2006 SOUTHEASTERN NATURALIST 5(4):573–586
Observations on the High Diversity of Native Ant Species
Coexisting with Imported Fire Ants at a Microspatial Scale
in Mississippi
Joe A. MacGown1,* and Richard L. Brown1
Abstract - A case study reporting a high diversity of native ants co-existing with
imported fire ants is presented. Thirty-six species of ants were collected within one
meter of the base and on the lower two meters of the trunk of a Quercus pagoda
(cherry bark oak) in Tombigbee National Forest, Winston County, MS, on five dates
during 2003 and 2004. Twenty-three of these species, including the imported fire ant
hybrid, Solenopsis invicta x richteri, were nesting in the same area. A list of all
species collected at the site is provided. Notes are given on the nesting habits of
Polyergus lucidus longicornis and its slave, Formica dolosa, and also for other
species in the area. Potential explanations for high diversity including the island
effect of an isolated habitat, differing diets and foraging behaviors, habitat partitioning,
and seasonal activity are discussed.
Introduction
The imported fire ants, Solenopsis richteri Forel, S. invicta Buren, and
their hybrid, S. richteri x invicta, have frequently been reported to adversely
affect species richness of native ants (Camilo and Phillips 1990, Holway et
al. 2002, Jusino-Altresino and Phillips 1994, Morris and Steigman 1993,
Porter and Savignano 1990). However, in a follow-up study 12 years after
that of Porter and Savignano, Morrison (2002) found that arthropod diversity
had returned to levels that were comparable to those prior to invasion of S.
invicta. Other studies in Florida and Texas have documented high diversity
of native ants coexisting with S. invicta (Helms and Vinson 2001, Morrison
and Porter 2003). Tschinkel (1988) speculated that predation by Solenopsis
molesta (Say) upon brood of S. invicta may restrict S. invicta to disturbed
habitats where S. molesta is absent.
Our knowledge of the ant fauna in Mississippi is based primarily on the
early inventory of M.R. Smith (1924a,b,c; 1927; 1928a,b; 1931a,b; 1932).
Smith’s inventory, subsequent records by other workers, and recent surveys
have documented 151 species of ants for Mississippi (Baroni Urbani and De
Andrade 2003; Bolton 2000; DuBois and Davis 1998; Hill and Brown 2005;
MacGown and Brown, in press; MacGown et al. 2005; MacKay 1993;
Shoemaker et al. 1994; Smith 1979; Snelling 1995; Trager 1984, 1988,
Trager et al., in press; Umphrey 1996; Ward 1985; Watkins 1985; Wilson
2003). An additional 14 new state records of ant species will be reported in
1Mississippi Entomological Museum, Box 9775, Department of Entomology and
Plant Pathology, Mississippi State, MS 39762. *Corresponding author -
jmacgown@entomology.msstate.edu.
574 Southeastern Naturalist Vol. 5, No. 4
the future, bringing the total to 165 species known to occur in Mississippi
(J.A. MacGown and R.L. Brown, unpubl. data).
This report documents a case study of high species richness of native ants
co-existing with the imported fire ant hybrid, S. richteri X invicta, at a
microspatial scale in Tombigbee National Forest, Winston County, MS. In
addition, a nest containing the rarely detected Polyergus lucidus longicornis
Smith, and its slave, Formica dolosa Buren, is described. Nesting habits and
behavior of other species are also described.
Study Area and Collection Methods
The study site was in the Tombigbee National Forest (Ackerman Unit) in
Winston County, MS at 33°12'30"N, 89°04'32"W and included the basal two
meters of a mature and partially decayed Quercus pagoda Rafinesque
(cherrybark oak; Fagaceae) and one meter of soil and litter around the
perimeter of the tree (Fig. 1). The oak tree was located at the edge of a sandy
gravel parking area next to an old cemetery (Noxubee Hill Cemetery) in a
relatively open and somewhat dry area on a ridge bordered by a rich mixed
forest on steep sloped ravines. The tree had a circumference of 3.21 meters
at 0.91 meters above ground level, and it was surrounded by a rich layer of
litter at its base.
The serendipitous checking of the oak tree by J.A. MacGown during an
afternoon rainstorm on 10 July 2003 resulted in discovery of a nest containing
Polyergus lucidus longicornis and its slave, Formica dolosa. In order to
collect specimens of the Polyergus, litter and soil from the nest area were
collected and sifted with a screen into a shallow pan from which specimens
of the Polyergus and additional species of ants were collected. A sample of
the sifted litter and soil was collected in a gallon-sized (3.79-L) plastic bag
for extraction with Berlese funnels, and the excess screened litter and soil
was placed back onto the nest. Other ants were collected as they were
sighted on the base of the tree or adjacent ground. All collecting was done
within one hour during which time there was continual rain.
