Pheidole navigans

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Pheidole navigans
Pheidole navigans 542238719.jpg
P. navigans minor worker
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Myrmicinae
Genus: Pheidole
Species:
P. navigans
Binomial name
Pheidole navigans
Forel, 1901

Pheidole navigans, also called the navigating big-headed ant or the wandering big-headed ant, is a species of big-headed ant in the Pheidole flavens species group. Native to the Neotropics, it is a widespread invasive species, however its ecological impacts are not considered significant as of 2022. Prior to 2015, it was classified as a synonym of Pheidole flavens , and it was misidentified in its invasive range as Pheidole moerens . [1] [2] As with the vast majority of species within the genus Pheidole, they possess a dimorphic caste system with a major and minor worker. They are small reddish-brown ants not larger than 2.5 millimetres (0.098 in) with a generalist ecology and flexible nest preferences. [3] [4]

Contents

Taxonomy

Pheidole navigans is traditionally considered to be part of the Pheidole flavens species group, a large assemblage of primarily Neotropical big-headed ants containing over 160 species characterized by small size, vestigial or absent mesosomal convexity, and 12-segmented antennae. [5] However, the P. flavens group was not phylogenetically recovered to be monophyletic in 2015 and therefore the grouping appears largely artificial. P. navigans appears to be closely related to P. moerens and P. flavens sensu stricto, though, as a relatively basal clade of Pheidole as a whole. [6] It was originally described as a subspecies of Pheidole flavens in 1901 by entomologist Auguste Forel, later fully synonymized under Pheidole flavens in 2003 by entomologist E. O. Wilson, and finally restored to full species status by Sarnat et al. in 2015. [1] Its type locality is peculiar in that it is located in Germany, part of its introduced range, as an interception record from orchids originating from Veracruz, Mexico. [2]

Identification

P. navigans is part of the subfamily Myrmicinae, which is characterized by a two-segmented waist consisting of a petiole and postpetiole, a fused promesonotum, the presence of frontal lobes over the antennal sockets, and generally triangular mandibles with a few exceptions. Within its subfamily, the genus is distinctly recognized by the presence of clearly dimorphic or rarely continuously polymorphic worker castes, a three- or four-segmented antennal club, and 12-segmented antennae except for two groups of small Neotropical Pheidole, the P. tachigaliae species group and P. perpusilla species group, with 11 and 10 antennal segments respectively. In the Neotropical components of its genus, the P. flavens species group may be distinguished with a combination of 12-segmented antennae, a three-segmented antennal club, a medium to small size, an absent or vestigial mesosomal convexity, a relatively thick antennal club, a short scape, a moderately curved head shape, relatively shallow antennal scrobes, a lack of ladder-like transverse carinae on the head of the major worker, and a lack of cephalic phragmotism and a very short metanotum in the major worker. [5] Their major workers may be separated from the closely related P. flavens and P. moerens by longitudinal rugae on the posterior lobes of the head, a longer, more distinct, and narrower antennal scrobe with a stronger and more continuous frontal carina, a more glossy scrobe depression, and transverse striae on the dorsal surface of the promesonotum. Their minor workers, however, are impossible to distinguish from other species in the P. flavens species complex. [2] [4]

Morphology

Pheidole navigans is a small, reddish-brown to dark brown, short-limbed species, with minor workers reaching 2.0 millimetres (0.079 in), major workers 2.4 millimetres (0.094 in), males 2.8 millimetres (0.11 in), and queens 3.5 millimetres (0.14 in) in total body length. [7] [8] They are morphologically similar to a number of other species in the Pheidole flavens species group like Pheidole flavens , Pheidole glomericeps , and Pheidole moerens , leading to frequent misidentifications in their invasive range prior to 2015. Their color variation was suspected in 1985 to be caused by dietary variations, as colonies fed with honey and seeds developed lighter in color than colonies fed with house flies. The queens of this species are small and blackish. [9] Otherwise, they share most traits with other members of its species group and are morphologically unremarkable. [2] [4]

