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The Ant (Hymenoptera: Formicidae) Fauna of Black Belt Prairie Remnants in Alabama and Mississippi
JoVonn G. Hill and Richard L. Brown

Southeastern Naturalist, Volume 9, Issue 1 (2010): 73–84

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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. Introduction “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 - 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. Methods 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 Mississippi 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 Alabama 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. Results 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. Discussion 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 2005 observations. 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. Acknowledgements 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 Station. Literature Cited Agosti, D., J.D. Majer, L.E. Alsonso, and T.R. Shultz. 2000. Ants: Standard Methods for Measuring and Monitoring Biodiversity. Smithsonian Press, Washington, DC. 280 pp. Barone, J.A. 2005. Historical presence and distribution of prairies in the Black Belt of Mississippi and Alabama. Castanea 70:170–183. Barone, J.A. and J.G. Hill. 2007. Flora of blackland prairie remnants in Mississippi and Alabama. Castanea.72:226–234. Bolton, B. 1995. A New General Catalogue of the Ants of the World. Harvard University Press, Cambridge, MA. 504 pp. Bolton, B. 2000. The ant tribe Dacetini. 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Davis. 2000 Exotic ants in Florida. Transactions of the American Entomological Society. 126:293–325. Gorb, E., and S. Gorb. 2003. Seed Dispersal by Ants in a Deciduous Forest Ecosystem: Mechanisms, Strategies, and Adaptations. Kluwer Academic Publishers, Dordrecht, The Netherlands. 225 pp. Gotelli, N.J., and A.E. Arnett. 2000. Biogeographic effects of red fire ant invasion. Ecology Letters 3:257–261. Graham, J.H., H.H. Hughie, S. Jones, K. Wrinn, A.J. Krzysik, J.J. Duda, D.C. Freeman, J.M. Emlen, J.C. Zak, D.A. Kovacic, C. Chamberlin-Graham, and H. Balbach. 2004. Habitat disturbance and the diversity and abundance of ants (Formicidae) in the southeastern fall-line sandhills. Journal of Insect Science 4:30–45. Hawkins, B. 1938. A sketch of the Creek Country in the years 1798 and 1799. Georgia Historical Society collections. Volume 18 reprint. Americus, GA. 2010 J.G. Hill and R.L. Brown 81 Hill, J.G. 2006. Field observations of Pheidole pilifera (Roger) and Paratrechina arenivaga (Wheeler) (Hymenoptera: Formicidae) in a Black Belt Prairie remnant in Mississippi. Journal of the Mississippi Academy of Sciences. 51:183–186. Hill, J.G. 2007. The grasshopper (Orthoptera: Romaleidae, Acrididae) fauna of Black Belt Prairie remnants in Mississippi and Alabama. Journal of Orthoptera Research 16:139–144. Hill, J.G., and R.L. Brown. 2005. The first record of the genus Polyergus (Hymenoptera: Formicidae) in Mississippi. Journal of the Mississippi Academy of Sciences 5:120. Hill, J.G., and J.A. MacGown. 2008. Survey of grasshoppers and ants from the Big Hammock, Ohoopee Dunes, and Fall Line Sandhills Natural Areas. Report submitted to the Georgia Department of Natural Resources. 30 pp. Hill, J.G., K.S. Summerville, and R.L. Brown. 2008. Habitat associations of ant species (Hymenoptera: Formicidae) in a heterogeneous Mississippi landscape. Environmental Entomology 37:453–463. Hölldobler, B., and E.O. Wilson. 1990. The Ants. Harvard University Press. Cambridge, MA. 732 pp. Kaspari, M. 2000. Do imported fire ants impact canopy arthropods? Evidence from simple arboreal pitfall traps. Southwestern Naturalist 45:118–122. Landry, B., and R.L. Brown. 