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I. Geographic Distribution and Paleobiogeography of Tachysphex pechumani (Hymenoptera: Crabronidae)
Frank E. Kurczewski

Northeastern Naturalist, Volume 15, Monograph 2 (2008): 1–32

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2008 NORTHEASTERN NATURALIST 15(Monograph 2):1–32 I. Geographic Distribution and Paleobiogeography of Tachysphex pechumani (Hymenoptera: Crabronidae) Frank E. Kurczewski* Abstract - Tachysphex pechumani, the antenna-waving wasp, has a disjunct geographic distribution in the Lower Peninsula of Michigan, Indiana Dunes, Oak Openings of Ohio, southern Ontario, and New Jersey pinelands. This distribution is tied to excessively/well-drained sandy soils, oak/pine-dominant barrens, savanna and woodland, climate moderation related to proximity to large bodies of water, habitat fragmentation from natural causes and human disturbance, and 20th-century land preservation. The 550-km gap between subpopulations in southwestern Ontario and southern New Jersey may be the consequence of Early Holocene pine-dominant barrens, savanna, and woodland being supplanted by deciduous and deciduousconiferous forest on loamy sand in the northeastern United States during the past 6000 years. A plausible scenario as to where T. pechumani lived during the past 18,000 years and what dispersal route(s) enabled it to attain its current geographic distribution is presented. Introduction Tachysphex, a very large genus of digger wasps, has 83 species in North America and the Caribbean Region and 444 species worldwide (Bohart and Menke 1976; Pulawski 1988, 2008). Many of the species, especially in the pompiliformis species group, are morphologically similar to one another and therefore difficult to define taxonomically (Pulawski 1988). The majority of nearctic Tachysphex inhabit areas of high temperature and low humidity in the southwestern United States and northwestern Mexico (Pulawski 1988). Species that occur in the Austral Faunal Zone are mostly bi- or multivoltine, whereas species that inhabit the Boreal Faunal Zone are univoltine (Pulawski 1988). These small, largely cursorial wasps are exceedingly rapid in their movements (Williams 1914), as inferred from the genus name (tachy = swift, sphex = wasp). The nearctic species nest in sandy, gravelly, or, rarely, loamy soils and provision shallow cells with orthopteroid insects, mainly nymphal grasshoppers (Acrididae) (Krombein 1979). Tachysphex pechumani Krombein, an apparent relict of the Pleistocene (Kurczewski and Elliott 1978), is an unusual acridid-hunting congener based on adult morphology, ecology, nesting behavior, and geographic and seasonal distribution (Kurczewski 1987). The species has a vicissitudinous taxonomic history, having been first described incorrectly as a subspecies of T. tarsatus from a single female (Krombein 1938) and then subtly elevated to species rank after examination of additional females (Bohart 1951). The *PO Box 15251, Syracuse, NY13215; 2 Northeastern Naturalist Vol. 15, Monograph 2 female is distinguished from other nearctic congeners in the pompiliformis species group by the wide space between the compound eyes, brassy-golden face, long orange apical antennal segments, and two or three orange-red basal abdominal segments on a predominantly black body (Kurczewski et al. 1970, Pulawski 1988). The all-black male was associated with the female and described nearly 20 years later (Kurczewski et al. 1970). Olive-green eyes are a distinguishing field characteristic in both sexes. Both the modern and historic geographic distributions of T. pechumani pose a variety of interesting yet perplexing paleobiogeographic questions. Where did T. pechumani reside during full and late glaciation? What dispersal route(s) did this species take in order to attain its present-day range? How extensive was its distribution during the peak warmth and aridity of the Mid-Holocene, when the preferred habitat may have been more expansive and contiguous? How did the current disjunct distribution arise, especially the wide gap between southwestern Ontario and southern New Jersey? What factors are responsible for fragmentation into populations and subpopulations? How did the pre-settlement geographic distribution differ from that of today? It is impossible to provide indisputable answers to these questions because fossil wasps remain undiscovered. Furthermore, phylogenetic study of living populations using molecular markers has not been attempted. However, indirect evidence gleaned from glacial recession chronology, paleo-water levels and paleoshorelines, climate reconstruction, palynology, fossil insects, post-glacial dispersal routes, and early land surveys can be useful in tracing the geographic distribution of the wasp’s habitat through the past 18,000 years. Tachysphex pechumani likely moved northward from its probable glacial refugium in the southeastern United States in similar fashion following its habitat. In drawing such inferences, several assumptions were made: (1) insects are intricately adapted to their environment; (2) their present-day climatic, edaphic, and ecological requirements are essentially the same as in times past (Coope 1967, Morgan 1987); (3) flying insects are often able to move from one suitable habitat to another if not prevented from doing so by insurmountable barriers; (4) such movement often results in temporary colonization of such habitats; (5) extirpation of source populations commonly occurs with changing environmental conditions; (6) geographic distributions fluctuated with changing glacial and interglacial conditions; and (7) geographic distributions fragmented as a result of natural causes and human disturbance. The goals of this study were: (1) delineate the modern (1991–2006) and historic (1902–1990) geographic distributions of T. pechumani; (2) define the climatic, ecological, and edaphic parameters that account for its highly fragmented range; and (3) determine where and how this species survived Pleistocene and Holocene climatic conditions in order to attain its current geographic distribution. 2008 F.E. Kurczewski 3 Methods Historic collection records were obtained from specimens in The American Museum of Natural History, National Museum of Natural History, University of California-Davis, Cornell University, Michigan State University, The University of Michigan, Commonwealth of Pennsylvania Department of Agriculture (Harrisburg), University of Guelph, and York University (Toronto). These records date from 1902 (Iona, Gloucester County, NJ) to 1990 (University of Michigan Biological Station, Cheboygan County) (Table 1). There were 167 specimens of T. pechumani known prior to the present study (Pulawski 1988; Mark O’Brien, University of Michigan, Ann Arbor, MI, 1991 pers. comm.). Table 1. Historic (1902–1990) geographic distribution of Tachysphex pechumani in Lower Michigan and New Jersey. Abbreviations: SF = State Forest, SGA = State Game Area, and SRA = State Recreation Area. State/County Locality (Collection year) Lower Michigan Allegan Allegan SGA (1975–1984) Cheboygan University of Michigan Biological Station (1981–1990) Crawford (1951–1953) Gladwin (1951) Gratiot (1947) Iosco (No year) Kalkaska (1951–1966) Midland (1941–1947) Missaukee (1945) Montcalm (No year) Montmorency (1966) Oakland Pontiac SRA (1972), Farmington (No year) Oscoda Luzerne (1966) Saginaw (1942–1948) Washtenaw Pinckney SRA (1981–1983) Wexford (1972) New Jersey Atlantic Elwood (1968) Weymouth (1926–1968) Burlington Browns Mills (1906–1921) Chatsworth (1923) Lebanon SF (No year) New Gretna (1968) Ong (1968) Rancocas Park (No year) Vincentown (1968) Camden Atco (1912?) Clementon (1912) Cape May 3 miles S Seaville (1972) Cumberland Manumuskin (1903–1923) Vineland (Maurice River) (No year) Gloucester Iona (1902) Ocean Lakehurst (Wrangle Brook Road) (No year) Manahawkin (1935) 4 Northeastern Naturalist Vol. 15, Monograph 2 Modern distribution records are based mainly on 41 collecting trips I made through oak/pine-dominant barrens, savanna, and woodland in northwestern Indiana, northwestern Ohio, Lower Peninsula of Michigan, southwestern Table 2. Modern (1991–2006) geographic distribution of Tachysphex pechumani in Indiana, Lower Michigan, Ohio, Ontario and New Jersey. Abbreviations: F&WMA = Fish and Wildlife Management Area, NF = National Forest, SF = State Forest, SGA = State Game Area, SRA = State Recreation Area, and WMA = Wildlife Management Area. State/County Locality (Section) Indiana Porter Pine Township Lower Michigan Alcona Huron NF (T26N, R7E, S17) Allegan Allegan SGA (T2N, R14W, S6,7; T3N, R15W, S36) Antrim Jordan River SF (T29N, R5W, S22-24) Arenac Alger Cemetery (T20N, R3E, S21) Barry Barry SGA (T3N, R10W, S13) Cass Wayne Township (T5S, R15W, S23) Cheboygan Hardwood SF (T35N, R2W, S22,27,28,34) University of Michigan Biological Field Station (T37N, R3W, S28,32,34) Clare Chippewa River SF (T20N, R4W, S3; T20N, R5W, S23) Crawford Huron NF (T25N, R1W, S11) Au Sable SF (T26N, R3W, S2,8,17; T27N, R4W, S25; T28N, R2W, S1,32; T28N, R3W, S31) Emmet Conway (T35N, R5W, S24) Hardwood SF (T37N, R4W, S25,33) Gladwin Tittabawassee River SF (T18N, R1E, S1,10,12; T18N, R2E, S1,7-10; T19N, R1E, S12) Iosco Huron NF (T23N, R6E, S22,34; T23N, R7E, S31,32; T24N, R6E, S3) Kalamazoo Oshtemo Township (T2S, R12W, S5,7,8) Kalkaska Kalkaska SF (T26N, R8W, S8; T27N, R5W, S27; T27N, R6W, S13; T27N, R7W, S13,24; T28N, R7W, S28) Lake Luther Valley Cemetery (T19N, R11W, S17) Livingston Island Lake SRA (T1N, R6E, S16) Manistee Manistee NF (T21N, R16W, S28) Mason Manistee NF (T18N, R15W, S21; T20N, R17W, S2). Midland Tittabawassee River SF (T16N, R1W, S5,8-10) Montmorency Thunder Bay River SF (T29N, R1E, S31; T32N, R2E, S4,9) Muskegon Manistee NF (T12N, R15W, S5,6,19-21,27; T12N, R16W, S19- 29,33) Newaygo Manistee NF (T15N, R13W, S24; T16N, R12W, S19) Oceana Pines Point SRA (T13N, R15W, S9) Manistee NF (T13N, R15W, S18,19,31,32) Ogemaw Ogemaw SF (T23N, R1E, S4,5,8,9; T23N, R2E, S19) Oscoda Huron NF (T26N, R1E, S11,25,26,30,33,35) Otsego Otsego SF (T29N, R1W, S16,21,32,33; T29N, R3W, S17; T29N, R4W, S17-20,22,23) Presque Isle Black Lake SF (T33N, R2E, S21) Roscommon Houghton Lake SF (T21N, R4W, S4, 10,15,22,28) Washtenaw Pinckney SRA (T1S, R4E, S6) Wexford Fife Lake SF (T21N, R9W, S1,2) 2008 F.E. Kurczewski 5 Ontario and southern New Jersey (Table 2). I traveled a total of 37,500 km by automobile mostly through national, state, and local refuges. State/province soil association maps, United States Department of Agriculture county soil surveys, pre-settlement vegetation maps, and aerial photographs were used to locate undeveloped, excessively/well-drained sandy soils with oak/pine-dominant vegetation. Locations that comprised ancestral oak savanna and pine barrens or had a history of periodic fires or prior wasp collections were targeted. Wasps from 156 sites were either hand-netted, frozen, pinned, identified, and counted, or hand-netted, put in vials on ice in a cooler, identified, counted, warmed to ambient temperature, and released at the collection site. Records Table 2, continued. State/County Locality (Section) Ohio Lucas Spencer Township (Kitty Todd Preserve [3 sites], Melke Road Savanna, Moseley Barrens) Springfield Township (Mescher Road, Miller Fireworks) Swanton Township (Oak Openings Preserve Metropark [7 sites], Sager Road Ponds*) Ontario Grey Hepworth Sand Dunes (Buck 2004) Halton Milton (Woodland Trails Camp)(Buck 2004) Lambton Bosanquet Township (Pinery Provincial Park, Karner Blue Sanctuary, Watson Property, Ausable-Bayfield Property, Thedford Conservation Area) Norfolk St. Williams Crown Forest (Manestar Tract, Turkey Point Tract), South Walsingham Sand Ridges, Turkey Point Provincial Park*, Simcoe Junction* Simcoe Canadian Forces Base Borden (3 sites, 2 sites*) Waterloo Erbsville (Buck 2004), Kitchener (Huron Natural Park) Wellington Guelph (Buck 2004) York Newmarket (Koffler Scientific Reserve) (Buck 2004) New Jersey Atlantic Wharton SF (Dutchman)* Hammonton Creek WMA Great Egg Harbor WMA Galloway Township Burlington Lebanon SF (Woodland Township, 2 sites) Woodland Township (Hedger House) Chatsworth Cemetery Chatsworth Lake Chatsworth Woods Bass River SF (Harrisville, Leektown) Cape May Belleplain SF (Powerline)* Ocean Lebanon SF (Manchester Township) Manchester F&WMA Greenwood Forest F&WMA (Howardsville, Webbs Mills) Barnegat Township (Cedar Bridge Heights) *Aggregations disappeared after 1996. 6 Northeastern Naturalist Vol. 15, Monograph 2 from Lucas County, OH (Robert Jacksy, Oak Openings Preserve Metropark, Toledo, OH, 2006 pers. comm.) and southern Ontario (Buck 2004; Matthias Buck, University of Guelph, Guelph, ON, Canada, 2006 pers. comm.) supplemented the modern records. Attempts to find T. pechumani in 1991–2006 at 552 other locations in Wisconsin (10 sites), Upper Peninsula of Michigan (16), Lower Peninsula of Michigan (112), Illinois (2), Indiana (7), Ohio (24), Pennsylvania (2), Ontario (59; Kurczewski 2000a), Quebec (2), upstate New York (47; Kurczewski 1998b), New England (3), Long Island (7), southern New Jersey (98), Delaware-Maryland (3), and the Carolinas (160; Kurczewski 2000b) were unsuccessful. Lower Michigan soil series were identified at the United States Department of Agriculture Soil Conservation Service, East Lansing, MI (Table 3). Soil series and acreage of sandy soil from non-Lower Michigan sites were extrapolated from United States Department of Agriculture Soil Conservation Service and Ontario County Soil Surveys (Table 3). Properties of soil samples from eight nesting sites in Lower Michigan and two nesting sites in New Jersey were analyzed at the State University of New York College of Environmental Science and Forestry, Syracuse, NY. Total acreage of sand, loamy sand, and loamy fine sand and percent sandy soils in Camden County, NJ were calculated by the Camden County Soil Conservation District. Common native plant species were identified from six T. pechumani nesting sites in the central Great Lakes Region (Table 4). The “Biological inventory and evaluation of the Manestar Tract, St. Williams Forest, Haldimand-Norfolk Regional Municipality and the Lambton Wildlife Inc., Karner Blue Sanctuary, Port Franks, Lambton County, Ontario” (Sutherland and Bakowsky 1995) was used as an aid to identify plant species from two of the locations. United States (1971–2000) and Canadian (1961–1990) climate normals provided annual mean temperature and annual mean amount of precipitation for localities near collection sites (Canadian Climate Program 1993, National Climatic Data Center 2005). The most recent Canadian climate normals (1971– 2000) were not used because many weather stations near the sites were obsolete and their records were unavailable or irregular. January and July mean, January extreme minimal, and July mean daily maximal temperatures were derived from modern and historic (1896/1899–1938) climate records (Canadian Climate Program 1993, United States Department of Agriculture 1941). Results Geographic distribution Tachysphex pechumani inhabited mainly very coarse and coarse sandy soils in northwestern Indiana (1 site), northwestern Ohio (14), Lower Michigan (120), southern Ontario (17), and southern New Jersey (16) (Fig. 1, Table 2). Elevation at historic and modern nesting sites ranged from 2 (New 2008 F.E. Kurczewski 7 Table 3. Soil series in which Tachysphex pechumani nested in Indiana, Lower Michigan, Ohio, Ontario, and