversely, T. apicalis, when nesting in dense aggregations, has the capability to excavate burrows in level sand, away from the sand cliffs in which the aggregation nests, albeit sometimes unsuccessfully so (Kurczewski 2009). Figure 32. Tachysphex terminatus female dismembering a maggot with her mandibles during a prey-cleaning episode following Sphixapata vigilans larviposition on the prey (Spofford et al. 1986). 2010 F.E. Kurczewski 49 Tachysphex pechumani females, when nesting in aggregations that are heavily cleptoparasitized, tend to remain below ground, during burrow excavation, and fling the sand backward while remaining mainly out of sight. Females of this species experiencing lesser amounts of cleptoparasitism at other locations tend to back farther from their burrows when removing the soil, often onto the proximal part of the tumulus, thereby exposing themselves (Kurczewski 2008b). Under presumably stressful circumstances, such as when being pursued by satellite-flies or with high sand surface temperatures, T. terminatus may enter its partly open nest directly, without pausing, maintaining the grasshopper underneath, instead of releasing it on the sand, entering, exiting, re-emerging from the burrow, and pulling it inside. Females of this species, under such adverse conditions, very rarely transport their prey into the burrow abdomen first rather than headfirst (Kurczewski 2009). This species and T. antillarum, when in hurry-up mode, may pull the prey into the nest by a leg or end of abdomen rather than by an antenna (Genaro and Sánchez 1992, Kurczewski 2009). Tachysphex terminatus, when nesting in steep sand slopes, is restricted to making an incomplete temporary closure of the nest entrance. In such cases, provisioning females enter the partly open burrow directly without pausing and releasing the grasshopper on the sand surface. Such behavior is reminiscent of that of T. apicalis, a related species that nests predominantly in sand cliffs and steep sand slopes (Kurczewski and Snyder 1968). Species in the terminatus group have the capability to prematurely close their nests under suboptimal conditions, including excessive cleptoparasitic pressure (Kurczewski 2009; F.E. Kurczewski, pers. observ.). Other conditions influencing premature nest closure, sometimes with only a single cell and one prey, include dusk, dense cloud cover, impending rainfall, and scarcity of prey. Tachysphex similis may oviposit and close nests prematurely with only a single cell and one prey when suitable-sized grasshoppers are rare and searching for additional prey is futile (Kurczewski 2009). Tachysphex inconspicuus often excavates several accessory burrows (Evans 1966b) around an entrance to distract or dissuade potential cleptoparasites, especially miltogrammine flies, from entering nests at sites where cleptoparasites are abundant (Buys 2007). They do not dig such burrows at other sites where the frequency of cleptoparasitism is reduced or negligible (Buys 2007). Tachysphex terminatus very rarely captures and stocks its cells with a single bush katydid, along with the usual grasshopper prey (Kurczewski 1966a). Tachysphex krombeini captures and stocks its cells with meadow katydids, along with grasshoppers, although such prey selection in this species may constitute typical behavior (Kurczewski 1971). Tachysphex mundus and T. inconspicuus rarely capture and stock in their cells tree crickets and crickets, respectively, along with the typical meadow katydid and cockroach prey (Buys 2007, Kurczewski 1979). 50 Northeastern Naturalist Vol. 17, Monograph No. 6 Tachysphex tarsatus very rarely lays its egg on the cervix (throat) of the grasshopper, rather than at the base of a foreleg, when prey individuals are exceedingly large and cumbersome to manipulate (Kurczewski 1991). Females of T. tarsatus may oviposit on a prey in the cell but omit final closure of the nest. Some wasps, nesting late in the season, hunt, capture and transport prey, and then final close their nests without laying an egg on the grasshopper, presumably having exhausted their egg supply (Kurczewski 1991). Discussion Some species with facultative diapause have only one or two generations per year in the Canadian and Transition faunal zones, respectively, but several generations per year in Austral faunal zones. Such species may have broad and often transcontinental geographic distributions (Pulawski 1988). Other species with facultative diapause and several generations per year are restricted geographically to the tropics or subtropics (Buys 2007; Elliott 1996; Elliott et al. 1979; Genaro 2004; Genaro and Sánchez 1992; Pulawski 1974, 1988). The number of generations per year in Europe for T. pompiliformis, a species that has only a single generation per year and a northern geographic distribution in North America (Pulawski 1988), is apparently one (Bonelli 1966, Pulawski 1971). Tachysphex laevifrons and T. pechumani, both in the pompiliformis group, have obligate diapause and one generation per year regardless of inhabiting more southern latitudes. Tachysphex laevifrons has a vernal flight season in the southeastern United States (Kurczewski 1971, 1987b, 2000b). The flight season of T. pechumani encompasses mainly late spring and early summer in southern New Jersey and the central Great Lakes Region (Kurczewski 2008b, Kurczewski and Elliott 1978, Kurczewski et al. 1970). The univoltine life cycle of T. pechumani is coincident with the period of maximal amount of daylight surrounding summer solstice, reduced fire frequency in late spring–early summer, and an abundance of suitable-sized potential prey grasshoppers at that time of year (Kurczewski 1998). Late emerging (early August), non-nesting T. pechumani females in northern Lower Michigan are exceptional and may represent stragglers from the first generation rather than a partial second generation of wasps (Kurczew-ski 2008b). Prey absence or insufficiency is not connected with the univoltine existence of T. pechumani and T. laevifrons. Suitable-sized grasshoppers of appropriate species are abundant during summer in the regions where these species occur. One generation per year and only a 4–5-weeks flight season in T. laevifrons and T. pechumani may seem unnecessarily restricted where, in both species, several additional months of seemingly favorable weather are available for nesting. Such a narrow time window may reflect a past geologic period when optimal climatic conditions were more abbreviated or climate was hotter, droughty, and unfavorable for nesting later in the summer (Kurczewski 1998, 2008b). 