Open access journal: http://periodicos.uefs.br/ojs/index.php/sociobiology
ISSN: 0361-6525
Sociobiology 60(1): 50-55 (2013)
Exotic spread of Solenopsis invicta Buren (Hymenoptera: Formicidae) beyond North America
JK Wetterer
Introduction
More than 100 years ago, Forel (1911) compiled a
list of 15 tramp ant species, spread by human commerce,
which had achieved or were in the process of achieving cosmopolitan
distributions. Eight of these have become major
ecological, agricultural, and/or household pests: Anoplolepis
gracilipes (Smith), Linepithema humile (Mayr), Monomorium
destructor (Jerdon), Monomorium pharaonis (L.),
Paratrechina longicornis (Latreille), Pheidole megacephala
(Fabricius), Solenopsis geminata (Fabricius), and Tapinoma
melanocephalum (Fabricius) (Wetterer 2005, 2007, 2008,
2009a, b, 2011, Wetterer et al. 2009). Over the past 100
years, many additional ant species, not on Forel’s (1911) list,
have begun to spread around the world. The most notorious
of these is the fire ant Solenopsis invicta Buren.
Abstract
The South America fire ant Solenopsis invicta Buren arrived in Mobile, Alabama by ship
sometime before 1945. Since then, S. invicta has spread in North America across the
southern US and northeastern Mexico. More recently, S. invicta has invaded the West
Indies and parts of the Old World. Here, I examine this more recent exotic spread of
S. invicta beyond North America, reporting new West Indian records and questioning
some Asian records. In 1981, S. invicta was first found in the West Indies, on Puerto Rico.
With my new records from Vieques, Aruba, and Jamaica, S. invicta is now known from
28 West Indian islands. In 2001, the first Old World populations of S. invicta were discovered
in New Zealand and Australia. Nascent populations of S. invicta in New Zealand
have been exterminated and Australia populations have been kept in check through
intensive control efforts. Populations of S. invicta in Taiwan and China first found in
2003-2004, however, have spread broadly. Published reports of S. invicta from Malaysia
and Singapore were based on misidentifications, presumably of the more widespread
Neotropical fire ant, Solenopsis geminata (Fabricius). Reports of S. invicta from
India and the Philippines seem questionable and need confirmation. Where S. invicta
has invaded, it has displaced S. geminata in open habitats, leaving remnant S. geminata
populations, primarily in forested areas. In working to limit the spread and impacts of
fire ants, it will be important to differentiate among the species, and recognize their
similarities and their differences.
SociobiologyAn international journal on social insects
Florida Atlantic University, Jupiter, Florida, USA
Research article - Ants
Article History
Edited by:
Kleber Del-Claro, UFU - Brazil
Received 	 07 November 2012
Initial acceptance 03 December 2012
Final acceptance 17 December 2012
Keywords
biological invasion, exotic species,
invasive species, stinging ants
Corresponding author
James K. Wetterer
Wilkes Honors College
Florida Atlantic University
5353, Parkside Drive, Jupiter, FL
33458, USA
E-Mail: wetterer@fau.edu
Solenopsis invicta is well known for its painful sting,
which in humans causes a burning sensation, usually followed
in a day or two by the appearance of a white pustule.
The venom can cause severe allergic responses and result in
secondary infections, sepsis, anaphylactic shock, and even
death (Prahlow & Barnard 1998). Solenopsis invicta poses a
threat to wildlife (Allen et al. 2004). For example, S. invicta
attacks and kills hatchling sea turtles (Allen et al. 2001, Parris
et al. 2002, Krahe et al. 2003, Krahe 2005).
Originally from the grasslands of South America, S.
invicta arrived in North America by ship at the port of Mobile,
Alabama apparently between 1933 and 1945 (Buren et
al. 1974) and has spread across the southern US and northeastern
Mexico, particularly in open disturbed areas, causing
ecological and economic damage. Many studies have documented
the current and potential range of S. invicta within
Sociobiology 60(1): 50-55 (2013) 51
North America (e.g., Callcott & Collins 1996, Korzukhin
et al. 2001). This dreaded ant is now spreading through the
West Indies (Wetterer & Davis 2010) and has recently arrived
in New Zealand, Australia, and parts of Asia (Chen
et al. 2005, Zhang et al. 2007, Ascunce et al. 2011). Climate
tolerance models predict that S. invicta should be able
to successfully invade many additional regions in both the
New World and Old World (Morrison et al. 2004, Sutherst
& Maywald 2005). Here, I examine this recent exotic spread
of S. invicta beyond North America, reporting new records
for the West Indies and questioning some published records
from Asia.
