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2010 SOUTHEASTERN NATURALIST 9(1):73–84
The Ant (Hymenoptera: Formicidae) Fauna of Black Belt
Prairie Remnants in Alabama and Mississippi
JoVonn G. Hill1,* and Richard L. Brown1
Abstract - Extensive areas of prairie were once found in the southeastern United
States; however, in the last 200 years much of this habitat type has been destroyed.
The largest of these prairie regions, the Black Belt Prairie, extends through portions
of Alabama, Mississippi, and Tennessee. The ant fauna of these endangered
grasslands has not been well documented; therefore, a survey was initiated, with
collections made at 23 Black Belt Prairie remnants in Alabama and Mississippi during
a four-year period. A total of 53 ant species, in six subfamilies and 25 genera
were collected. Six exotic species were collected, and 11 species were found to be
restricted to trees within the prairie, resulting in 36 species that likely represent the
true prairie fauna.
“In the fields the grass is short, no bush; the soil in places is a lead color, yellow
underneath, within the abode of the ants, and very stiff.”
- Benjamin Hawkins (1938) on traveling through a Black
Belt Prairie in Lowndes County, AL in 1798–99.
Prior to European settlement, the southeastern United States had several
regions dominated by grasslands or prairie (DeSelm and Murdock 1993,
Peacock and Schauwecker 2003). One such region, called the “Black Belt,”
extends in an arc from McNairy County, TN to Russell County, AL. Instead
of being a continuous swath of open grassland, the historic landscape of the
Black Belt was a heterogeneous landscape of prairies and several types of
forest (see Barone 2005 for extensive references). Surveys conducted by the
General Land Office in the 1830s suggest that prairies once covered at least
144,000 hectares of the Black Belt (Barone 2005). Since that time, more than
99% of these prairies have been lost to agriculture and urban development
(Noss et al. 1995). Remaining remnants of prairie are threatened by further
development, erosion, and the encroachment of Juniperus virginiana L.
(Cupressaceae) (Eastern Red Cedar), the latter probably as a result of fire
suppression. The Mississippi Natural Heritage Program gives Black Belt
Prairie remnants a ranking of S1, meaning they are “critically imperiled”
within the state due to extreme rarity or factors making their biota vulnerable
to extirpation (Mississippi Museum of Natural Science 2005).
The presence of the prairies in the Black Belt is due, in part, to the
region’s distinctive soils that are derived from a Selma chalk bedrock
1Mississippi Entomological Museum, Box 9775, Mississippi State University, MS
39762. *Corresponding author - email@example.com.
74 Southeastern Naturalist Vol. 9, No. 1
that was formed from Cretaceous marine deposits laid down when the
Mississippi embayment occupied the region (Lowe 1921). These calcareous
loamy soils have a pH in the 7.5–8.5 range, and in some areas have a
dark color, hence the name “Black Belt.” However, much of the land with
these rich, dark soils has been converted to agriculture, and remaining
prairie remnants are often found on marginal light gray soils where the
chalk is close to the surface.
Today, small remnants of these prairies may be found along roadsides,
fencerows, gas, rail, and powerline right-of-ways, and on marginal lands not
suitable for agriculture. These prairie remnants support a diverse and distinct
fl ora and fauna, including species of plants and insects that otherwise are
found primarily in the Great Plains as well as several endemic species of
insects. These disjunct distributions, along with endemic species and fossil
evidence, suggest that Black Belt prairies may have had a prehistoric link to
the Great Plains and served as a refugium for the prairie biota during the last
glacial period (Brown 2003).
Floristic surveys of Black Belt prairie remnants have revealed a distinct
plant community similar to that of the Great Plains of central North America,
with several rare or critically imperiled plants (Leidolf and McDaniel 1998,
Schuster and McDaniel 1973). This community is dominated by grasses
(Poaceae), with Andropogon gerardii Vitman (Big Bluestem), Bouteloua
curtipendula (Michx.), Panicum virgatum L. (Switch Grass), Schizachyrium
scoparium (Michx.) (Little Bluestem), Sorghastrum nutans (L.) Nash
(Indian Grass), and Sporobolus vaginifl orus (Torr. ex Gray) being the most
common. The most prominent forbs include those in the Asclepidaceae
(Asclepias spp. [milkweeds]), Asteraceae (Chrysopsis camporum (Green)
[Golden Aster], Ratibida pinnata (Vent.) [Prairie Conefl ower], Liatris spp.