2010 F.E. Kurczewski 51 Tachysphex laevifrons and T. pechumani enter diapause with increased soil temperatures brought on by higher June (T. laevifrons) or July–August (T. pechumani) air temperatures. Tachysphex pechumani is ill-suited to withstand the higher temperatures of mid-summer, as evidenced by its bimodality in nesting on hot days with maximal activity in late morning and a secondary peak in mid–late afternoon (Kurczewski 2008b). This species does not inhabit the southern Atlantic Coastal Plain today probably because of that region’s high summer and mild winter temperatures, thermic soil temperature regime (annual mean soil temperature range is from 15 to <22 °C), large amount of summer precipitation, and seasonal high soil moisture values. Tachysphex pechumani needs an extended cold period in winter to break diapause (F.E. Kurczewski, pers. observ.). Most North American, Central American, and Caribbean species of Tachysphex nest in friable, level to moderately sloping, sandy or gravelly soils, as implied by the extensive foretarsal digging rake of the female consisting of long, slender, flexible, close-set setae (Bohart and Menke 1976, Pulawski 1988, Williams 1914). Psammophilous species tend to have a more extensive foretarsal rake than species that nest in less friable soils (Fig. 33). Tachysphex antennatus is genus-atypical in excavating nests beneath flat stones in gravelly and loamy, often hard-packed soils (Buck 2004, Kurczewski and Kurczewski 1987). Tachysphex acutus digs nests in sandy and gravelly loam of fields (Kurczewski 1989). Tachysphex apicalis, in the terminatus group, excavates nests in vertical sand cliffs and steeply sloping sandbanks, an unusual Tachysphex habitat (Kurczewski 2009, Kurczewski and Snyder 1968). Figure 33. Comparison of foretarsal digging rakes of Tachysphex propinquus (= T. ashmeadii), a psammophilous species, and Tachysphex iridipennis, which nests in finer-grained soils (Bohart and Menke 1976). 52 Northeastern Naturalist Vol. 17, Monograph No. 6 Table 1. Nesting behavior characteristics and prey type for North American, Central American, and Caribbean species of Tachysphex. No. No. Soil Temp Type Prey prey/ cells/ Prey Prey Species1 surf2 clos3 trans4 size5 cell6 nest7 stage8 type9 pompiliformis group pompiliformis + O G M, L 1–2 1 A, N Acrid montanus + O G M, L 1–2 1 A, N Acrid aethiops +, O O G L 1 2–5 N Acrid orestes G? M N Acrid acutus +, O O G M, L 1 1–2 N Acrid punctifrons +, O O G L 1-2 3–6+ A, N Acrid semirufus +? O G M 1 1 N Tetti pauxillus +? O G M 1 1 N Acrid hopi +? O G M, L 1–2? 1 N Acrid pechumani + + G M, L 1 1 A, N Acrid laevifrons + O G, F S–L 1–4 1 N Acrid tarsatus + O G, F S–L 1–4 1 A, N Acrid williamsi M N Acrid crenulatus N Acrid miwok F? S N Acrid krombeini + O F S 7 1 N Acrid, Tetti antennatus + O F S, M 3–9 1 N Acrid crassiformis + O G, F S–L 1–6 1 N Acrid occidentalis 2? N Acrid apricus10 L N Heter tipai N Tetti psammobius + O F S 11 1 N Acrid yolo O G,F S,M N Acrid texanus O F S, M 4 1 N Acrid ashmeadii + O G, F M, L 1–2 1 A, N Acrid terminatus group clarconis + + F S 3–6 2–3 N Acrid antillarum + + F S 6–11 1–5 N Acrid alpestris + + F S 2–13 1–3 N Acrid linsleyi + + F S 7 1 N Acrid terminatus + + F11 S, M 1–14 1–5 N Acrid, Tetti similis + + F11 S 1–15 1–5 N Acrid apicalis +, O O F S, M 2–12 1–3 A, N Acrid Species in the brullii group and T. aethiops and T. punctifrons, both in the pompiliformis group, renovate and use for nesting the pre-existing burrows of other wasps and bees (Table 1; Alcock 1973; Kurczewski 1979, 1987a; O’Brien 1987; Pulawski 1988). The latter two species often nest in sandy soils. Tachysphex pompiliformis may modify and nest in pre-existing subterranean burrows in Europe (Crèvecoeur 1951, Maneval 1932, Kurczewski and O’Brien 1988, Pulawski 1988). During excavation, many species of Tachysphex remove soil from the burrow in a common manner, loosening it with the mandibles and then using 2010 F.E. Kurczewski 53 the forelegs in unison to rake it backward beneath the synchronously lifting abdomen. Such a manner of soil removal may, in fact, apply to most North American, Central American, and Caribbean species, including T. mundus in the brullii group and ancestral species in the pompiliformis group that modify pre-existing burrows for use as nesting sites (Alcock 1973; Kurczewski 1979, 1987a; O’Brien 1987). Most species in the pompiliformis group make little or no attempt to level the soil removed from burrow excavation, resulting in a convex mound or tumulus in front of the entrance. The wasps probably use this mound of soil and open entrance to locate the burrow upon return with or without prey. The females also use the tumulus as ready-made fill for final closure of the nest (Kurczewski 1987b, 1989, 1991). Tachysphex pechumani may start excavating its burrow in typical Tachysphex manner but, once the wasp disappears below ground level, she only occasionally appears on the surface to remove the loosened sand. When she does resurface to remove soil from the burrow, she rarely backs farther than the length of her body. Females usually remain below ground Table 1, continued. No. No. Soil Temp Type Prey prey/ cells/ Prey Prey Species1 surf2 clos3 trans4 size5 cell6 nest7 stage8 type9 brullii group mundus O O F S 3–6 1 N Tetti, Gryll belfragei O? O G, F? S?, M 4 1 N Tetti menkei N Tetti obscuripennis group inconspicuus + + G, F S, M 1–5 1–2 A, N Blatt, Gryll iridipennis G, F? S, M? A, N Blatt alayoi + O G, F S, M 2 1 A, N Blatt julliani group coquilletti F? M? N Manti cockerellae N Manti 1Species listed phylogenetically following Pulawski (1988). 2Soil surf: + indicates burrow excavated from soil surface, O indicates burrow started from pre-existing burrow. 3Temp clos: + indicates species temporarily fills entrance with soil when hunting, O indicates species leaves entrance open when hunting. 4Type trans: G indicates ground transport of prey, F indicates flight transport of prey. 5Prey size: S indicates prey smaller than wasp, M indicates prey moderate-sized (same size as wasp or slightly larger), L indicates prey noticeably larger than wasp. 6No. prey/cell = number of prey per fully provisioned cell. 7No. cells/nest = number of cells per completed nest. 8Prey stage: A indicates adult, N indicates nymph. 9Prey type abbreviations: Acrid = Acrididae, Tetti = Tettigoniidae, Gryll = Gryllidae, Blatt = Blattellidae, Manti = Mantidae, Heter = Heteronemiidae. 10Single prey available for T. apricus is four times longer than wasp and of questionable authenticity. 11Vast majority of prey (>98%) were transported in flight. 54 Northeastern Naturalist Vol. 17, Monograph No. 6 level, inside their burrow, and fling an arc-shaped “spray” of sand backward from the entrance, resulting in a well-distributed, very shallow, dark-colored (charcoal-laced), fan-shaped sand deposit on the lighter sand surface. Such a dark-colored, fan-shaped tumulus and depressed entrance from the rudimentary temporary closure probably function as guides to enable the wasp to more readily find its burrow (Kurczewski 2008b). Species in the terminatus group, except T. apicalis, and T. inconspicuus, in the obscuripennis group, level the sand removed from burrow excavation by raking it in various directions with the forelegs. They then make a temporary closure of the burrow by filling the entrance with soil (Table 1; Buys 2007; Elliott 1996; Elliott and Kurczewski 1985; Genaro and Sánchez 1992; Krombein 1964; Kurczewski 2009; Kurczewski et al., in press). Leveling the surface and temporarily closing the entrance with soil serve to make the nest area visibly indistinguishable from the surrounding sand and may function as a deterrent to cleptoparasites and other wasp enemies attempting to find the nest. During leveling, the wasp marks the area with her mandibular glands perhaps to assist her in finding the entrance. Temporary closure of the entrance is usually associated with species that stock several or many prey