Ant community composition in a citrus grove reveals eastern expansion in Florida of the South American big-headed ant (Hymenoptera: Formicidae)

Abstract

Florida has become a worldwide hotspot for introduced and invasive ants (Hymenoptera: Formicidae). Furthermore, studies have shown that non-native ants support other invasive insects in Florida, such as the Asian citrus psyllid, Diaphorina citri Kuwayama (Hemiptera: Liviidae), which is the vector of Candidatus Liberibacter asiaticus, the causal pathogen of citrus greening, the worst citrus disease in the world. The Asian citrus psyllid establishes beneficial interactions with invasive, introduced, and native ants. In this study, we described the ant diversity in a citrus grove in south Florida as a first step to assess ant-Asian citrus psyllid interactions for future studies. During the summer of 2018, 25 pitfall traps were set in a citrus grove. The red imported fire ant, Solenopsis invicta (Buren), was present in all the traps (100 %), and 19 traps (76 %) had Dorymyrmex bureni (Trager), the pyramid ant, which is native to the southeast United States. Two introduced ants, Cardiocondyla emeryi (Forel) and Brachymyrmex obscurior (Forel), were found in 15 (60 %) and 11 (44 %) traps, respectively. The South American big-headed ant, Pheidole obscurithorax (Naves), was found in 14 traps (56 %). This introduced ant has been found to co-exist with S. invicta, and to our knowledge, this is the first report of the species in this eastern part of peninsular Florida. Finally, Cardiocondyla venustula (Wheeler), an introduced ant from Africa, was found in one of the traps. Although, this ant is found in south Florida, it seems uncommon, and this is its first report in this county. Our study highlights the continued spread of introduced and invasive ants in Florida.

Full text

Research Article
Marina S. Ascunce*, Katherine Casey Carroll, Joe Aufmuth, Lauren Justice, Patricia Perez,
Andrew Nisip, Jenny M. Gavilanez-Slone, Jawwad A. Qureshi, Sanford Porter and Andrea Lucky
Ant community composition in a citrus grove
reveals eastern expansion in Florida of the South
American big-headed ant (Hymenoptera:
Formicidae)
https://doi.org/10.1515/flaent-2024-0011
Received February 28, 2024; accepted February 29, 2024;
published online March 29, 2024
Abstract: Florida has become a worldwide hotspot for
introduced and invasive ants (Hymenoptera: Formicidae).
Furthermore, studies have shown that non-native ants
support other invasive insects in Florida, such as the Asian
citrus psyllid, Diaphorina citri Kuwayama (Hemiptera:
Liviidae), which is the vector of Candidatus Liberibacter
asiaticus, the causal pathogen of citrus greening, the worst
citrus disease in the world. The Asian citrus psyllid estab-
lishes beneficial interactions with invasive, introduced, and
native ants. In this study, we described the ant diversity in a
citrus grove in south Florida as a first step to assess ant-Asian
citrus psyllid interactions for future studies. During the
summer of 2018, 25 pitfall traps were set in a citrus grove.
The red imported fire ant, Solenopsis invicta (Buren), was
present in all the traps (100 %), and 19 traps (76 %) had
Dorymyrmex bureni (Trager), the pyramid ant, which is
native to the southeast United States. Two introduced ants,
Cardiocondyla emeryi (Forel) and Brachymyrmex obscurior
(Forel), were found in 15 (60 %) and 11 (44 %) traps, respec-
tively. The South American big-headed ant, Pheidole
obscurithorax (Naves), was found in 14 traps (56 %). This
introduced ant has been found to co-exist with S. invicta, and
to our knowledge, this is the first report of the species in this
eastern part of peninsular Florida. Finally, Cardiocondyla
venustula (Wheeler), an introduced ant from Africa, was
found in one of the traps. Although, this ant is found in south
Florida, it seems uncommon, and this is its first report in this
county. Our study highlights the continued spread of intro-
duced and invasive ants in Florida.
Keywords: invasive ants; fire ants; Pheidole obscurithorax;
pitfall traps; Asian citrus psyllid; citrus greening
Sumario: Floridasehaconvertidoenun“hot-spot”para
especies introducidas e invasoras de hormigas
Current address: Katherine Casey Carroll, Department of Ecology and
Evolutionary Biology, University of Colorado at Boulder, Boulder, CO 80309,
USA, E-mail: Casey.Carroll@colorado.edu.