Castes

As with the vast majority of ants within the genus Pheidole , they have a dimorphic caste system consisting of major and minor workers. Minor workers are more versatile and more numerous, and typically perform most of the tasks in the colony. In contrast, the more specialized major workers are mainly used for defense and food dissection, although they may occasionally venture out to forage. Minor workers have relatively normal appearances, however major workers possess hypertrophied heads with powerful muscles. The minor workers allocate more space for the nervous system than major workers, suggesting they perform more complex tasks, while major workers tend to have more integrated brains than minor workers and more muscle mass in the head. [4] [10] [11]

Minor worker

P. navigans AntWeb minor worker specimen Pheidole navigans casent0005743 profile 1.jpg
P. navigans AntWeb minor worker specimen

The minor workers of P. navigans are small, with head width between 0.44 millimetres (0.017 in) and 0.47 millimetres (0.019 in), head length between 0.49 millimetres (0.019 in) and 0.51 millimetres (0.020 in), scape length between 0.42 millimetres (0.017 in) and 0.43 millimetres (0.017 in), eye length between 0.08 millimetres (0.0031 in) and 0.10 millimetres (0.0039 in), mesosoma length, also known as Weber's length, between 0.67 millimetres (0.026 in) and 0.74 millimetres (0.029 in), spiracle length between 0.08 millimetres (0.0031 in) and 0.10 millimetres (0.0039 in), petiole width between 0.13 millimetres (0.0051 in) and 0.14 millimetres (0.0055 in), postpetiole width between 0.18 millimetres (0.0071 in) and 0.22 millimetres (0.0087 in), and pronotum width between 0.42 millimetres (0.017 in) and 0.45 millimetres (0.018 in). They are generally reddish-brown to dark brown with their gasters sometimes slightly darker and their legs orange-brown. Their heads are slightly longer than wide and possess foveolate sculpture, which is characterized by numerous small pits, and rugae running from the front margin of the head to behind the eyes. There are scattered suberect to erect, flexuous setae over the entire cephalic and mandibular surfaces, and the antennal scapes. Their eyes are relatively small, with 6-8 ommatidiae from the anterior to posterior margins, located at the middle of the head on the lateral margins. They lack ocelli and possess 12-segmented antennae with three-segmented antennal clubs, and their scapes slightly surpass the posterior corners of their heads in full-face view. Their mandibles are triangular, with 6 to 7 small teeth along the inner border plus a longer apical tooth. Their mesosoma has foveolate sculpture and appears continuously arced, while the dorsum contains numerous flexuous setae of varying length. The promesonotum is fused, while the metanotal groove is noticeable. Their propodeum is angular, with short, slightly upturned, triangular propodeal spines, and distinctly set below the promesonotum. As with all myrmicines, they possess a two-segmented waist consisting of a petiole and a postpetiole. Their petiole is approximately twice as long as they are wide, thinly attached to the propodeum. The petiolar node is raised and conicaL in lateral view, while appearing nearly circular in dorsal view. The petiolar dorsum is shiny, while the sides are foveolate. The postpetiole is about as long as it is wide, appearing globular with some foveolate sculpture appearing on the sides of its posterior portion, although it is primarily glabrous and shiny. Some flexuous, elongate setae are present on both the petiole and postpetiole. The gaster is primarily smooth and shiny without obvious sculpture, and the same elongate, flexuous setae on the other parts of the body are present here and primarily facing backwards. As with most other species in the genus Pheidole, their sting is atrophied and nonfunctional. [4]

Major worker

P. navigans AntWeb major worker specimen Pheidole navigans casent0005742 profile 1.jpg
P. navigans AntWeb major worker specimen