[blazing stars], Silphium laciniatum L. [Compass Plant], S. terebinthinaceum
(Jacq.) [Prairie Rosinweed]), and Fabaceae (Dalea candida (Mich.) [White
Prairie Clover] and D. purpurea Vent. [Purple Prairie Clover]). Barone and
Hill (2007) provide a more complete fl oral description.
Previous faunal surveys of these prairies have been limited to a few
taxa, but they have revealed an interesting fauna, including: 1160 species of
moths, with 57 disjunct and one endemic species; 33 species of Acrididae
(grasshopper), with one disjunct species; four disjunct species of Cerambycidae
(long-horned beetle), two of which feed on Asclepias spp. (milkweed);
an endemic Carabidae (fl ightless ground beetle); and a disjunct species of
Apidae (bee) that forages only on Purple Prairie Clover (R.L. Brown, unpubl.
Data; Brown 2003; Hill 2007; Landry and Brown 2005; MacGown and
Schiefer 1992; Schiefer 1998).
Ants are typically the most dominant and infl uential force in terrestrial
ecosystems, and as such, may be among the most promising groups of animals
for inclusion in community-based studies due to their contributions to
ecosystem function (Agosti et al. 2000, Hölldobler and Wilson 1990). They
directly and indirectly affect faunal and fl oral groups by predation, scaveng2010
J.G. Hill and R.L. Brown 75
ing, tending homopterans, protecting certain plants, dispersing seeds, and
aiding in nutrient and soil turnover (Agosti et al. 2000, Gorb and Gorb 2003,
Hölldobler and Wilson 1990, Shultz and McGlynn 2000, Wheeler 1910). Ants
are also of interest because of the putative deleterious effects caused by nonnative
species, especially the imported fire ants (Solenopsis invicta Buren, S.
richteri Forel, and their hybrid S. invicta x richteri), which negatively affect
human endeavors and may reduce biodiversity in the communities they invade
(Gotelli and Arnett 2000, Kaspari 2000, Morris and Steigman 1993, Porter
and Savignano 1990, Vinson 1997). Given the ecological importance of ants,
documenting the ant fauna of endangered ecosystems, such as the Black Belt
Prairie, could provide useful information for guiding current management and
future conservation/restoration projects in those systems.
Study sites throughout the Black Belt were sampled to ensure that a comprehensive
picture of the fauna, including any latitudinal variations, was
developed (Fig. 1). Sites were chosen based on their accessibility and relative
fl oral intactness. The level of past disturbance did vary, but most of the sites
experienced relatively little human disturbance during this study. Sites experiencing
disturbance consisted of several roadside prairies being partially
mowed, and the site in the Tombigbee National Forest was burned during
the early spring of each sample year. Almost all of the sites are within or near
Figure 1. Map showing historical prairie remnants, based on General Land Office
Surveys in the 1930s (from Barone 2005), location, site number, and number of ant
species collected during this survey.
76 Southeastern Naturalist Vol. 9, No. 1
the boundaries of known historic prairies, as described by Barone (2005)
based on General Land Office plat maps from the 1830s and 1840s (Fig. 1).
Twenty-three prairie remnants (Fig. 1, Table 1) were surveyed from June
2003 through July 2007. Three sites–Crawford, Osborn, and Tombigbee
National Forest–were studied more intensively during the initial phase of
the study. These sites were sampled twice a month from June to October in
2003 and from April to December in 2004, then revisited every three months
throughout the study period. The remaining sites were sampled monthly
from April to October for at least one year subsequent to their discovery.
After the first sampling year of a site, it was re-visited several times across
seasons and years to ensure that it had been sampled effectively.
From June to October, 2003 and from April to December, 2004, ants were
collected at the Crawford, Osborn, and Tombigbee sites using a variety of
collecting techniques including baiting, hand collecting, and pitfall traps.
Hill et al. (2008) provide a more detailed explanation of the procedures used
at these sites. On subsequent dates, these sites as well as the other 20 sites
were sampled by baiting with cookies, (Keebler Sandies Pecan Shortbread®),
by sifting through soil, grass duff, and cedar litter, and by active searching.