per fully provisioned cell and nest in barren or sparsely vegetated sand. Tachysphex apicalis does not temporarily close its nest entrance, having apparently lost or never acquired this highly characteristic nesting behavior component of the terminatus group likely due to its atypical nesting habitat—sand cliffs and steep slopes (Kurczewski 2009, Kurczewski and Snyder 1968). Running around the entrance on the ground or flying above it in increasingly larger circles as a form of orientation to the nest surroundings is connected with degree of soil barrenness, presence or absence of temporary closure, size of prey, and method of transport. Species such as T. acutus and T. pechumani that nearly always return to the nest on the ground with larger prey, walk or run around the entrance before hunting (Kurczewski 1989, 2008b). Species in the terminatus group, T. mundus, and T. inconspicuus fly in increasingly larger circles prior to hunting and return to the nest with small prey nearly always in extended flight (Buys 2007; Elliott 1996; Elliott and Kurczewski 1985; Kurczewski 1979, 2009; Kurczewski et al., in press). Females of T. ashmeadii, T. tarsatus, T. crassiformis, and T. laevifrons, all in the pompiliformis group, run around the entrance or make low, circular flights above the area before searching for prey (Elliott and Kurczewski 1985; Kurczewski 1987b, 1991). Females that run around the entrance often return carrying larger grasshoppers on the ground, but infrequently return to their nests with smaller prey in a low series of flights. Wasps that orient in flight almost always return with smaller prey in longer, more extended flights. Species in the terminatus group in temperate regions are better adapted to nest in spring–early summer than in mid–late summer. There is more rainfall, fresher and larger quantity of vegetation for the young grasshoppers, 2010 F.E. Kurczewski 55 and an abundance of small nymphal prey. Tachysphex pechumani, T. laevifrons, T. antennatus, and, perhaps, T. psammobius are restricted to nesting in spring–early summer or nest more successfully during that time of year (Kurczewski 1987a, 1987b, 2000b, 2008b; Kurczewski and Elliott 1978; Kurczewski and Kurczewski 1987). Tachysphex acutus and T. tarsatus, on the other hand, nest more successfully in mid–late summer in the northeastern US, when Melanoplus femurrubrum, their primary prey at that time of year, is in abundant supply. Many species in the pompiliformis group hunt relatively large grasshoppers (especially Oedipodinae and Melanoplinae), usually restrict their hunting forays to the ground or low vegetation, and hunt rather close (<10 m) to their nests (Alcock 1973; Kurczewski 1987b, 1989, 1991, 2008b; Kurczewski and Elliott 1978; Williams 1914). Species in the terminatus group and T. mundus often search for smaller prey on upright vegetation, often at some distance (15–60 m) from their nests (Elliott 1996; Kurczewski 1979, 2009). Species in the obscuripennis group hunt for small cockroaches in high vegetation above their nesting area. Tachysphex inconspicuus flies into the tropical wet forest canopy in search of mainly adult females of certain genera of blattellid cockroaches (“canopy indicators”; Fisk 1983; Kurczewski et al., in press; Schal et al. 1984). Elliott et al. (1979) observed provisioning females of T. alayoi flying to the ground from nearby plants with paralyzed cockroaches. Under laboratory conditions, prey of T. pompiliformis are subdued by four successive stings in the soft intersegmental membrane encircling a hindleg, foreleg, midleg, and cervix (throat), in that order (Steiner 1981a, b). The critical insertion of the first sting in the corium surrounding a hindcoxa effectively prevents the grasshopper from leaping away (Steiner 1976, 1981b). Four to eight stings made by T. tarsatus and up to five stings made by T. laevifrons and T. crassiformis, all in the pompiliformis group, are sometimes required to paralyze larger grasshoppers in the field (Kurczewski 1987b, 1991). Capture of small prey by species in the terminatus group sometimes involves only one, two, or three stings in some of the same areas of the prey’s body as for T. pompiliformis (Kurczewski 2009). However, it is not known whether this represents the full arsenal of wasp stings in these species. Prey capture in T. pechumani is unlike that in other Tachysphex. Females capture only Gomphocerinae and Melanoplinae. They circle the grasshopper once or twice in clockwise and counterclockwise directions, always facing it, while constantly waving their mostly orange antennae. Females pounce on the acridid from the side and, maintaining their bodies at about a 90° angle to the longitudinal axis of the prey, insert the sting into the hindcoxal corium followed by successive other sting insertions (Kurczewski 2008b, Kurczewski and Elliott 1978). The circling behavior in which the antennae are waved up and down is reminiscent of courtship behavior in certain grasshoppers and may function in holding the acridid’s attention (F.E. Kurczewski, pers. observ.). 56 Northeastern Naturalist Vol. 17, Monograph No. 6 Females of T. pechumani totally ignore species of Oedipodinae (band-winged grasshoppers) when hunting, and either walk around such individuals or fly away from them (Kurczewski 2008b, Moan and Tramer 2008). Nymphal oedipodines of the genera Dissosteira and Spharagemon of favorable prey size are never captured, although they are often abundant in the nesting area. Tachysphex pechumani females may be unable to straddle such grasshoppers appropriately for stinging purposes because of their stiff mid-dorsal thoracic crest (Moan and Tramer 2008). Yet T. aethiops, T. tarsatus, T. crassiformis, and T. ashmeadii prey on sizeable nymphs of Dissosteira or Spharagemon. Adults of these grasshopper genera, too large to be captured by T. pechumani, lack this prominent thoracic crest (Moan and Tramer 2008). Malaxation often follows prey capture in Tachysphex species. It may occur intermittently during prey transport or perhaps below ground inside the cell and out of view (Kurczewski 1987b, 1989, 1991, 2008b, 2009; Kurczewski and Elliott 1978; Kurczewski and O’Brien 1988; Williams 1914). The function of malaxation has not been ascertained. Softening the intersegmental membrane surrounding the base of a foreleg with the mandibles may facilitate egg affixation. Applying the mouthparts to the stung area enables the wasp to obtain a source of nourishment (hemolymph) exuding from the puncture wound. Grasshoppers that are malaxated extensively are sometimes abandoned, suggesting they do not provide a satisfactory source of food for the developing wasp larva (Kurczewski 1987b, 1991). Mutilation of the prey’s appendages has been noted in a number of Tachysphex species (Genaro 2004; Kurczewski 1987a, 1989, 1991, 2008b, 2009; Kurczewski et al., in press). Some of the missing appendages on the prey are the result of accidental mishaps in the field not involving the wasps. Other instances of missing appendages or segments thereof are probably due to excessive handling of the prey during capture or transport or extensive malaxation following prey capture. Evans et al. (1976) and Genaro (2004) observed missing appendages or parts thereof on the cockroach prey of