The major workers of P. navigans are small, but larger than the minor workers. They have a head width between 0.80 millimetres (0.031 in) and 0.84 millimetres (0.033 in), head length between 0.74 millimetres (0.029 in) and 0.83 millimetres (0.033 in), scape length between 0.44 millimetres (0.017 in) and 0.46 millimetres (0.018 in), eye length between 0.11 millimetres (0.0043 in) and 0.13 millimetres (0.0051 in), mesosoma length, also known as Weber's length, between 0.52 millimetres (0.020 in) and 0.55 millimetres (0.022 in), spiracle length between 0.05 millimetres (0.0020 in) and 0.06 millimetres (0.0024 in), petiole width between 0.08 millimetres (0.0031 in) and 0.09 millimetres (0.0035 in), postpetiole width between 0.11 millimetres (0.0043 in) and 0.13 millimetres (0.0051 in), and pronotum width between 0.28 millimetres (0.011 in) and 0.31 millimetres (0.012 in). Their head, mesosoma, waist, and antennae are dark reddish-brown, while their gaster is darker brown to somewhat infuscated at their bases to darker brownish black further back. Their legs are lighter in color, presenting as orangish brown. Their head is greatly enlarged, characteristic of Pheidole, almost as wide as long, and larger than their mesosoma. The posterior margin of their head is deeply concave, and the corners of their head strongly rounded. The posterior corners of their head are shiny and lack sculpture, while the remainder of head are patterned with strong longitudinal rugae and some rugoreticulation between their eye and frontal carinae. The interrugal spaces are mostly shiny, while their clypeus and mandibles mostly lack sculpture. Their frontal triangle is prominent and shiny, and setae are present on their entire head. Scattered, but regular short, erect setae, mostly direct forwards and away from the head on and near cephalic margins, but setae located on the central portion of head are directed towards the midline. Their eyes are small and located below the midline of their head. They lack visible ocelli and have 12-segmented antennae with a three-segmented club. Their antennal scapes are short, about half of their head length. Their mandibles are triangular, enlarged, and lacking teeth except for the apical tooth. On the underside of their heads, five hypostomal teeth are present, with the central tooth wide and short, and the inner and outer teeth stout and triangular. The sides and dorsum of the mesosoma are foveolate, while transverse carinae are present on the pronotal dorsum. A few weak carina are present on the mesopleura and sides of the propodeum, and scattered erect setae are present on the promesonotal dorsum. A distinct promesonotal groove is present, as well as strong metanotal groove. The propodeum is set distinctly below the height of the promesonotum, while the propodeal declivity is concave. The propodeal spines are strong, somewhat finger-like, and directed upwards and slightly posteriorly. The propodeal spiracle is circular. Their waist is two-segmented, with scattered elongate, flexuous setae present on both segments. The petiole is about twice as long as wide in dorsal view, thinly attached to the propodeum. The petiolar node is raised and conical in lateral view, while it is somewhat rectangular in dorsal view. The sides of the petiole present foveolate sculpturing, while the petiolar dorsum is shiny. Their postpetiole is wider than long in dorsal view, with an elliptical postpetiolar node. Some foveolate sculpturing is present appearing on the sides of its posterior portion, while the remainder is shiny. Their gaster is smooth and shiny, lacking obvious sculpture. Scattered elongate, flexuous setae that are directed posteriorly are present on then gaster, while the sting is highly reduced and nonfunctional. [4]