Searching consisted of looking for nests and foragers on the open ground and
on or inside stems and other plant parts. As Eastern Red Cedars are common
components of unburned prairies, and often the main target of management
activities, an effort was also made to collect ants that occurred on these trees
Table 1. List of study sites with their latitude and longitude.
Site County Latitude, longitude
Tupelo Powerline Lee 34°14'26''N, 88°48'39''W
Trace State Park Pontotoc 34°15'24"N, 88°53'12"W
Natchez Trace National Park Lee 34°15'24"N, 88°45'22"W
Peacely Ferry Road Monroe 34°00'20"N, 88°34'23"W
Tombigbee National Forest Chickasaw 33°55'39"N, 88°51'18"W
Clay County Clay 33°43'52"N, 88°49'44"W
Osborn Oktibbeha 33°30'41"N, 88°44'08"W
MSU North Farm Oktibbeha 33°27'43"N, 88°45'31"W
HWY 82 X 45A Lowndes 33°29'17"N, 88°39'38"W
Brooksville Noxubee 33°15'52"N, 88°33'19"W
Crawford Lowndes 33°18'01"N, 88°36'38"W
Plum Creek Noxubee 33°06'01"N, 88°31'14"W
Kemper County Kemper 32°52'24"N, 88°29'28"W
Cochrane Recreational Area Greene 33°04'21”N, 88°15'52"W
Aliceville Pickens 33°14'09”N, 88°24'15"W
Emelle Sumter 32°46'38"N, 88°18'50"W
Epes Sumter 32°41'49"N, 88°07'11"W
University of West Alabama Sumter 32°36'17"N, 88°11'35"W
Sparks Road Marengo 32°29'18"N, 87°44'44"W
Old Cahawba Dallas 32°18'44"N, 87°07'33"W
Hayneville Lowndes 32°08'51"N, 86°32'30"W
Letohatchee Lowndes 32°08'29"N, 86°30'35"W
County Road 37 Lowndes 32°08'56"N, 86°29'44"W
2010 J.G. Hill and R.L. Brown 77
and in their litter to investigate their effect on ant species composition in
the prairie. Specimens were collected in 90% ethanol, then were pinned and
labeled, and identified. Ants were also sorted from stored pitfall trap samples
taken by the Mississippi Entomological Museum (MEM) at the Osborn and
Crawford sites in the early 1990s. Voucher specimens have been deposited
in the MEM.
A total of 52 ant species including the hybrid fire ant Solenopsis invicta
x richteri, representing 6 subfamilies and 25 genera, were collected
(Appendix 1). The most diverse subfamilies included the Myrmicinae with
30 species and the Formicinae with 12 species. The most diverse genera were
Pyramica with seven species and Pheidole with five species. Six exotic species
were collected including the imported fire ants (S. invicta and S. invicta
x richteri), Brachymyrmex patagonicus Mayr, Cyphomyrmex rimosus, Pyramica
margaritae, and P. membranifera. Eleven native species were found to
only be associated with cedar or other trees within the remnants. The remaining
35 native species likely represent the true prairie ant fauna. Imported
fire ants were collected at all the sites, with the hybrid fire ant, S. invicta x
richteri, collected at the sites from Plum Creek northward (16 sites), while
the red imported fire ant S. invicta, was collected at the sites south of the
Plum Creek site (7 sites). The native species collected at the most sites were
Monomorium minimum (23 sites), followed by Forelius mccooki (17 sites),
and Pheidole tysoni and Solenopsis c.f. molesta (16 sites both). Two species
found in this survey, Hyponera inexorata and Pheidole lamia, represent new
state records for Alabama.
The number of species at individual sites ranged from a low of three at
the County Road 37 site, a small (0.011 ha) roadside remnant, to highs of
28, 26, and 24, represented by the Osborn, Crawford, and Tombigbee sites,
respectively. All three of these latter remnants are found in Mississippi and
are large (more than 6 ha.) and protected. The Letohatchee site, a roadside
remnant (0.11 ha), was the most diverse site in Alabama with 18 species. The
mean number of species per site was 12.35, and 10 of the sites contained at
least this many species.
