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Worldwide spread of the tropical fire ant, Solenopsis geminata (Hymenoptera: Formicidae)


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The tropical fire ant, Solenopsis geminata (FABRICIUS, 1804), is a well-known pest in many parts of the world, where it is notorious for its potent sting. To evaluate the worldwide spread of S. geminata, I compiled and mapped published and unpublished specimen records from > 2100 sites. I documented the earliest known S. geminata records for 122 geo-graphic areas (countries, island groups, major Caribbean islands, US states, and Canadian provinces), including several areas for which I found no previously published records, e.g., Anguilla, Barbuda, Curaçao, Madagascar, Montserrat, Nebraska, Nevis, St Martin, and Vanuatu. Several New World Solenopsis species were once considered junior synonyms of S. geminata (e.g., S. gayi (SPINOLA, 1851), S. saevissima (SMITH, 1855), S. virulens (SMITH, 1858), and S. xyloni MCCOOK, 1880). Therefore, I did not map unconfirmed New World S. geminata records published before CREIGHTON's (1930) re-vision of Solenopsis from areas where these species occur, because some of these early records were likely to be based on misidentifications. Solenopsis geminata records are common through much of the New World tropics as well as parts of subtropical North America. Whereas S. geminata is certainly native to South and Central America, it may well be exotic to the southeastern US and the West Indies, introduced several hundred years ago. By 1900, S. geminata had also spread through many parts of the Old World, notably tropical Asia and Oceania.
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Myrmecological News 14 21-35 Vienna, January 2011
Worldwide spread of the tropical fire ant, Solenopsis geminata (Hymenoptera:
The tropical fire ant, Solenopsis geminata (FABRICIUS, 1804), is a well-known pest in many parts of the world, where it
is notorious for its potent sting. To evaluate the worldwide spread of S. geminata, I compiled and mapped published and
unpublished specimen records from > 2100 sites. I documented the earliest known S. geminata records for 122 geo-
graphic areas (countries, island groups, major Caribbean islands, US states, and Canadian provinces), including several
areas for which I found no previously published records, e.g., Anguilla, Barbuda, Curaçao, Madagascar, Montserrat,
Nebraska, Nevis, St Martin, and Vanuatu. Several New World Solenopsis species were once considered junior synonyms of
S. geminata (e.g., S. gayi (SPINOLA, 1851), S. saevissima (SMITH, 1855), S. virulens (SMITH, 1858), and S. xyloni MCCOOK,
1880). Therefore, I did not map unconfirmed New World S. geminata records published before CREIGHTON's (1930) re-
vision of Solenopsis from areas where these species occur, because some of these early records were likely to be based
on misidentifications.
Solenopsis geminata records are common through much of the New World tropics as well as parts of subtropical North
America. Whereas S. geminata is certainly native to South and Central America, it may well be exotic to the southeastern
US and the West Indies, introduced several hundred years ago. By 1900, S. geminata had also spread through many parts
of the Old World, notably tropical Asia and Oceania.
In the mid-20th century, another invader from the Neotropics, the "unvanquished" fire ant, Solenopsis invicta BUREN, 1974,
began to spread around the world. Solenopsis invicta has displaced S. geminata in open and disturbed habitats in many
parts of the southern US, leaving only remnant S. geminata populations, primarily in forested areas that S. invicta does
not invade. Although S. geminata still has a much broader worldwide range, I expect S. invicta will continue to spread
and displace S. geminata in open habitats through many other parts of the tropics and subtropics.
Key words: Biogeography, biological invasion, exotic species, invasive species, stinging ant.
Myrmecol. News 14: 21-35 (online 24 June 2010)
ISSN 1994-4136 (print), ISSN 1997-3500 (online)
Received 14 January 2010; revision received 23 March 2010; accepted 29 March 2010
Prof. Dr. James K. Wetterer, Wilkes Honors College, Florida Atlantic University, 5353 Parkside Drive, Jupiter, FL 33458,
USA. E-mail:
The tropical fire ant, Solenopsis geminata (FABRICIUS, 1804)
has long been one of the world's most dreaded ants due to
its powerful sting, "which it uses on the slightest provoca-
tion" (WHEELER 1910). Solenopsis geminata is originally
from the New World, where "its colonies are populous and
so numerous that it may be said to be in possession of a
large portion of the soil of the American tropics" (WHEE-
LER 1910). Like many other "tramp" ants (e.g., Paratrechina
longicornis (LATREILLE, 1802) and Tapinoma melanoce-
phalum (FABRICIUS, 1793); see WETTERER 2008, 2009),
S. geminata has spread through human commerce to many
other parts of the world.
Reports on S. geminata are replete with superlatives.
WHEELER (1914) wrote that the behavior of S. geminata
"exhibits a more bewildering diversity than that of any other
known ant, for it is not only highly carnivorous, but attends
scale insects and leaf hoppers for the sake of their saccha-
rine excrement and even collects and stores seeds in its nests
ike a true harvesting ant. In Texas I have known it to sting
young chickens to death when they happened to be con-
fined in a coop near its nest. In the same State it is help-
ful in destroying the boll weevil and other insects injurious
to cotton, but is sometimes a pest in gardens when it takes
to gnawing holes in strawberries." WHEELER (1910) wrote
that S. geminata "attacks and eats almost everything that
comes its way," and SMITH (1965) reported that S. gemi-
nata "is considered to be one of the most important pre-
dators of all the ants." Solenopsis geminata may be found
in a wider range of habitats than any other fire ant species
(TSCHINKEL 1988, 2006). For example, in Surinam, KEMPF
(1961) reported S. geminata from gardens, fields, pas-
tures, woodland, and primary forest. In addition, WHEE-
LER (1922) called S. geminata "perhaps the most dreaded
of the house ants."
In recent years, a new invader from the Neotropics, the
"unvanquished" fire ant, Solenopsis invicta BUREN, 1974,
has begun to spread around the world, displacing S. gemi-
nata both in the field and in the popular press. Nonetheless,
Figs. 1 - 4: Solenopsis geminata. (1) Head of minor worker from the Philippines; (2) lateral view of the same minor
worker; (3) head of major worker from the Philippines; (4) lateral view of the same major worker (photos by G. Alpert).
S. geminata remains a dominant global pest. In the pre-
sent study, I examine the worldwide spread of S. geminata.
Identification and taxonomy: Solenopsis geminata is
a highly polymorphic species, with a wide range of wor-
ker size within a colony (head width = 0.55 - 2.30 mm;
Figs. 1 - 4). Solenopsis geminata shows considerable vari-
ation in coloration. As a result of this variability, com-
bined with some poor taxonomic work, S. geminata has
been described repeatedly under many different names,
now designated as junior synonyms. As CREIGHTON (1930)
wrote: "S. geminata, the type of the genus, has been de-
scribed a surprising number of times. There are no less
than eleven names for this species which have been aban-
doned on the synonymic junk-heap." For example, in just
one paper, BUCKLEY (1867), a geologist untrained in ento-
mology, described S. geminata under three or more differ-
ent names. WHEELER (1902) wrote that BUCKLEY'S (1867)
"descriptions are, indeed, fearfully and wonderfully made.
With a persistency, which at times seems almost inten-
tional, the author selects for description the worthless, in-
significant features of the ant's body."
Encouragingly, CREIGHTON (1930) and TRAGER (1991)
made detailed taxonomic re-evaluations of S. geminata
nd its close relatives and their analyses largely agree on
the definition of S. geminata. CREIGHTON (1930) wrote: "It
is inevitable that such a widely distributed and abundant
ant should repeatedly appear in the literature under new
names and it is remarkable that, despite this plethora of de-
scriptions, the taxonomy of geminata has largely escaped
confusion. At various times related species have been re-
garded as variants of geminata but, during the past sixty
years there has been little doubt as to what constitutes the
typical form. Truth compels one to add that this unusual
situation is more a result of the characteristic cephalic struc-
ture of the major worker than an outcome of taxonomic
acumen." Indeed, although S. geminata minors are very dif-
ficult to distinguish from minors of related fire ants, large S.
geminata majors have several distinctive cephalic charac-
teristics that make them simple to identify. These include:
(1) a disproportionally large, almost square head with par-
allel sides, (2) a deep longitudinal groove on the front of
the head extending from a distinct medial indentation in the
vertex, (3) black mandibles, often with all teeth worn off
from use, and (4) short antennal scapes extending only about
halfway to the occiput in the largest majors (Figs. 3 - 4).