Queen

P. navigans AntWeb queen specimen Pheidole navigans casent0104413 profile 1.jpg
P. navigans AntWeb queen specimen

The queens of P. navigans are small, but larger than the major workers. They have a head width between 0.81 millimetres (0.032 in) and 0.89 millimetres (0.035 in), head length between 0.74 millimetres (0.029 in) and 0.83 millimetres (0.033 in), scape length between 0.45 millimetres (0.018 in) and 0.49 millimetres (0.019 in), eye length between 0.24 millimetres (0.0094 in) and 0.27 millimetres (0.011 in), mesosoma length, also known as Weber's length, between 1.16 millimetres (0.046 in) and 1.22 millimetres (0.048 in), spiracle length between 0.08 millimetres (0.0031 in) and 0.11 millimetres (0.0043 in), petiole width between 0.23 millimetres (0.0091 in) and 0.25 millimetres (0.0098 in), postpetiole width between 0.37 millimetres (0.015 in) and 0.40 millimetres (0.016 in), pronotum width between 0.71 millimetres (0.028 in) and 0.75 millimetres (0.030 in), and front wing length between 3.80 millimetres (0.150 in) and 4.20 millimetres (0.165 in). Their head, including mandibles and antennae, mesosoma, and waist are dark reddish brown in coloration, their gaster dark brownish black, legs orangish brown, and wings clear with a dark stigma. Their head is slightly wider than long, widest at the back before gradually converging anteriorly, with the posterior margin of the head broadly concave. The entire head in full face and lateral view presents strong longitudinal rugosity, with transverse rugae on the posterior region of the head, and between eye and frontal carina forming a rugoreticulate pattern. The spaces between rugae are shiny in the central region of the head in full-face view and foveolate laterally approximately from the edge of the frontal carina to the margins of the head. The entire head bears numerous scattered erect setae, with the setae on sides of the head directed rearwards away from the head and setae on the central region of head directed towards the midline of the head. The scapes possess numerous erect setae, and the eyes are large. The length of the eyes is less than the distance between the eye and the posterior edge of the head. The ocelli are conspicuous and the frontal triangle is obvious, with cephalic rugae extending through it. The antennae are 12-segmented with a three-segmented club. The scapes are short with a length about two-thirds that of the head length. The mandibles are large, triangular, and edentate except for the apical tooth. The mesosoma is rounded, rectangular, dorsally flattened, overall shiny, and mostly lacking dorsal sculpture. The pronotum and propodeum present rugulate sculpture, while the smooth mesopleura mostly lack sculpture. The entire mesosoma bears numerous short, semierect curved setae that are directed posteriorly. The wings are translucent with a light brown tint and light brown venation. The forewing venation is typical of myrmicines, with a pterostigma and costal, basal, subbasal, discal, marginal, and two submarginal cells closed. The hindwing lacks a jugal lobe, and contains closed costal, basal, and subbasal cells. Its waist is two-segmented, with scattered elongate, flexuous setae present on both segments. The pedunculate petiole is about twice as long as wide in dorsal view. The petiolar node raised and conical in lateral view and somewhat rectangular in dorsal view. The sides of the petiole are foveolate with the dorsum shiny. The postpetiole is wider than long in dorsal view, almost twice as long as the petiole, with the postpetiolar node elliptical in dorsal view. Some foveolate sculpture is present posterolaterally, while the remainder of the postpetiole is shiny. The gaster is smooth and shiny, lacking obvious sculpture, and bearing scattered elongate, flexuous setae that are directed posteriorly. There is no functional sting apparatus as with the major and minor workers. [4]

Male

P. navigans AntWeb male specimen Pheidole navigans casent0104414 profile 1.jpg
P. navigans AntWeb male specimen

The males of P. navigans are also relatively small, similar in size to the workers. They have a head width between 0.59 millimetres (0.023 in) and 0.60 millimetres (0.024 in), head length between 0.47 millimetres (0.019 in) and 0.50 millimetres (0.020 in), scape length approximately 0.12 millimetres (0.0047 in), eye length between 0.29 millimetres (0.011 in) and 0.32 millimetres (0.013 in), mesosoma length, also known as Weber's length, between 1.12 millimetres (0.044 in) and 1.16 millimetres (0.046 in), petiole width between 0.15 millimetres (0.0059 in) and 0.16 millimetres (0.0063 in), postpetiole width between 0.20 millimetres (0.0079 in) and 0.23 millimetres (0.0091 in), pronotum width between 0.65 millimetres (0.026 in) and 0.68 millimetres (0.027 in), and front wing length between 2.80 millimetres (0.110 in) and 3.04 millimetres (0.120 in). Their head is dark brown, mesosoma, waist and gaster light brown, antennae and legs pale yellow brown in color, and wings clear and membranous. Their head is longer than wide, in full-face view widest at the rear. Measuring from outer edge of eye to eye and excluding the eyes, the head is somewhat hexagonal in shape. The clypeal area possesses some longitudinal carinae, while the remainder of the head contains strong, tightly woven rugoreticulation. Numerous erect setae of varying lengths are present on the head. The antennal scapes bear short, semierect setae, while the funiculus is densely pubescent. The mesosoma, waist, and gaster are mostly shiny and lacking sculpture, with numerous short, semierect setae present. The eyes are huge, more than half the length of the head, and placed on the lower half of head. The ocelli large and protruding, while the antennae are 13-segmented and beaded. The antennal scapes are short and rectangular, while the pedicel is globular. The mesosoma is rounded, rectangular, shiny with limited sculpture, and bears numerous, short, erect setae. The pronotum is shaped somewhat trapzoidally, the mesoscutum is broadly rounded towards the front and overhanging the pronotum, and the mesoscutellum has a declivious face that is distinctly elevated above the propodeum. The propodeal declivity of P. navigans males is irregular. The wings are pale, translucent, and hyaline with the veins light brown. The forewing venation is typical of myrmicines with a pterostigma and costal, basal, subbasal, discal, and two submarginal cells closed. The hindwing lacks a jugal lobe and contains closed costal, basal, and subbasal cells. The waist is two-segmented with limited sculputure and numerous erect setae present. The petiole is pedunculate and elongate, with the petiolar node barely raised. The postpetiole is short, rounded, and somewhat circular. The gaster is shiny, with numerous erect setae present. [4]