Subsequent visits to this site were made on 15 July 2003 (J.A. MacGown),
21 October 2003 (J.A. MacGown), 11 November 2003 (J.A. MacGown and
R.L. Brown), and 26 June 2004 (J.A. MacGown). Visits on 15 July and 21
October were limited to less than a half hour, and collecting on these days
was limited to use of baits. On both 11 November and 26 June, ants were
collected for about an hour by visual examination of the basal two meters of
the tree trunk and surrounding ground within one meter of its base, sifting
litter, and baiting. Baits included cookies (Keebler Sandies Pecan
Shortbread®), tuna (StarKist® chunk light in water), and peanut butter (generic
brand). Cookie bait on six white cards (15 July) and tuna bait on 12
white cards (21 October) were evenly spaced around the perimeter of the tree.
Both tuna and cookie baits on cards were used in the same pattern on 11
November. Cookie and peanut butter baits were placed on the tree trunk
approximately 1.5 to 2 meters high on 26 June. Soil and litter samples were
2006 J.A. MacGown and R.L. Brown 575
taken on 11 November (5 one-gallon bags) and 26 June (1 one-gallon bag) for
extraction of ants with Berlese funnels.
Results
During the five visits to this site over the duration of a year, 36 species of
ants, including the hybrid fire ant, Solenopsis invicta x richteri, were collected
on the lower trunk of the oak tree or within one meter from its base
(Table 1). These 36 species, collected in less than four hours, represent more
than half of the species that were collected at 21 sites in the Tombigbee
National Forest from 1999 to 2004 (MacGown and Brown, in press).
Figure 1. Joe MacGown searching for ants at the base of a Quercus pagoda
(cherrybark oak ) in Tombigbee National Forest, Winston County, MS.
576 Southeastern Naturalist Vol. 5, No. 4
Twenty-three species were collected on the first visit on 10 July, one species
was added on 21 October, and six species were added on each of the last two
visits when collecting was more intense. Based on the presence of colonies
or the collection of both workers and dealate queens, 22 species were
determined to be nesting at the site. This number of nesting species is
probably conservative because small species, such as Pyramica spp. and
Brachymyrmex depilis Emery, do not forage far from their nests, and therefore
likely had colonies in the area.
Table 1. Ant species collected at an individual cherrybark oak tree in Tombigbee National
Forest with collection dates and indication if they were foraging (F), collected at bait (cookies-
C, peanut butter-P, or tuna-T), found nesting (N), or extracted from soil and litter in Berlese
funnels (B).
2003 2004
Jul Jul Oct Nov Jun
Species 10 15 21 11 26 F C P T N B
Aphaenogaster carolinensis X X X X
Aphaenogaster fulva X X X X X X X X X
Aphaenogaster lamellidens X X X X X
Aphaenogaster mariae X X X X X
Brachymyrmex depilis X X
Camponotus nearcticus X X X
Camponotus pennsylvanicus X X X X X X
Camponotus snellingi X X X X X X
Crematogaster lineolata X X X X
Formica dolosa X X X X
Formica pallidefulva X X X X X
Hypoponera opacior X X X X X
Monomorium minimum X X X X X
Myrmecina americana X X X X X
Paratrechina arenivaga X X X
Paratrechina faisonensis X X X X X X X
Paratrechina vividula X XX
Pheidole bicarinata X X X X X X
Pheidole dentata X X X X X X
Pheidole dentigula X X X X X
Pheidole tysoni X X X X X X
Polyergus lucidus longicornisX X
Ponera pennsylvanica X X X X X
Prenolepis imparis X X X X X
Proceratium silaceum X X
Pyramica angulata X X
Pyramica dietrichi X XX
Pyramica ornata X X X
Pyramica pulchella X X
Solenopsis invicta x richteri X X X X X X X X X
Solenopsis molesta X X X X X X
Strumigenys louisianae X X X X X
Temnothorax curvispinosus X X
Temnothorax pergandei X X
Temnothorax schaumii X X X
Trachymyrmex septentrionalis X X X
2006 J.A. MacGown and R.L. Brown 577
The initial detection of the Polyergus lucidus longicornis workers on
10 July led to an investigation of its nest in the soil beneath the leaf litter.
The nest was somewhat trapezoidal in shape, with its greatest width of
approximately 50 cm at a point farthest from the tree and its narrowest
width of approximately 39 cm wide nearest the tree. A large Pheidole
dentata Mayr nest was adjacent to the Polyergus nest, and the line of
separation between them was not well defined. The Polyergus nest extended
into the soil at least 20 cm deep to a level where the roots of the
tree prevented further digging. The nest contained hundreds of Polyergus
and Formica dolosa workers in approximately equal numbers. After the
nest was excavated, the Polyergus and Formica ran erratically on the
ground near the base of the tree. A large number of phorid flies, which
were not collected nor identified, were observed flying around the ants
when the nest was disturbed, but the phorids appeared to be flying at P.