New Jersey. State/County Soil series Indiana Porter Oakville fine sand Lower Michigan Alcona Rubicon sand Allegan Oakville fine sand Antrim Kalkaska, Rubicon sands Arenac Rubicon sand Barry Coloma loamy sand Cass Ormas loamy sand Cheboygan Grayling, Rubicon sands Clare Grayling, Rubicon sands Crawford Grayling, Rubicon sands; Kalkaska loamy sand Emmet Kalkaska, Rubicon sands Gladwin Croswell, Rubicon sands Iosco Grayling, Rubicon sands Kalamazoo Coloma loamy sand Kalkaska Kalkaska, Rubicon sands Lake Graycalm sand Livingston Oshtemo loamy sand Manistee Grattan sand Mason Grattan sand Midland Covert sand Montmorency Grayling, Rubicon sands Muskegon Grattan sand Newaygo Coloma, Grattan sands Oceana Grattan sand Ogemaw Graycalm, Grayling sands Oscoda Grayling, Rubicon sands Otsego Grayling, Kalkaska, Rubicon sands Presque Isle Cheboygan loamy sand Roscommon Grayling, Rubicon sands Washtenaw Boyer loamy sand Wexford Rubicon sand Ohio Lucas Oakville, Ottokee fine sands Ontario Grey Tioga sand Halton Grimsby loamy sand Norfolk Plainfield sand Lambton Plainfield sand Simcoe Tioga sand Waterloo Camilla sand Wellington Sand overlying Guelph loam York Pontypool sand New Jersey Atlantic Evesboro, Lakehurst sands; Downer loamy sand Burlington Evesboro, Lakehurst sands Cape May Hammonton loamy sand Ocean Lakehurst, Lakewood, Woodmansie sands; Downer loamy sand 8 Northeastern Naturalist Vol. 15, Monograph 2 Table 4. Common native vascular plant species at selected Tachysphex pechumani locations in the central Great Lakes Region. Abbreviations for locations: AGA = Allegan State Game Area, MI; UMB = University of Michigan Biological Station, MI; OOP = Oak Openings Preserve Metropark, OH; SWF = St. Williams Crown Forest, ON; KBS = Karner Blue Sanctuary, ON; CFB = Canadian Forces Base Borden, ON. Species* AGA UMB OOP SWF KBS CFB Pteridium aquilinum (L.) Kuhn X X X X X X Pinus banksiana Lambert X X X X X Pinus resinosa Aiton X X X X X X Pinus strobus L. X X X X X X Sassafras albidum (Nutt.) Nees X X X X Comptonia peregrina (L.) J.M. Coulter X X X X X Quercus alba L. X X X X X X Quercus rubra L. X X X X X X Quercus velutina Lamarck X X X X Helianthemum divaricatus L. X X X X X X Populus tremuloides Michx. X X X X X X Arabis lyrata L. X X X X X X Fragaria virginiana Duchesne X X X X X X Potentilla simplex Michx. X X X X X X Rubus flagellaris Willd. X X X X X X Prunus serotina Ehrh. X X X X X X Lupinus perennis L. X X X X X Lespedeza capitata Michx. X X X X X Comandra umbellata (L.) Nutt. X X X X X X Euphorbia corollata L. X X X X X Ceanothus americana L. X X X X X X Polygala polygama Walter X X X X X X Asclepias tuberosa L. X X X X X X Lithospermum canescens (Michx.) Lehm. X X X X X Aster laevis L. X X X X X X Erigeron philadelphicus L. X X X X X X Carex pensylvanica Lamarck X X X X X X Bromus kalmii A. Gray X X X X X X Danthonia spicata (L.) F. Beauv. X X X X X X Panicum virgatum L. X X X X X X Andropogon gerardii Vitman X X X X X X Schizachyrium scoparium (Michx.) Nash X X X X X X *Species listed according to Gleason and Cronquist (1991). Gretna, Burlington County, NJ) to 398 m (Cadillac, Wexford County, MI). Tachysphex pechumani occurred mostly in areas currently or previously occupied by oak/pine-dominant barrens, savanna, and woodland. The species was located at the edges of man-made sand and gravel pits, but not in large tracts of barren sand such as wind-blown dunes, unstable blowouts, and beaches near shorelines and coasts. Although appropriate prey species occur 2008 F.E. Kurczewski 9 in intervening areas, soil and habitat limitations have produced a wide gap (550 km) between the southernmost Ontario and northernmost New Jersey subpopulations (Figs. 1, 2). Of the 168 T. pechumani sites, 159 (94.6%) are located in national and state forests, state parks, game and recreation areas, regional preserves and sanctuaries, provincial forests and parks, military reservations, boy scout camps, and cemeteries. The Huron and Manistee National Forests in Lower Michigan and the New Jersey Pinelands National Reserve have more aggregations of T. pechumani than the other areas as they contain the largest acreage of contiguous, excessively/well-drained sandy soils and pine/oakdominant barrens, savanna, and woodland. I did not find T. pechumani in Oakland, Saginaw, Gratiot, and Montcalm counties, MI, although the species was collected there in the 1940s and 1972 (Oakland County; Table 1). All four counties have moderate amounts of land set aside in state game and recreation areas, and three of the four counties have a slow rate of commercial and residential development. Tachysphex pechumani probably inhabits sandy sections of these counties bringing the total number of counties in Lower Michigan occupied by this species to 34. I did not locate T. pechumani in Camden, Cumberland, and Gloucester counties, NJ, where it occurred in the very early 1900s (Table 1) or, in 2006, Figure 1. Modern (black circles) and historic (open circles) collection locations for Tachysphex pechumani superimposed on well-drained sandy soils (gray areas) in Lower Michigan, Indiana, Ohio, Ontario, and New Jersey. (Sandy soils from Acton and Harkes 1988, Markley 1979, Ohio STATSGO Database 2006, US Department of Agriculture Soil Conservation Service 1981, 1982). 10 Northeastern Naturalist Vol. 15, Monograph 2 Cape May County, NJ, where it was found in the 1990s (Table 2). Suitable habitat in these counties has largely been converted to agricultural, commercial, industrial, recreational, and residential uses. The likelihood of T. pechumani occurring in accessible areas there is slight. Soils Tachysphex pechumani aggregations occupied level, mildly compacted, moderately well- to excessively-drained acidic sand and loamy sand of 26 different soil types (Table 3). Aggregation is defined herein as a group of conspecific individuals occurring together at a site. Soil samples from eight nesting sites in Lower Michigan ranged from 85.4 (Au Sable SF, Crawford County) to 94.4 (University of Michigan Biological Station) percent sand content. Except for the Pinckney SRA, Washtenaw County (7.73), the samples varied in pH from 5.29 (Allegan SGA, Allegan County) to 6.23 (Houghton Lake SF, Roscommon County). Percent organic matter, mainly charcoal, was as high as 3.73% (Roscommon County). Nesting site soils in Newaygo, Roscommon, and Crawford counties had burned repeatedly based on the relatively large amount of charcoal. The very dark or dark gray to gray soils had hue and chroma values as high as 10YR 5/1–5/2. Study site soils in southern Lower Michigan contained less charcoal and were lighter in color than those in central and northern Lower Michigan: very dark grayish brown to dark brown, Figure 2. Modern (black circles) and historic (open circles) collection locations for Tachysphex pechumani superimposed on pre-settlement oak/pine-dominant vegetation (gray areas) in Lower Michigan, Indiana, Ohio, Ontario, and New Jersey. (Presettlement vegetation from W.D. Bakowsky, Ontario Natural Heritage Information Centre, Peterborough, ON, Canada, 2005 pers. comm.; Gordon 1966; Lindsey et al. 1965; Markley 1979; Stearns and Guntenspergen 1987). 