Ecology

A P. navigans dealate queen found in Texas. Pheidole navigans 389462680.jpg
A P. navigans dealate queen found in Texas.

In Florida, they are rather common and nest in various microhabitats, including under boards, at the bases of fence posts and oak trees, in rotten wood, wall crevices, hollow twigs, nuts, leaf littler, and occasionally arboreally. They are rather generalist and consume a wide variety of foods, from small arthropods and seeds to sugars and human food. However, they are not reported to be major house pests. [3] Mature colonies contain over 100 majors and 500 workers, while nuptial flights mostly occur in July. They are monogynous but undergo pleometrosis, in which multiple queens may cooperate to found a colony yet only one survives past colony founding, and also practice dependent colony foundation, also termed budding, in which existing colonies split to form autonomous daughter groups which separate to form new colonies. Their nest chambers have small openings and are built with soil and debris. Major workers often forage alongside workers, and foraging is done very close to the nest. [2] [9] They are attracted to both sugar and protein baits, and respond extremely quickly to tuna baits by recruiting both major and minor workers. [12] [13] Although they are widespread in Florida and other parts of the southeastern United States, their ecological effects are not well-known. In Florida, they appear to be partially successful in displacing their native congeners Pheidole dentigula and Pheidole bilimeki . They may compete for space for nesting sites with native species in the southeastern United States, like those in the genera Aphaenogaster , Camponotus , Nylanderia , Solenopsis , Hypoponera , Strumigenys , Brachymyrmex , and others. [4]

Distribution

Pheidole navigans is native to the Neotropics and established as invasive in numerous regions like California, the southeastern United States, Madeira, Tenerife, Spain, Hawaii, and Vanuatu, however the species has possibly spread further since then. They were often misidentified in their invasive range, most notably as Pheidole moerens before 2015. They are classified as category D2 invasives, meaning that they are capable of surviving, reproducing, and sustaining a stable population in introduced regions. They are capable of exerting strong propagule pressure, appearing frequently in interception records worldwide. In their presumed native range, they have been recorded from Brazil, Colombia, Venezuela and Paraguay. [2] [14] [15] They were first recorded as an established population outside of their native range in Alabama in 1967 [16] and Florida in 1975, [3] and have since been slowly expanding their range throughout the southeastern United States, later spreading to Georgia, Mississippi, North Carolina, and Texas by 2018. Elsewhere, they have spread to southern California by 1995, Hawaii by 2003, and Vanuatu in 2011. [17] More recently, the species was first recorded in Madeira in 2014, where they may be successful in displacing other widespread invasive species in the region like Pheidole megacephala and Linepithema humile near the city of Funchal. [13] In Bermuda, they were first recorded as an established population in 2016, although interception reports date back to 2004. [18] The species was first recorded as an established population in the Canary Islands by 2020 in a farming area in the northern part of Tenerife [19] and continental Europe in 2021, represented by a series of minor workers collected in Málaga, Spain. [12]