dentata major workers rather than the Polyergus or Formica workers.
Winged queens of the Polyergus were found in the nest, and a chamber
containing cocoons of F. dolosa was found about 15 to 20 cm deep.
Berlese funnel extraction of ants from one gallon of the litter and soil
mixture of the Polyergus and Pheidole nest included three additional nesting
species: Paratrechina vividula (Nylander), Strumigenys louisianae Roger,
and Solenopsis molesta. Other species found in the same soil sample included
Brachymyrmex depilis, Paratrechina faisonensis (Forel),
Camponotus pennsylvanicus (DeGeer), Hypoponera opacior (Forel),
Ponera pennsylvanica Buckley, Pyramica angulata (Smith), P. ornata
(Mayr), Aphaenogaster carolinensis Wheeler, A. fulva Roger, Pheidole
dentigula Smith, and Crematogaster lineolata (Say).
Ants found within one meter of the Polyergus nest included Solenopsis
invicta x richteri, Pheidole bicarinata Mayr, P. tysoni Forel, and Myrmecina
americana Emery, all on the ground; Camponotus pennsylvanicus on the
tree trunk; and Paratrechina vividula, C. snellingi Bolton, Monomorium
minimum (Buckley), and Solenopsis molesta under bark of a decayed portion
of the tree.
A second visit to the site on 15 July revealed that the nest of Polyergus
and Formica had been abandoned and reoccupied by Aphaenogaster fulva.
The only other collections made at the site that day were at cookie bait,
which was placed on six white cards evenly spaced around the tree one meter
from its base. The bait was observed for approximately 30 minutes, and
collections were made of Camponotus pennsylvanicus, Pheidole bicarinata,
Crematogaster lineolata, and Solenopsis invicta x richteri, all of which were
at separate bait stations.
The imported fire ant hybrid, Solenopsis invicta x richteri, was the most
numerous species that came to tuna bait on 21 October. Paratrechina
faisonensis, Prenolepis imparis (Say), and Aphaenogaster fulva also were
collected at baits, but each of the baited cards attracted only a single species.
578 Southeastern Naturalist Vol. 5, No. 4
On 11 November, Prenolepis imparis, Solenopsis invicta x richteri,
Aphaenogaster lamellidens Mayr, Pheidole bicarinata, P. dentata, P.
tysoni Forel, and Temnothorax pergandei Emery were collected while they
foraged on the ground. Pheidole tysoni was the only ground-foraging species
whose nest was found and also the only species collected at the tuna
bait. Prenolepis imparis was the most abundant species seen on the ground
as well as at the cookie bait. The nest of Camponotus snellingi was found
in a low-hanging, dead branch of the tree. A large colony of Aphaenogaster
lamellidens was collected under bark in a dead section of the tree approximately
1.6 meters above ground. Species extracted from five gallons of
soil/litter in Berlese funnels included Paratrechina faisonensis,
Hypoponera opacior, Ponera pennsylvanica, Pyramica dietrichi (Smith),
P. ornata, P. pulchella (Emery), Strumigenys louisianae, Solenopsis
invicta x richteri, S. molesta, Aphaenogaster carolinensis, A. fulva, A.
lamellidens, A. mariae Forel, Pheidole dentigula, P. tysoni, Temnothorax
curvispinosus Mayr, and Myrmecina americana. The only two A. mariae
specimens collected from the soil samples were dealate queens. This rare
species has been speculated to be a temporary parasite of A. fulva (Smith
1979), which was also collected in the soil. The imported fire ant hybrid,
Solenopsis invicta x richteri, was reduced in numbers compared to previous
visits, and only two specimens were collected. Twenty-two total
species were collected on this visit, and species collected for the first time
included Pyramica dietrichi, P. pulchella, Aphaenogaster lamellidens, A.
mariae, Temnothorax curvispinosus, and T. pergandei.