species in the obscuripennis group and believed this was the result of malaxation of the prey or rough handling of the cockroach during capture. Manner of prey transport is related to prey size in Tachysphex. Larger prey are transported on the ground, and smaller prey are carried in flight (Table 1; Alcock 1973; Elliott 1996; Evans 1970; Kurczewski 1979, 1987a, 1987b, 1989, 1991, 2009; Kurczewski and Evans 1986; Williams 1914). In T. pechumani, however, even smaller grasshoppers probably capable of being carried in flight are nearly always transported on the ground (Kurczewski 2008b). Only one of >250 observations in this species involved prey transport of a small grasshopper in short, low flights (Kurczewski 2008b, Kurczewski and Elliott 1978). In the pompiliformis group, larger prey are straddled more anteriorly than smaller prey during prey transport, held farther out on the antennae with the mandibles, and grasped more forward on the body with the hindlegs 2010 F.E. Kurczewski 57 (Kurczewski 1987b, 1989, 1991; Kurczewski and Elliott 1978). Prey transport in some of these species involves grasshoppers that weigh six to nine times the weight of the wasp. Prey transport in the terminatus, brullii, and obscuripennis groups is nearly always in flight and usually involves small prey (Table 1; Buys 2007; Elliott 1996; Elliott et al. 1979; Genaro and Sánchez 1992; Kurczewski 1979, 2009; Kurczewski and Evans 1986; Kurczewski et al., in press; Rau 1933). The ratio of weight of prey to wasp at which T. terminatus and T. similis switch from flight to ground transport is >2.1–2.3 (Kurczewski 2009). Tachysphex alayoi and T. inconspicuus females fly with prey weighing twice to nearly three times their own weight (Elliott et al. 1979; Kurczewski et al., in press). In the terminatus group, prey transport in the dorsal side upward position or on the side instead of the usual ventral side upward position is probably related to the small sizes of the grasshoppers (Kurczewski 2009). The duration between bringing consecutive prey to the nest is often related to prey size—smaller prey being provisioned more rapidly than larger prey— and distance from point of capture (Kurczewski 1979, 1987b, 1991, 2008b, 2009). It is also connected with the relative abundance and availability of suitable-sized individuals in the nesting area, as exemplified by T. antennatus in the pompiliformis group (Kurczewski and Kurczewski 1987). Tachysphex terminatus and T. similis spend significantly less time obtaining first than second and later generation prey individuals (Kurczewski 2009). There are more prey species, especially nymphs of small size, available to the wasps earlier in the nesting season (A.B. Gurney 1963, pers. comm.). Tachysphex similis, T. terminatus, and T. apicalis, all in the terminatus group, fly with smaller prey, stock several or many individuals per cell, and complete up to 5, 4, and 3 cells, respectively, in a single day (Kurczewski 2009). The first two species complete more cells per day early in the nesting season than later in the year due largely to an abundance of small, readily available prey. Tachysphex tarsatus, T. pechumani, and T. acutus, all in the pompiliformis group, mainly transport relatively large prey on the ground, stock one or a few individuals per cell, and complete up to 3, 2, and 1.5 cells, respectively, in a single day (Kurczewski 1989, 1991, 2008b). Tachysphex tarsatus excavates burrows, hunts, transports prey, and closes nests about twice as fast as T. acutus when the two species are nesting side by side in the same soil (Kurczewski 1989, 1991). Manner of nest entry with prey is associated with presence or absence of temporary closure and size of prey in Tachysphex species. In the case of the nest entrance being left open during provisioning, smaller prey are usually taken directly into the burrow while larger prey are released outside or just within the opening and dragged in from inside the nest (Kurczewski 1987a, 1987b, 1989, 1991). In the terminatus group and T. pechumani, the prey is released on the ground near the nest entrance prior to removing the soil fill (Kurczewski 2008b, 2009; Kurczewski and Elliott 1978). 58 Northeastern Naturalist Vol. 17, Monograph No. 6 Regardless of whether or not the species makes a temporary closure, most Tachysphex release the partly paralyzed prey in front of the entrance with its head near the opening and its body in a direct line with the orientation of the burrow. Females of T. pechumani, unlike other congeners, release their prey with its head near the entrance, but position the longitudinal axis of the prey’s body variously with respect to the direction of the burrow. The grasshoppers are often placed at right angles or even behind the entrance, rather than in front of it (Kurczewski 2008b, Kurczewski and Elliott 1978). Tachysphex inconspicuus, in the obscuripennis group, is genus-atypical in retaining its grasp of the prey’s antennae with the mandibles and its body with a hindleg when entering the nest. The wasp holds the prey to one side as it removes the temporary sand fill from the entrance with the forelegs and enters headfirst without releasing the cockroach (Buys 2007; Kurczewski et al., in press). Holding the prey to one side may enable the female to remove the soil from the entrance unimpededly, thus ensuring a more rapid entry. Such an entry would have positive implications for the wasp considering its endless encounters and interactions at the nest with foraging ants and cleptoparasitic flies (Kurczewski et al., in press). Two palearctic species, T. albocinctus (Lucas) in the albocinctus group, a mantid-hunter (Asis et al. 1989), and T. mediterraneus Kohl in the plicosus group, a tree-cricket hunter (Ferton 1923), exhibit similar nest entry behavior. These two species and T. inconspicuus nest in sandy soils, level the tumulus following burrow excavation, and make a temporary closure of the nest entrance prior to hunting for prey. Many genera in the Tribe Larrini make deep, multi-celled nests (Bohart and Menke 1976). Tachysphex is exceptional in excavating relatively short burrows and shallow cells (Williams 1914). Tachysphex aethiops and T. punctifrons use pre-existing burrows of other wasps and bees and make longer and deeper multi-celled nests than species that excavate their own burrows from the ground surface (Table 1; Kurczewski 1987a, O’Brien 1987). However, T. aethiops and T. punctifrons are two of the largest North American Tachysphex (Pulawski 1988). Larger species, e.g., Tachytes, Crabro, Philanthus, Cerceris, tend to excavate longer burrows and deeper cells than smaller congeners when nesting in the same soils (F.E. Kurczewski, pers. observ.). Larger T. similis females in Florida dig longer burrows and deeper cells than smaller T. similis females from other regions (Kurczewski 2009). Species in the terminatus group typically make multi-celled nests (Table 1). Multi-celled nests are associated with dense aggregations and nesting in close proximity. Multi-celled nests are also associated with barren or sparsely vegetated sandy soils. Conserving space by making a single multi-celled nest instead of several one-celled nests is advantageous where the amount of suitable soil is limited (Kurczewski 2009). Tachysphex