Methods
Using published and unpublished records, I documented
the worldwide range of S. invicta. I obtained unpublished
site records from museum specimens in the collections
of Archbold Biological Station (ABS, identified by M.
Deyrup) and the Museum of Comparative Zoology (MCZ,
identified by S. Cover). In addition, I used on-line databases
with collection information on specimens by Antweb (www.
antweb.org), and the Global Biodiversity Information Facility
(www.gbif.org).
Geographic coordinates for collection sites came
from published references, specimen labels, maps, or geography
web sites (e.g., earth.google.com, www.tageo.com,
and www.fallingrain.com). 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 islands and
natural areas, the center of the region. I made one exception,
for Peru. Trager (1991) and Pitts (2002) both listed S.
invicta from Peru with no site, but Peru is a large country, so
I mapped the record to Esperanza, the city in Peru closest to
a known S. invicta population (in Brazil). Taber (2000) presented
a range map with S. invicta only in this part of Peru.
In total, I plotted >1600 site records for S. invicta (Fig. 1).
Results
I collected Solenopsis invicta in the southeastern US
and on West Indian islands (vouchers deposited in Harvard
University’s Museum of Comparative Zoology). I documented
S. invicta records from 28 West Indian islands (Table
1), including its first records from Vieques (seven sites:
Colonia Lujan, trees by pasture, 18.116N, 65.440W, 13
October 2005; Montealta, forest patch, 18.143N, 65.454W,
13 October 2005; west of Puerto Martineau, Route 200 at
4 km mark, 18.141N, 65.479W, 18 October 2005; east of
airport, Route 200 at 5 km mark, 18.137N, 65.487W, 18
October 2005; pier turn-off, Route 200, 18.130N, 65.513W,
18 October 2005; Mosquito Pier, Acacia near end, 18.148N,
65.513W, 7 June 2006; Santa Maria, baseball field, 18.154,
65.432W, 8 June 2006), Aruba (two sites: Malmok, Tierra
del Sol Golf Course, 12.607N, 70.043W, 3 August 2007;
Oranjestad, Divi Village Golf Course, 12.538N, 70.058W,
3 August 2007), and Jamaica (one site: Cinnamon Hill, Cinnamon
Hill Golf Course, 18.517N, 77.813W, 20 December
2010).
The ISSG (2010) website reported records of S. invicta
from three of the Cayman Islands (Table 1), citing “Burton,
2003 in Varnham, 2006,” though Varnham (2006) gives
no specific island records. F. Burton (pers. comm.) could not
recall the original source of the S. invicta records, however,
in 2008, I collected S. invicta on Grand Cayman. It would
be useful to confirm records from Cayman Brac and Little
Cayman.
For published Old World records of S. invicta, site
records and species identity are well documented for exotic
populations in New Zealand, Australia, Taiwan, and China
(Table 2). There have been three successive populations of
S. invicta discovered in New Zealand, and each has been
successfully exterminated (MAF Biosecurity 2010).
I was unable to verify reports of S. invicta from Malaysia,
Singapore, India, and the Philippines (Na & Lee
2001, Rajagopal et al. 2005, Vanderwoude et al. 2006, Sarty
2007, Kuo 2008, Wikipedia 2012).
An and Lee (2001) reported S. invicta from Malaysia,
Sarty (2007) listed S. invicta from Singapore, and Vanderwoude
et al. (2006) mentioned S. invicta from both Malaysia
and Singapore. I e-mailed C.Y Lee, M. Sarty, and C.
Vanderwoude asking about the bases of these records. C.Y.
Lee (pers. comm.) wrote: “S. invicta is not found in Malaysia
and Singapore so far. It was a mistake in identification in
the Na & Lee (2001) paper.” C. Vanderwoude (pers. comm.)
sent me copies of Na & Lee (2001) and Sarty (2007) and
wrote “I’m scratching around for a better ref for RIFA [S.
invicta] in Singapore but can’t seem to lay my hands on it.”
Rajagopal et al. (2005) surveyed ants in Sattur Taluk,
Tamil Nadu, India, reporting S. invicta from all three
habitats studied: riverine, cultivated, and industrial areas. It
seems improbable to find high saturation of S. invicta in this
one small area, but no S. invicta anywhere else in India. I
e-mailed T. Rajagopal and R. Gadagkar (whom Rajagopal
et al. 2005 credited with ant identification). R. Gadagkar
(pers. comm.) replied: “I have now confirmed from my colleague
Dr. Thresiamma Varghese that she did identify some
ant specimens for Rajagopal et al but Solenopsis invicta was
certainly not one of them.”