Twenty-three species were collected on 26 June, either on the tree or in
the soil at its base. Paratrechina arenivaga (Wheeler), P. faisonensis,
Hypoponera opacior, Trachymyrmex septentrionalis (McCook),
Solenopsis invicta x richteri, Aphaenogaster fulva, A. lamellidens, and
Pheidole dentigula were all found nesting at the base of the tree. Ponera
pennsylvanica, Proceratium silaceum Roger, Strumigenys louisianae,
Solenopsis molesta, Pheidole tysoni, and Myrmecina americana were
found in a sample of soil and litter taken at the base of the tree and were
extracted from a Berlese funnel. Formica dolosa was collected crawling on
the tree trunk. Camponotus nearcticus Emery, C. pennsylvanicus, C.
snellingi, Formica pallidefulva (Latreille), Monomorium minimum,
Solenopsis invicta x richteri, and Aphaenogaster lamellidens were collected
at cookie bait that had been placed at various points on the tree at a
height between 1.5 and 2 meters. Formica pallidefulva, Monomorium minimum,
Aphaenogaster mariae, Pheidole dentata, and Temnothorax
schaumii Roger were collected at peanut butter bait in the same area of the
tree trunk. Workers of A. mariae were observed climbing into the upper
reaches of the tree, presumably toward their nest. A nest of F. pallidefulva
was located in the soil in the open parking area approximately 5 meters
from the tree. Hybrid fire ants were more common than on any previous
2006 J.A. MacGown and R.L. Brown 579
visit, and their nests with simple holes for entrances also were found
beneath the soil in the parking area. Much of the parking area was covered
with leaf litter, and when leaves were moved away, the fire ants were seen
foraging in great abundance. Six of the 23 species collected on 26 June
were new for the site and included Paratrechina arenivaga, Camponotus
nearcticus, Formica pallidefulva, Proceratium silaceum, Trachymyrmex
septentrionalis, and Temnothorax schaumii.
Discussion
Much anecdotal information is available from myrmecologists to support
the finding that large number of species can be found at a
microspatial scale. The largest number of ant species reported from a
single point sample was 43 species in 21 genera obtained by fogging a
single tree in Amazonia (Wilson 1987). This point sample included about
32% of the total diversity of species collected by fogging canopies of
trees in four forest types in 14 sites during two months. However,
Wilson’s study dealt primarily with arboricolous species, rather than species
that reside in soil or litter. A study documenting soil and leaf-litter
dwelling ants in a Malaysian rain forest (Agosti et al 1994) reported
finding 37 species of ants in an 8-m2 plot and 104 species in a 20-m2 plot.
The Malaysian study differed from the present study in that the size of
the areas sampled was considerably larger (although still relatively
small). A study of ant communities in Pinus palustris Miller (longleaf
pine) flatwoods in northern Florida (Lubertazzi and Tschinkel 2003)
yielded 72 total species of ants, with a high of 55 species reported from
one of the 70-x 80-m plots. Similar to the results found in these studies,
J.A. MacGown has collected up to 40 species of ants in relatively small
areas (less than 100 m2) within single habitats in Mississippi on numerous
occasions. The 23 species collected in the present study at the base and
periphery of the oak tree during approximately one hour on both 10 July
and 26 June represent 32% of the 72 species reported from Tombigbee
National Forest (MacGown and Brown, in press). For the five sampling
dates at the same site, the total of 36 species collected on the lower two
meters of the tree trunk and within one meter of its base represents 50%
of the species richness documented in Tombigbee National Forest during
a five-year period. This is the highest number of ant species to have been
reported from this small of an area in North America.
The question can be raised as to how a community of so many ant
species can exist in such a small area, especially if the introduced fire ant
adversely affects species richness as previously reported. Hölldobler and
Wilson (1990) considered interspecific competition to be the most obvious
interaction affecting community organization. Andersen (2000)
emphasized environmental stress and disturbance as determinants of ant
580 Southeastern Naturalist Vol. 5, No. 4
community organization in Australia, and he defined seven functional
groups in relation to stress and disturbance. In the three-tiered classification
of dominance by Vepsäläinen and Pisarski (1982), the lowest level of
dominance includes species that defend only their nests, and these appear
to be the same as the “opportunistic” species found at food baits (Wilson
1971). The intermediate level includes species that defend their nests and
food finds. The highest level of dominance includes species that defend
their nests and all their foraging area, which is the same as the dominant
Dolichoderinae functional group recognized by Andersen (2000). Various
authors have shown that ant communities can be influenced by several
factors of niche differentiation, including feeding habits, habitat preferences,
temporal partitioning, and other aspects (Brown 1975, Hölldobler
and Wilson 1990, Torres 1984).