terminatus excavates longer burrows and deeper cells in loose, coarse2010 F.E. Kurczewski 59 textured sand and shorter burrows and shallower cells in compact, loamy fine sand (Kurczewski 2009). Nests of T. tarsatus in loose, coarse-textured sand are often longer and deeper than nests in sandy loam, gravelly loam, or loamy fine sand (Kurczewski 1991). First-generation females of T. tarsatus, T. crassiformis, T. terminatus, and T. similis usually excavate longer burrows and deeper cells than second- and later-generation females of these species (Kurczewski 1987b, 1991, 2009). A decrease in mean burrow length and mean cell depth from spring to late summer at many sites is inexplicable. The different generations of wasps often nest in the same soils, thus eliminating a difference in soil type or texture as an explanation. Longer and deeper first generation Tachysphex nests in Kansas and New York usually coincide with higher soil moisture values at that time of year (Kurczewski 1987b, 1991, 2009). In the Bahamas, longer and deeper June nests of T. similis coincide with much more precipitation than shorter and shallower April nests of this species (Elliott 1996). Yet, longer and deeper nests in other digger wasp genera are often associated with drier soil (Alcock and Ryan 1973, Hager and Kurczewski 1986, Kurczewski 2002, Kurczewski and Wochadlo 1998). Surface light levels or soil moisture may control burrow length and cell depth in the sphecid Sphex ichneumoneus (L.) (Brockmann 1980). Variability in burrow length in this species may also result from local surface conditions such as amount of exposure to sunlight and sand temperature or from presence or absence of plants and soil moisture intake (Brockmann 1980). Longer burrows and deeper cells of first generation Tachysphex in the Northeast coincide with increased soil moisture, longer day length, and high sand surface temperature. Other factors to consider in longer and deeper first generation nests are small prey, less individual prey surface area, decreased chance of prey dessication, and the insulating effect of snow cover on larval wasp survival in second and later generation cells. Genera in the Tribe Larrini prey almost exclusively on orthopteroid insects (Bohart and Menke 1976). The vast majority of North American, Central American, and Caribbean Tachysphex in the pompiliformis and terminatus groups prey on Acrididae (Table 1 and Appendix 1; Krombein 1979, Kurczewski 1987a, Pulawski 1988). Tachysphex krombeini, in the pompiliformis group, captures and stocks acridids and tettigoniids mixed in its cells (Kurczewski 1971). The propensity for preying on Tettigoniidae exists in T. semirufus and T. tipai, both in the pompiliformis group (Kurczewski and Evans 1986, Pulawski 1988). There is one questionable record of T. apricus, in the pompiliformis group, pinned with a much larger nymphal walkingstick (Elliott and Kurczewski 1985, Pulawski 1988). Tachysphex terminatus, in the terminatus group, very rarely stocks its cells with a single bush katydid in addition to its usual grasshopper prey (Kurczewski 1966a). Tachysphex mundus, in the brullii group, sometimes uses gryllid as well as tettigoniid prey (Kurczewski 1979), and T. inconspicuus, in the obscuripennis group, very rarely captures Gryllidae in addition to 60 Northeastern Naturalist Vol. 17, Monograph No. 6 its usual blattellid prey (Buys 2007). Tachysphex species probably have an inherent capability allowing females to switch from typical to atypical prey when typical prey numbers are scarce or rare in the field. There may be a degree of prey specificity in some species of the pompiliformis group (Appendix 1). Tachysphex aethiops preys only on Oedipodinae (Alcock 1973; Evans 1970, 1973; O’Brien 1987), T. crassiformis captures mainly Oedipodinae (Krombein 1963, Kurczewski 1987b), and T. punctifrons and T. acutus prey on the genus Melanoplus (Melanoplinae) (Kurczewski 1987a, 1989, 2000b). Tachysphex pechumani stocks only Gomphocerinae and Melanoplinae in its cells (Kurczewski 2008b, Kurczewski and Elliott 1978, Moan and Tramer 2008). Tachysphex tarsatus captures and stocks different species of Acrididae in its cells according to nesting habitat (Kurczewski 1991). This species, T. terminatus, and T. similis stock different species of Acrididae in late spring– early summer versus mid–late summer in connection with varied grasshopper phenology and seasonal prey availability (Kurczewski 1991, 2009). There is a gradual decrease in the number of potential small prey available to members of the terminatus group throughout the nesting season, except in the tropics and subtropics (Kurczewski 2009). In T. tarsatus in late summer in the Northeast, larger prey individuals are readily available for capture due to an abundance of appropriate-sized Melanoplus femurrubrum (Kurczewski 1991). Very large sample sizes for T. terminatus and T. similis, two species with similar nesting behavior, reveal overall equivalent percentages of subfamilies of Acrididae as prey. Females of these species are evidently unselective of their prey species and capture whatever small nymphal grasshoppers are abundant and available in the nesting area (Kurczewski 2009; A.B. Gurney, 1963 pers. comm.). The high degree of prey specificity in the brullii (Tettigoniidae) and obscuripennis (Blattellidae) groups is a valid reason for separating them (Table 1; Bohart and Menke 1976). Another logical reason for maintaining the distinctness of the brullii and obscuripennis groups is the dissimilarity in nesting behavior (Table 1). Species in the brullii group renovate pre-existing burrows for use as nest sites, leave the entrance open when hunting for prey, and dive into the open burrow holding the prey underneath. They lay their egg transversely across the prey’s pro- and mesosterna between the bases of the fore- and midcoxae (Kurczewski 1979). Tachysphex inconspicuus, in the obscuripennis group, excavates its nest from the sand surface, levels the tumulus, makes a temporary closure of the entrance, and retains its grasp of the prey with the mandibles and a hindleg when removing the temporary closure and entering the burrow. Tachysphex inconspicuus, T. alayoi, and other species in the obscuripennis group lay their egg longitudinally posteriad on the thoracic venter of the cockroach (Buys 2007; Evans et al. 1976; Genaro 2004; Kurczewski et al., in press). There appears to be a degree of prey specificity among species in the obscuripennis group (Appendix 1). Tachysphex inconspicuus stocks its 2010 F.E. Kurczewski 61 cells with cockroaches belonging to the genera Chorisoneura and Riatia (family Blattellidae, subfamily Pseudophyllodromiinae; Appendix 1; Buys 2006, 2007; Callan 1954, 1993; Kurczewski et al., in press; Pulawski 1974; G. Beccaloni, Natural History Museum, London, UK, 2009 pers. comm.). Species of Chorisoneura and Riatia are common inhabitants in the canopy of trees in tropical wet forests (Fisk 1983; Kurczewski et al., in press; Schal et al. 1984). Tachysphex iridipennis and T. alayoi provision their nests with other genera and species of Blattellidae, including some not restricted to the tropical wet forest canopy (Appendix 1; Elliott et al. 1979, Genaro 2004, Pulawski 1988, Vesey-FitzGerald 1956). In T. tarsatus