2008 F.E. Kurczewski 11 10YR 3/2–3/3 (Allegan SGA); and light brownish gray to grayish brown, 2.5YR 5/2–6/2 (Pinckney SRA). Two nesting-site soil samples from the Bass River SF, Burlington County, NJ had 95% sand content, very high C/N values due to inclusive charcoal, pH values as low as 4.6–5.0, and were gray (10YR 6/1) to dark grayish brown (10YR 4/2) in color. Tachysphex pechumani had more and larger nesting aggregations in sand (>85% sand, <15% silt and clay) than in loamy sand (>70 to <85% sand, >15 to <30% silt and clay). Of the 168 sites, 138 (82.2%) comprised very coarse (1.0–2.0 mm in diameter), coarse (0.5–1.0 mm), or medium (0.25–0.50 mm) sand; 14 (8.3%) were characterized by fine (0.10–0.25 mm) or very fine (0.05–0.10 mm) sand; and 16 (9.5%) were loamy sand. Available water capacity in the A (uppermost) horizon, expressed as inches of water per inch of soil, ranged from 0.04–0.09 (excessively-drained sand) to 0.10–0.16 (moderately well-drained sand, well-drained loamy sand). Pre-settlement vegetation The modern and historic geographic distributions of T. pechumani are superimposed mainly on pre-settlement oak/pine-dominant barrens, savanna, and woodland (Fig. 2). Most (147/168 or 87.5%) sites were within the ranges of these ecological communities, including all (16) sites in southern New Jersey. Twenty-one (12.5%) locations were outside of the pre-settlement ranges of oak/pine-dominant vegetation and involved human disturbance of the original habitat. In south-central Ontario, where most of its typical ecological communities are absent, this species was able to colonize degraded anthropogenic habitat (Matthias Buck, 2005–2006 pers. comm.). Ecological communities In northern and central Lower Michigan and Simcoe County, ON, T. pechumani inhabited pine-heath barrens, pine-oak-heath savanna and pine- and pine-oak-dominant open woodland. Tachysphex pechumani occupied oak savanna and oak-dominant open woodland in the Indiana Dunes, Ohio Oak Openings, extreme southern Lower Michigan and Lambton and Norfolk counties, ON. This species nested in pitch pine-heath barrens, pitch pine-oak-heath savanna and pine-oak- and oakpine- dominant open woodland in the New Jersey pinelands. Twenty-one sites in Lower Michigan and southern Ontario comprised edges of deciduous- coniferous woodland, openings in abandoned agricultural fields, and recently bulldozed areas. The ecological communities in which T. pechumani occurred in the central Great Lakes Region contained native plant species characteristic of sandy places, prairies, and dry open woods (Table 4). Many of these sites held several species of Ericaceae: Gaultheria procumbens L. (wintergreen), Arctostaphylos uva-ursi (L.) Sprengel (bearberry), Vaccinium angustifolium Aiton (lowbush blueberry), V. myrtilloides Michx. (velvetleaf blueberry), V. pallidum Aiton (hillside blueberry), and Gaylussacia baccata (Wangenh.) K. Koch (black huckleberry). Southern New Jersey T. pechumani locations had unique plant 12 Northeastern Naturalist Vol. 15, Monograph 2 assemblages that included Pinus rigida Miller (pitch pine), P. echinata Miller (shortleaf pine), Quercus ilicifolia Wangenh. (scrub oak), Q. marilandica Muenchh. (blackjack oak), Q. alba L. (white oak), Q. velutina Lamarck (black oak), Q. stellata Wangenh. (post oak), Q. coccinea Muenchh. (scarlet oak), Q. prinus L. (chestnut oak), Hudsonia ericoides L. (golden heather), Kalmia angustifolia L. (sheep laurel), K. latifolia L. (mountain laurel), Gaultheria procumbens, Arctostaphylos uva-ursi, Vaccinium angustifolium, V. pallidum, Gaylussacia baccata, and G. frondosa (L.) T. & G. (dangleberry). Climate Annual mean temperature near historic and modern T. pechumani nesting sites ranged from 5.4 ºC (University of Michigan Biological Station) to 12.4 ºC (Belleplain SF, Cape May County, NJ). January mean temperature, which may influence the northern range extent of T. pechumani, varied from -8.7 ºC (Otsego SF, Otsego County, MI) to +1.1 °C (Wharton SF, Atlantic County, NJ). Extreme minimal temperature, which may also affect the northern range extent of this species, ranged from -43.9 ºC (Huron NF, Oscoda County, MI) to -25.0 ºC (Wharton SF, NJ). July mean temperature varied from 19.0 ºC (Otsego SF, MI) to 24.4 °C (Hammonton Creek WMA, Atlantic County, NJ). July mean daily maximal temperature was 31.1 °C (Hammonton Creek WMA, NJ). Annual mean amount of precipitation ranged from 679 mm (Huron NF, Oscoda County, MI) to 1197 mm (Lebanon SF, Burlington County, NJ). Paleobiogeography Much of the Upper Midwest and Northeast was glaciated during the Full- Wisconsinan (Dyke and Prest 1987). Although southern New Jersey was not covered by ice, the periglacial effects nevertheless made this area cold, windy, and dry (July mean temperature = 14 °C, January mean temperature = -14 °C, annual average amount of precipitation = 600 mm; Prentice et al. 1991). Such conditions would have been unsuitable for T. pechumani habitation based on its present-day climatic requirements. During full glaciation, sea level off the Atlantic Coast fell to 121 m below the present level as water became locked in the continental glaciers (Fairbanks 1989). This exposed a broad, gently sloped continental shelf consisting mainly of medium and coarse sands (Duane and Stubblefield 1988, Knebel 1981). During the height of the Wisconsinan glaciation, this shelf was more than 100 km wide throughout much of its length (Edwards and Emery 1977). The region had the most equable climate and deepest, most uniform sandy sediments in the southeastern United States and was likely colonized early by plants and animals (Dincauze and Mulholland 1977). Tachysphex pechumani probably inhabited this expanded Atlantic Coastal Plain during glacial and quasi-glacial periods. Upon climate amelioration and glacial recession, T. pechumani likely dispersed northward through the Atlantic Coastal Plain and what is now northern New Jersey and southeastern New York, then westward through 2008 F.E. Kurczewski 13 the Mohawk Valley or northward through the Champlain Valley to the Erie- Ontario Lake Plain. From this lake plain, the species could have readily moved westward into what is now Ontario, Ohio, Lower Michigan, Indiana, and Illinois. Such a dispersal route would have covered a distance of more than 2000 km. A relict population remained behind in southern New Jersey, eventually to be separated from the central Great Lakes geographic variant by Mid- to Late Holocene deciduous and deciduous-coniferous forest expansion (Kurczewski 1998a). Unfortunately, there is no fossil record or DNA markers for the two disjuncts to substantiate this pathway. However, several arguments support this post-glacial dispersal route: 1) Tachysphex pechumani occurs only in the central Great Lakes Region and southern New Jersey. 2) The species inhabits excessively and well-drained, coarse sandy soils at relatively low to low elevation near large bodies of water. 3) The temperate climate of the Southeast, Mid-Atlantic, and central Great Lakes regions coincided time-transgressively with the climatic requirements of this species. 4) Favorable habitat in the form of pine barrens, savanna, and woodland existed in the Atlantic Coastal Plain, Hudson-Mohawk/Champlain Valleys, and Erie-Ontario Lake Plain from the Early to Mid-Holocene 9000–6000 years ago. Maximal northern pine dispersal accompanied a warmer, drier climate from south to north along the Atlantic Seaboard and in the Northeast (Prentice et al. 1991, R. Webb et al. 1993, T. Webb et al. 1994). 5) The enlarged Atlantic Coastal Plain extended eastward for 100–150 km throughout much of its length during the Late Pleistocene-Early Holocene. Level to undulating relief, insignificant amount of erosion, estuarine development and flooding, and lack of major watercourse obstacles posed no insurmountable physiographic barriers to dispersal. 6) Sandy soils were readily available for colonization in the Middle Atlantic Region and Northeast during the Early to Mid-Holocene due to drier climate and lower water levels. Prior existing coarse sandy deposits were augmented by oceanic shoreline shifts, fluvial-sedimentological activity, and morainal and proglacial lake effects of the Wisconsinan glaciation. 