The results of this survey indicate that Black Belt Prairie remnants support
a relatively diverse ant fauna. Typically, the most abundant ant species within
a remnant are Forelius mccooki (McCook) followed by several Myrmicines
such as Monomorium minimum (Buckley), imported fire ants (Solenopsis
spp.), Solenopsis c.f. molesta (Say), Pheidole tysoni Forel, and Crematogaster
lineolata (Say); although species composition and abundance varies
depending on the amount of disturbance (based on area, native/exotic plant
ratio, etc.) at the site, with imported fire ants apparently being more dominant
78 Southeastern Naturalist Vol. 9, No. 1
on highly disturbed sites (J.G. Hill, pers. observ.). The ant fauna of Black Belt
Prairies are similar to that of the southern portion of the tall grass prairie biome
of Arkansas, Texas, and Oklahoma where the subfamily Myrmicinae and the
genus Forelius also make up dominant components of the fauna (Trager 1998;
M. Warriner and J.G. Hill, unpubl. data). However, Hill et al. (2008) indicated
that the ant fauna of prairie remnants was significantly different and more diverse
than pastures on similar soils in the Black Belt of Mississippi, and that
prairie remnants supported a significantly different but less diverse ant fauna
than the forests of the region. The ant fauna of Black Belt Prairies also differs
from the fauna of other naturally occurring open habitats in the Southeastern
United States, such as coastal dunes or sandhill/pine savannah habitats, based
on faunal lists from these habitats compiled from MacGown and Hill (2007),
Hill and MacGown (2008), and Graham et al. (2004). Notable ants that occur
on Black Belt prairie remnants include Dolichoderus pustulatus, Polyergus
montivagus, Hypoponera inexorata, Pyramica bimarginata, Pheidole lamia,
and Pheidole pilifera. Dolichoderus pustulatus was found apparently nesting
in a clump of Indian Grass at the Letohatchee site. Specimens of D. pustulatus
in the MEM were collected from the Osborn and Crawford sites in pitfall
traps run in the late 1990s, but no recent specimens were found at those sites
despite extensive searches. Polyergus montivagus was collected in pitfall
traps from the Tombigbee prairie site (Hill and Brown 2005). Hypoponera
inexorata is a subterranean species that can be collected by sifting soil taken
from the base of grass clumps or cedar trees. This species was not listed as
occurring in Alabama in MacGown and Forester (2005), thus the collections
made during this survey from the Aliceville, Sparks Road, and Old Cahawba
sites represent new state records for Alabama. Smith (1927) lists H. inexorata
from A and M College, MS (now Mississippi State University), the campus
of which lies at the edge of the Black Belt and still contains prairie remnants.
One colony of this species, containing several queens, was found underneath
a hybrid fire ant colony. Pyramica bimarginata was found in pitfall samples
taken in the early 1990s from the Osborn site, and despite extensive searches
during this survey no additional specimens were discovered. Three specimens
of Pheidole lamia were collected at the Aliceville site, a small (0.107 ha)
roadside remnant. Specific microhabitat information for this collection could
not be determined; however, this species has been found to inhabit the soil in
large stands of Indian Grass and Big Bluestem in the similar Jackson Prairie
further south in Mississippi. (J.G. Hill, pers. observ.). This species is a new
state record for Alabama based on MacGown and Forester (2005). Pheidole
pilifera was only found at the three largest and most fl oristically intact sites—
Crawford, Osborn, and Tombigbee—indicating that this species may inhabit
only the less disturbed remnants. Hill (2006) reported observations of this
species feeding on seeds of Switch Grass during the fall of 2005; however, in
the fall of 2006, P. pilifera was observed ignoring seeds of that species completely
in favor of seeds of an undetermined grass a few weeks earlier than the
2010 J.G. Hill and R.L. Brown 79
Several ant species inhabit the soil and litter that accumulates around the
bases of clump grasses such as Bib Bluestem, Switch Grass, Little Bluestem,
and Indian Grass. These include Brachymyrmex depilis, Myrmecina americana,
Pyramica creightoni, P. dietrichi, P. margaritae, P. membranifera,
P. talpa, and Pheidole lamia. Most of these species are typically found in
forest leaf litter; however, they also seem to be adapted to this microhabitat
of open prairies. All of the above mentioned litter-dwelling species were
absent from the Tombigbee prairie, which is burned in entirety annually
resulting in the loss of the microhabitat that these species require. Most of
these species may also be found in the litter under Eastern Red Cedar trees,
but are not limited to it.