FABRICIUS (1804) described Atta geminata (= S. gemi-
nata) based on a queen from Central America. Taxa cur-
rently considered junior synonyms of S. geminata (with type
Fig. 5: Worldwide distribution records of Solenopsis geminata. In many areas, however, S. geminata populations have
been greatly reduced or eliminated (see text).
locality in parentheses) include: Myrmica paleata LUND,
1831 (Brazil); Solenopsis mandibularis WESTWOOD, 1840
(Tropical America); Atta rufa JERDON, 1851 (India); Atta
clypeata SMITH, 1858 (Mexico); Solenopsis cephalotes
SMITH, 1859 (Aru Islands, Indonesia); Myrmica mellea
SMITH, 1859 (Aru Islands); Crematogaster laboriosus
SMITH, 1860 (Bacan, Indonesia); Myrmica laevissima
SMITH, 1860 (Bacan); Diplorhoptrum drewseni MAYR, 1861
(Italy); Myrmica glaber SMITH, 1862 (Panama); Myrmica
polita SMITH, 1862 (Panama); Atta coloradensis BUCKLEY,
1867 (Texas); Atta lincecumii BUCKLEY, 1867 (Texas);
Myrmica saxicola BUCKLEY, 1867 (Texas); Solenopsis ge-
minata diabola WHEELER, 1908 (Texas); Solenopsis gemi-
nata nigra FOREL, 1908 (Costa Rica); Solenopsis eduardi
FOREL, 1912 (Colombia); Solenopsis geminata medusa
MANN, 1916 (Brazil); Solenopsis geminata galapageia
WHEELER, 1919 (Galapagos); Solenopsis edouardi bahia-
ensis SANTSCHI, 1925 (Brazil and Venezuela); and Sole-
nopsis edouardi perversa SANTSCHI, 1925 (Brazil). In ad-
dition, there is one remaining subspecies, Solenopsis gemi-
nata micans STITZ, 1912 (Seram Island, Indonesia); taxo-
nomists need to evaluate whether this taxon deserves syno-
nymy (see BOLTON 1995 for a taxonomic history of these
Some taxa were designated junior synonyms of S. gemi-
nata, but were later revived from synonymy, including:
Solenopsis gayi (SPINOLA, 1851) (Chile; synonymized by
MAYR 1886; revived by EMERY 1895); Solenopsis saevis-
sima (SMITH, 1855) (Brazil; synonymized by MAYR 1865;
revived by WHEELER 1915); Solenopsis virulens (SMITH,
1858) (Brazil; synonymized by MAYR 1886; revived by
TRAGER 1991); and Solenopsis xyloni MCCOOK, 1880 (US;
synonymized by MAYR 1886; revived by WHEELER 1910).
GOETSCH (1933) synonymized Aphaenogaster santiagu-
ensis G
OETSCH, 1930 (Chile) with S. geminata, but I
have been unable to find this taxon in any reference except
GOETSCH (1930, 1933). Because SNELLING & HUNT (1975)
recognized no records of S. geminata from Chile and con-
sidered published S. geminata records from Chile (MAYR
1865) to be misidentifications of S. gayi, it seems very likely
that A. santiaguensis is a junior synonym of S. gayi. Al-
though WHEELER (1902) and others speculated that Atta
brazoensis BUCKLEY, 1867 (Texas) and Myrmica sabeana
BUCKLEY, 1867 (Texas) were also junior synonyms of S.
geminata, TRAGER (1991) conjectured that they were actu-
ally S. xyloni.
TRAGER (1991) and BOLTON (1995) considered Sole-
nopsis geminata innota SANTSCHI, 1915 (described from
Gabon, Liberia, and Zaire) to be a junior synonym of S. ge-
minata, but other researchers disagree with this designa-
tion. COLLINGWOOD & VAN HARTEN (1993) reported S.
innota from Cape Verde Islands off the coast of West Af-
rica and stated "the enlarged petiole and postpetiole link it
with S. globularia." In 2003, I collected ants in Cape Verde
and found no Solenopsis geminata. Instead, at 78 sites on
all nine inhabited islands, I collected a Solenopsis cf. glo-
bularia, a species with an enlarged petiole and post-petiole,
apparently corresponding to S. innota (J.K. Wetterer &
X. Espadaler, unpubl.). TAYLOR (2007) reported both S.
geminata and S. innota from Cameroon. TAYLOR (2007)
noted that the S. innota major had numerous characters dis-
tinguishing it from the S. geminata major, including sev-
eral pronounced teeth on the mandible and longer scapes
"almost attaining the occiput." I therefore consider S. in-
nota to be a distinct taxon from S. geminata. One possibi-
lity is that some S. innota records are actually Solenopsis
virulens (SMITH, 1858). This species, known from Brazil,
Guyana, and Peru, is "yellow, monomorphic, and about the
size of small majors of S. saevissima or S. geminata, but the
postpetiole is subglobose and very large" (KEMPF & BROWN
A third fire ant taxon has been reported from Africa as
well: Solenopsis saevissima itinerans (FOREL, 1911), de-
scribed from Tanzania and later reported from Guinea
(SANTSCHI 1914). FOREL (1911) described this taxon as
"very similar to var. Richteri FOREL, but entirely brown.
The head is rather narrow, the nodes are also narrow and
the hair is a little less abundant." It seems possible that these
ants could be S. saevissima, S. xyloni, or S. invicta.
Some taxa have changed their taxonomic status relative
to S. geminata many times. For example, FOREL (1904) de-
scribed Solenopsis geminata pylades FOREL, 1904 (Mex-
ico), but FOREL (1909) raised S. pylades to a full species.
WHEELER (1915) considered it a junior synonym of S. saevis-
sima, but FOREL (1916) again raised S. pylades to a full spe-
cies and WHEELER (1919c) again considered it a junior syno-
nym of S. saevissima. EMERY (1922) designated S. pylades
a junior synonym of S. geminata, but SANTSCHI (1923) con-
sidered it a variety of S. saevissima. TRAGER (1991) de-
signated S. pylades a junior synonym of S. xyloni, but PITTS
(2002) made S. pylades a junior synonym of S. saevissima.
Previous accounts of the New World and worldwide
distribution of S. geminata: There are many previous ac-
counts describing the geographic range of S. geminata (e.g.,
1979, TRAGER 1991, TABER 2000, TSCHINKEL 2006). These
accounts, however, vary considerably and are based on data
of unknown detail and questionable accuracy.
FOREL (1899) wrote that S. geminata "is the most com-
mon species of tropical America, where it is not lacking, I
think, on one square kilometer of lowland from Virginia
south to the Argentine Republic." In 1899, however, S. ge-
minata was considered the senior synonym of several other
species, including S. xyloni in North America and S. sae-
vissima and S. virulens in South America.
Using a revised, narrower concept of S. geminata,
CREIGHTON (1930) wrote that its distribution "may be sum-
marized as uniform throughout the West Indies and on the
Continent from Florida to Costa Rica. In the eastern Gulf
States its range extends inland only about a hundred to a
hundred and fifty miles [160 - 240 km] and this also ap-
pears to be true for the greater part of Texas. From Mexico
southward, however, the distribution is from coast to coast."
CREIGHTON (1930) wrote that one color variant of S. ge-
minata "extends northward from Peru through Ecuador and
western Colombia into Central America and thence through
Mexico to the Southern United States. That of the second
group begins in Northeastern Colombia and passes through
Venezuela, the Guianas and into eastern Brazil apparently
terminating about the latitude of Bahia... Except for their
greater variation in color and sculpture the members of the
first group agree closely in structure with the insular [West
Indian] geminata."
CREIGHTON (1950) wrote of S. geminata: "the main
range of this insect lies in Central America and the An-
tilles. In the United States it occurs from Texas to South
Carolina. The majority of these records come from areas on
or near the coast. As one goes inland the incidence usually
decreases except in Florida, where the insect seems to be
uniformly distributed over the entire state."
SMITH (1951) wrote that S. geminata ranged "Fla. to
Tex.; W. Indies, Mexico, Cent. Amer. and Pacific Coastal
Plain through Columbia, Ecuador, and Peru," and S. ge-
minata rufa ranged: "Fla. to Panama; also in Oriental and
Australian Regions." SMITH (1965) wrote that S. geminata
ranged "from Texas to South Carolina and Florida and
south to at least Costa Rica. It also occurs in the West In-
dies." SMITH (1979) wrote that S. geminata ranged "S.C. s.
to Fla., w. to Tex. and s. to Peru; West Indies; tropical Asia
and Pacific Islands."
The Commonwealth Agricultural Bureaux (COMMON-
map of the worldwide distribution of S. geminata. In the
New World, S. geminata was shown ranging across the
southern US from southern Virginia to southern California,
through all of Mexico and Central America. In South Ame-
rica, the range of S. geminata was shown extending down
the west coast to the Tropic of Capricorn in Chile, and down
the east coast to São Paulo, except for a gap from south-
ernmost Bahia to Rio de Janeiro. In the Old World, the map
had S. geminata records scattered in Africa, Asia, Austra-
lia, and the Pacific. HILL (1974, 2008) reprinted this map
TRAGER (1991) wrote that S. geminata range included
the "coastal plain of the Carolinas and Georgia, Florida
west to Texas, Central America, Antilles," and northern
South America, "including the coastal areas of Northeast-
ern Brazil, west through the Guianas to the Orinoco Basin,
the western Amazon Basin and coastal areas of Peru." TRA-
GER (1991) wrote that in the Old World "S. geminata has
been introduced into both tropical Asia and Africa. The
first of these populations is now distributed from Taiwan
and India in the North, throughout the Malay archipelago
and Polynesia in the south, but the population is highly
uniform throughout this vast range, with the light reddish
coloration, relatively weak sculpture, and well-developed
mesoplural process typical of the form rufa, and may re-
sult from the successful spread of a single original intro-
duction." TRAGER (1991) wrote "the only population which
might, in my opinion, be a distinct species among the mat-
erial called S. geminata here is the western South Ameri-
can population of Colombia and Peru. This form averages
smaller in all castes than S. geminata from elsewhere, and
its sting is reputed to be more painful and to cause a pus-
tule as do stings of the S. saevissima complex. The western
population is apparently the source of the rather small S.
geminata typical of the Galapagos Islands."