7) Eastern grasses, sedges, and forbs furnished a plentiful food supply for T. pechumani prey in the Mid-Atlantic Region and Northeast as the grasshoppers followed the plants northward from ancestral refugia in the Southeast (Hubbell 1960, Wells 1970). During full glaciation the expanded southern Atlantic Coastal Plain would have provided favorable habitat in the form of low relief, well-drained sandy soils, moderate climate, northern pine/oak-dominated barrens, savanna, and woodland, abundant graminoid and forb understory, high incidence of lightning strikes and periodic natural fires (Grimm et al. 1993; Jacobson et al. 1987; Watts 1979, 1980, 1983; Watts and Hansen 1994; Watts and Stuiver 1980) (Fig. 3). South Carolina to northern Florida July mean temperature ranged from 19 to 22 °C, January mean temperature varied from -4 to 4 °C, and annual mean amount of precipitation was 700–850 mm, as inferred from a study of pollen types (Prentice et al. 1991). 14 Northeastern Naturalist Vol. 15, Monograph 2 Other full and late glacial areas and post-glacial invasion routes fall far short of maintaining the climatic, edaphic, and ecological requirements Figure 3. Plausible Tachysphex pechumani dispersal route(s) (arrows) correlated time-transgressively with glacial recession chronology and maximal northern pine migration. Age estimates are in thousands of years. (Quaternary surficial sand/fine gravel [gray areas] and land boundaries from Barnett et al. 1991, Curray 1965, Dincauze and Mulholland 1977, Farrand and Bell 1982, Fullerton et al. 2003, Kurczewski 1998a, Lewis and Anderson 1989). 2008 F.E. Kurczewski 15 necessary to support T. pechumani. Much of the Gulf Coastal Plain was inappropriate for full and late glacial habitation because of a preponderance of clayey, silty, and loamy soils and suboptimal paleoclimate governed by cool ancestral Mississippi River meltwater (Overpeck et al. 1989, Watts et al. 1992). The post-glacial Mississippi River Valley was unsuitable for habitation by T. pechumani because of excessive meltwater runoff, cold-air drainage, and advection fogs (Delcourt and Delcourt 1977, Porter and Guccione 1994, Teller 1990). Governed by a cooler climate, this region was dominated by spruce and deciduous hardwood forest in the south and spruce, fir, and tamarack woodlands to the north (Delcourt et al. 1980, Jackson and Givins 1994, Royall et al. 1991). Predominantly clayey-silty alluvial soils (Saucier 1974), thick aeolian loess (Ruhe 1983), and a paucity of pine barrens and other fire dependent natural communities probably would have discouraged T. pechumani habitation of the Mississippi River Valley. Fragmented sandy soils occupied the High Plains during the Late Pleistocene and Holocene (Ahlbrandt et al. 1983, Ruhe 1983), but they were discounted as a post-glacial dispersal pathway because of unsuitable climate, inappropriate vegetation, and reduced fire frequency (Fredlund 1995, Fredlund and Jaumann 1987, Holliday 1986, Wells and Stewart 1987). Much of the Central Plains and Midwest was mantled with unfavorable, fine loessal deposits of various thicknesses (Ruhe 1983, Wright et al. 1985). Considerable aeolian activity responsible for extensive dune development in the upper Great Plains during the Late-Wisconsinan and Early to Mid-Holocene (Ahlbrandt et al. 1983) would have been an additional obstacle for this species. Tachysphex pechumani does not inhabit active sand dunes because its shallow cells are readily exposed by wind erosion. At first, high and then very low soil moisture, lake, and water levels characterized much of the Great Plains and northern Midwest from the Late-Wisconsinan through the Mid-Holocene (Digerfeldt et al. 1992, Harrison 1989, Webb et al. 1994, Winkler et al. 1986). A boreal spruce forest covered much of the Great Plains during the cool Late-Wisconsinan. This forest was broken into a mosaic of spruce and spruce-deciduous woodlands with aspen savanna to the west and south (Fredlund and Jaumann 1987). Climatic warming at the end of the Pleistocene resulted in unsuitable steppe vegetation in the west-central and cool-temperate, mesic deciduous forest in the eastern Great Plains (Ahlbrandt et al. 1983, Fredlund and Jaumann 1987). In the Early Holocene, increased flow of dry Pacific air initiated the development of prairie from west to east throughout much of the westcentral Great Plains (Webb et al. 1983). In the eastern Great Plains, Pacific air masses were blocked by a strong monsoonal flow of maritime warm air from the Gulf of Mexico (Baker et al. 1992). Because of the relative proximity of the wasting Laurentide ice sheet, cold fronts may have provided the mechanism for increased precipitation from this moist southern airflow (Wright 1992). Eastern Iowa, southern Wisconsin, and northern Illinois 16 Northeastern Naturalist Vol. 15, Monograph 2 were kept warm and moist while areas of the Great Plains to the north and west were warm and very dry (Baker et al. 1996). As the glacier dissipated, the blocking effect of the Gulf of Mexico monsoonal airflow gradually relaxed, and the prairie governed by dry Pacific air edged eastward displacing savanna and deciduous forest about 5500 yr BP (Baker et al. 1992, Wright 1992). If a Great Plains-northern Midwest dispersal route was used by T. pechumani, one might expect to find remnant populations persisting in fire-prone sections of Minnesota, Wisconsin, and the Upper Peninsula of Michigan. There is no historic or present-day record of this species ever inhabiting such areas. The southern Atlantic Coastal Plain represents the primary geographic range of T. pechumani since glacial or quasi-glacial periods account for about 85,000 years of each Late-Quaternary, l00,000 year-long Milankovitch cycle (Schoonmaker and Foster 1991). Tachysphex pechumani probably dispersed north- and southward through this region in synchrony with warming and cooling trends, rising and lowering of sea level, and narrowing and widening of the continental shelf as it oscillated in size with the waning and waxing of the glaciers. Tachysphex pechumani does not inhabit the southern Atlantic Coastal Plain today probably because of high summer and mild winter temperatures, thermic soil temperature regime (annual mean soil temperature ranges from 15 to <22 °C), large amounts of summer precipitation, seasonal high soil moisture values, and an abundance of potential avian, reptilian, and invertebrate predators. This species does not nest during the hottest hours of the day. It needs an extended cold period to break diapause. The slow and jerky movements of the females would seemingly make them easy targets for possible predators. In addition, estuarine erasure of the low elevation, predominantly sandy continental shelf that occurred after 7000 yr BP (Kraft 1977, Whitehead 1972) eliminated considerable habitat for this psammophilous species. Present-day Atlantic Coastal Plain soils from the Neuse River in central North Carolina to Delaware Bay are mostly clayey, silty, and loamy (United States Department of Agriculture 1972). During the l5,000 or so years of climate amelioration or interglacial period, including the Holocene, T. pechumani lived in the northeastern US and southern Ontario. This secondary range is inexorably bound to extensive sand deposits, the result of oceanic regressive and transgressive processes and glacial reworking of the landscape, and climate moderation associated with large bodies of water. As climate started to ameliorate and the ice sheet began to recede about 14,000 years ago (Jacobson et al. 1987, Schoonmaker and Foster 1991), sea level gradually rose, but did not breach the continental shelf until about 9500 yr BP (Fairbanks 1989). Founder populations of T. pechumani could have readily moved northward through a potentially