Current address: Patricia Perez, Florida Department of Agriculture and
Consumer Services, Department of Plant Industry, Gainesville, FL 32608,
USA, E-mail: patricia.perez@fdacs.gov.
Current address: Andrew Nisip, Department of Biology, University of
Louisville, Louisville, KY 40292, USA; E-mail: andrew.nisip@louisville.edu.
Current address: Jenny M. Gavilanez-Slone, Entomology Laboratory,
Faculty of Natural Resources, Escuela Superior Politécnica de Chimborazo
(ESPOCH), Riobamba, Ecuador; E-mail: jenny.gavilanez@espoch.edu.ec.
*Corresponding author: Marina S. Ascunce, USDA-ARS Center for
Medical, Agricultural, and Veterinary Entomology, Gainesville, FL 32608,
USA; and Emergency Pathogens Institute, Department of Plant
Pathology, University of Florida, Gainesville, FL 32610, USA,
E-mail: marina.ascunce@usda.gov. https://orcid.org/0000-0002-5239-
3585
Katherine Casey Carroll, Patricia Perez and Jenny M. Gavilanez-Slone,
Emergency Pathogens Institute, Department of Plant Pathology, University
of Florida, Gainesville, FL 32610, USA
Joe Aufmuth, George A. Smathers Libraries, University of Florida,
Gainesville, FL 32611, USA, E-mail: mapper@uflib.ufl.edu
Lauren Justice and Sanford Porter, USDA-ARS Center for Medical,
Agricultural, and Veterinary Entomology, Gainesville, FL 32608, USA,
E-mail: lauren.justice@usda.gov (L. Justice), sdporter22@gmail.com
(S. Porter)
Andrew Nisip, USDA-ARS Center for Medical, Agricultural, and Veterinary
Entomology, Gainesville, FL 32608, USA; and Emergency Pathogens
Institute, Department of Plant Pathology, University of Florida, Gainesville,
FL 32610, USA
Jawwad A. Qureshi, Entomology and Nematology Department, Southwest
Florida Research and Education Center (SWFREC), University of Florida,
Immokalee, FL 34142, USA, E-mail: jawwadq@ufl.edu
Andrea Lucky, Entomology and Nematology Department, University of
Florida, Gainesville, FL 32611, USA, E-mail: alucky@ufl.edu
Fla. Entomol. 2024; 107(1): 20240011
Open Access. © 2024 the author(s), published by De Gruyter on behalf of the Florida Entomological Society. This work is licensed under the Creative
Commons Attribution 4.0 International License.

(Himenóptera: Formicidae). A su vez, estudios han mos-
tradoquelashormigasno-nativasayudanaotrasespecies
invasoras en Florida como el psílido asiático de los cítricos,
Diaphorina citri Kuwayama (Hemíptera: Liviidae), el cual
es vector de Candidatus Liberibacter asiaticus, el agente
causal de la enfermedad del “enverdecimiento”de los
cítricos, la peor enfermedad de los cítricos a nivel mundial.
El psílido asiático de los cítricos establece relaciones
beneficiosas con hormigas invasoras, introducidas y nati-
vas. En este estudio, describimos la diversidad de hormigas
en una huerta de cítricos en el sur de la Florida, como un
primer paso para evaluar las interacciones entre las hor-
migas y el psílido asiático de los cítricos en estudios futu-
ros. En el verano del 2018 se pusieron 25 trampas de caída
enunahuertadecítricos.Lahormigainvasorarojade
fuego, Solenopsis invicta (Buren), estuvo presente en todas
las trampas (100 %), mientras que 19 trampas (76 %) tenían
Dorymyrmex bureni (Trager), la hormiga piramidal, la cual
es nativa del sureste de los Estados Unidos. Dos hormigas
introducidas: Cardiocondyla emeryi (Forel) y Brachymyr-
mex obscurior (Forel) se encontraron en 15 (60 %) y 11
(44 %) de las trampas, respectivamente. La hormiga Sud-
americana de cabeza grande, Pheidole obscurithorax
(Naves), fue hallada en 14 trampas (56 %). Esta hormiga
introducida ha sido hallada coexistiendo con S. invicta ya
nuestro entender, este sería el primer reporte de esta
hormiga en el este de la península de Florida. Finalmente,
Cardiocondyla venustula (Wheeler), una hormiga intro-
ducida del Viejo Mundo fue hallada en una de las trampas.