TABER (2000) included a map of the worldwide distri-
bution of S. geminata. In the New World, the map showed
a range that roughly followed the accounts of CREIGHTON
(1930) and TRAGER (1991) with a few deviations: (1) great-
er inland penetration in the southeastern US (~ 400 km)
with a continuous distribution from southern North Caro-
lina across to Texas, (2) greater extension down the west
coast of South America beyond Peru, extending ~ 500 km
into Chile, (3) absence in western Mexico north of Aca-
pulco, and (4) absence in the western Amazon basin. TA-
BER (2000) also mapped scattered records of S. geminata in
the Old World in rough agreement with TRAGER's (1991)
general report, with most records spread between India
and New Guinea, including large population records in
Bangladesh, Malaysia, and New Guinea.
Most recently, TSCHINKEL (2006) included a map of the
New World range of S. geminata that was similar to the
range shown in TABER (2000), but with (1) only ~ 300 km
inland penetration in the southeastern US from southern
South Carolina to Texas, (2) only ~ 250 km extension into
Chile, and (3) less extension into western Mexico.
Because few actual specimen records accompany all
previously published reports on the range of S. geminata,
it is unclear what is based on actual data and what is sup-
position. Here, I document in detail the known worldwide
distribution of S. geminata based on specimen records.
Using published and unpublished records, I documented
the worldwide range of S. geminata. I obtained unpublished
site records from museum specimens in the collections of
the Smithsonian Institution (SI; identified by M.R. Smith),
Archbold Biological Station (ABS; identified by M. Dey-
rup), the British Natural History Museum (BMNH; iden-
tified by B. Bolton), the Museum of Comparative Zool-
ogy (MCZ; identified by S. Cover), and the Northern Ter-
ritory Economic Insect Reference Collection (NTEIRC; pro-
vided by D. Chin). In addition, I used on-line databases
with collection information on specimens by Antweb (www., the Chinese Ant Database (
english.html), the Essig Museum (, the
Global Biodiversity Information Facility (,
and the Nebraska State Insect Records (
I also received unpublished records from H. Axen (Belize,
Brazil, Costa Rica, Dominican Republic, Florida, French
Guiana, Galapagos, Guatemala, Honduras, India, Madagas-
car, Mascarene Islands, Mexico, Nicaragua, Texas, Trini-
dad, Turks & Caicos Islands, and Venezuela), M. DaSilva
(Grenadine islands), B. Hoffmann (Australia), J. Lattke (Ve-
nezuela), J. MacGown (Georgia, Mississippi), P.D. Rajan
(India), and S. Sonthichai (Thailand). Finally, I collected S.
geminata specimens in Central America, Florida, and on
numerous Pacific and Caribbean Islands.
Geographic coordinates for collection sites came from
published references, specimen labels, maps, or geography
web sites (e.g.,,, and For older references and specimens, many
site names, particularly in Asia, are no longer in use or are
now spelled differently and I searched, not always success-
fully, to determine current names. If a site record listed a
geographic region rather than a "point locale," and I had
no other record for this region, I used the coordinates of the
largest town within the region or, in the case of small is-
lands and natural areas, the center of the region. Often, if
one source had many sites less than 10 - 20 km apart (e.g.,
STARR & al. 2007), I did not plot every site. I did not map
records of S. geminata on boats, found in newly imported
goods, or intercepted in transit by quarantine inspectors.
Published records usually included collection dates. In a
number of cases, publications did not include the collection
dates for specimens, but I was able to determine the date
based on information on the collector's travel dates or limit
the date by the collector's date of death. For example,
MAYR (1865) published S. geminata records collected by
the Novara expedition, which visited Sri Lanka in 1858 and
Tahiti in 1859. MAYR (1862) identified many specimens as
S. saevissima that MAYR (1865) re-identified as S. geminata.
Through much of its worldwide range, S. geminata is
the only fire ant known and thus records are likely to be re-
liable. In the New World, this includes all of Central Am-
erica and (before 1981, when S. invicta was first recorded)
the West Indies. In parts of the North and South America,
other fire ants occur, including S. amblychila WHEELER,
1915, S. aurea WHEELER, 1906, and S. xyloni in the south-
ern US and northern Mexico, and S. gayi, S. invicta, S.
saevissima, and S. virulens in South America. Due to mis-
identifications and transitory synonymies, some specimen
records from these areas reported as S. geminata may ac-
tually be other Solenopsis species and vice versa. In addi-
tion, worker hybrids between S. geminata and S. xyloni
(HUNG & VINSON 1977) further complicate taxonomy. To
minimize the risk of identification errors, I did not map pub-
lished S. geminata records predating CREIGHTON's (1930)
revision from New World regions where other fire ant spe-
cies occur, unless the records were confirmed in a post-
1930 publication.
I compiled published and unpublished specimen records
from > 2100 sites worldwide (Fig. 5). I documented the
earliest known S. geminata records for 122 geographic ar-
eas (countries, island groups, major Caribbean islands, and
US states; Tabs. 1 - 6), including several areas for which I
found no previously published records: Anguilla, Barbuda,
Curaçao, Madagascar, Montserrat, Nebraska, Nevis, St Mar-
tin, and Vanuatu.
Unconfirmed New World records that I did not map in-
cluded all published S. geminata records from Chile (e.g.,
MAYR 1862, 1865, BERG 1890, IHERING 1894), Paraguay
(e.g., FOREL 1908), California (e.g., MAYR 1886, WHEELER
1908, ANDREWS 1916), and Virginia (e.g., FOREL 1899), as
well as a subset of records from other areas, including Ar-
gentina (e.g., BAUR 1897, BRUCH 1914), Arizona (e.g.,
WHEELER 1908), Bolivia (e.g., EMERY 1894), Brazil (e.g.,
MAYR 1865, WASMANN 1890, 1896, IHERING 1894, FOREL
1907, 1908, MANN 1916), Mexico (e.g., ANDRÉ 1893, PER-
GANDE 1893, FOREL 1899, WHEELER 1901), New Mexico
(e.g., TINSLEY 1898, CREIGHTON 1930), and North Caro-
lina (e.g., WITHERS 1898, FOREL 1899). Although WHEE-
LER (1908) described S. geminata diabola (= S. geminata)
from Texas, with additional records from Arizona, and Cali-
fornia, I did not map the records from Arizona and Cali-
fornia. KEMPF (1972) has the only post-1930 record of S.
geminata from Bolivia, but with no site listed. Rather than
mapping this record to the largest city, La Paz, in the cool
highlands, I mapped this to Cobija in the northern lowlands,
placing it nearest to records just across the border in Peru
and Brazil. Although CREIGHTON (1930), SMITH (1951),
and TRAGER (1991) all mention S. geminata in coastal Ec-
(1958), TABER (2000), and TSCHINKEL (2006) all mapped
populations there, I found no site records for this region.
I mapped no point in this region because I suspect that
earlier authors may have simply inferred that S. geminata
populations in coastal Colombia and coastal Peru were con-
tinuous through Ecuador.
I omitted CREIGHTON'S (1930) New Mexico record be-
cause CREIGHTON (1950) omitted it. I mapped only one post-
1930 S. geminata record from New Mexico and one from
Arizona. COLE (1953) reported "one colony of what I be-
lieve represents this species was nesting beneath a stone in
very dry, level, stony soil 13 mi. N of Carlsbad, 3,300 ft,"
though this high elevation desert record seems very likely
to be a misidentification. PLOWES & al. (2009) reported
phorid flies that B. Brown collected 5 km E Portal, Ari-
zona, attacking S. geminata; unfortunately, Brown (pers.
comm.) could not locate any voucher ant specimens to ver-
ify this record.
In the case of several South American countries and US
states (Tabs. 1 - 2), the earliest record was unconfirmed and
Tab. 1: Earliest known records for Solenopsis geminata
from South and Central America and neighboring Pacific
islands. Unpublished records include collector, museum
source, and site. BMNH = Natural History Museum in Lon-
don. * = earliest record unconfirmed and may be misiden-
tification of another Solenopsis species.
Earliest record
Central America 1804 (FABRICIUS 1804 – type locality)
Brazil 1831 (LUND 1831 as M. paleata)
Venezuela 1857 (H. Clark, BMNH): Tejuca
Mexico 1858 (SMITH 1858 as A. clypeata)
Panama 1861 (SMITH 1862 as M. glaber & M.
Guyana 1862 (ROGER 1862*)
Uruguay 1862 (ROGER 1862*)
Surinam 1863 (ROGER 1863*)
French Guiana 1868 (RADOSZKOWSKY 1884*)
Nicaragua 1872 (T. Belt, BMNH): Chontales
Galapagos 1891 (EMERY 1893)
Bolivia 1894 (EMERY 1894*)
Cocos Islands 1898 (EMERY 1919)
Guatemala 1899 (Champion, BMNH): Guatemala
Belize 1899 (FOREL 1900)
Colombia 1900 (FOREL 1900*)
Costa Rica 1900 (FOREL 1900)
Honduras 1920 (MANN 1922)
Ecuador 1930 (CREIGHTON 1930)
Peru 1930 (CREIGHTON 1930)
Islas Revillagigedo 1935 (WHEELER 1935)
El Salvador 1957 (BERRY & SALAZAR 1957)
not mapped, but many later records verified the presence
of S. geminata. For example, MAYR (1886) published the
earliest S. geminata records from Alabama, Florida, and
Louisiana, and though these records were not confirmed,
later records verified the presence of S. geminata in these
states (e.g., CREIGHTON 1930).
For the Old World, I mapped all published records of
S. geminata. However, I did not map African records ori-
ginally identified as S. innota because these records ap-
pear to represent a distinct taxon: Cameroon (TAYLOR
2007), Cape Verde (COLLINGWOOD & VAN HARTEN 1993),
Congo (Zaire) (SANTSCHI 1915), Gabon (SANTSCHI 1915),
and Liberia (SANTSCHI 1913).