hospitable Atlantic Coastal Plain. Dispersal would have been facilitated by low relief and lack of deeply incised fluvial valleys due to minimal erosion and absence of glaciofluvial 2008 F.E. Kurczewski 17 activity in the region. Based on climate reconstruction (Watts 1980, 1983; Whitehead 1981; Whitehead and Oaks 1979), T. pechumani might have lived as far north as North Carolina by l2,000 yr BP and Virginia before 10,000 yr BP (Fig. 3). Tachysphex pechumani could have been permanently established on the Delmarva Peninsula and interconnected southern New Jersey before 9000 yr BP (Fig. 3). Climate reconstruction for the Chesapeake Bay Region at 9000 yr BP indicates a July mean temperature of 22 °C, January mean temperature of -4 to -2 °C, and annual average amount of precipitation of 800–900 mm (Prentice et al. 1991). The Delmarva Peninsula was broadly expanded to nearly twice its current width at 10,000–9500 yr BP due to lowered sea level (Edwards and Merrill 1977, Kraft 1977). Delaware Bay would have posed only a slight obstacle to dispersal as it did not attain its modern estuarine form until 8000–7000 yr BP (Kraft 1977), probably well after the entry of T. pechumani into southern New Jersey. The inferred Delmarva Peninsula subpopulation has since been eliminated by erasure of the sandy continental shelf, deciduous forest expansion, and/or extensive habitat destruction. An ecological setting comparable to that of the present day was in place in southern New Jersey by 9600 yr BP according to fossil plant assemblages (Florer 1972, Watts 1979). The coastline of New Jersey reached 45–90 km beyond the present shoreline depending on the extent of estuarine inland penetration (Edwards and Emery 1977, Edwards and Merrill 1977). A Mid-Holocene temperate broadleaf community replacement of oak/pinedominant vegetation never occurred in southern New Jersey due to the development of an impoverished podzol on coarse sand and gravel, high soil acidity, and frequent fires (Florer 1972). Fires were perhaps strongly influenced in some areas by Native Americans. From southern New Jersey northward, it is difficult to trace the speculative post-glacial movement of T. pechumani due to size reduction and eventual disappearance of the Atlantic Coastal Plain. Northward dispersal could have taken place only under the most optimal climatic, edaphic, and ecological conditions. According to palynological and stratigraphic evidence (Harrison 1989, Prentice et al. 1991, R. Webb et al. 1993, T. Webb et al. 1994), an environment conducive to T. pechumani habitation existed in the Northeast beginning about 9000 yr BP. Regionally drier and warmer conditions, as inferred from near maximal summer insolation, higher summer temperatures, reduced summer precipitation, decreased soil moisture, lowered lake and stream levels, prevalent convectional thunderstorms, increased incidence of lightning strikes, and periodic natural fires, were coincident with a peak pine period at about that time. The Early to Mid-Holocene dry conditions were time-transgressive along the Eastern Seaboard, occurring first at sites farther from the wasting ice sheet, e.g., 10,000 yr BP in Delaware (Webb et al. 1993). The most plausible, seemingly least disruptive dispersal route with favorable habitat would have been via the low-elevational, low-relief 18 Northeastern Naturalist Vol. 15, Monograph 2 Hackensack (NJ) Lowland, Neversink/Wallkill/Hudson-Mohawk/Champlain Valleys, and Ontario-Erie Lake Plains. This pathway was similar to the Atlantic Coastal Plain in containing a nearly uninterrupted series of abandoned lacustrine bars, beaches, deltas, ridges, terraces, and aeolian modifications of these sandy and gravelly deposits (Barnett et al. 1991, Cadwell et al. 1986 [1986–1991], Stanford and Harper 1991). Presumably this was the route taken by many vascular plants (Catling and Catling 1993, Dirig 1994, McLaughlin 1932, Peattie 1922), psammophilous grasshoppers (Hubbell 1960, Thomas 1951), savanna leafhoppers (Hamilton 1994), various beetles (Schwert et al. 1985), low-altitude butterflies and skippers (Shapiro 1971), and probably many sand-nesting wasps and bees. Whether the dispersal route of T. pechumani included Long Island and Cape Cod is unknown, but these areas were interconnected 9000 years ago due to lowered sea level and they contained a continuum of potentially suitable habitat (Dincauze and Mulholland 1977, Dyke and Prest 1987) (Fig. 3). Governed by a warmer, drier climate and frequent fires, pitch pine/oak barrens, savanna, and woodland were dominant on Cape Cod 8500–6000 years ago (Tzedakis 1992, Winkler 1985). However, both Long Island and Cape Cod became largely forested during the Late Holocene under a cooler, wetter weather regimen heavily influenced by maritimal climate (Tzedakis 1992, Winkler 1985). One fossil insect study demonstrated the movement of an eastern coastal lowland fauna including present-day species into southern Ontario about 8400–7900 yr BP (Schwert et al. 1985). July mean temperature was estimated to have been 1–2 °C higher than present-day. By 7500 yr BP, the insect assemblages in southern Ontario were similar to those found there today (Morgan 1987). Clearly, environmental conditions in this region were favorable for T. pechumani habitation by about 8000 yr BP (Kurczewski 2000a). Such a dispersal route would have incorporated ancestral sand and gravel plains and abandoned sandy and gravelly beaches associated with lower-than-present water levels in Lakes Ontario and Erie (Fig. 3). It would have included a substantially widened land bridge below the then low water stage of Lake Stanley-Nipissing, the predecessor of Lake Huron (Anderson and Lewis 1985, 1992; Coakley 1992; Coakley and Karrow 1994; Lewis et al. 1994) (Fig. 3). Water levels in the Lake Huron Basin were significantly lower from isostatic downwarping and northeastward outflow through North Bay via the Ottawa River to the St. Lawrence Valley (Dyke and Prest 1987). The new shoreline of Lake Huron, especially to the south, had hundreds of square kilometers of recently exposed sandy and gravelly soils. Tachysphex pechumani could have moved virtually unobstructed from southern Ontario to Lower Michigan, Ohio, and Indiana. Had T. pechumani been able to move into northern Lower Michigan at that time, wasps would have had access to the Upper Peninsula of Michigan, Wisconsin, and western Ontario across a narrow Mackinac River (Lewis and 2008 F.E. Kurczewski 19 Anderson 1989, Lewis et al. 1994). But such northward dispersal was probably impeded by delayed summer warming and reduced growing season, the result of a regionally cool climate governed by substantial flow of cold meltwater from proglacial lakes into the western Great Lakes (Anderson and Lewis 1992). After 8000 yr BP, the ice sheet collapsed and retreated from Hudson Bay, thereby permitting northward diversion of most meltwater away from the Great Lakes (Lewis et al. 1994). At that time, July mean temperature averaged 20–21 °C (Prentice et al. 1991), climatic conditions were drier and fires were relatively frequent (Jacobson et al. 1987). By 7500 yr BP, gradually rising water levels in the Great Lakes had begun to form the Straits of Mackinac that persisted through the Holocene (Lewis et al. 1994). Tachysphex pechumani does not inhabit the Upper Peninsula of Michigan. The Straits of Mackinac present a 6–7 km-wide obstacle to this short-distance flying species. Should individuals be accidentally wind swept across the straits, then unstable lakeside dunes, exposed bedrock, poorly drained soils, northern hardwood forests, and coniferous swamps provide unsuitable habitat. Lower average winter and extreme minimum temperatures, cooler summers, increased summer