The species composition at the Tombigbee National Forest site was
possibly affected by the act of collecting at the site, in particular by the
disturbance of the Polyergus nest. However, several of the species not
collected on the first date (10 July) were collected on subsequent trips
and were likely also present on 10 July. Collections on this date were
made in the rain, no baits were used near or on the tree, and only one soillitter
sample was taken from the Polyergus nest at the base of the tree.
Camponotus nearcticus, A. mariae, T. schaumii (all arboreal species), F.
pallidefulva (a ground-nesting species), and A. lamellidens (later found
nesting in the tree) were collected at baits on the tree at later dates. The
cryptic soil species, Proceratium silaceum, Pyramica pulchella, and
Temnothorax curvispinosus, were collected later in litter and soil samples
from different locations around the base of the tree than where the first
litter sample was taken. Paratrechina arenivaga was found nesting in the
soil near the tree on a later date, but that immediate area was not searched
on the first collection date. Only two species were not likely there on 10
July: Prenolepis imparis, a cool-weather species, and Trachymyrmex
septentrionalis, whose distinctive nest was not seen until after the
Polyergus vacated their nest. There is no reason to doubt that all four
species of Aphaenogaster could have coexisted, because they were all
collected on 11 November. Likewise, all three species of Camponotus
were collected together on 26 June, as were both species of Formica.
The study area was a disturbed site with an oak tree in a gravel
parking area beside a cemetery surrounded by relatively undisturbed forest,
but the diversity of ants was quite high. Typically, diversity of ant
species is low in disturbed habitats (Andersen 2000), but in this situation,
the large tree with its rich soil and humus layer at its base may have acted
as an island for ant species that were more dispersed in the adjacent
forest. This does not appear to be an isolated situation as J.A. MacGown
has observed many similar instances in a variety of habitats, where isolated
large trees in disturbed areas that were near undisturbed forests had
2006 J.A. MacGown and R.L. Brown 581
high numbers of ant species. In addition, the openness of the immediate
area provided nesting habitat for species not usually found in the forested
areas, which added to the total number of species. Another significant
factor was that no dominant species of Dolichoderinae, such as Forelius
spp. or Linepithema humile (Mayr), was present at the study site although
these species are widespread throughout Mississippi. The introduced fire
ants Solenopsis invicta, S. richteri, and their hybrid are often dominant
species in open areas, but they do not appear to exhibit the same level of
dominance in forested areas.
Another potential explanation for the high diversity of ant species
collected at this oak tree may be differences in diets and foraging behavior.
Many of the species at this oak tree were opportunistic species that are
generalists in their diets, including Brachymyrmex depilis, Paratrechina
spp., Prenolepis imparis, Camponotus spp., Formica spp., Monomorium
minimum, Solenopsis spp., Aphaenogaster spp., Pheidole spp.,
Crematogaster lineolata, and Temnothorax spp. Many of these species
were attracted to baits, in agreement with observations of Wilson (1971),
whereas other species of ants were much more specialized in their diet.
For example, Trachymyrmex septentrionalis grow fungus in vegetative
matter and insect feces, whereas other species are specialized predators.
Proceratium silaceum and Myrmecina americana are specialized predators
on spider eggs and mites, respectively (Brown 1958, 2000). Pyramica
spp. and Strumigenys louisianae are predators of various species of
Collembola, and it has been speculated that the spongiform bodies on
these dacetine ants may be a lure for their prey (Bolton 2000). These
dacetine ants also have highly modified elongate mandibles that vary
greatly in their length, dentition, and setation. Species with longer mandibles
are typically more mobile and hunt by stealth, whereas those with
shorter mandibles simply lie in wait for their prey (Hölldolber and Wilson
1990). Due to the unique structural modifications of mandibles and
spongiform bodies that differentiate dacetine species from one another, it
is unlikely that their prey selection is the same.