Seven species were limited to inhabiting the litter or soil under Eastern
Red Cedar trees, including Camponotus castaneus, Proceratium pergandei,
Pyramica ornata, Strumigenys louisianae, Solenopsis picta, Aphaenogaster
carolinensis Wheeler, and Temnothorax curvispinosus. Two ant species
were limited to other species of trees, Camponotus decipiens on Diospyros
virginiana L. (Ebenaceae) (Persimmon), and Camponotus mississippiensis
on Fraxinus americana L. (Oleaceae) (White Ash). Crematogaster ashmeadi
occurred on both cedar and ash trees. A single alate queen of Camponotus
impressus, a species typically associated with trees, was collected in open
prairie by J. Trager and J.G. Hill on 3 June 2003. No other specimens of this
species were collected, suggesting that this individual was possibly migrating
through this habitat.
Exotic ant species reached their highest population densities in remnants
that had anthropogenically disturbed areas or were wholly disturbed. In
remnants where only a portion of the area is disturbed, exotic ants primarily
reached their highest densities in the disturbed areas. An exception to
this may be Brachymyrmex patagonicus, which is a recent invader to these
remnants, due to its range expansion (MacGown et al. 2007). Brachymyrmex
patagonicus seemingly reached one of the remnants (site 13) during the
course of this study. It was likely already established at the other three sites
(21, 22, and 23); although it was not found in the initial two surveys of site
22, it was extremely abundant throughout the site on the third survey.
In conclusion, the faunal list produced in this study should provide some
baseline data on the ant species composition of prairies for current and future
conservation and restoration efforts. Such efforts are important given
the endangered status of Black Belt Prairie remnants. Future investigations
will focus on relatedness of individual remnants, how various habitat variables
(e.g., plant diversity and remnant size) infl uence ant assemblages,
and the relationship of grasslands in the southeastern United States to those
of the Great Plains.
We thank John A. Barone for his assistance in the field and producing the figure.
We thank Joe MacGown for verifying several identifications. We thank Libby
80 Southeastern Naturalist Vol. 9, No. 1
Hartfield, Mississippi Museum of Natural Science, for assisting in obtaining collecting
permits for state parks, John Baswell for allowing collections to be made
in Tombigbee National Forest, and Doug Wymer and Robby Limerick for access to
the UWA prairie. We also express our gratitude to Sarah McClure, Rebecca Jones,
and Savannah Duckworth for their assistance in specimen preparation. This research
was supported by the Mississippi Forestry Experiment Station Project MIS-311080,
USDA-ARS Areawide Management of Imported Fire Ant Project, National Science
Foundation grants DSR-90204 and DEB-9200856, and a grant from Prairie Biotic
Inc. This is publication J-11468 of the Mississippi Agricultural and Forestry Experiment
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Harvard University Press. Cambridge, MA. 794 pp.
2010 J.G. Hill and R.L. Brown 83
Appendix 1. The following list is arranged by subfamily, tribe, and genus according
to Bolton (2003). Names follow Bolton (1995), except Dorymyrmex, which
follows Snelling (1995); Pyramica, which follows Bolton (2000), Pheidole,
which follows Wilson (2003), and Polyergus, which follows Smith (1947). Following
species names is the site number at which they occurred. The superscript
T denotes a species that is associated with trees, an * denotes that this species was
found in the collection of the MEM from pitfall traps run in the early 1990s, and
Exotic indicates a species not native to the United States based on Deyrup (1992),
McGlynn (1999), and Deyrup et al. (2000).
Dolichoderus pustulatus Mayr: Sites 7*, 11*, and 22
Dorymyrmex smithi McCook: Sites 17 and 21.