A pesar de que esta hormiga se encuentra en el sur de la
Florida, es menos común, y este sería el primer reporte de
esta hormiga en este condado. Nuestro estudio remarca la
continua expansión de las hormigas introducidas e inva-
soras en Florida.
Palabras Claves: hormigas invasoras; hormigas de fuego;
Pheidole obscurithorax; trampas de caída; psílido asiático de
los cítricos; enverdecimiento de los cítricos
More than 520 ant species (Hymenoptera: Formicidae) have
been transported outside their native range and established
in new areas worldwide (Bertelsmeier et al. 2018; Lach et al.
2010; Wong et al. 2023). Five of them are among the 100 most
concerning invasive species including the red imported fire
ant, Solenopsis invicta (Buren), the Argentine ant, Line-
pithema humile (Mayr), the African big-headed ant, Pheidole
megacephala (Fabricius), the little fire ant, Wasmannia
auropunctata (Roger), and the longlegged ant, Anoplolepis
gracilipes (Smith) (ISSG 2015). All these five ants are in the
United States, and all but A.gracilipes are in Florida (Deyrup
2003; King and Porter 2007; Ohyama et al. 2020). The total ant
fauna in Florida encompasses 218 ant species, with 52 being
introduced or invasive (Deyrup 2003; Deyrup et al. 2000).
This non-native ant fauna is the largest among any of the
states in the United States (Deyrup 2003) and the largest
regional non-native ant fauna in the world (King and Porter
2007), making Florida a hot spot for introduced and invasive
ants. Even though non-native ant species are known in
Florida, the understanding of ant community composition
has not received enough attention, and the knowledge of the
distributions of even well-established species is limited. A
clear understanding of how these non-native ants get
established in Florida may eventually allow formal meta-
analyses of invasion impacts, which could help pinpoint
problematic taxa and prioritize early responses. Mapping
the new reports of invasive species is central to reporting
expansions of invasive species, guiding effective early re-
sponses to limit their further geographical expansion, and
determining if control efforts are working (Elith and
Leathwick 2009). This study aimed to characterize the ant
fauna at one agricultural site in Florida, particularly at a
citrus grove, due to its importance in Florida agriculture.
In Florida, citrus is a major part of the agricultural
landscape and a high-value crop of USD $438 million
(USDA-NASS 2022). In 1998, a serious pest of citrus, the Asian
citrus psyllid, Diaphorina citri Kuwayama (Hemiptera: Livii-
dae), was discovered in Florida (Halbert 1998). This invasive
psyllid is the vector of Candidatus Liberibacter asiaticus, a
phloem-limited α-proteobacterium, that is the cause of
the devastating citrus disease Huanglongbing or citrus
greening. About seven years after D. citri arrived in Florida,
Huanglongbing was detected in 2005 (Bové 2006) and since
this date has spread throughout Florida. Currently, 100 % of
the citrus groves in Florida are positive for Huanglongbing
(Graham et al. 2020). Because no cure is available, infected
trees die within a few years, resulting in an 85 % reduction in
citrus production since the onset of Huanglongbing in the
state (Singerman and Rogers 2020; USDA-NASS 2022). Man-
agement of Huanglongbing with antimicrobial agents and
nutritional treatments has not prevented major production
losses in Florida (Blaustein et al. 2018; Wang et al. 2017).
Classical biological control of D. citri using a parasitic wasp,
Tamarixia radiata (Waterston) (Hymenoptera: Eulophidae)
(Hoy et al. 1999) has not been effective in controlling the
psyllid presumably due to its sensitivity to insecticides and
low temperatures during winter (Chong et al. 2010; Qureshi
et al. 2009). The limited effect of biological control agents gets
exacerbated by ants in citrus groves, as ants help D. citri by
protecting them against natural enemies (predators and
parasitoids) in return for the psyllid’s sugary excretions
(i.e., honeydew) as their food. Ants do this by physically
attacking predators and parasitoids and driving them away
2M.S. Ascunce et al.: Ant community composition in a citrus grove in Florida

from D. citri (Michaud 2004; Navarrete et al. 2013; Oliver et al.