and T. crassiformis, a decrease in the mean number of prey per cell from late spring to late summer is correlated with an increase in the mean size and weight of prey individuals, although this correlation is not perfectly reciprocal (Kurczewski 1987b, 1991). In T. tarsatus, cells with the largest amount of biomass in September produce the smallest and lightest wasps in June of the following year, implying that overwintering may have a deleterious effect on the developing wasp larva (Kurczewski 1991). There is a positive correlation between wasp size and prey size in T. pechumani and T. tarsatus (Kurczewski 1991, 2008b). Species in the terminatus group stock more and smaller prey per cell early in the nesting season and fewer larger grasshoppers later in the nesting season (Kurczewski 2009). The number of prey per cell is approximately halved from May–June to August in T. terminatus. Not only are fewer grasshoppers stocked in the cells later in the year, but there is also a reduction in total biomass of prey per cell resulting in slightly smaller first generation wasps the following spring (Elliott and Kurczewski 1975, Kurczewski 2009). In Tachysphex, most prey are placed in the cells in a head inward and ventral side upward position. Prey found in other positions are either taken into the nest in an atypical manner or are accidentally rearranged by the wasps maneuvering to oviposit within the confines of the prey-filled cell (Kurczewski 2009). The wasp’s egg is not affixed to a prey at the front end of the cell as this individual might be readily discovered and destroyed by enemies entering the nest (Kurczewski 2009). Tachysphex eggs laid on acridid or tettigoniid prey are affixed to the soft intersegmental membrane surrounding the base of the prey’s forecoxa (Elliott 1996, Evans 1970, Genaro and Sánchez 1992, Krombein 1964, Kurczewski 2009, Kurczewski and Kurczewski 1987, Newton 1956, Williams 1914). The egg extends transversely across the ventral thorax between the bases of the foreand midlegs. Similarity in egg affixation site in these species and several other genera of Larrini is connected with provisioning with Caelifera (grasshoppers) and Ensifera (katydids, crickets; Bohart and Menke 1976). Eggs laid by species in the obscuripennis group are also attached to a forecoxal corium, but they extend longitudinally posteriad on the ventral thorax of the cockroach (Evans et al. 1976; Genaro 2004; Kurczewski et al., in press). Eggs laid by exotic species of Tachysphex that prey on mantids are 62 Northeastern Naturalist Vol. 17, Monograph No. 6 placed variously: T. costae (De Stefani Perez), in the erythropus group, at the base of an anterior leg (Deleurance 1945, Grandi 1930); T. albocinctus, in the albocinctus group, outside the jointure of one of the anterior legs (Ferton 1912); and, T. multifasciatus Pulawski, also in the albocinctus group, “across the joint of the femur and tibia of a middle leg” (Evans et al. 1976). Tachysphex wasps are almost never without their associated cleptoparasites, particularly miltogrammine flies. Certain Tachysphex behavior patterns are probably the result of a long history of conjoined activities between the wasps and flies and repeated attempts on the part of the wasps to avoid or thwart fly attacks (Evans 1966a, 1970; Evans and West Eberhard 1970). Miltogrammine flies utilize many aspects of the wasp’s nesting sequence for deposition of their maggots (Spofford and Kurczewski 1990, Spofford et al. 1986). The cleptoparasitic flies produce successive generations at a much faster rate over a much shorter time period than Tachysphex wasps (Spofford and Kurczewski 1992). Such advantages on the side of the flies undoubtedly account for the relatively high frequency of cleptoparasitism of species in the terminatus group (Genaro and Sánchez 1992, Spofford et al. 1986). Some Tachysphex behavioral components represent adaptations that reduce the incidence of cleptoparasitic fly attacks (Buys 2007, McCorquodale 1986, Spofford et al. 1986). Prey and cell abandonment is an effective preventative against miltogrammine fly attacks. The wasp mitigates her losses by not wasting an egg on a cleptoparasitized prey or in a cell containing maggots. Prey cleaning, including maggot removal or dismemberment, is an effective defense against fly larviposition in the terminatus group (Genaro and Sánchez 1992, Spofford et al. 1986). Acknowledgments Julio A. Genaro kindly agreed to serve as Guest Editor for this manuscript. Julio A. Genaro, Keith Goldfarb, Allan Hook, and Wojciech Pulawski read the manuscript critically. Richard Archbold, Leonard Brass, Terry Crabe, Thomas Eisner, Denise Gehring, Michelle Grigore, Bill Huff, John Lerg, Rob Line, Charles Michener, Verne Pechuman, Mary Rabe, and Bill Westrate aided with equipment, supplies, facilities, or funding. Hugh Boyle, Sean Brady, Dana Bruington, Matthias Buck, George Byers, Peter Carson, Nancy Burdick Elliott, Howard Evans, David Furth, Grant Gaumer, A. R. Gittins, Barbara Harris, Brian Harris, Paul Hurd, Jr., Bob Jacksy, Lynn Kimsey, Edmund Kurczewski, Keith Kurczewski, H. B. Leech, George Matuza, W.W. Middlekauff, Richard Miller, Mary Morlando, Mark O’Brien, Laurence Packer, Ann Peck, Wojciech Pulawski, Jerome Rozen, Jeff Skevington, Noel Snyder, Margery Spofford, Fred Stehr, and Ed Stanton furnished collection information or field assistance. Howard Evans initially proposed a comparative nesting behavior study of Tachysphex tarsatus and T. terminatus. Noel Snyder added numerous research suggestions and took many of the photographs. Rollin Coville and Coby Schal sent me their research notes on T. inconspicuus nesting in Costa Rica. Coby Schal furnished the photographs of nest entry with prey and cockroach prey and eggs of T. inconspicuus. Grant Gaumer, Bob Jacksy, and Art Weber provided photographs of T. pechumani or T. tarsatus. John Alcock is responsible for Figure 31 (T. pompiliformis) and David Peckham for Figure 32 (T. terminatus). George Snyder photographed the egg and prey of T. antennatus. Frances McKittrick drew Figure 8 (T. apicalis). Joe Stoll 2010 F.E. Kurczewski 63 re-touched and image-scanned the photographs. Don Artz formatted Figure 1. John Simeone translated Bonelli’s (1966) study of T. pompiliformis and Grandi’s (1930) study of T. costae from Italian to English. Julio A. Genaro translated his 2004 paper on T. alayoi from Spanish to English. George Beccaloni, M.A. Brusven, Irving Cantrall, George Eickwort, Frank Fisk, Ashley Gurney, T.H. Hubble, Norman Marston, David Nickle, Mark O’Brien, Dan Otte, and D.C.F. Rentz identified Tachysphex prey or associated insects. W.L. Downes, Jr., R.J. Gagne, H.J. Reinhard, and Margery Spofford named the cleptoparasitic Miltogrammini. Richard Bohart, Karl Krombein, and Wojciech Pulawski identified or confirmed my identifications of Tachysphex species. Funds and assistance for travel, per diem, field materials, equipment, and manuscript preparation were provided by the Archbold Biological Station, Sigma Xi-RESA Grants-in-Aid of Research, Cornell University Department of Entomology and Faculty Research Fund, National Science Foundation Postdoctoral Fellowship and Undergraduate Research Participation Programs, National Institutes of Health Postdoctoral Fellowship, The University of Kansas Department of Entomology, State University of New York-RESA Grants-in-Aid of Research, Michigan Chapter of The Nature Conservancy, Michigan Department of Natural Resources, The University of Michigan Museum of Zoology, Toledo Area Metroparks, State University of New York College of Environmental Science and Forestry Office of Research Programs, Biden Environmental Center at Cape Henlopen State Park, Delaware, and New York State United University Professions. Literature Cited Adlerz, G. 1904. 