Kuo (2008) listed S. invicta from the Philippines and
Wikipedia (2012) included a report of S. invicta in the Philippines,
which has been paraphrased on many other web
sites: “There have also been reports of colonies in metro
Manila and the Province of Cavite in the Philippines since
July 2005; however, since early 2007, they have spread now
as far as the Bicol Region.” I could not find an e-mail contact
for K.C. Kuo, nor the source of the Wikipedia reports,
JK Wetterer - Exotic spread of Solenopsis invicta beyond North America52
which were posted anonymously from two IP addresses in
Houston, Texas. G. Alpert and D. General do not include
S. invicta on their list of ants of the Philippines on antweb.
org. D. General (pers. comm.), who worked with S. invicta
for six years in Arkansas, has been actively searching for
S. invicta in the Philippines, but so far has not found any.
Although I cannot be certain, it seems likely that reports of
S. invicta from India and the Philippines were based on mis-
identifications.
Discussion
	
Sometime before 1945, S. invicta arrived in Alabama
from South America. Solenopsis invicta has since become
widespread across the southern US and into northeastern
Mexico, with scattered indoor records in more temperate
areas (Fig. 1).
Starting about 1980, S. invicta began spreading
through the West Indies, with the earliest records from Puerto
Rico and the US Virgin Islands (Table 1, Wetterer & Snelling
2006). Solenopsis invicta is now known from 28 West
Indian islands (Table 1). The most recent first records, from
Aruba and Jamaica, all come from golf courses. Cinnamon
Hill golf course in Jamaica, where I found S. invicta, imports
sod from Florida. Such importation of sod from Florida may
be an important mode of spreading S. invicta in the West
Indies.
Solenopsis invicta was first recorded in the Old World
in 2001 (Nattrass & Vanderwoude 2001, Harris 2001). Nascent
populations of S. invicta have been exterminated in
Table 1. Earliest known records of Solenopsis invicta from West Indian islands. MCZ = Museum of Comparative Zoology. + = no previously
published records.
				
Earliest record
Puerto Rico 1981 (Buren 1982)
St Croix, USVI 1988 (Wetterer & Snelling 2006)
Trinidad 1991 (G.L. White, pers. comm.): several sites
San Salvador, Bahamas 1993 (Deyrup 1994)
New Providence, Bahamas 1995 (Deyrup et al. 1998)
North Andros, Bahamas 1996 (Deyrup et al. 1998)
Guana Island, BVI 1996 (Davis et al. 2001)
Gorda Cay, Bahamas 1997 (Davis et al. 2001)
Antigua 2000 (Davis et al. 2001)
Abaco, Bahamas 2000 (Davis et al. 2001)
Grand Bahama, Bahamas 2000 (Davis et al. 2001)
Providenciales, TCI 2001 (Davis et al. 2001)
Grand Cayman ≤2003 (ISSG 2010)
Cayman Brac ≤2003 (ISSG 2010)
Little Cayman ≤2003 (ISSG 2010)
Berry Islands, Bahamas 2005 (Wetterer & Snelling 2006)
St Thomas, USVI 2005 (Wetterer & Snelling 2006)
St John, USVI 2005 (Wetterer & Snelling 2006)
Tortola, BVI 2005 (Wetterer & Snelling 2006)
+Vieques, PR 2005 (J.K. Wetterer, MCZ): Colonia Lujan
Anguilla 2006 (Wetterer & Davis 2010)
St Martin 2006 (Wetterer & Davis 2010)
Barbuda 2007 (Wetterer & Davis 2010)
Montserrat 2007 (Wetterer & Davis 2010)
St Kitts 2007 (Wetterer & Davis 2010)
Nevis 2007 (Wetterer & Davis 2010)
+Aruba 2007 (J.K. Wetterer, MCZ): Tierra del Sol golf course
+Jamaica 2010 (J.K. Wetterer, MCZ): Cinnamon Hill golf course
Table 2. Earliest known records of Solenopsis invicta from the Old World. * = needs confirmation.
