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; Fkurczewski@twcny.rr.com.
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. Funds for this study were partly provided by
the Michigan Chapter of The Nature Conservancy, Michigan Department of Natural
Resources, Toledo Area Metroparks, State University of New York College of Environmental
Science and Forestry Office of Research Programs, and New York State
United University Professions.
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