Table

Established invasion history of Pheidole navigans
RegionSub-regionFirst published recordReferenceNotes
Atlantic Islands Bermuda 2016 [18]
Madeira 2014 [13]
Tenerife 2020 [19]
Continental Europe Spain 2021 [12]
Eastern North America Alabama 1967? [16] as P. moerens, exact date not specified
Florida 1975? [3] as P. moerens, exact date not specified
Georgia 2018 [20]
Louisiana 2002-2004 [21] as P. moerens, exact date not specified
Mississippi 2005 [22] as P. moerens
North Carolina 2006-2011 [23] as P. moerens, exact date not specified
Texas 2003? [24] as P. moerens, exact date not specified
Mexico Hidalgo 1993? [25] exact date not specified
Veracruz 1901 [25]
Pacific Islands Hawaii 2000 [26] as P. moerens
Vanuatu 2011 [17] as P. moerens
Western North America California 1995 [17] as P. moerens

References

  1. 1 2 Bolton, Barry. "Pheidole navigans Forel, 1901 valid". AntCat. antcat.org. Retrieved 19 December 2024.
  2. 1 2 3 4 5 6 Sarnat, Eli M.; Fischer, Georg; Guénard, Benoit; Economo, Evan P. (9 December 2015). "Introduced Pheidole of the world: taxonomy, biology and distribution". ZooKeys (543): 1–109. Bibcode:2015ZooK..543....1S. doi: 10.3897/zookeys.543.6050 . PMC   4714327 . PMID   26798286.
  3. 1 2 3 4 Deyrup, Mark; et al. (2000). "Exotic Ants in Florida". Transactions of the American Entomological Society. 126 (3+4): 293–326. Retrieved 30 May 2025.
  4. 1 2 3 4 5 6 7 8 9 MacGown, Joe A. "Subfamily MYRMICINAE Tribe ATTINI Pheidole navigans Forel, 1901 (=Pheidole moerens Wheeler, 1908, auct.) Navigating big-headed ant". Mississippi State University: Mississippi Entomological Museum. Mississippi State University. Retrieved 23 January 2026.
  5. 1 2 Wilson, Edward O. (March 2003). Pheidole in the New World: A dominant, hyperdiverse ant genus. Harvard University Press. pp. 1–794. ISBN   9780674002937 . Retrieved 23 January 2026.
  6. ↑ Economo, Evan P.; Klimov, Pavel; Sarnat, Eli M.; Guénard, Benoit; Weiser, Michael D.; Lecroq, Beatrice; Knowles, L. Lacey (22 October 2014). "Global phylogenetic structure of the hyperdiverse ant genus Pheidole reveals the repeated evolution of macroecological patterns". Proc Biol Sci: 1–10. doi:10.1098/rspb.2014.1416. PMID   25429013 . Retrieved 26 January 2026.{{cite journal}}: Cite journal requires |journal= (help)
  7. ↑ "Species: Pheidole navigans Forel, 1901". AntWeb. California Academy of Sciences. Retrieved 30 May 2025.
  8. ↑ Shattuck, Steven O. "Pheidole navigans". AntWiki. antwiki.org. Retrieved 30 May 2025.
  9. 1 2 Naves, Marcio A. (March 1985). "A monograph of the genus Pheidole in Florida (Hymenoptera: Formicidae)". Insecta Mundi. 1: 53–90. Retrieved 24 January 2026.
  10. ↑ Wilson, Edward O. (20 March 1984). "The Relation between Caste Ratios and Division of Labor in the Ant Genus Pheidole (Hymenoptera: Formicidae)". Behavioral Ecology and Sociobiology. 16 (1): 89–98. Retrieved 26 January 2026.
  11. ↑ Lillico-Ouachour, Angelica; et al. (5 January 2018). "Internal head morphology of minor workers and soldiers in the hyperdiverse ant genus Pheidole". Canadian Journal of Zoology. 96 (5): 383–392. doi:10.1139/cjz-2017-0209 . Retrieved 26 January 2026.
  12. 1 2 3 Reyes-López, Joaquin-Luis; Rodriguez Reyes, Marta (March 2022). "An unstoppable process: The first record of Pheidole cfr navigans Forel, 1901 (Hymenoptera, Formicidae) for continental Spain (Southern Europe)". Boletin - Asociacion Espanola de Entomologia. 46 (1–2): 147–149. doi:10.70186/baeeYVFH2901 . Retrieved 24 January 2026.