Earliest record
New Zealand 2001 (Harris 2001)
Australia 2001 (Nattrass & Vanderwoude 2001)
*India 2001-2002 (Rajagopal et al. 2005)
Taiwan 2003 (Chen et al. 2005)
Guangdong Province, China 2004 (Zeng et al. 2005)
Hong Kong 2004 (Zeng et al. 2005)
Fujian Province, China 2005 (Zhang et al. 2007)
Guangxi Province, China 2005 (Zhang et al. 2007)
Hunan Province, China 2005 (Zhang et al. 2007)
Macau 2005 (Zhang et al. 2007)
*Philippines 2005 (Kuo 2008)
Sociobiology 60(1): 50-55 (2013) 53
New Zealand (MAF Biosecurity 2010) and have been largely
kept in check in Australia through intensive control efforts
(Vanderwoude et al. 2004). However, S. invicta populations
in Taiwan and China, first identified in 2003 and 2004
respectively, have spread broadly (Fig. 1, Table 2). Lu et
al. (2008) estimated that, based on its geographic spread, S.
invicta first arrived in China around 1995. There seems to be
no geographic or climatic barriers to prevent S. invicta from
spreading throughout tropical and subtropical Asia (Morrison
et al. 2004, Sutherst & Maywald 2005).
Published reports of S. invicta from Malaysia and
Singapore were based on misidentifications (see above),
presumably of the much more widespread fire ant, Solenopsis
geminata (Fabricius), which invaded Asia from South
America more than 160 years ago (Jerdon 1851). Reports of
S. invicta from India (Rajagopal et al. 2005) and the Philippines
(Kuo 2008) are questionable. It would be prudent to
try to confirm whether or not S. invicta has actually invaded
these countries.
Ascunce et al. (2011) analyzed DNA from S. invicta
populations in Australia, Taiwan, and China and found they
all originated from exotic S. invicta populations in the US.
It seems most likely that S. invicta populations in the West
Indies also originated in the US, imported on plant products,
such as grass sod grown in Florida.
Despite its continued spread, S. invicta is still much
less widespread globally than is S. geminata (see Wetterer
2011). When S. invicta invades, it typically displaces S.
geminata, particularly in habitats preferred by both species:
open, grassy areas. As a result, S. geminata has largely
disappeared from much of its former exotic range in North
America (Wojcik et al. 1976). 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.
Models predict that S. invicta should be able to invade
many tropical and subtropical regions now occupied by S.
geminata (Morrison et al. 2004, Sutherst & Maywald 2005).
In working to mitigate the spread and negative impact of S.
invicta, S. geminata, and other fire ants around the world,
it will be important to distinguish between these ants, and
recognize both their similarities and their differences.
Acknowledgments
	 I thank M. Wetterer for comments on this manuscript,
S. Cover and L. Davis for ant identification, S. Cover
(MCZ) and M. Deyrup (ABS) for help with their respective
ant collections, D. General his search for S. invicta in the
Philippines, C. Vanderwoude, C.Y Lee, D. General, R. Gadagkar,
and F. Burton for their personal communications, W.
O’Brien for GIS help, D.P. Wojcik and S.D. Porter for compiling
their FORMIS bibliography, C. Scheid and R. Pasos
of the FAU library for processing so many interlibrary loans,
Florida Atlantic University, the National Science Foundation
(DEB-0515648) for financial support.
Figure 1. Worldwide distribution records of Solenopsis invicta. Records from India and the Philippines are questionable and need to be confirmed
(see text).
JK Wetterer - Exotic spread of Solenopsis invicta beyond North America54
References
Allen, C.R., Epperson, D.M. & Garmestani, A.S. (2004)
Red imported fire ant impacts on wildlife: a decade of research.
Am. Midl. Nat. 152:88-103. doi: 10.1674/0003-0031-
(2004)152[0088:RIFAIO]2.0.CO;2
Allen, C.R., Forys, E.A., Rice K.G. & Wojcik, D.P. (2001)
Effects of fire ants on hatching sea turtles and the prevalence
of fire ants on sea turtle nesting beaches in Florida. Florida
Entomol. 84:250-253.
Ascunce, M., Yang, C.C., Oakey, J., Calcaterra, L., Wu,
W.J., Shih, C.J., Goudet, J., Ross K. & Shoemaker, D. (2011)
Global invasion history of the fire ant Solenopsis invicta. Science
331: 1066-1068 DOI: 10.1126/science
Buren, W.F. (1982) Red imported fire ant now in Puerto Rico.
Florida Entomol. 65:188-189.
Buren, W.F., Allen, G.E., Whitcomb, W.H., Lennartz, F.E.