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Wasp species1 Prey species2,3 Source of information pompiliformis group pompiliformis Aulocara elliotti (Thomas) Newton 1956, Pulawski 1988 Camnula pellucida (Scudder) Krombein 1967 Melanoplus viridipes Scudder (adult male) Kurczewski and O’Brien 1988 Melanoplus sp., possibly bivittatus (Say) Kurczewski and O’Brien 1988 Melanoplus sp., probably sanguinipes (Fabricius) Kurczewski and O’Brien 1988 Melanoplus spp. Krombein 1967; Kurczewski and O’Brien 1988; Newton 1956; Pulawski 1988; M. Buck, 2009 pers. comm. Oedaleonotus enigma (Scudder) Kurczewski and O’Brien 1988, Newton 1956, Pulawski 1988 montanus Oedaleonotus enigma Newton 1956 Oedaleonotus sp. Krombein 1967 aethiops Camnula pellucida M. Buck, 2009 pers. comm. Spharagemon sp., probably bolli Scudder O’Brien 1987 Trimerotropis sp., probably verruculata suffusa Scudder Evans 1973 Trimerotropis sp. Evans 1970 orestes Gomphocerinae, ensiform antennae L.S. Kimsey, University of California, Davis, CA, 2009 pers. comm.; D.A. Nickle, Smithsonian Institution, Washington, DC, 2009 pers. comm. acutus Melanoplus femurrubrum (DeGeer) Kurczewski 1989 Melanoplus sanguinipes Kurczewski 1989 Melanoplus sp. Kurczewski 1989 punctifrons Melanoplus bivittatus (+ adult) Kurczewski 2000b Melanoplus differentialis (Thomas)(+ adults) Kurczewski 1987a Melanoplus infantilis Scudder (adult) Kurczewski 1987a, Pulawski 1988 Melanoplus sp., probably bivittatus Kurczewski 1987a, Pulawski 1988 Melanoplus sp. (+ adult female) Kurczewski 1987a semirufus Anabrus simplex Haldeman (TETTIGONIIDAE) Kurczewski and Evans 1986 Capnobates sp., probably occidentalis (Thomas) (TETTIGONIIDAE) Pulawski 1988 pauxillus Melanoplus sp. Evans 1970 2010 F.E. Kurczewski 71 Wasp species1 Prey species2,3 Source of information hopi Aulocara elliotti Pulawski 1988 Oedaleonotus enigma Newton 1956, Pulawski 1988 pechumani Chloealtis conspersa (Harris) Kurczewski 2008b Melanoplus bivittatus Kurczewski 2008b Melanoplus eurycerus Hebard Kurczewski 2008b Melanoplus fasciatus (F. Walker) Kurczewski 2008b, Kurczewski and Elliott 1978 Melanoplus impucidus Scudder (+ adult male) Kurczewski 2008b, Kurczewski and Elliott 1978 Melanoplus sanguinipes (+ adult males) Kurczewski 2008b, Kurczewski and Elliott 1978 Melanoplus viridipes Kurczewski 2008b Melanoplus spp. Kurczewski 2008b; L.S. Kimsey, 2009 pers. comm.; D.A. Nickle, 2009 pers. comm. Orphulella pelidna (Burmeister) (+ adult) Kurczewski and Elliott 1978 laevifrons Dissosteira carolina (Linnaeus) Kurczewski 2000b Melanoplus puer group Kurczewski 1987b Melanoplus sp. Kurczewski 1987b, 2000b tarsatus Ageneotettix deorum Scudder Kurczewski 1991 Aulocara elliotti Kurczewski 1991 Camnula pellucida Rust et al. 1985 Chloealtis conspersa Kurczewski 1999 Chorthippus curtipennis (Harris) Kurczewski 1991 Chorthippus sp. Kurczewski 1991 Chortophaga viridifasciata (DeGeer) Kurczewski 1991 Dissosteira carolina Elliott and Kurczewski 1985; Kurczewski 1991, 1999 Encoptolophus sordidus (Burmeister) Kurczewski 1991 Melanoplus bivittatus (+ adult males) Kurczewski 1991 Melanoplus complanatipes Scudder Elliott and Kurczewski 1985 Melanoplus devastator Scudder Kurczewski 1991 Melanoplus femurrubrum (+ adult males) Kurczewski 1991, 1999 Melanoplus sanguinipes (+ adult male and female) Kurczewski 1991 Melanoplus spp. Elliott and Kurczewski 1985; Evans 1970; Kurczewski 1991, 1999; Williams 1914; M. Buck, 2009 pers. comm. Oedaleonotus enigma Kurczewski 1991 72 Northeastern Naturalist Vol. 17, Monograph No. 6 Wasp species1 Prey species2,3 Source of information Orphulella pelidna (adult male) Elliott and Kurczewski 1985 Orphulella speciosa (Scudder) (adult male) Kurczewski 1991 Pseudopomala brachyptera (Scudder) Elliott and Kurczewski 1985 Psinidia fenestralis (Serville) Kurczewski 1991 Trimerotropis maritima interior (E.M. Walker) Kurczewski 1991, 1999 Trimerotropis m. maritima (Harris) Kurczewski 1991 Trimerotropis spp. Elliott and Kurczewski 1985, Evans 1970, Kurczewski 1991 williamsi Melanoplus sanguinipes4 Elliott and Kurczewski 1985 Trimerotropis occidentalis (Bruner) Elliott and Kurczewski 1985 crenulatus Melanoplus marginatus (Scudder) Pulawski 1988 Trimerotropis sp. Pulawski 1988 miwok Conozoa sulcifrons (Scudder) Pulawski 1988 krombeini Melanoplus sp. Kurczewski 1971 Odontoxiphidium apterum Morse (TETTIGONIIDAE) Kurczewski 1971 antennatus Chorthippus curtipennis Buck 2004 Melanoplus bivittatus Kurczewski and Kurczewski 1987 Melanoplus f. femurrubrum Kurczewski and Kurczewski 1987 Melanoplus keeleri luridus (Dodge) Kurczewski and Kurczewski 1987 Melanoplus s. sanguinipes Kurczewski and Kurczewski 1987 Melanoplus sp. Elliott and Kurczewski 1985, Kurczewski and Kurczewski 1987 crassiformis Chortophaga sp. Krombein 1967, Kurczewski 1987b Psinidia fenestralis Krombein 1963, Kurczewski 1987b Spharagemon marmorata picta (Scudder) Krombein 1963 Trimerotropis sp. Kurczewski 1987b occidentalis Melanoplus sp., probably cinereus Scudder Pulawski 1988 Schistocerca shoshone (Thomas) Elliott and Kurczewski 1985 apricus Parabacillus hesperus Hebard (HETERONEMIIDAE)? Elliott and Kurczewski 1985 tipai Ateloplus notatus Scudder (TETTIGONIIDAE) Pulawski 1988 Pediodectes sp. (TETTIGONIIDAE) Pulawski 1988 2010 F.E. Kurczewski 73 Wasp species1 Prey species2,3 Source of information psammobius Aerochoreutes (= Circotettix) maculatus (Scudder) Kurczewski 1987a Melanoplus sp., either differentialis or sanguinipes Kurczewski 1987a yolo Melanoplus foedus Scudder Elliott and Kurczewski 1985 Melanoplus sp. Kurczewski 1987a texanus Arphia sp. Kurczewski 1987a, Williams 1914 Encoptolophus robustus Rehn and Hebard Rust et al. 1985 Melanoplus sp. near flavidus Scudder Kurczewski 1987a Orphulella pelidna Kurczewski 1987a ashmeadii Ageneotettix deorum Lavigne and Pfadt 1966, Williams 1914 Arphia sp. Elliott and Kurczewski 1985 Aulocara sp. Pulawski 1988 Bruneria sordida (McNeill) Elliott and Kurczewski 1985 Conozoa texana (Bruner) (+ adult male) Elliott and Kurczewski 1985 Cordillacris crenulata (Bruner) (adult) Krombein 1979, Williams 1914 Cordillacris occipitalis (Thomas) Lavigne and Pfadt 1966, Pulawski 1988 Dissosteira pictipennis Bruner Pulawski 1988 Melanoplus bivittatus ? Elliott and Kurczewski 1985 Melanoplus lakinus Scudder Elliott and Kurczewski 1985 Melanoplus sanguinipes Elliott and Kurczewski 1985 Melanoplus spp. Elliott and Kurczewski 1985, Kurczewski 1987a Metator sp. Krombein 1967 Opeia sp., probably obscura (Thomas) Krombein 1967, Williams 1914 Orphulella sp. (adult) Elliott and Kurczewski 1985 Orphulella pelidna Pulawski 1988 Phlibostroma sp. (adult) Krombein 1967 Schistocerca sp. Pulawski 1988 Trachyrhachis kiowa (Thomas) (+ adult male) Lavigne and Pfadt 1966, Williams 1914 Trimerotropis pallidipennis (Burmeister) (+ adult) Elliott and Kurczewski 1985 Trimerotropis spp. Elliott and Kurczewski 1985, Pulawski 1988 74 Northeastern Naturalist Vol. 17, Monograph No. 6 Wasp species1 Prey species2,3 Source of information terminatus group clarconis Chloealtis sp., probably conspersa Evans 1970 Melanoplus foedus Elliott and Kurczewski 1985 Melanoplus sanguinipes Elliott and Kurczewski 1985 Melanoplus sp. Evans 1970 antillarum Chortophaga cubensis Saussure Genaro and Sánchez 1992 Parachloebata (=Orphulella) scudderi Bolivar Genaro and Sánchez 1992 alpestris Chorthippus curtipennis Evans 1970 Melanoplus sp., probably sanguinipes Pulawski 1988 Melanoplus spp. Pulawski 1988 Microtes sp., probably helferi (Strohecker) Pulawski 1988 Oedaleonotus sp. Elliott and Kurczewski 1985 linsleyi Clematodes larreae Scudder Kurczewski 2009 Eremiacris (=Paropomala) pallida (Bruner) Pulawski 1988 Psoloessa sp., possibly texana Scudder Kurczewski 2009 terminatus Ageneotettix deorum Kurczewski 2009 Arphia simplex Scudder Kurczewski 2009 Arphia sulphurea (Fabricius) Rau 1946 Arphia xanthoptera (Burmeister) Kurczewski 2009 Boopedon gracile Rehn Kurczewski 2009 Chloealtis conspersa Kurczewski 1966a, 2009 Chorthippus curtipennis Kurczewski 1966a, 2009 Chortophaga viridifasciata Ashmead 1894, Kurczewski 2009, Rau 1926 Dichromorpha viridis (Scudder) Kurczewski 2000b, 2009; Rau 1946; Rau and Rau 1918 Dissosteira carolina Kurczewski 1966a, 2000b, 2009 Encoptolophus sordidus Kurczewski 2009 Melanoplus bivittatus Kurczewski 1966a, 2009 Melanoplus differentialis Kurczewski 2009 Melanoplus femurrubrum Kurczewski 1966a, 2000b, 2009 Melanoplus keeleri (Thomas) Kurczewski 1966a, 2009 Melanoplus s. sanguinipes Kurczewski 2009 2010 F.E. Kurczewski 75 Wasp species1 Prey species2,3 Source of information Melanoplus spp. Kurczewski 2009, Rau 1946, Rau and Rau 1918, Strandtmann 1953 Mermiria sp. Kurczewski 2009 Orphulella pelidna Kurczewski 2000b Orphulella speciosa Kurczewski 2009 Pardalophora apiculata (Harris) Kurczewski 1966a, 2009 Phlibostroma quadrimaculatum (Thomas) Kurczewski 2009 Schistocerca americana(Drury) Kurczewski 2009 Scudderia sp., probably furcata Brunner von Wattenwyl Kurczewski 1966a, Pulawski 1988 (TETTIGONIIDAE) Syrbula admirabilis (Uhler) Kurczewski 2009, Rau and Rau 1918 similis Achurum carinatum (F. Walker) Krombein and Evans 1955, Kurczewski 2009 Amblytropidia mysteca (Saussure) Kurczewski 2009 Aptenopedes sphenarioides Scudder Kurczewski 2009 Aptenopedes sp. Krombein 1964 Arphia granulata Saussure Kurczewski 2009 Arphia simplex Kurczewski 2009 Arphia xanthoptera Kurczewski 2009 Arphia sp. Kurczewski 2000b Chloealtis conspersa Kurczewski 2009 Chorthippus curtipennis Kurczewski 2009 Dichromorpha viridis Kurczewski 2000b Dissosteira carolina Kurczewski 2000b, 2009 Melanoplus bivittatus Kurczewski 2009 Melanoplus devastator Pulawski 1988 Melanoplus differentialis Kurczewski 2009 Melanoplus femurrubrum Kurczewski 2000b, 2009 Melanoplus keeleri Kurczewski 2009 Melanoplus puer group Kurczewski 2009 Melanoplus s. sanguinipes Kurczewski 2009 Melanoplus sanguinipes vulturnus Gurney and Brooks Kurczewski 2009 Melanoplus seminole Hubbell Kurczewski 2009 76 Northeastern Naturalist Vol. 17, Monograph No. 6 Wasp species1 Prey species2,3 Source of information Melanoplus viridipes Kurczewski 2009 Melanoplus spp. Elliott and Kurczewski 1985, Krombein 1964, Kurczewski 2009 Mermiria sp. Elliott and Kurczewski 1985 Opeia obscura Elliott and Kurczewski 1985 Orphulella pelidna Kurczewski 2000b, 2009 Pseudopomala brachyptera Elliott and Kurczewski 1985 Psinidia fenestralis Kurczewski 2009 Schistocerca americana Kurczewski 2009 Schistocerca sp. Kurczewski 2000b, Pulawski 1988 Spharagemon marmorata picta Kurczewski 2009 Syrbula admirabilis Kurczewski 2009 apicalis Achurum carinatum Kurczewski 2009 Amblytropidia mysteca Kurczewski 2009 Aptenopedes sphenarioides Kurczewski 2009 Arphia granulata Kurczewski 2009 Arphia simplex Kurczewski 2009 Chortophaga australior Rehn and Hebard Kurczewski 2009 Chortophaga viridifasciata Kurczewski 2009 Dichromorpha viridis Kurczewski 2009 Melanoplus femurrubrum Kurczewski 2009 Melanoplus keeleri Kurczewski 2009 Melanoplus puer group Kurczewski 2009 Melanoplus s. sanguinipes Elliott and Kurczewski 1985 Melanoplus sanguinipes vulturnus Kurczewski 2009 Melanoplus seminole (+ adult males) Kurczewski 2009 Melanoplus sp., possibly puer (Scudder) Krombein 1964 Melanoplus spp. Kurczewski 2000b, 2009; Williams 1914 Orphulella pelidna Kurczewski 2000b, 2009 Oxya chinensis (Thunberg) Williams 1932 Psinidia fenestralis Kurczewski 2009 Spharagemon marmorata picta Kurczewski 2009 Trimerotropis maritima Kurczewski 2009 2010 F.E. Kurczewski 77 Wasp species1 Prey species2,3 Source of information brullii group mundus Conocephalus sp. (TETTIGONIIDAE) Elliott and Kurczewski 1985, Kurczewski 1979 Odontoxiphidium apterum (TETTIGONIIDAE) Kurczewski 1979 Orchelimum sp. (TETTIGONIIDAE) Kurczewski 1979 Oecanthus argentinus Saussure (GRYLLIDAE) Kurczewski 1979 belfragei Conocephalus sp. (TETTIGONIIDAE) Krombein 1967, Kurczewski 1979 menkei Eremopedes sp. (TETTIGONIIDAE) Pulawski 1988 obscuripennis group inconspicuus Chorisoneura cabimae Hebard (BLATTELLIDAE) (adults) Kurczewski et al., in press Chorisoneura diaphana Princis (BLATTELLIDAE) (adults) Kurczewski et al., in press Chorisoneura excelsa Rocha e Silva & Lopes (BLATTELLIDAE) Buys 2006, 2007 (+ adults) Chorisoneura fuscipennis Hebard (BLATTELLIDAE) (adults) Callan 1993; Kurczewski et al., in press Chorisoneura gemmicula Hebard (BLATTELLIDAE) (adults) Callan 1993; Kurczewski et al., in press Chorisoneura lopesi Rocha e Silva (BLATTELLIDAE) (+ adults) Buys 2007 Chorisoneura sp. near flavipennis Saussure & Zehntner Kurczewski et al., in press (BLATTELLIDAE) (adults) Chorisoneura sp. near gemmicula (BLATTELLIDAE) (adults) Kurczewski et al., in press Chorisoneura sp., probably parishi Rehn (BLATTELLIDAE) (+ adults) Kurczewski et al., in press Chorisoneura spp. (BLATTELLIDAE) (+ adults) Buys 2007; Callan 1954; Kurczewski et al., in press; S. Brady, 2009 pers. comm.; G. Beccaloni, 2010 pers. comm. Riatia fulgida (Saussure) (BLATTELLIDAE) (adult) Kurczewski et al., in press Riatia orientis (Hebard) (BLATTELLIDAE) (adults) Callan 1993 Riatia spp. (BLATTELLIDAE) (+ adult) Buys 2007, Pulawski 1974 Undetermined species (GRYLLIDAE) Buys 2007 iridipennis Anaplecta asema Hebard (BLATTELLIDAE)5 Rau 1933 Cariblatta tobagensis Hebard (BLATTELLIDAE) (adult)5 Vesey-FitzGerald 1956 Euthlastoblatta abortiva (Caudell) (BLATTELLIDAE) (adult) Pulawski 1988 Ischnoptera rufa debilis Hebard (BLATTELLIDAE) Pulawski 1988 78 Northeastern Naturalist Vol. 17, Monograph No. 6 Wasp species1 Prey species2,3 Source of information alayoi Cariblatta sp. (BLATTELLIDAE) (+ adult female) Genaro 2004 Symploce sp. near munda Gurney (BLATTELLIDAE) (+ adult male) Elliott et al. 1979 julliani group coquilletti Litaneutria minor (Scudder) (MANTIDAE) Alcock and Gamboa 1975 cockerellae Litaneutria minor (MANTIDAE) Elliott and Kurczewski 1985 1Wasp species listed phylogenetically following Pulawski (1988). 2Prey species arranged alphabetically; Acrididae unless indicated otherwise. 3Nymphs unless indicated otherwise. 4Prey record pertains to T. williamsi or T. tarsatus. 5Prey record pertains to T. iridipennis or T. inconspicuus.