Tab. 2: Earliest known records for Solenopsis geminata
from the USA and Canada. Abbreviations as in Tab. 1, and
MCZ = Museum of Comparative Zoology. + = no known
published record.
Earliest record
+ Texas 1862 (ROGER 1862*)
+ Alabama 1886 (MAYR 1886*)
+ Florida 1886 (MAYR 1886*)
+ Louisiana 1886 (MAYR 1886*)
+ New Mexico 1897 (TINSLEY 1898*)
+ North Carolina 1899 (FOREL 1899*)
+ South Carolina 1899 (FOREL 1899*)
+ Georgia 1899 (FOREL 1899*)
+ Mississippi 1899 (FOREL 1899*)
+ Arkansas 1964 (WHITCOMB & BELL 1964 in
+ Manitoba 1977 (AYRE 1977)
+ Nebraska 1992 (D Ferraro, NIR): Omaha Zoo –
Lied Jungle
Only 12 S. geminata site records came from latitudes
at or above 35° (Fig. 5; five New World and seven Old
World). The high latitude New World records are prob-
ably either introduced or misidentifications: the Tropical
House of Assiniboine Park Zoo in Winnipeg, Canada (49.9°
N; AYRE 1977), the Lied Jungle (billed as the "World's
Largest Indoor Rainforest") at the Omaha Zoo (41.2° N;
Tab. 2), Maquinchao, Argentina (~ 41° S; DONISTHORPE
1933), Sherman County, Texas (~ 36.3° N; O'KEEFE &
al. 2000), and Uruguay (~ 35° S; BRANDÃO 1991). The high
latitude Old World records are certainly all introduced:
Netherlands (~ 52.3° N; BOER & VIERBERGEN 2008), Kew
Gardens greenhouses, England (51.5° N; DONISTHORPE
1943), Italy (~ 42° N; MAYR 1861), Beijing, China (39.9°
N; WHEELER 1927a), Zante Island, Greece (37.7° N; COL-
LINGWOOD 1993), Mount Maunganui, New Zealand (37.6°
N; HARRIS 2005), and Cyprus (~ 35.2° N; COLLING-
WOOD & al. 1997). Most or all of these high latitude popu-
lations of S. geminata have probably been eradicated or
have died out.
Worldwide distribution: In the New World, Solenopsis
geminata has been reported from all South and Central
American countries, the southern US from California to
Virginia, and every island group in the West Indies. How-
ever, a number of these records were identified when then
the definition of S. geminata included several other species.
Thus, a number of unconfirmed records may be based on
misidentification of these related species. Such likely mis-
identifications include all published S. geminata records
from areas beyond the northern and southern ends of its
confirmed range, including California, Virginia, and prob-
Tab. 3: Earliest known records for Solenopsis geminata
from the West Indies. Abbreviations as in Tabs. 1 and 2.
Earliest record
+ Cuba 1862 (ROGER 1862)
+ Puerto Rico 1863 (ROGER 1863)
+ Virgin Islands 1878 (FOREL 1881)
+ St Lucia 1889 (G.A. Ramge, BMNH): site un-
+ St Vincent 1892 (FOREL 1893)
+ Jamaica 1893 (ANDRÉ 1893)
+ Barbados 1896 (FOREL 1912)
+ Grenada 1897 (FOREL 1897)
+ Bahamas 1904 (WHEELER 1905)
+ Haiti 1907 (FOREL 1907)
+ Guadeloupe 1911 (WHEELER 1913)
+ Dominica 1911 (WHEELER 1913)
+ Martinique 1911 (WHEELER 1913)
+ Trinidad 1915 (URICH 1915)
+ Tobago 1918 (WHEELER 1919a)
+ Antigua 1920 (WHEELER 1923)
+ Dominican Re-
+ public
1930 (MENOZZI & RUSSO 1930)
+ Bonaire 1936 (WEBER 1948)
+ Aruba 1936 (WEBER 1948)
+ St Kitts 1937 (WEBER 1948)
+ Statia 1937 (WEBER 1948)
+ Montserrat 1991 (J.P.E.C. Darlington, MCZ):
Paradise Yard
+ Cayman Islands 2004 (VARNHAM 2006)
+ Turks & Caicos 2004 (VARNHAM 2006)
+ Curacao 2004 (J.K. Wetterer, MCZ): De Savaan
+ Anguilla 2006 (J.K. Wetterer, MCZ): Meads Bay
+ St Martin 2006 (J.K. Wetterer, MCZ): Pic Paradis
+ Barbuda 2007 (J.K. Wetterer, MCZ): Codrington
+ Nevis 2007 (J.K. Wetterer, MCZ): Cane Garden
ably Arizona and New Mexico in the north and Chile, Pa-
raguay, and probably Argentina and Uruguay in the south.
My specimen record map for S. geminata (Fig. 5) in
the New World differs from the range maps of COMMON-
Tab. 4: Earliest known records for Solenopsis geminata from
Asia and neighboring islands. Abbreviation as in Tab. 1.
Earliest record
India 1851 (JERDON 1851 as S. geminata rufa)
Sri Lanka 1858 (MAYR 1865)
Indonesia 1859 (SMITH 1859 as S. cephalotes &
M. mellea)
Philippines 1862 (MAYR 1862 misidentified as S.
Burma/Myanmar 1885 - 1887 (EMERY 1889)
Singapore 1897 (MAYR 1897)
1915 (CRAWLEY 1915 as S. geminata rufa)
Malaysia 1919 (WHEELER 1919b as S. geminata rufa)
Laos 1920 (SANTSCHI 1920 as S. geminata rufa)
China 1927 (WHEELER 1927a as S. geminata rufa)
Vietnam 1927 (WHEELER 1927b as S. geminata rufa)
Hong Kong 1928 (WHEELER 1928)
Macao 1928 (WHEELER 1928)
Thailand 1928 (Hillman, BMNH): Bangkok
Papua New
1935 (WHEELER 1935 as S. geminata rufa)
Japan 1967 (KIRITANI & MORIMOTO 2004)
Taiwan ~ 1985 (CHEN & al. 2005)
UAE 1995 (COLLINGWOOD & al. 1997)
Hainan Island 1998 (FELLOWES & HAU 2001)
Brunei 1999 - 2000 (EGUCHI & YAMANE 2003)
Cocos (Keeling)
2005 (NEVILLE & al. 2008)
and TSCHINKEL (2006) in many respects. The COMMON-
to approximate the combined range of S. geminata, S. xy-
loni, S. saevissima, and S. gayi, no doubt due to misidenti-
fications in the southwestern US and Mexico in the north
and in Argentina, Brazil, and Chile in the south. The maps
of TABER (2000) and TSCHINKEL (2006) omit published
records from western Mexico north of Acapulco, e.g.,
CREIGHTON's (1930) record from Guadalajara, Jalisco. On
the west coast of South America, I found S. geminata rec-
ords extending only into northern Peru, in agreement with
CREIGHTON (1930), SMITH (1979), and TRAGER (1991),
not down into Chile as in TABER (2000) and TSCHINKEL
(2006). Also, the TABER (2000) and TSCHINKEL (2006)
Tab. 5: Earliest known records for Solenopsis geminata
from Oceania. Abbreviations as in Tabs. 1 and 2.
Earliest record
+ Society Islands 1859 (MAYR 1865)
+ Australia 1863 (ROGER 1863)
+ New Zealand 1876 (MAYR 1876)
+ Hawaii 1879 (BLACKBURN & KIRBY 1880)
+ New Caledonia 1882 (EMERY 1883)
+ Mariana Islands 1894 (collector unknown, BMNH):
+ Cook Islands 1914 (WILSON & TAYLOR 1967)
+ Samoa 1916 (H. Swale, BMNH): Apia
+ Tuamotu Islands 1925 (CHEESMAN & CRAWLEY 1928)
+ Austral Islands 1934 (WILSON & TAYLOR 1967)
+ Line Islands 1934 (WHEELER 1936)
+ Palau 1936 (CLOUSE 2007)
+ FSM 1950 (CLOUSE 2007)
+ Marshall Islands 1953 (CLOUSE 2007)
+ Tonga 1956 (WILSON & TAYLOR 1967)
+ Banaba 1957 (CLOUSE 2007)
+ Solomon Islands 1962 (GREENSLADE 1969)
+ Tuvalu 1976 (P. Maddison, BMNH): Funafuti
+ Gambier Islands 1996 (MORRISON 1997)
+ Marquesas
+ Islands
1996 (MORRISON 1997)
+ Fiji 1997 (WATERHOUSE 1997)
+ Gilbert Islands 1999 (CLOUSE 2007)
+ Vanuatu 1999 (E. Tabi, MCZ): Sama
maps did not include records from Peru and Ecuador east of
the Andes (e.g., PARDO VARGAS 1964, ESCALANTE-GUTI-
ÉRREZ 1975, TOBIN 1995, PLOWES & al. 2009). In eastern
South America, I found coastal records extended farther
south than indicated by TABER (2000) and TSCHINKEL
(2006), with many records not only throughout Bahia (e.g.,
REIS & SANTOS 2001, PITTS 2002), but also farther south,
in Espiritu Santo (PITTS 2002), Minas Gerais (FOWLER & al.