Opportunistic species appear to have a variety of methods to deal with
competition including speed, size, aggressiveness, and the ability to forage
far from their colonies. The large Camponotus species are very fast moving
ants that can forage (both diurnally and nocturnally) many meters from
their colonies (Hansen and Klotz 2005). At the study site, both
Camponotus spp. and Formica spp. were observed moving rapidly about
baits, but never taking over a bait with large numbers of individuals. This
behavior was contrary to the actions of other opportunistic species that
monopolized the bait after it was discovered. Two of the arboreal species,
A. mariae and T. schaumii, exhibited different behavior from the other
generalists by moving slowly and cryptically along the bark of the tree near
the baits. Smaller sized opportunists, e.g., B. depilis and S. molesta, may
582 Southeastern Naturalist Vol. 5, No. 4
avoid competition by their diminutive size and need for less food than
larger species. Some species, such as S. invicta x richteri, are much more
aggressive than others and out-compete other species at baits. However,
when observing ants at baits on the ground at this study site, it appeared to
be “first come, first serve,” with species that got to the bait first maintaining
control of the bait. The rules of engagement at baits on the ground did
not seem to apply to the baits placed on the tree, and it was not unusual to
see several species together on the bait on the tree including C. nearcticus,
C. snellingi, C. pennsylvanica, and F. pallidefulva at cookie bait and F.
pallidefulva, A. mariae, and T. schaumii at peanut butter bait.
Another factor allowing for many species to live in a small area is
habitat partitioning of related species in different microhabitats within
the same area. This is not a major issue for minute species or those with
small colony size, but it is more important for larger species that occupy
more space. In the case of colonies of Paratrechina spp., P. faisonensis
was nesting in the rich litter layer at the base of the tree, P. vividula was
found nesting under bark of the tree, and P. arenivaga was nesting in the
open and somewhat sandy soil. Camponotus nearcticus, whose nests were
not found, is known to nest in twigs and branches in trees (Smith 1965),
and C. snellingi nests were found in a dead branch and under bark in a
rotting section of the tree about one meter high. No confirmed colony of
C. pennsylvanicus was found, but many individuals were collected from
the lower area of the tree in a somewhat hollowed section with rotting
wood present. Workers of Formica pallidefulva were collected crawling
on the tree, but the only nest that was found was located approximately
10 meters from the tree in an open area in the ground. Solenopsis invicta
x richteri nests were found scattered around the entire open area under a
light layer of leaf litter, although no visible mounds were seen.
Solenopsis molesta was only found nesting at the base of the tree in the
soil and leaf litter. Four species of Aphaenogaster were collected at
the site, three of which were nesting in slightly different areas. Although
the nest of Aphaenogaster mariae was not found, workers were observed
carrying bait up high into the tree where they were apparently nesting. A.
lamellidens was nesting under bark and in the rotting wood of the tree, A.
fulva was nesting in the soil and packed litter at the base of the tree with
the nest extending into the roots of the tree, and A. carolinensis was
nesting in the soil itself. Four species of Pheidole were found nesting at
the site including: Pheidole dentata, which had a large colony at the base
of the tree; P. dentigula, which was nesting in the rich litter layer at the
base of the tree; and P. bicarinata and P. tysoni, both of which had
colonies in open soil surrounding the tree. Three species of Temnothorax
were collected: T. curvispinosus, T. pergandei, and T. schaumii; however,
no colonies were found. Temnothorax schaumii workers were only collected
at bait on the bark of the tree. This species is known to nest under
2006 J.A. MacGown and R.L. Brown 583
and in the bark of trees, and it is assumed that they were nesting somewhere
in the tree. Temnothorax curvispinosus colonies have been
collected elsewhere in the National Forest (MacGown and Brown, in
press) nesting in twigs or nuts of Carya glabra (P. Mill.) Sweet
(Juglandaceae), and T. pergandei has been found in the National Forest
nesting in soil and in nuts of C. glabra.
Seasonal differences in species composition and relative abundance were
evident among the various species during the sampling period. Prenolepis
imparis, which is known to be more common in the cooler months of the
year, was first seen in small numbers on 21 October and then in large
numbers on 11 November. In contrast, the hybrid fire ant, S. invicta x
richteri, which is most active during warmer months, was the most common
ant seen on 21 October, but only one individual was seen on 11 November.
Acknowledgments
This research was funded by Mississippi Agricultural and Forestry Experiment
Station, Project MIS-311080, the US Forest Service (Tombigbee National Forest),
and USDA-ARS Areawide Management of Imported Fire Ant Project (Richard L.
Brown, Principal Investigator).
Literature Cited
Agosti, D., M. Mohamed, and C.Y.C. Arthur. 1994. Has the diversity of tropical ant
fauna been underestimated? An indication from leaf litter studies in a west
Malaysian lowland rain forest. Tropical Biodiversity 2:270–275.