& Williams, R.N. (1974) Zoogeography of the imported fire
ants. J. N. Y. Entomol. Soc. 82:113–124.
Callcott, A.A. & Collins, H.L. (1996) Invasion and range expansion
of imported fire ants (Hymenoptera: Formicidae) in
North America. Florida Entomol. 79:240-251.
Chen, Y.-F., H.-C. Chang & C.-M. Chao 2005. Fire ant, a
new hazard to military camps in Taiwan. J. Med. Sci. 25:161-
166.
Davis, L.R., Jr., Vander Meer, R.K. & Porter, S.D. (2001)
Red imported fire ants expand their range across the West
Indies. Florida Entomol. 84:735-736.
Deyrup, M. (1994) Biogeographical survey of the ants of the
island of San Salvador, Bahamas. pp. 21-28. in Kass, L.B.
(ed.) Proceedings of the 5th Symposium on the Natural History
of the Bahamas. Bahamian Field Station, San Salvador,
Bahamas.
Deyrup, M., Davis, L. & Buckner, S. (1998) Composition of
the ant fauna of three Bahamian islands. pp. 23-31 in Proceedings
of the 7th Symposium of Natural History. Bahamian
Field Station, San Salvador, Bahamas.
Forel, A. (1911) Aperçu sur la distribution géographique et la
phylogénie des fourmis. Memoires 1er Congrès International
d’Entomologie 2:81-100.
Harris, R. (2001) Blatant breaches of the border. Stowaways
1:11.
ISSG (Invasive Species Specialist Group) 2010. Solenopsis
invicta http://www.issg.org/database/species/distribution_
display.asp?si=77&ri=19159&pc=*&sts=&status=Alien&la
ng=EN#Alien
Jerdon, T.C. (1851) A catalogue of the species of ants found
in southern India. Madras J. Lit. Sci. 17:103-127.
Korzukhin, M.D., Porter, S.D., Thompson, L.C. & Wiley, S.
(2001) Modeling temperature-dependent range limits for the
fire ant Solenopsis invicta (Hymenoptera: Formicidae) in the
United States. Environ. Entomol. 30:645-655.
Krahe, H. B. (2005) Impact of the red imported fire ant (Solenopsis
invicta) on two species of sea turtle hatchlings. M.S.
Thesis. Florida Atlantic Univ., Boca Raton.
Krahe, H., Wetterer, J.K. & Wood, L.D. (2003) Impact of fire
ant stings on sea turtle hatchling survival. Proceedings of the
22nd Annual Symposium on Sea Turtle Biology and Conservation.
NOAA Technical Memorandum NMFS-SEFSC
503:211-212.
Kuo, K.C. (2008) Management of red invasive fire ants and
fruit flies - the Taiwan experience. Food and Fertilizer Technology
Center, Taipei, Taiwan. 5 pp.
Lu, Y.Y., Liang, G.W. & Zeng, L. (2008) Study on expansion
pattern of red imported fire ant, Solenopsis invicta Buren, in
South China. Sci. Agric. Sin. 41:1053-1063.
MAF Biosecurity (2010) Red Imported Fire Ant, Solenopsis
invicta.
Morrison, L.W., Porter, S.D., Daniels, E. & Korzukhin, M.D.
(2004) Potential global range expansion of the invasive fire
ant, Solenopsis invicta. Biol. Invas. 6:183–191.
Na, J.P.S. & Lee, C.Y. (2001) Identification key to common
urban pest ants in Malaysia. Trop. Biomed. 18:1-17.
Nattrass, R. & Vanderwoude, C. (2001) A preliminary investigation
of the ecological effects of Red Imported Fire Ants
(Solenopsis invicta) in Brisbane. Ecol. Manag. Restor. 2:220-
223.
Parris, L.B., Lamont, M.M. & Carthy, R.R. (2002) Increased
incidence of red imported fire ant (Hymenoptera: Formicidae)
presence in loggerhead sea turtle (Testudines: Cheloniidae)
nests and observations of hatchling mortality. Florida
Entomol. 85:514-517.
Pitts, J.P. (2002) A cladistic analysis of the Solenopsis saevissima
species-group (Hymenoptera: Formicidae). Ph.D. dissertation.
University of Georgia, Athens, GA, 266 pp.
Porter, S.D. (1992) Frequency and distribution of polygyne
fire ants (Hymenoptera: Formicidae) in Florida. Florida Entomol.
75:248-257.