  13. 1 2 3 Guillem, Rhian; Bensusan, Keith (30 June 2022). "Three new exotic species of ants (Hymenoptera, Formicidae) for Madeira, with comments on its myrmecofauna". Journal of Hymenoptera Research. 91: 321–333. doi: 10.3897/jhr.91.81624 . Retrieved 23 January 2026.
  14. ↑ Economo, Evan; Guénard, Benoit. "antmaps.org Pheidole navigans". antmaps.org. Okinawa Institute of Science and Technology. Retrieved 19 December 2024.
  15. ↑ Violante, Guilherme M.; Lira, Paula K.; Delabie, Jacques H. C.; Queiroz, Jarbas M. (23 September 2025). "First records of two non-native ant species in Brazil: the starting point for avoiding biological invasions". Studies on Neotropical Fauna and Environment: 1–5. doi:10.1080/01650521.2025.2548565 . Retrieved 25 January 2026.
  16. 1 2 Smith, Marion R. (February 1967). "Hymenoptera of America north of Mexico. Synoptic catalog. Second supplement. United States Department of Agriculture". Agriculture Monograph 2. suppl. 2: 1–584. Retrieved 24 January 2026.
  17. 1 2 3 Sarnat, Eli M.; Rabeling, Christian; Economo, Evan P.; Wilson, Edward O. (20 October 2014). "First record of a species from the New World Pheidole flavens-complex (Hymenoptera: Formicidae) introduced to the southwestern Pacific". BioInvasions Records. 3 (4): 301–307. doi:10.3391/bir.2014.3.4.13 . Retrieved 24 January 2026.
  18. 1 2 Wetterer, James K. (27 February 2017). "Invasive ants of Bermuda revisited". Journal of Hymenoptera Research. 54: 33–41. doi: 10.3897/jhr.54.11444 . Retrieved 23 January 2026.
  19. 1 2 Hernández-Teixidor, David; et al. (28 March 2020). "Six new non-native ants (Formicidae) in the Canary Islands and their possible impacts". Journal of Applied Entomology. 144 (6): 434–441. doi:10.1111/jen.12751. hdl:10261/206341 . Retrieved 24 January 2026.
  20. ↑ Braman, Charles A.; Forschler, Brian T. (2018). "Survey of Formicidae Attracted to Protein Baits on Georgia's Barrier Island Dunes". Southeastern Naturalist. 17 (4): 645–653. doi:10.1656/058.017.0412 . Retrieved 24 January 2026.
  21. ↑ Hooper-Bùi, Linda M.; Dash, Shown T. (November 2008). "Species Diversity of Ants (Hymenoptera: Formicidae) in Louisiana". Annals of the Entomological Society of America. 101: 1056–1066. doi:10.1603/0013-8746-101.6.1056 . Retrieved 24 January 2026.
  22. ↑ MacGown, Joe A.; Hill, JoVonn G. (December 2010). "A New Species of Pyramica (Hymenoptera: Formicidae) from Mississippi, U.S.A." Florida Entomologist. 93 (4): 571–576. doi:10.1653/024.093.0415 . Retrieved 24 January 2026.
  23. ↑ Guénard, Benoit; McCaffrey, Katherine A.; Lucky, Andrea; Dunn, Rob R. (16 November 2012). "Ants of North Carolina: an updated list (Hymenoptera: Formicidae)". Zootaxa. 3552: 1–36. Retrieved 24 January 2026.
  24. ↑ Wilson, E. O. (2003). Pheidole in the New World. A dominant, hyperdiverse ant genus. Cambridge, Mass.: Harvard University Press. pp. 561–562. ISBN   9780674002937.
  25. 1 2 Rosas-Mejía, Madai; et al. (6 November 2020). "Alien ants (Hymenoptera: Formicidae) in Mexico: the first database of records". Biol Invasions. 23: 1669–1680. doi:10.1007/s10530-020-02423-1 . Retrieved 26 January 2026.
  26. ↑ Gruner, Daniel S.; et al. (2003). "Two ant species (Hymenoptera: Formicidae) new to the Hawaiian Islands" (PDF). Bishop Museum Occasional Papers. 73: 35–40. Retrieved 24 January 2026.

Further reading

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