1995, NASCIMENTO 2005), and São Paulo (H. Axen, pers.
comm.). TABER (2000) included a record from Rio de Ja-
neiro, and though there are published records from here
(MAYR 1865) and further south (IHERING 1894, FOREL
1908), I did not map these unconfirmed records because
they are likely to be based on misidentifications.
Tab. 6: Earliest known records for Solenopsis geminata
from Europe, Africa, and neighboring islands. Abbrevia-
tion as in Tab. 1.
+ Europe Earliest record
+ Italy 1861 (MAYR 1861 as D. drewseni)
+ England 1932 (DONISTHORPE 1943)
+ Greece 1982 - 1988 (COLLINGWOOD 1993)
+ Cyprus 1997 (COLLINGWOOD & al. 1997)
+ Netherlands 1992 (BOER & VIERBERGEN 2008)
+ Africa
+ Senegal 1862 (ROGER 1862)
+ Mascarene
+ Islands
in MOUTIA & MAMET 1946)
+ Congo Republic) 1909 (SANTSCHI 1909)
+ Sierra Leone 1915 (BACOT 1916)
+ Guinea 1935 (SANTSCHI 1939 as S. geminata
+ Equatorial Guinea 1939 - 1940 (MENOZZI 1942)
+ Nigeria 1969 (BOLTON 1973)
+ Madagascar 1981 (J.M. Wilson, BMNH): Ambilobe
+ South Africa 1990 (PRINS & al. 1990)
+ Canary Islands 1997 (HÔGMO 2003)
+ Cameroon 2001 (TAYLOR 2007)
In the Old World, S. geminata is widespread through
tropical and subtropical Asia, Australia, and Oceania. The
documented range of S. geminata in Africa is much more
limited, and many S. geminata records from Africa appear
to be a different species. Finally, there are a few indoor rec-
ords from temperate Europe and North America. My new
S. geminata distribution map (Fig. 5) includes many Old
World records not found on TABER's (2000) map, e.g.,
England, Greece, Cyprus, Italy, Nigeria, Cameroon, Repub-
lic of Congo, Zaire, Vietnam, Burma, New Caledonia,
Tuvalu, Solomon Islands, Fiji, Caroline Islands, Marshall
Islands, and New Zealand. Many S. geminata records from
Australia and New Zealand came from port areas and from
plant nurseries, e.g., the Illparpa Nursery in Alice Springs
(2005; NTEIRC). Control efforts in Australia and New
Zealand have successfully exterminated numerous isolated
local populations of S. geminata (B. Hoffmann, pers. comm.).
In Australia, S. geminata populations are widespread only
in the northern "Top End" region of Northern Territory.
Native versus exotic New World range: Solenopsis
geminata is originally from the New World tropics and
subtropics. However, the extent of the native range of S.
geminata in the New World remains unclear. Solenopsis
geminata is almost certainly native to South America, Cen-
tral America, and Mexico, and most authors consider S.
geminata as native to the southeastern US (e.g., WOJCIK
& al. 1976, JOUVENAZ & al. 1977, HÖLLDOBLER & WILSON
1990, TSCHINKEL 2006). SMITH (1930) listed S. geminata
as an exotic in Florida, but later SMITH (1965) considered
S. geminata to be a native species in the eastern US. TRA-
GER (1991) wrote: "S. geminata is apparently native to the
southeast coastal plain and Florida to Texas ... south through
Central America to Northern South America ... Popula-
tions of the Antilles and Galápagos (and possibly the South-
eastern U.S.A.) are probably introduced, but have been in
these areas for several centuries." DEYRUP & al. (2000)
proposed that S. geminata in Florida might be a mix of
native and exotic populations. In fact, this may be true in
many other parts of the world as well.
In the southeastern US, populations of S. geminata
appear to have more parasites than do populations of the
exotic Solenopsis invicta, supporting the hypothesis that S.
geminata is native to this region. For example, JOUVENAZ
& al. (1977) surveyed Solenopsis parasites in the southe-
astern US, and found a 140 times higher rate of micro-
sporidia infection in S. geminata colonies than in S. in-
victa colonies. In northern Florida, MCINNES & TSCHINKEL
(1996) found mermithid nematode infection rate of 16%
among S. geminata queens and 0% among S. invicta queens.
In Florida, however, no parasitoid phorid flies naturally at-
tack S. geminata, in contrast to Texas where 6 - 8 species
of phorids attack S. geminata (S. Porter, pers. comm.), sug-
gesting that S. geminata is not native to Florida.
Some authors have considered S. geminata as exotic to
the West Indies. WHEELER (1908) reported S. geminata as
"transported to the Antilles." TRAGER (1991) considered
the West Indian populations as "probably introduced." New-
ly introduced species may show population explosions, so
population explosions of S. geminata in the West Indies
could indicate that this species is exotic to this region. WHEE-
LER (1926) proposed that Formica omnivora LINNAEUS,
1758, a voracious ant that besieged several tropical West
Indian islands in the 16th to 18th centuries, was most prob-
ably S. geminata. In summarizing early reports on F. om-
nivora in the West Indies, MORLEY (1953) was less cir-
cumspect concerning the identity of this pest: "In 1760 it
was seriously debated whether the island of Barbados should
be abandoned, because of the sudden and disastrous ap-
pearance of a plague of "Fire Ants," Solenopsis geminata.
Their large aphids and scale insect herds did such damage
to the sugar-cane crops that the island, formerly a flourish-
ing colony, was driven to the verge of bankruptcy. The
same ants appeared in Martinique in 1763, the year after its
capture from the French by the British Admiral Rodney.
Seven years later the island of Grenada, 170 miles to the
south, suffered a similar fate." Population explosions of this
ant, however, may have been related not to recent intro-
duction, but instead to habitat changes associated with hu-
man settlement or to the introduction of a new mutualist
aphid, scale insect, or other Hemiptera.
On-going research on the genetic diversity of S. gemi-
nata at sites around the world should help to resolve ques-
tions concerning the native range and invasion history of
this species. For example, preliminary genetic results sug-
gest that S. geminata populations in the southeastern US
are exotic (AXEN 2008, H.J. Axen, pers. comm.). LONGINO
(2005) noted that in Costa Rica, S. geminata has "a "red
form" that is more abundant in open areas and a "black
form" that prefers forested areas."
Community dominance: In many parts of its range, S.
geminata is a dominant species in both natural and dis-
turbed ecosystems, tending plant-feeding Hemiptera and
acting as an important predator and seed-disperser (e.g.,
JEANNE 1979). For example, in Mexico, RISCH & CARROLL
(1986) considered S. geminata to be a keystone species,
because its presence or absence had a tremendous effect
on both the arthropod community through predation and
on the plant community through seed dispersal (RISCH &
CARROLL (1986) found that S. geminata in Mexico har-
vested small seeds and when common, S. geminata could
reduce the abundance of many weedy plants, particularly
grasses. In northeastern Mexico, FLORES-MALDONADO &
al. (1999) found that S. geminata dominated disturbed oak
and pine forests, making up the vast majority of ants col-
lected in these habitats (91.7% and 96.2%, respectively).
In Honduras, WYCKHUYS & O'NEIL (2007) found that S.
geminata made up more than 50% of ants collected in sub-
sistence maize fields.
Solenopsis geminata is also known to attack the hatch-
lings of birds & reptiles (e.g., STODDARD 1931, TRAVIS
1938, 1941, KROLL & al. 1973, MRAZEK 1974, MAES &
MACKAY 1993). WETTERER (2006) found that S. geminata
was the most common ant on an important sea turtle nesting
beach in Tortuguero, Costa Rica, where earlier research-
ers documented ants preying on sea turtle eggs and hatch-
lings. Solenopsis geminata was the only species present
that was likely to pose a significant threat to sea turtles.
Because of the very high numbers of S. geminata on Tor-
tuguero beach, the impact of these ants on hatchling sea
turtles could be substantial, particularly on hatchlings stung
as they exit their nests.
In agricultural areas, S. geminata may be a serious pest.
For example, on Java and Mauritius, ORDELHEIDE (1929)
and D'EMMEREZ DE CHARMOY (1930) reported S. gemi-
nata as a serious pest harvesting planted tobacco seeds.
CHERIAN (1933) reported S. geminata destroying seedlings
and burrowing into the roots of marijuana plants. STAHL &
SCARAMUZZA (1929) reported S. geminata tending mealy-
bugs on sugar.
In some cases, S. geminata is considered beneficial.
For example, in Sri Lanka, ROTHNEY (1889) reported that
S. geminata were deliberately introduced into warehouses
to control termites. In Malaysia, SIVAPRAGASAM & CHUA
(1997) found S. geminata was the major predator on cab-
bage pests.
Competition with other ants: Solenopsis geminata of-
ten shows a mutually exclusive distribution with other do-
minant ant species. For example, WHEELER (1910) reported
this pattern with S. geminata and the African big-headed
ant, Pheidole megacephala on two Caribbean islands: "I
devoted ten days to a careful study of the ant-fauna of the
little island of Culebra off the eastern coast of Porto Rico
without seeing a single specimen of Ph. megacephala. This
island is, however, completely overrun with a dark vari-
ety of the vicious fire-ant (Solenopsis geminata). One day,
on visiting the island of Culebrita, which is separated by
a shallow channel hardly a mile in width from the eastern
coast of Culebra, I was astonished to find it completely
overrun with Ph. megacephala. This ant was nesting un-
der every stone and log, from the shifting sand of the sea-
beach to the walls of the light-house on the highest point
of the island. The most careful search failed to reveal the
presence of any other species, though the flora and physical
conditions are the same as those of Culebra. It is highly
probable that Ph. megacephala, perhaps accidentally intro-
duced from the island of St. Thomas a few miles to the
east, had exterminated all the other ants which must have
previously inhabited Culebrita. The absence of megacepha-
la on Culebra is perhaps explained by the presence of the
equally prolific and pugnacious fire-ant" (WHEELER 1910).