Forelius mccooki (McCook): Sites 2, 3, 4, 5, 6, 7, 10, 11, 12, 14, 15, 16, 17, 18,
20, 21, and 22
Tapinoma sessile (Say): Sites 3, 8, 10, and 11
Lasius neoniger Emery: Site 11
Brachymyrmex depilis Emery: Sites 2, 14, 15, 20, 22
Brachymyrmex patagonicus Mayr: Sites 13, 21, 22 and 23. Exotic
Paratrechina arenivaga (Wheeler): Sites 5, 7, 11, 12, 14, and 16
Paratrechina vividula (Nylander): Sites 1, 2, 3, 4, 7, 8, 11, 12, 13, 14, 15, 16,
17, 18, 19, 20, 21, and 22
Prenolepis imparis Emery: Sites 5 and 12
Camponotus castaneus (Latreille) T: Sites 5, 7, and 11
Camponotus decipiens Emery T: Sites 5 and 15
Camponotus impressus (Roger) T: Site 7
Camponotus mississippiensis Smith T: Sites 12, 13, and 19
Formica dolosa Buren: Sites 1, 2, 4, 5, 12, 13, 15, 16, and 18
Polyergus lucidus montivagus Wheeler: Site 5
Pseudomyrmex pallidus (Smith): Sites 17, 20, and 22
Pseudomyrmex seminole Ward: Sites 20 and 21
Hypoponera inexorata (Wheeler): Sites 6, 7, 13, 15, 19, and 20
Hypoponera opaciceps (Mayr): Sites 1, 11, 15, 16, 19, and 21
Hypoponera opacior (Forel): Sites 3, 5, 7, 10, 11, 12, 13, 14, 15, 16, 17, 18, 19,
Ponera pennsylvanica Buckley: Sites 2, 4, 5, 7, 8, and 11
Proceratium pergandei (Emery) T: Site 7
Pyramica bimarginata (Wesson & Wesson): Site 7*
Pyramica creightoni (Smith): Sites 11, 12, and 22
Pyramica dietrichi (Smith): Sites 11, 14, and 17
Pyramica margaritae (Forel): Site 21. Exotic
Pyramica membranifera (Emery): Site 20. Exotic
Pyramica ornata (Mayr) T: Site 7
Pyramica talpa (Weber): Sites 7, 11, 12, 13, 17, and 22
84 Southeastern Naturalist Vol. 9, No. 1
Strumigenys louisianae Roger T: Sites 7, 8, and 11
Cyphomyrmex rimosus (Spinola): Sites 21 and 22. Exotic
Trachymyrmex septentrionalis (McCook): Sites 5, 7, 11
Monomorium minimum (Buckley): Sites1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13,
14, 15, 16, 17, 18, 19, 20, 21, 22, and 23
Solenopsis invicta Buren: Sites 16, 17, 18, 19, 20, 21, 22, and 23. Exotic
Solenopsis invicta x richteri: Sites 1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15.
Solenopsis c.f. molesta (Say) : Sites 1, 2, 5, 7, 8, 9, 10, 11, 12, 13, 14, 15, 18,
20, 21, and 22
Solenopsis picta Emery T: Site 7
Aphaenogaster carolinensis Wheeler T: Sites 2 and 11
Aphaenogaster fl emingi Smith: Site 5
Aphaenogaster treatae Forel: Sites 5, 7, 11, and 15
Pheidole bicarinata Mayr: Sites 6, 12, 15, and 18
Pheidole dentata Mayr: Sites 1, 2, 5, 6, 7, 8, 11, 12, 15, and 22
Pheidole lamia Wheeler: Site 15
Pheidole pilifera Roger: Sites 5, 7, and 11
Pheidole tysoni Forel: Sites 1, 2, 4, 5, 6, 7, 8, 9, 11, 12, 13, 16, 18, 20, 21, and
Crematogaster ashmeadi Mayr T: Sites 7 and 13
Crematogaster lineolata (Say): Sites 2, 5, 6, 7, 11, 15, 17, 21, and 22
Crematogaster missuriensis Emery: Sites1, 5, 8, 11, 12, 13, and 17
Crematogaster pilosa Emery: Sites 5, 21, and 22
Temnothorax curvispinosus (Mayr) T: Sites 7, 12, and 15
Temnothorax pergandei (Emery): Sites 2, 5, 6, 7, and 11
Myrmecina americana Emery: Sites 5, 7, 11, 13, 18, 20, and 22