Anderson, A.N. 2000. Global ecology of rainforest ants: Functional groups in relation
to environmental stress and disturbance. Pp. 25–34, In D. Agosti, J.D. Majer,
L.E. Alonso, and T.R. Schultz (Eds.). Ants: Standard Methods for Measuring and
Monitoring Biodiversity. Smithsonian Institution Press, Washington, DC. xix +
280 pp.
Baroni Urbani, C., and M.L. De Andrade. 2003. The ant genus Proceratium in the
extant and fossil record (Hymenoptera: Formicidae). Museo Regionale di
Scienze Naturali, Monografie 36:1–480.
Bolton, B. 2000. The ant tribe Dacetini. Memoirs of the American Entomological
Institute 65:1–1028.
Brown, R.L. 1975. Behavioral observations on Aethalion reticulatum (Hem.,
Aethalionidae) and associated ants. Insectes Sociaux 23:99–107.
Brown, W.L. 1958. Predation of arthropod eggs by the ant genera Proceratium and
Discothyrea. Psyche 64:115.
Brown, W.L. 2000. Diversity of ants. Pp. 45–79, In D. Agosti, J.D. Majer, L.E.
Alonso, and T.R. Schultz (Eds.). Ants: Standard Methods for Measuring and
Monitoring Biodiversity. Smithsonian Institution Press, Washington, DC. xix +
280 pp.
Camilo, G.R., and S.A. Phillips, Jr. 1990. Evolution of ant communities in response
to invasion by the fire ant Solenopsis invicta. Pp. 190–198, In R.K. Vander Meer,
K. Jaffe, and A. Cedeno (Eds.). Applied Myrmecology: A World Perspective.
Westview Press, Boulder, CO.
584 Southeastern Naturalist Vol. 5, No. 4
DuBois, M., and L.R. Davis. 1998. Stenamma foveolocephalum (= S. carolinense)
rediscovered (Hymenoptera: Formicidae: Myrmicinae). Sociobiology 32:125–138.
Hansen, L.D., and J.H. Klotz. 2005. Carpenter Ants of the United States and Canada.
Cornell University Press, Ithaca, NY. xii + 204 pp.
Helms, K.R., and S.B. Vinson. 2001. Coexistence of native ants with the red imported
fire ant, Solenopsis invicta. Southwestern Naturalist 46:396–400.
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.
Hölldobler, B., and E.O. Wilson. 1990. The Ants. Belknap Press of Harvard University
Press, Cambridge, MA. 732 pp.
Holway, D.A., L. Lach, A.V. Suarez, N.D. Tsutsui, and T.J. Case. 2002. The causes
and consequences of ant invasions. Annual Review of Ecology and Systematics
33:181–233.
Jusino-Altresino, R., and S.A. Phillips, Jr. 1994. Impact of red imported fire ants
on the ant fauna of central Texas. Pp. 259–268, In D.F. Williams (Ed.). Exotic
Ants: Biology, Impact, and Control of Introduced Species. Westview Press,
Boulder, CO.
Lubertazzi, D., and W.R. Tschinkel. 2003. Ant-community change across a groundvegetation
gradient in north Florida’s longleaf pine flatwoods. Journal of Insect
Science 3(21):1–17.
MacGown, J.A., and R.L. Brown. In Press. Survey of ants (Hymenoptera:
Formicidae) of the Tombigbee National Forest in Mississippi. Journal of the
Kansas Entomological Society.
MacGown, J.A., R.L. Brown, and J.G. Hill. 2005. An annotated list of the Pyramica
(Hymenoptera: Formicidae: Dacetini) of Mississippi. Journal of the Kansas Entomological
Society 78:285–289.
MacKay, W.P. 1993. A review of the New World ants of the genus Dolichoderus
(Hymenoptera: Formicidae). Sociobiology 22:1–148.
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 38:126–140.
Morrison, L.W. 2002. Long-term impacts of an arthropod-community invasion by
the imported fire ant, Solenopsis invicta. Ecology 83:2337–2345.
Morrison, L.W., and S.A. Porter. 2003. Positive association between densities of the
red imported fire ant, Solenopsis invicta (Hymenoptera: Formicidae), and generalized
ant and arthropod diversity. Environmental Entomology 32:548–554.
Porter, S.D., and D.A. Savignano. 1990. Invasion of polygyne fire ants decimates
native ants and disrupts arthropod community. Ecology 71:2095–2106.
Shoemaker, D.D., K.G. Ross, and M.L. Arnold. 1994. Development of RAPD
markers in two introduced fire ants, Solenopsis invicta and S. richteri, and their
application to the study of a hybrid zone. Molecular Biology 3:531–539.