WETTERER & O'HARA (2002) found a similar pattern on
the Dry Tortugas, the outermost Florida Keys. On Garden
Key, S. geminata was the dominant ant, occurring at 70%
of bait stations, usually with several hundred workers on
each bait card. Many visitors on Garden Key reported be-
ing stung on the feet by these ants. Although P. mega-
cephala may have occurred at one time on Garden Key,
we found none. In contrast, we found no S. geminata on any
other key in the Dry Tortugas. Instead, on Loggerhead
Key and Bush Key, P. megacephala was the dominant ant.
In Laguna del Tigre National Park, BESTELMEYER &
al. (2000) collected S. geminata in all surveyed areas, but
"in the grassland transect, Solenopsis geminata was ex-
tremely dominant." In the grasslands, Pheidole sp. was the
most common ant at tuna baits 10 and 30 min after place-
ment. From 50 min on, however, S. geminata was the in-
creasingly dominant species present and by 130 min, Phei-
dole sp. was completely displaced and no longer found at
the baits.
Solenopsis invicta versus Solenopsis geminata: In
the 20th century, another Solenopsis fire ant has begun to
spread around the world: S. invicta. This species has pro-
ven to be an even greater pest than S. geminata. For exam-
ple, TSCHINKEL (2006) noted that the stings of S. invicta
"have more severe and longer-term effects than those of
S. geminata." Originally from South America, S. invicta
arrived in Alabama by ship sometime before 1945 (BUREN
& al. 1974). Since then, S. invicta has spread across the
US from Texas to Virginia in the southeast and California
in the west, causing widespread ecological and economic
damage (WOJCIK & al. 2001). This dreaded ant is now
spreading through the West Indies (WETTERER & DAVIS
2010) and has recently arrived in Asia (e.g., CHEN & al.
2005, ZHANG & al. 2007).
When it invades, S. invicta often displaces S. geminata,
particularly in habitats preferred by both species: open, gras-
sy areas. As a result, S. geminata has largely disappeared
from much of its former range in the southeastern US (WOJ-
CIK & al. 1976). For example, TSCHINKEL (2006) wrote that
S. geminata "before S. invicta spread into Florida, was one
of the most common ants there. It tolerated a wider range
of habitats... including some types of woodland not pre-
ferred by S. invicta. As S. invicta spread, the two species
occurred side by side for a while, but within a few years, S.
geminata could be found only as enclaves in more wooded
locations." In Florida, PORTER (1992) found S. geminata
at 83% of roadside sites where S. invicta were absent, but
only 7% of sites where S. invicta was present. Many cli-
mate models predict that S. invicta should be able to in-
vade many additional regions in both the New World and
Old World now occupied by S. geminata (e.g., see KOR-
ZUKHIN & al. 2001, MORRISON & al. 2004, SUTHERST &
MAYWALD 2005). It seems likely that Solenopsis gemi-
nata is also excluded from many parts of South America
by S. invicta and other resident fire ants.
With S. invicta receiving so much media attention, it
appears that some recent reports in the popular press of S.
invicta from parts of Asia (e.g., Malaysia, Singapore, Phil-
ippines) may actually be based on misidentifications of the
long-resident S. geminata. In working to mitigate the nega-
tive impacts of S. geminata, S. invicta, and other fire ants
around the world, it will be important to distinguish between
these ants, and recognize both their similarities and their
I thank M. Wetterer, J. Trager, S. Porter, B. Hoffmann, and
H. Axen for comments on this manuscript; S. Cover for
help, encouragement, and ant identification; B. Bolton
(BMNH), S. Cover (MCZ), M. Deyrup (ABS), and T.
Schultz (SI) for help with their respective ant collections;
H. Axen, D. Chin, M. DaSilva, B. Hoffmann, J. Lattke, J.
MacGown, P.D. Rajan, and S. Sonthichai for supplying un-
published records; W. O'Brien for GIS help; D.P. Wojcik
and S.D. Porter for compiling their FORMIS bibliogra-
phy; C. Scheid and R. Pasos of the FAU library for pro-
cessing so many interlibrary loans; Florida Atlantic Univer-
sity, the National Science Foundation (DEB-0515648), and
the National Geographic Society for financial support.
Die tropische Feuerameise, Solenopsis geminata (FABRI-
CIUS, 1804), ist eine in vielen Teilen der Welt als Lästling
und Schädling bekannte Art und wird wegen ihres schmerz-
haften Stichs gefürchtet. Um die weltweite Verbreitung von
S. geminata abzuschätzen, habe ich veröffentlichte und un-
veröffentlichte Nachweise von > 2100 Fundorten zusam-
mengetragen. Ich dokumentiere die frühesten bekannten
Nachweise der Art für 122 geographische Gebiete (Län-
der, Inselgruppen, große Karibische Inseln, US-Bundes-
staaten und kanadische Provinzen), einschließlich einiger,
für die ich keine bisher veröffentlichten Nachweise gefun-
den habe, beispielsweise Anguilla, Barbuda, Curaçao, Ma-
dagaskar, Montserrat, Nebraska, Nevis, St Martin und Va-
nuatu. Mehrere Solenopsis-Arten der Neuen Welt wurden
früher als Juniorsynonyme von S. geminata betrachtet, z.B.
S. gayi (SPINOLA, 1851), S. saevissima (SMITH, 1855), S.
virulens (SMITH, 1858) und S. xyloni MCCOOK, 1880. Des-
halb habe ich unbestätigte Meldungen von S. geminata von
vor der Gattungsrevision von CREIGHTON (1930) für Regi-
onen der Neuen Welt, in denen jene Arten vorkommen,
nicht berücksichtigt – einige dieser frühen Meldungen ba-
sieren wahrscheinlich auf Fehlbestimmungen.
Zahlreiche Nachweise von Solenopsis geminata liegen
für weite Teile der Tropen der Neuen Welt sowie auch für
Teile des subtropischen Nordamerika vor. Die Art ist sicher-
lich in Süd- und Mittelamerika heimisch, aber könnte in den
südöstlichen USA und den Westindischen Inseln exotisch
sein, wobei die Einschleppung mehrere Jahrhunderte zu-
rückliegen könnte. Bereits 1900 war die Art auch in vielen
Teilen der Alten Welt verbreitet, einschließlich des tropi-
schen Asiens und Ozeaniens.
In der Mitte des Zwanzigsten Jahrhunderts begann eine
andere invasive Art aus den Neotropen, die "unbesiegte"
Feuerameise, Solenopsis invicta BUREN, 1974, ihre glo-
bale Ausbreitung. Solenopsis invicta hat S. geminata in of-
fenen und gestörten Habitaten in großen Teilen der südli-
chen USA ersetzt; Restpopulationen von S. geminata konn-
ten vor allem in bewaldeten Gebieten überleben, die S.
invicta nicht besiedelt. Obwohl S. geminata derzeit noch
eine weitaus größere globale Verbreitung hat, erwarte ich,
dass S. invicta seine Ausbreitung fortsetzen und S. gemina-
ta in offenen Habitaten vieler Teile der Tropen und Sub-
tropen ersetzen wird.
ANDRÉ, E. 1893: Description de quatre espèces nouvelles de four-
mis d'Amérique. – Revue d'Entomologie Caen 12: 148-152.
ANDREWS, H. 1916: Ants caught on a trip to California (Hym.).
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... Anoplolepis gracilipes (Smith, F., 1857) Yes Yes (Gillespie and Reimer 1993, Hill et al. 2003, Hoffmann and Saul 2010, Plentovich et al. 2018 (Kennedy 1998, Gómez and Oliveras 2003, TOUYAMA et al. 2003, Carpintero et al. 2005, Menke et al. 2018, Devenish et al. 2021) Myrmica rubra (Linnaeus, 1758) Yes Yes (Naumann and Higgins 2015) Pheidole megacephala (Fabricius, 1793) Yes Yes (Hoffmann et al. 1999, Hoffmann and Parr 2008, Gaigher et al. 2013, Williams 2021 Solenopsis geminata (Fabricius, 1804) Yes Yes (Risch and Carroll 1982, Wetterer 2011, Wauters et al. 2014, Erickson and Baccaro 2016, Rojas and Fragoso 2021 Solenopsis invicta Buren, 1972 Yes Yes (Porter and Savignano 1990, Allen et al. 2001, Castellanos et al. 2016, Allen et al. 2017, Wang et al. 2019, Wong et al. 2020 Solenopsis papuana Emery, 1900 Yes Yes (Gillespie andReimer 1993, Krushelnycky et al. 2017, Ogura-Yamada and Krushelnycky 2020) Technomyrmex albipes (Smith, F., 1861) Yes Yes (Samways et al. 1982, Hansen and (Wetterer andKeularts 2008, Gotzek et al. 2012, Williams and Lucky 2020) Solenopsis richteri Forel, 1909 No Yes Likely to have been confused with Solenopsis invicta and S. geminata Tapinoma melanocephalum (Fabricius, 1793) No Yes Cases of ant-Hemipteran mutualisms reported with some impacts on aphidophagous predators/parasitoids but in native range mainly ( Abstract: Same as above. ...