precipitation (Albert et al. 1986), added soil moisture (Webb et al. 1993), and reduced amount of summer sunshine (Eichenlaub 1979) in Upper Michigan compared to Lower Michigan would probably inhibit wasp daily activities and deleteriously impact embryonic development. In contrast, Lower Michigan is the stronghold for T. pechumani. Based on its climatic, ecological, and edaphic requirements, this wasp may have entered Lower Michigan just after the arrival of white pine, perhaps 8000– 7500 years ago (Jacobson 1992), but probably in advance of the deciduousconiferous forest expansion 7000–6000 yr BP (Davis et al. 1986). Pine reached its maximum in Lower Michigan and southwestern Ontario during a warm and dry period 8000 to 7000 years ago (Bernabo and Webb 1977). Tachysphex pechumani presumably dispersed into northern Indiana and northeastern Illinois, where it inhabited oak savanna on sandy soils shortly after it moved into Lower Michigan (Fig. 3). However, the species was probably eliminated from much of this area in the 19th and 20th centuries by habitat destruction and subsequent land alteration. By the 1900s, this region had the least amount of natural vegetation remaining in the conterminous US (Klopatek et al. 1979). Paleohydrologic and palynological data indicate that conditions as wet as today, or perhaps even wetter, persisted in southern Wisconsin from the Early Holocene until about 6300–5500 yr BP (Baker et al. 1992, Winkler et al. 1986). A mesic deciduous forest was in place in the area as maritime tropical air from the Gulf of Mexico blocked dry Pacific air masses from reaching there (Baker et al. 1992), perhaps deterring farther westward dispersal of T. pechumani during the Mid-Holocene. Tachysphex pechumani does not inhabit west-central Wisconsin despite substantial acreage of pine barrens, savanna, and woodland on well-drained 20 Northeastern Naturalist Vol. 15, Monograph 2 sandy soils. Approximately 400 km with no annectent sandy soils separate this area from the nearest suitable habitat for this species, the Indiana Dunes. West-central Wisconsin winters are colder with lower extreme temperatures than in Lower Michigan (National Climatic Data Center 2005). Less snow cover in west-central Wisconsin than in Lower Michigan (United States Department of Agriculture 1941) might expose the shallow wasp nests to frost penetration and potentially dire consequences. West-central Wisconsin receives 65–70 mm more precipitation in May–July than central-northern Lower Michigan (Wendland et al. 1992). Wetter soils in the former region might deleteriously affect development of the immature wasp stages. Slightly more cloud cover and fewer hours of sunshine in June–July in west-central Wisconsin compared to central-northern Lower Michigan could conceivably limit reproductive and nesting activities. The absence of T. pechumani from upstate New York, where it probably once occurred, may be connected with extensive land alteration. But a more likely scenario for its disappearance from the region includes the nearly total replacement of Early to Mid-Holocene, open, pine-dominant woodland with Mid- to Late Holocene, closed, primarily deciduous forest (Bernabo and Webb 1977, Miller 1973, Webb 1988). Land surveys from the 1790s and Late Holocene palynological studies indicate that most of upstate New York was heavily forested (Marks and Gardescu 1992, Russell 1996, Seischab 1992). Dense forest even grew on the loamy sand presumably once inhabited by T. pechumani. Similarly, a closed, pre-settlement forest in mainland New England occupied sandy soils that should have supported edaphically controlled, pine-scrub oak barrens (Patterson 1993). Closed, deciduous-coniferous and deciduous forests in upstate New York and mainland New England flourished from about 6000 yr BP to present-day in connection with regionally wetter and cooler climate. Their growth and expansion resulted from increased summer precipitation, higher soil moisture levels, decreased evapotranspiration, infrequent convectional thunderstorms, diminished incidence of lightning strikes, and fewer natural fires (Harrison 1989, Patterson 1993, Prentice et al. 1991, R. Webb et al. 1993, T. Webb et al. 1994). The absence of T. pechumani from Long Island is an apparent enigma (Kurczewski and Boyle 2000). Many New Jersey pine barrens species of Hymenoptera are likewise missing from this region. Pitch pine-scrub oak-heath vegetation once extended nearly continuously on coarse sandy soils from southern New Jersey through Staten Island, NY and western Long Island to the barrens of Suffolk County (Britton 1880, Cryan 1985, Taylor 1915). The Suffolk County pine barrens persist today under climatic, ecological, and edaphic conditions somewhat similar to those in southern New Jersey, except they are largely disturbed and have a lower water table due to human use (Olsvig et al. 1979). Suffolk County sand and loamy sand have a lower available water capacity (0.02–0.04) and are 2008 F.E. Kurczewski 21 more xeric than comparable southern New Jersey sand (0.04–0.09) and loamy sand (0.10–0.16). Perhaps T. pechumani and many other species of fossorial Hymenoptera were eliminated from Long Island during the extensive deciduous-coniferous and deciduous forest expansion that characterized the Northeast during the Mid- to Late Holocene. Such forest expansion did not affect southern New Jersey because of the extensive acreage of acid, coarse sandy soils and periodic fires (Florer 1972). The Long Island maritime climate may have maintained this forest for 6000 or so years while restricting the frequency and intensity of natural and human-set fires. Pine barrens on Long Island evidently came into prominence only after early settlement destruction of the ancestral, predominantly deciduous forest (Black and Pavacic 1997, Kurczewski and Boyle 2000). Discussion Tachysphex pechumani is at present a central Great Lakes species with a relict population in southern New Jersey. Kurczewski (2000a) noted that the range of this species in the central Great Lakes Region is governed by: (1) extensive acreage of level, nutrient-impoverished sandy soils; (2) frigid/ mesic soil temperature regime (annual mean soil temperature of <8 to <15 °C); (3) climate moderation associated with temperate latitude, peninsular land masses and large bodies of water; (4) maximum coefficient of continentality of 44% (Kopec 1965); (5) annual average precipitation minus evapotranspiration <300 mm (Winter and Woo 1990); (6) maximum average soil moisture as a function of precipitation, evapotranspiration, and temperature of 40/150 mm (Webb et al. 1993); (7) periodic disturbance, especially fire; (8) presence of pre-settlement oak/pine-dominant barrens, savanna, and woodland; and (9) abundance of prey grasshoppers of suitable size, stage, and species. The historic geographic distribution of T. pechumani is superimposed mainly on pre-settlement oak/pine-dominant “woods” (woodland), “plains” (savanna), and “openings” (barrens), as defined in early land surveys (Kurczewski 2000a). Human disturbance has altered the ecological communities in southern Lower Michigan and south-central Ontario enabling this wasp to inhabit areas that are not oak/pine-dominant. Early logging, land clearance, and human-caused fires followed later by bulldozers and motorized vehicles produced openings, sand pits, and trails that facilitated the wasp’s dispersal into atypical habitats. The geographic range of T. pechumani is closely tied to well- and excessively drained, nutrient-poor sand. Acid sandy soils with low organic matter content and reduced available water capacity are inhospitable for agriculture and therefore largely ignored by humans. However, such soils are conducive to high temperature, drought, and fire—all requirements for aspects of the wasp’s ecology and life history. Tachysphex pechumani prefers very coarse and coarse sand over fine sand because large size grains expedite burrow excavation and provide added cohesion. 