Smith, D.R. 1979. Superfamily Formicoidea. Pp. 1323–1467, In K.V. Krombein,
P.D. Hurd, Jr., D.R. Smith, and B.D. Burks (Eds.). Catalog of Hymenoptera in
America North of Mexico, Vol. 2: Apocrita (Aculeata). Smithsonian Institution
Press, Washington, DC. xvi + 1199–2209 pp.
Smith, M.R. 1924a. An annotated list of the ants of Mississippi (Hym.). Entomological
News 35:47–54.
2006 J.A. MacGown and R.L. Brown 585
Smith, M.R. 1924b. An annotated list of the ants of Mississippi (Hym.). Entomological
News 35:77–85.
Smith, M.R. 1924c. An annotated list of the ants of Mississippi (Hym.). Entomological
News 35:121–127.
Smith, M.R. 1927. An additional annotated list of the ants of Mississippi with a
description of a new species of Pheidole (Hym.: Formicidae). Entomological
News 38:308–314.
Smith, M.R. 1928a. An additional annotated list of the ants of Mississippi with a
description of a new species of Aphaenogaster (Hym.: Formicidae). Entomological
News 39:242–246.
Smith, M. R. 1928b. An additional annotated list of the ants of Mississippi with a
description of a new species of Aphaenogaster (Hym.: Formicidae). Entomological
News 39:275–279.
Smith, M.R. 1931a. An additional annotated list of the ants of Mississippi (Hym.:
Formicidae). Entomological News 42:16–24.
Smith, M.R. 1931b. A revision of the genus Strumigenys of America, north of
Mexico, based on a study of the workers. Annals of the Entomological Society of
America 24:384–387.
Smith, M.R. 1932. An additional annotated list of the ants of Mississippi (Hym.:
Formicidae). Entomological News 42:157–160.
Smith, M.R. 1965. House-infesting ants of the eastern United States: Their recognition,
biology, and economic importance. United States Department of Agriculture
Technical Bulletin No. 1326: ii + 1–105.
Snelling, R.R. 1995. Systematics of Nearctic ants of the genus Dorymyrmex (Hymenoptera:
Formicidae). Contributions in Science (Los Angeles) 454:1–14.
Torres, J.A. 1984. Niches and coexistence of ant communities in Puerto Rico:
Repeated patterns. Biotropica 16:284–295.
Trager, J.C. 1984. A revision of the genus Paratrechina (Hymenoptera: Formicidae)
of the continental United States. Sociobiology 9:49–162.
Trager, J.C. 1988. A revision of the Conomyrma (Hymenoptera: Fomicidae) from the
southeastern United States, especially Florida, with keys to the species. Florida
Entomologist 71:11–29.
Trager, J.C., J.A. MacGown, and M.D. Trager. In Press. Revision of the Nearctic
endemic Formica pallidefulva group (Hymenoptera: Formicidae: Formicinae).
Memoirs of the American Entomological Institute.
Tschinkel, W.R. 1988. Distribution of the fire ants Solenopsis invicta and S.
geminata (Hymenoptera: Formicidae) in northern Florida in relation to habitat
and disturbance. Annals of the Entomological Society of America 81:76–81.
Umphrey, G.J. 1996. Morphometric discrimination among sibling species in the
fulva-rudis-texana complex of the ant genus Aphaenogaster (Hymenoptera:
Formicidae). Canadian Journal of Zoology 74:528–559.
Vepsäläinen, K., and B. Pisarski. 1982. The taxonomy of the Formica rufa group:
Chaos before order. Pp. 27–35, In P.E. Howse and J.-L. Clément (Eds.). Biosystematics
of Social Insects, Systematics Association Special Volume No. 19.
Academic Press, NY.
Ward, P.S. 1985. The Nearctic species of the genus Pseudomyrmex (Hymenoptera:
Formicidae). Quaestiones Entomologicae 21:209–246.
586 Southeastern Naturalist Vol. 5, No. 4
Watkins, J.F. 1985. The identification and distribution of the army ants of the United
States of America (Hymenoptera, Formicidae, Ecitoninae). Journal of the Kansas
Entomological Society 58:479–502.
Wilson, E.O. 1971. The Insect Societies. Belknap Press of Harvard University Press,
Cambridge, MA. x + 548 pp.
Wilson, E.O. 1987. The arboreal ant fauna of Peruvian forests: A first assessment.
Biotropica 19:245–251.
Wilson, E.O. 2003. Pheidole in the New World: A Dominant, Hyperdiverse Ant
Genus. Harvard University Press, Cambridge, MA. 794 pp.