... Anoplolepis gracilipes (Smith, F., 1857) Yes Yes (Gillespie and Reimer 1993, Hill et al. 2003, Hoffmann and Saul 2010, Plentovich et al. 2018 (Kennedy 1998, Gómez and Oliveras 2003, TOUYAMA et al. 2003, Carpintero et al. 2005, Menke et al. 2018, Devenish et al. 2021) Myrmica rubra (Linnaeus, 1758) Yes Yes (Naumann and Higgins 2015) Pheidole megacephala (Fabricius, 1793) Yes Yes (Hoffmann et al. 1999, Hoffmann and Parr 2008, Gaigher et al. 2013, Williams 2021 Solenopsis geminata (Fabricius, 1804) Yes Yes (Risch and Carroll 1982, Wetterer 2011, Wauters et al. 2014, Erickson and Baccaro 2016, Rojas and Fragoso 2021 Solenopsis invicta Buren, 1972 Yes Yes (Porter and Savignano 1990, Allen et al. 2001, Castellanos et al. 2016, Allen et al. 2017, Wang et al. 2019, Wong et al. 2020 Solenopsis papuana Emery, 1900 Yes Yes (Gillespie andReimer 1993, Krushelnycky et al. 2017, Ogura-Yamada and Krushelnycky 2020) Technomyrmex albipes (Smith, F., 1861) Yes Yes (Samways et al. 1982, Hansen and (Wetterer andKeularts 2008, Gotzek et al. 2012, Williams and Lucky 2020) Solenopsis richteri Forel, 1909 No Yes Likely to have been confused with Solenopsis invicta and S. geminata Tapinoma melanocephalum (Fabricius, 1793) No Yes Cases of ant-Hemipteran mutualisms reported with some impacts on aphidophagous predators/parasitoids but in native range mainly ( Abstract: Same as above. ...
Island systems are known to harbor disproportionate amounts of geographically restricted biodiversity, but also for experiencing high rates of species loss, ultimately representing critical systems with significant conservation values. Knowledge of the biodiversity value of insular systems remains, however, highly fragmented and incomplete for many groups of organisms, especially insects. This gap limits our understanding of their global significance for biodiversity and inhibits prioritization for future exploration and conservation efforts. Here, we developed a new database introducing current knowledge of ants on 2678 islands globally, based on nominal species, and providing information on their native or exotic status. In total, this database contains 7,010 ant species (44.6% of the known global ant fauna) that have been recorded on islands globally from 449,232 records. In addition, this database identifies 108 large islands (area > 200km2) that have received no ant sampling efforts globally. This new data set provides the most comprehensive understanding of ant diversity and composition on islands globally, opening new opportunities to address questions on a multitude of research questions and fields related to biogeography, ecology, and evolution. This data set also provides a roadmap for future exploration and conservation actions of ants on islands as well as for overall ant diversity, with updates available as new records and taxonomic updates are published. There are no copyright restrictions on this database and users should cite this data paper in publications when using the data. This article is protected by copyright. All rights reserved.
... Our results increase the complexity of S. geminata alkaloids, suggesting that S. geminata workers have indeed responded to evolutionary pressure with alkaloid diversity more complicated than the ancestral cis/trans 2-methyl-6-undecyl-piperidines found in Solenopsis female sexuals. What makes our results especially intriguing is that S. geminata occupied coastal areas of the southern USA, and with the advent of worldwide trade routes in the mid-sixteenth century, S. geminata accidentally hitchhiked to many tropical/ subtropical areas of the world (Wetterer 2011;Gotzek et al. 2015). The ecological habitats that S. geminata found itself in and survived in are extraordinarily diverse, e.g., Senegal, Canary Islands, Madagascar, Australia, Fiji, Gilbert Islands, India, Laos, and Italy (Wetterer 2011). ...
... What makes our results especially intriguing is that S. geminata occupied coastal areas of the southern USA, and with the advent of worldwide trade routes in the mid-sixteenth century, S. geminata accidentally hitchhiked to many tropical/ subtropical areas of the world (Wetterer 2011;Gotzek et al. 2015). The ecological habitats that S. geminata found itself in and survived in are extraordinarily diverse, e.g., Senegal, Canary Islands, Madagascar, Australia, Fiji, Gilbert Islands, India, Laos, and Italy (Wetterer 2011). The specific focus for this paper is on S. geminata in the USA and specifically Florida, but the results will be of interest to researchers throughout the tropical/subtropical areas of the world where S. geminata is found. ...
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South American fire ants, Solenopsis richteri and Solenopsis invicta , were accidently introduced into the southern USA in the 1900s and 1930s, respectively. The rapid spread and high population densities of S. invicta , and its potent sting, resulted in broad economic impacts and a variety of research efforts. In the 1970s, their venom alkaloids were identified as a complex blend of trans -2-methyl-6-alkyl- and alkenyl-piperidines. Solenopsis geminata is a worldwide tramp species but a native of the southern coastal regions of the USA. It was found to only produce cis - and trans -2-methyl-6-undecyl-piperidines. These alkaloids were considered the Solenopsis ancestral alkaloid profile since they were identified from female sexuals (potential queens) of all Solenopsis species in South and North America. The dramatic modification of alkaloids in Solenopsis invicta was attributed to their response to evolutionary pressure and the lack of change in S. geminata alkaloids due to no response to evolutionary pressure. Here we report the unexpected discovery of 6-undecyl-pyridine, 2-methyl-6-undecyl-pyridine and 2-methyl-6-(1)-undecenyl-pyridine as components of S. geminata worker venom, suggesting that S. geminata like its South American relatives have responded to evolutionary pressures. Our results will stimulate future research on S. geminata populations throughout the tropical/subtropical world.
... Cryptic invasions are likely more widespread than currently understood as a result of the difficulty in detecting them (Morais and Reichard 2018). A classic example of this in ants is the misidentification of the tropical fire ant, Solenopsis geminata (Fabricius) (Hymenoptera: Formicidae), as the red-imported fire ant, Solenopsis invicta Buren (Hymenoptera: Formicidae) (Wetterer 2013(Wetterer , 2011, in Malaysia and Singapore, and possibly elsewhere (e.g., India, the Philippines). The former species is native to southern North America, Central America, South America, and the Caribbean, while the latter is native to South America (distributions from AntMaps: Janicki et al. 2016). ...
... In this case, the inability to correctly identify S. geminata meant that its distribution was underestimated and the invaded range of S. invicta was overestimated. Both of these species are known for endangering native wildlife and domestic animals, for being costly to control, and for having a painful sting that can cause a variety of health issues, including secondary infection, anaphylactic shock, or even death (Wetterer 2013(Wetterer , 2011. This type of misidentification is likely in taxa with many undescribed species and multiple globetrotting species. ...
Full-text available
The ant genus Nylanderia Emery has a cosmopolitan distribution and includes 150 extant described species and subspecies, with potentially hundreds more undescribed. Global taxonomic revision has long been stalled by strong intra- and interspecific morphological variation, limited numbers of diagnostic characters, and dependence on infrequently collected male specimens for species description and identification. Taxonomy is further complicated by Nylanderia being one of the most frequently intercepted ant genera at ports of entry worldwide, and at least 15 globetrotting species have widespread and expanding ranges, making species-level diagnoses difficult. Three species complexes (‘bourbonica complex’, ‘fulva complex’, and ‘guatemalensis complex’) include globetrotting species. To elucidate the phylogenetic positions of these three complexes and delimit species boundaries within each, we used target enrichment of ultraconserved elements (UCEs) from 165 specimens representing 98 Nylanderia morphospecies worldwide. We also phased the UCEs, effectively doubling sample size and increasing population-level sampling. After recovering strong support for the monophyly of each complex, we extracted COI barcodes and SNPs from the UCE data and tested within-complex morphospecies hypotheses using three molecular delimitation methods (SODA, bPTP, and STACEY). This comparison revealed that most methods tended to over-split taxa, but results from STACEY were most consistent with our morphospecies hypotheses. Using these results, we recommend species boundaries that are conservative and most congruent across all methods. This work emphasizes the importance of integrative taxonomy for invasive species management, as globetrotting occurs independently across at least nine different lineages across Nylanderia.
... The diverse climatic and topographic conditions found in South Africa, together with anthropogenic introductions and changes to disturbance regimes, offers ample opportunity for invasive species to establish (Wilson et al., 2020). Established and Lepisiota capensis (black sugar ant) have been recorded in South Africa (Devenish et al., 2021, Sinu et al., 2017Slingsby, 2017;Wetterer, 2009Wetterer, , 2011Wetterer, , 2013, which means indigenous ant diversity is already under pressure by these invasive ants. ...
Full-text available
Ants are important ecosystem engineers and invasive ants pose serious threats to biodiversity of southern Africa. Biogeography databases are important for studying biodiversity, understanding invasion ecology and in conservation management. This study aimed to create a biogeographical database of epigeic ants within the Mogale's Gate located in Gauteng, South Africa. A species inventory of epigeic ants was created, the distributions of invasive species were identified, and the stability and homogeneity of ant communities where asses by calculating Shannon Diversity Index and Shannon Equitability Index scores. Ants were sampled using pitfall trapping along a 200 m elevation gradient and in grassland, wetland and woodland habitats in Mogale’s Gate from July to September 2022. A total of 60 species were identified, six of which were invasive. The highest species diversity was recorded at mid and high elevations and the lowest diversity was recorded in wetlands. Invasive species were present in all samples, with Pheidole megacephala and Lepisiota capensis being the most abundant. The Shannon Diversity Index scores indicated that the epigeic ant assemblage of Mogale’s Gate is partially degraded, with woodlands having degraded ant assemblages. Low Shannon Equitability Index scores for all sampled sites, especially wetlands and woodlands, indicated high species homogeneity in communities. An ant monitoring programme would greatly benefit ant ecology research and biodiversity management at Mogale’s Gate.