2008; Shrestha et al. 2022; Vanek and Potter 2010; Yoo et al.
2013). In Florida citrus groves, invasive and introduced ants,
including the red imported fire ant, the African big-headed
ant, the rover ant, Brachymyrmex obscurior (Forel), as well as
the native pyramid ant, Dorymyrmex bureni (Trager) have
been observed tending and protecting D. citri (Michaud 2004;
Navarrete et al. 2013; Shrestha et al. 2022). This study focused
on ant species diversity ina Florida citrus grove as a first step
to gain insights into the different ant species that can poten-
tially provide protection for D. citri against its predators.
On 5 July 2018, we surveyed a citrus grove at the Uni-
versity of Florida Indian River Research and Education
Center (IRREC) in Fort Pierce (27.43 °N, 80.41 °W). Ants were
collected using pitfall traps with ethanol (70 %), as described
in Laub et al. (2009). A “mini-roof”was designed to protect
the traps from rain and reduce evaporation (Supplementary
Material 1). Traps were placed along three transects sepa-
rated by 25 m. Along each transect, 7–10 pitfall traps were
placed at about 5 m intervals for a total of 25 traps. Traps
were collected after 48 h. Ants were sorted, transferred
into vials with 95 % ethanol, and identified under a stereo-
microscope using taxonomic keys from Deyrup (2003, 2016)
and MacGown (2022).
After taxonomic identification, the presence of each ant
species per pitfall was summarized in Table 1. All 25 traps
(100 %) contained specimens of red imported fire ant, and 19
(76 %) contained D. bureni.Cardiocondyla emeryi (Forel) and
B. obscurior were found in 15 (60 %) and 11 (44 %) traps,
respectively. The South American big-headed ant, Pheidole
obscurithorax (Naves), was found in 14 traps (56 %). Finally,
Cardiocondyla venustula (Wheeler) was found in one pitfall
(4 %).
A map was created to show the co-distribution of red
imported fire ant and D. bureni, as well as P.obscurithorax
and C. venustula (Figure 1). We used the Global Biodiversity
Information Facility (GBIF) to create this map. Detailed in-
formation on the process used to create this map is in Sup-
plementary Material 2. Presence-only data for each species
was downloaded individually, combined and evaluated for
occurrences and duplications. The species records outside of
Florida were removed. A final presence or absence matrix
Table :Presence of ant species per pitfall: Solenopsis invicta,Dorymyrmex bureni,Cardiocondyla emeryi,Pheidole obscurithorax,Brachymyrmex obscurior,
and Cardiocondyla venustula.
Pitfall S. invicta D. bureni C. emeryi P. obscurithorax B. obscurior C. venustula
xxxx
xxx x
xx
xx
xx x x
xx x
xxx x
xxx x x
xx
 xxx
 xx
 x
 xxx x
 x x
 xx x
 xxx x x
 x
 xxx x
 xxx x x
 xxxx
 xxx
 xxx x x
 xxx x
 xxx x x
 xxxx
# Positive pitfalls      
Prevalence  % % % % %%
Status Inv N Int Int Int Int
Status: Inv: invasive; Int: introduced; N: native.
M.S. Ascunce et al.: Ant community composition in a citrus grove in Florida 3

based on latitude, longitude, and species was imported into
ArcGIS Pro v2.2 (Esri, Redlands, California), where separate
layers were created for each species.
Most agricultural lands are highly disturbed, facilitating
the establishment of invasive species that outcompete and
exclude native species (Mooney and Cleland 2001). This could
be the case in this citrus grove, in which five out of the six
species collected were non-native (Table 1). However, long-
term studies need to be done to confirm this hypothesis. It
was not surprising to find red imported fire ant in 100 % of
the traps (Table 1). This invasive ant from South America was
first detected in Mobile, Alabama, in the late 1930s. Since
then, it has rapidly spread throughout the southeast US as
well as other countries, becoming a global invasive species in
the 2000s (Ascunce et al. 2011). Several features make the red
imported fire ant a successful invader, including high
reproducibility, ability to outcompete native ants for re-
sources, and social structure (Arnan et al. 2018; Bertelsmeier
et al. 2017; Holway et al. 2002; Passera 1994). Despite the red
imported fire ant’s ability to outcompete native ants in
general, the native Dorymyrmex are able to coexist with
S. invicta (Helms and Vinson 2001; LeBrun et al. 2012; Stuble
et al. 2010), and in this study, D. bureni was in fact found in
70 % of the traps. Both species have been found to tend to
D. citri; however, S. invicta provides greater protection to the
psyllid against its natural enemies, resulting in a larger
economic cost to citrus growers in Florida (Shrestha et al.