Prahlow, J.A. & Barnard, J.J. (1998) Fatal anaphylaxis due to
fire ant stings. Amer. J. Foren. Med. Pathol. 19:137-142.
Rajagopal, T., Sevarkodiyone, S.P. & Sekar, M. (2005) Ant
species richness, diversity and similarity index at five selected
localities of Sattur Taluk. Indian J. Environ. Educ. 5:7-
12.
Sarty, M. (2007) Fire ant eradicated at Port of Napier. Biosecurity
73:10.
Sociobiology 60(1): 50-55 (2013) 55
Sutherst, R.W. & G. Maywald 2005. A climate model of the
red imported fire ant, Solenopsis invicta Buren (Hymenoptera:
Formicidae): implications for invasion of new regions, particularly
Oceania. Environ. Entomol. 34:317-335.
Taber, S.W. (2000) Fire ants. Texas A&M University Press,
College Station, Texas, 308 pp.
Trager, J.C. (1991) A revision of the fire ants, Solenopsis
geminata group (Hymenoptera: Formicidae: Myrmicinae). J.
New York Entomol. Soc. 99:141-198.
Vanderwoude, C., Elson-Harris, M., Hargreaves, J.R., Harris,
E. & Plowman, K.P. (2004) An overview of the red imported
fire ant (Solenopsis invicta Buren) eradication plan for Australia.
Rec. S. Aust. Mus. Monog. Ser. 7:11-16.
Vanderwoude, C., Numbuk, S. & Camilosi, C. (2006) Preliminary
report on infestation of little fire ant (Wasmannia
auropunctata) at Kreer Heights, Wewak. Aliens 24/25:6-7.
Varnham, K. (2006) Non-native species in UK Overseas Territories:
a review. Joint Nature Conservation Committee Report
372:1-135.
Wetterer, J.K. (2005) Worldwide distribution and potential
spread of the long-legged ant, Anoplolepis gracilipes. Sociobiol.
45:77-97.
Wetterer, J.K. (2007) Biology and impacts of Pacific islands
invasive species. 3. The African big-headed ant, Pheidole
megacephala (Hymenoptera: Formicidae). Pac. Sci. 61:437-
456.
Wetterer, J.K. (2008) Worldwide spread of the longhorn
crazy ant, Paratrechina longicornis (Hymenoptera: Formicidae).
Myrmecol. News 11:137-149.
Wetterer, J.K. (2009a) Worldwide spread of the ghost ant,
Tapinoma melanocephalum (Hymenoptera: Formicidae).
Myrmecol. News 12:23-33.
Wetterer, J.K. (2009b) Worldwide spread of the destroyer
ant, Monomorium destructor (Hymenoptera: Formicidae).
Myrmecol. News 12:97-108.
Wetterer, J.K. (2011) Worldwide spread of the tropical fire
ant, Solenopsis geminata (Hymenoptera: Formicidae).
Myrmecol. News 14:21-35.
Wetterer, J.K. & Davis, L.R. Jr. (2010) The red imported fire
ant, Solenopsis invicta, (Hymenoptera: Formicidae) in the
Lesser Antilles. Florida Entomol. 93:128-129.
Wetterer, J.K. & Snelling, R R. (2006) The red imported fire
ant, Solenopsis invicta, in the Virgin Islands (Hymenoptera:
Formicidae). Florida Entomol. 89:431-434.
Wetterer, J.K., Wild, A.L., Suarez, A.V., Roura-Pascual, N. &
Espadaler, X. (2009) Worldwide spread of the Argentine ant,
Linepithema humile (Hymenoptera: Formicidae). Myrmecol.
News 12:187-194.
Wikipedia (2012) Red imported fire ant. http://en.wikipedia.
org/wiki/Solenopsis_invicta
Wojcik, D.P., Buren, W.F., Grissell, E.E. & Carlysle, T.
(1976) The fire ants (Solenopsis) of Florida (Hymenoptera:
Formicidae). Florida Dept. Agric. Cons. Serv. Div. Plant Indust.
Entomol. Circ. 173:1-4.
Zeng L, Lu, Y.Y., He, X.F., Zhang, W.Q. & Liang, G.W.
(2005) Identification of the red imported fire ant Solenopsis
invicta to invade mainland China and infestation in Wuchuan,
Guangdong. Chinese Bull. Entomol. 42, 144–148.
Zhang, R., Li, Y., Liu, N. & Porter, S.D. (2007) An overview
of the red imported fire ant (Hymenoptera: Formicidae) in
Mainland China. Florida Entomol. 90:723-731.