... These ants attack children and farmers, sometimes causing anaphylactic shock [8] . Solenopsis geminata is native of Central and South America, but has spread through human commerce to many parts of the world [9] . Like other fire ants, Solenopsis geminata has dramatic impact on many plants such as tomato, corn, sorghum and soybean particularly. ...
Full-text available
In Ndog-bong and Mboppi, soybean was planted at three replication on three different plots: one of these plots was non-infested, while the other two were infested with 124 190 and 186 285 workers of Solenopsis geminata respectively. Comparison was done between the infested plots and the non-infested one on seed germination, plant density, nodule number and yield. The data collected showed that, the four studied parameters decreased with the increasing of Solenopsis geminata foragers in the infested plots. The germinating rate of soybean seeds was 48.82%, 28.77% and 22.41% in the non-infested, slightly infested and heavily infested plots respectively. Plant density was 40.04%, 33.07% and 26.89% in the non-infested, slightly infested and heavily infested plots respectively. Nodule number was 38.36%, 32.63% and 29.01% in the non-infested, slightly infested and heavily infested plots respectively. Yield was 1.36t/ha, 0.82t/ha and 0.60t/ha in the non-infested, slightly infested and heavily infested plots respectively. In conclusion, direct feeding of Solenopsis geminata on seeds, seedlings and plant root reduced plant density and caused an important reduction of soybean yield.
... Among these invader ants, the tropical fire ant Solenopsis geminata is considered one of the most damaging and widespread introduced species [6] . Solenopsis geminata is native of Central and South America, but has spread through human commerce to many parts of the world [7] . In it introduced area, this tropical fire ant has often become one of the dominant pest decreasing biodiversity and displacing other ants and arthropods via competition behavior [8,9] . ...
Full-text available
In Cameroon, we compared recruitment and competition behavior between Solenopsis geminata and two resident ant species i.e. Paratrechina longicornis and Pheidole megacephala. For the recruitment essays, a 35 cm long and 30 cm wide plywood were placed in contact with three nest and fried fish was deposited at 15 cm from the nest. The mean number of Solenopsis geminata foragers collected on the fish was compared with that of the two resident ants. On the other hand, individual and group aggression assays were used to competition evaluation between Solenopsis geminata and the two resident ants. In individual aggression essays, one medium-sized of Solenopsis geminata worker and one medium-sized worker of each resident ant species were separately placed in a Petri dish. Aggression between the two competitor ants was recorded and a comparison was done between ant species on the attacks initiation by the competitors. In the group essay, 150 workers of each resident ant species were separately allowed to competition with 150 workers of Solenopsis geminata and a comparison was done between the two groups of competitors on the mean number of ant species found on the baits and the mean mortality. Our result showed that the mean number of Pheidole megacephala recruited on the fish (156.47±16.05) was the highest, follow by Paratrechina longicornis (140.53±9.21) and Solenopsis geminata (127.54±8.54). During individual aggression essays, Solenopsis geminata workers initiated more attacks (67.33%) than those of Paratrechina longicornis (32.67%), while Pheidole megacephala workers initiated more attacks (62.38%) than those of Solenopsis geminata (37.62%). In the group essay for the species pair, the mean number of Solenopis geminata workers on the bait (30.30±0.60) was higher than that of Paratrechina longicornis (12.16±1.10), while the mean number of Pheidole megacephala workers on the bait (22.50±1.50) was higher than that of Solenopsis geminata (18.60±2.00). Moreover, the mean mortality of Paratrechina longicornis (61.94±0.48) was higher than that of Solenopsis geminata (38.06±0.48), while the mean mortality of Solenopsis geminata (58.64±1.25) was higher than that of Pheidole megacephala (41.36±1.25). The data led us to conclude that Pheidole megacephala is more competitive than Solenopsis geminata and can then be used to his control.
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Elucidating invasion pathways of invasive species is often challenging because invasive populations frequently have low genetic diversity caused by genetic bottlenecks during introduction events. Genome-wide sequencing such as Restriction Site-Associated DNA Sequencing (RADseq) can overcome these challenges by generating thousands of genome-wide single nucleotide polymorphic (SNP) markers. The tropical fire ant, Solenopsis geminata, is a global invader with low genetic diversity in its introduced range, making RADseq one of the best available methods to investigate its population genetics. We used double digest RADseq to generate 3,834 SNPs to compare the genetic diversity of S. geminata in its introduced range to its most likely source of introduction, determined the invasion pathways among populations at an unprecedented level of detail for this species, and determined the social structure of S. geminata workers collected in 13 locations worldwide. We found that introduced S. geminata went through a strong genetic bottleneck. We also identified multiple secondary introduction events among S. geminata populations, indicating that the bridgehead effect is an important driver in the global spread of this species. We found that all colonies in the introduced range were polygyne (i.e., with more than one queen) which may increase their invasion success and potential to cause adverse effects.
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Urban pests are organisms that constantly interfere with human interests and transmit diseases causing microorganisms. Among urban pests, ants and cockroaches are categorized under household and public health insect pests. Urban pests live near or sometimes live in human habitats; therefore, pest control strategies should be susceptible. To control urban pests, environmental, human, and nontargeted animal-friendly control methods are always in high demand instead of synthetic toxic chemicals. Botanicals are better alternatives to the conventional synthetic toxic pesticides against urban pests. Clove (Syzygium aromaticum) has been used in several forms, such as clove oil derived from dried clove flower bud, eugenol, eugenyl acetate, and β-caryophyllene that are major oil constituents, and clove bud powder. Essential oils from clove, clove powder, or its constituents have repellent, toxic, and anesthetic properties against many urban pest insects. In this chapter, significant studies yet on ants and cockroaches control applying different forms of clove are reported.
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Despite a legacy of extensive deforestation, the 720 km2 city state of Singapore still harbours impressively diverse flora and fauna. Given increasing evidence of global insect declines, we urgently need to better document and protect local insect diversity. Numerous species of ants (Hymenoptera, Formicidae) have been recorded or described from Singapore since its founding in 1819. However, it has been over a century since Hugo Viehmeyer (1916) documented a total of 159 species found in the country. Here, we present an updated comprehensive checklist of all named species and subspecies of ants found in Singapore, with specimen collection data, and notes on taxonomy and ecology in the local context. We compiled the list based on museum collections material (the Zoological Reference Collection), primary literature sources, and verified records from known overseas repositories. We documented a total of 409 nominal species and subspecies, also a few notable morphospecies, from 10 subfamilies and 100 genera. These include new records for 121 species and 10 genera. Another 96 species and subspecies have types designated from Singapore; of these, 34 are currently considered as endemic. We also raised nine subspecies to species and synonymized two species, providing reasons justifying each status change: 1) Camponotus (Tanaemyrmex) carinifer stat. n., 2) Camponotus (Tanaemyrmex) tinctus nom rev., 3) Paraparatrechina malaccana stat. n., 4) Aphaenogaster simulans Forel, 1915 stat. n., 5) Myrmicaria adpressipilosa stat. n., 6) Vollenhovia minuta stat. n., 7) Vollenhovia brevicornis (Emery, 1893) = V. fridae Forel, 1913 syn. n., 8) Hypoponera javana stat. n., 9) Hypoponera singaporensis stat. n., 10) Mesoponera javana stat. n. Most species are considered native to Indomalaya, including 13 cosmopolitan tramps. Only 10 other species are presumed exotic to the region. At the time of writing, Singapore can be deemed the city with the highest recorded ant diversity in the world. Despite the sheer numbers, this list remains incomplete, with more species awaiting discovery or taxonomic resolution in future. The immense diversity of ants in Singapore is mainly threatened by continued decimation of remnant forest habitats and encroaching urban developments.
Human activity has facilitated the introduction of many exotic species via global trade. Asia-Pacific countries comprise one of the most economically and trade-active regions in the world, which makes it one of the most vulnerable regions to invasive species, including ants. There are currently over 60 exotic ant species in the Asia-Pacific, with the red imported fire ant, Solenopsis invicta, among the most destructive. Exotic ants pose many economic and ecological problems for the region. Countries in the Asia-Pacific have dealt with the problem of exotic ants in very different ways, and there has been an overall lack of preparedness. To improve the management of risks associated with invasive ants, we recommend that countries take action across the biosecurity spectrum, spanning prevention, containment, and quarantine. The creation of an Asia-Pacific network for management of invasive ants should help prevent their introduction and mitigate their impacts. Expected final online publication date for the Annual Review of Entomology, Volume 67 is January 2022. Please see for revised estimates.
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Patterns of dietary and foraging strata utilization by the ground ant community of a Brazilian cocoa planatation were examined using sugar, meat and flour baits on the surface as well as buried. A total of 49 ant species was recorded, including 10 species of Pheidole. Strong dominance was exercised by Solenopsis geminata at both surface and subterranean strata, and at all food resources. The dissimilarity matrix of the epigaeic ant fauna was much greater than that for the hypogaeic species. At least four guilds were identified: the fungus-growing ants, epigaeic nectivores, epigaeic carnivores, and hypogaeic foragers. Niche breadth reduction, leading to the formation of guilds, permits the coexistence of many species in the this ground ant community from a tropical cocoa plantation. -Authors