2005. Two new species of Neodactria Landry (Lepidoptera: Pyralidae: Crambinae) from the United States of America. Zootaxa 1080:1–16. Leidolf, A., and S. McDaniel. 1998. A fl oristic study of Black Prairie plant communities at Sixteen-Section Prairie, Oktibbeha County, Mississippi. Castanea 63:51–62. Lowe E.N. 1921. Plants of Mississippi: A list of fl owering plants and ferns. Mississippi State Geological Survey Bulletin 17:1–292. MacGown, J.A., and J.A. Forster. 2005. A preliminary list of the ants (Hymenoptera: Formicidae) of Alabama. Entomological News 116:61–74. MacGown, J.A., and J.G. Hill. 2007. Ants of Bon Secour National Wildlife Refuge, Baldwin County, Alabama. Available online at mississippientmuseum/Researchtaxapages/Formicidaepages/faunal.lists/Bon- Secour.Ants.htm. Accessed September 2007. MacGown, J.A., J.G. Hill, and M.A. Deyrup. 2007. Brachymyrmex patagonicus (Hymenoptera: Formicidae), an emerging pest in the southeastern United States. Florida Entomologist 90:457–467. MacGown M.W., and T.L. Schiefer. 1992 Disjunct distribution and new record for an anthophorid bee, Xenoglossodes albata (Hymenoptera: Anthophoridae), in the southeastern United States. Entomological News 103:81–82. McGlynn, T.P. 1999. The worldwide transfer of ants: Geographical distribution and ecological invasions. Journal of Biogeography 26:535–548. Mississippi Museum of Natural Science. Natural Heritage Inventory: Global and State Rankings. Available online at ranking.asp. Accessed April 2005. Morris, J.R., and K.L. Steigman 1993. Effects of polygyne fire ant invasion on native ants of a blackland prairie in Texas. Southwestern Naturalist 38:136–140. 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Ants: Standard Methods for Measuring and Monitoring Biodiversity. Smithsonian Institution Press, Washington, DC. 280 pp. Schuster, M.F., and S. McDaniel. 1973. A vegetative analysis of a black prairie relict site near Aliceville, Alabama. Journal of the Mississippi Academy of Science 19:153–159. Smith, M.R. 1927. An additional annotated list of the ants of Mississippi with a description of a new species of Pheidole (Hymenoptera: Formicidae). Entomological News 38:308–314. Smith M.R. 1947. A study of Polyergus in the United States, based on the workers (Hymenoptera: Formicidae). American Midland Naturalist 38:150–161. Snelling, R.R. 1995. Systematics of Nearctic ants of the genus Dorymyrmex (Hymenoptera: Formicidae). Contributions in Science (Los Angeles) 454:1–14. Trager, J.C. 1998. An introduction to ants (Formicidae) of the tallgrass prairie. Missouri Prairie Journal 18:4–8. Northern Prairie Wildlife Research Center, Jamestown, ND. Available online at ants/index.htm. (Version 17MAY99). Accessed September 2007. Vinson, B.S. 1997. Invasion of the red imported fire ant (Hymenoptera: Formicidae): spread, biology, and impact. American Entomologist 43:23–39. Wheeler, W.M. 1910. Ants: Their Structure, Development, and Behavior. Columbia University Press, New York, NY. 663 pp. Wilson, E.O. 2003. Pheidole in the New World: A Dominant, Hyperdiverse Ant Genus. 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). Family Formicidae Subfamily Dolichoderinae 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 Subfamily Formicinae 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 Subfamily Pseudomyrmecinae Pseudomyrmex pallidus (Smith): Sites 17, 20, and 22 Pseudomyrmex seminole Ward: Sites 20 and 21 Subfamily Ponerinae 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, and 22 Ponera pennsylvanica Buckley: Sites 2, 4, 5, 7, 8, and 11 Subfamily Proceratiinae Proceratium pergandei (Emery) T: Site 7 Subfamily Myrmicinae 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. Exotic 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 22 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