22 Northeastern Naturalist Vol. 15, Monograph 2 Tachysphex pechumani nests in sand more often than loamy sand. Sand provides a better nesting substrate than loamy sand by restricting the amount and type of plant growth. If left undisturbed, wasp aggregations in loamy sand are frequently eliminated by succession to field and, later, woodland. Closed forest is an eventual outcome of this succession. Loamy sand is much more abundant than sand in northeastern Ohio, Pennsylvania, eastern Ontario, upstate New York, mainland New England, and the Delmarva Peninsula—all regions from which T. pechumani is absent. Large tracts of sand blanket sections of the Lower Peninsula of Michigan and southern New Jersey, and, less so, southern Ontario, northwestern Indiana and northwestern Ohio—all areas where T. pechumani occurs. Sand comprises no less than 19 and 16% of the surficial soils in Lower Michigan and southern New Jersey, respectively. Sand constitutes at least 5% of the surficial soils in 40 counties in Lower Michigan, 9 in northern Indiana, 2 in northwestern Ohio, 13 in southern Ontario, and 10 in southern New Jersey (Fig. 4). Forty-five counties in Lower Michigan, 10 in northern Indiana, 2 in northwestern Ohio, 17 in southern Ontario, and 8 in southern New Jersey have more than 5000 ha of sand. In the 20th century, T. pechumani inhabited 26 of the 40 (65.0%) counties in Lower Michigan that have more than 5% of their surface area in sand and 25 of the 45 (55.6%) counties that hold more than 5000 ha of sand. During the 1900s, this species occupied 7 of the 10 (70%) counties in southern New Figure 4. Percent well-drained sand of total soil acreage by individual county in Lower Michigan, northern Indiana, northwestern Ohio, southern Ontario, southern New Jersey, and the Delmarva Peninsula (percent sand from United States Department of Agriculture and Ontario County Soil Surveys; Camden County, NJ Soil Conservation District). 2008 F.E. Kurczewski 23 Jersey that have more than 5% of their acreage in sand and 6 of the 8 counties (75%) that contain over 5000 ha of sand. Fire and other disturbance are also important in shaping the geographic distribution of T. pechumani, especially in Lower Michigan and New Jersey (Whitney 1986, Niering 1982). Native Americans burned woodlands during the Late Holocene for a variety of reasons, in the process expanding barrens, savanna, and openings. In recent times, the Huron-Manistee National Forests in Lower Michigan demonstrate the highest annual acreage burns in the north-central and northeastern US (Patterson and Backmann 1988). High acreage burns characterized the New Jersey pinelands through the 1800s and early 1900s. A century ago, fires were more frequent and larger than they are today in southern New Jersey (Little 1979). With the advent of motorized fire-fighting equipment and increased surveillance, much less acreage was lost in the late 1900s from fires (New Jersey Department of Environmental Protection 1991). Fire suppression is especially evident in New Jersey’s Lebanon State Forest. In the 1950s, pine-heath barrens and sunlit openings in pine/oak-dominant woodland were visibly part of the landscape (McCormick and Jones 1973). Half a century later, these barrens and woodland have largely been supplanted by pine/oak-dominant forest, resulting in a marked reduction in potential nesting habitat for T. pechumani and other fossorial Hymenoptera. The ancestral New Jersey pinelands included fire-dependent pine/oakdominant woodland with savanna-like, grassy and herbaceous openings (Niering 1982). Recurrent fires prevented the growth of deciduous trees that would enclose the canopy, shade the area, and discourage or prevent nesting by T. pechumani. Periodic fires sustained the pines, oaks, evergreen shrubs, forbs, and grasses whose components fueled the system that established and maintained the barrens and savannas. The heliophilic grasses and forbs, in particular, were vital constituents of this fire-dependent ecosystem because they constituted food for the prey grasshoppers. Tachysphex pechumani perseveres today in the central Great Lakes Region and New Jersey pinelands because of past protection and maintenance of appropriate habitat in state, regional, and local refuges. Nearly all of the present-day nesting sites are located in such areas. This land was available for purchase and conservation in the 1900s because it constituted impoverished sandy soils unsuitable for agriculture and other anthropogenic uses. Fire suppression and excessive use of off-road motorized recreational vehicles are presently eliminating these unique natural communities and endangering the status of T. pechumani and other fossorial Hymenoptera. Currently, T. pechumani subpopulations appear safe in the long-term from extirpation only in sections of Lower Michigan and in the short-term, in the Oak Openings Preserve Metropark, Lucas County, OH and a few small preserves in southwestern Ontario. The continued existence of T. pechumani throughout the lengthy glacial and quasi-glacial periods of the Pleistocene probably can be attributed 24 Northeastern Naturalist Vol. 15, Monograph 2 mainly to the stable, favorable environment of the southeastern Atlantic Coastal Plain. During the geologically brief interglacial periods, this species inhabits areas with conditions that simulate the glacial/quasi-glacial environment of the Southeast, namely the sandy peninsulas of the central Great Lakes and the Middle-Atlantic regions. In order to move from one region to another over many millennia, T. pechumani must utilize the expanded Atlantic Coastal Plain and other extensive sand deposits, the result of oceanic regressive and transgressive processes and glacial reworking of the landscape and inclusive bodies of water. During the 100,000-or-so-years-long period of alternative stability and change incorporating a 4000–5000-kmlong round trip journey, the climatic, ecological, and edaphic requirements of the species may remain essentially the same. Acknowledgments Matthias Buck, K.G.A. Hamilton, Elliot Tramer, and two anonymous reviewers suggested changes that improved the manuscript. Matthias Buck kindly agreed to serve as manuscript editor. Terry Crabe, Denise Gehring, Michelle Grigore, Bill Huff, John Lerg, Rob Line, Mary Rabe, and Bill Westrate assisted with funding or facilities. Matthias Buck, Peter Carson, Ralph Grundel, Steve Krauth, Steve Marshall, Judi Maxwell, Mark O’Brien, Laurence Packer, Gary Parsons, Jeff Skevington, and Bob Vande Kopple furnished collection information. Dick Baker, George Jacobson, Mike Lewis, and Tom Webb answered paleoclimate questions. Wasyl Bakowsky sent maps and early land surveys for southern Ontario. Hugh Boyle, Dana Bruington, Bob Jacksy, Keith Kurczewski, and Ed Stanton assisted with field studies. Bob Jacksy, Dan Krause, and Dudley Raynal identified plant species. Bill Frederick and Mike Guthrie determined Lower Michigan soil series. Dan Dindal and Don Bickelhaupt analyzed New Jersey and Lower Michigan soil samples, respectively. Craig Fisher calculated acreage of sand, loamy sand, and loamy fine sand and percent sandy soils for Camden County, NJ. Bryan Smith and Colleen Totton sent climate information for southern Ontario. Erin Moan and Elliot Tramer agreed to publish their paper concurrently with mine in monograph format. Bob Jacksy provided the cover photograph. Joe Stoll produced Figures 1–4. 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