2022).
Cardiocondyla are native to Africa and Asia, but some
Cardiocondyla species have become common cosmopolitan
introduced ants, including C. emeryi. In the case of C. venus-
tula, this ant has been recently introduced to the Caribbean,
Florida and Hawaii and its distribution seems to be
expanding (Seifert 2002; Starr et al. 2007), with this study
being its first report in this county (Table 1, Figure 1). For B.
obscurior, a native ant from the Neotropics, the nature of its
presence in Florida as introduced or native species is not
clear (Deyrup et al. 2000). Finally, P. obscurithorax, an ant
native to South America, was first found in Mobile, Alabama
in 1949, and then spread to other parts of the southeast US
(Naves 1985). Its current distribution in the US extends from
Texas to Florida, where it co-exists with the red imported fire
ant. It reached the panhandle of Florida in the late 1990s
(Storz and Tschinkel 2004; Wetterer et al. 2015; Wilson 2003).
This study is the first report of this species in the eastern part
of peninsular Florida and in a citrus grove. Although the
negative impact of its introduction is unclear, it continues to
spread through Florida and is expected to eventually occupy
a range similar to the red imported fire ant (King and
Tschinkel 2007). Our study highlights the continued spread
Figure 1: Distribution of S. invicta,D. bureni,
P. obscurithorax, and C. venustula in Florida.
S. invicta (shaded light red, data from
USDA-APHIS-PPQ https://www.aphis.usda.
gov/plant_health/plant_pest_info/fireants/
downloads/federal-imported-fire-ant-
quarantine.pdf), D. bureni (gray stripes, data
from Deyrup (2016)), P. obscurithorax (filled
circles, data from GBIF https://doi.org/10.
15468/dl.bc4w7h) and C. venustula (open cir-
cles, data from GBIF https://doi.org/10.15468/
dl.npp4zj). The black star shows the location of
the collection site for this study. The distribu-
tion of the introduced ants, C. emeryi and B.
obscurior is in Supplementary Material 2.
4M.S. Ascunce et al.: Ant community composition in a citrus grove in Florida

of introduced and invasive ants in Florida. The erosion of
native ant diversity in this citrus grove may likely be the
result of “invasional meltdown”, which was defined as: “the
process by which a group of [introduced] species facilitate
one another’s invasion in various ways, increasing the
likelihood of survival and/or of ecological impact, and
possibly the magnitude of impact”(Simberloffand Von Holle
1999, p 22). However, to test this hypothesis, long-term
studies are needed. In addition, management practices
directed to control the Asian citrus psyllid and other citrus
insect pests could also affect native ant diversity.
Acknowledgments: We thank the University of Florida, IFAS
Seed Grant on Interdisciplinary Research on Invasive Spe-
cies (IRIS) for initial funding of this project, Erica Goss
(University of Florida, Plant Pathology Department) for her
encouragement about the project and comments on early
versions of this manuscript, Rachel Atchison (USDA-ARS-
CMAVE) for help with taxonomic identification and Sara
Alvarez (Education and Volunteer Specialist for Manatee
County, Florida) who assisted with the ant collection. We
would like to thank two anonymous reviewers and the editor
for their valuable comments on this manuscript.
Research ethics: Not applicable.
Author contributions: The authors have accepted
responsibility for the entire content of this manuscript and
approved its submission.
Competing interests: The authors state no conflict of
interest.
Research funding: This research work was supported by
IFAS Seed Grant on Interdisciplinary Research on Invasive
Species (IRIS) and partial funding was obtained from USDA-
ARS-CMAVE-Fire Ant Unit based funds.
Data availability: The raw data is provided in the manuscript.
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Supplementary Material: This article contains supplementary material
(https://doi.org/10.1515/flaent-2024-0011).
6M.S. Ascunce et al.: Ant community composition in a citrus grove in Florida