There once was a time when, in Texas, the little black ant (Monomorium minimum Buckley 1866) did not assume momentous significance as a pest of human dwellings. Leland Howard, in his 1914 monograph on the insects, indicated that — even then — the little black ant was much inferior as a pest to what he called the “little red ant” (Monomorium pharaonis Linn. 1758).
That latter species — named by Linnaeus in the 18th century for the plague-like manner in which it invades habitations, hospitals, nursing homes, and commercial kitchens (some believe the Master Taxonomist of Olde thought this ant represented one of the biblical plagues cited in Exodus ch. 8-13) — was in the late 1970’s the ant scourge du jour, not only in Texas but throughout the United States. By the early 1980’s, though, the red imported fire ant, or RIFA (Solenopsis invicta Buren 1972) ascended the highest throne of ant infamy. The RIFA had spread steadily outward from the docks of Mobile, Alabama, where it first arrived in the U.S. from Brazil in the 1930’s. The destructive nature of its relentless advance, unchecked here by the natural predators that kept it under control in its native setting, was already legendary (though its northward progress appears to have been blocked by the winter frost line).
Still, among the problematic ants of the 1980’s and most of the 1990’s, the pharaoh ant remained an important a pest of homes and medical facilities. Its importance as a pest was bolstered by suspicions that this ant is capable of transmitting disease, via ordinary cross-contamination and other mechanisms, within hospitals and nursing homes. Though little in the way of proof ever emerged to bolster that suspicion, the notion made enough sense to stand on its own. For that and a host of other reasons, tens of millions of dollars were invested by the pesticide industry in research programs, both internally and as research grants to the halls and labs of academia, to find effective control measures that could be employed by pest managers and homeowners alike.
Of the various pesticides under analysis, one of the most promising was a rather non-exotic synthetic terpenoid — an isoprenoid derived initially from botanical sources, usually with a pleasant, somewhat floral fragrance and a well-established mode of action — known as methoprene (11-methoxy-3,7,11-trimethyl-2,4-dodecadienoic acid 1-methylethyl ester). Methoprene functions in arthropod biology as a juvenile growth hormone analog. When applied to the nymphs of developing larvae, particularly of insects that undergo complete metamorphosis (such as ants and a number of other social insects), development is permanently arrested before the adult stage is reached. Thus methoprene is capable of bringing the propagation of entire ant colonies to a halt.
The ZOËCON Corporation (now Wellmark International), which was founded in 1968 by a group of prominent scientists — among them Dr. Carl Djerassi, one of the developers of the first birth control pills for women — first registered methoprene for horn fly and mosquito control in 1975. Later, in 1980, the company registered the same molecule for flea control, and by the mid-1980’s, in a unique formulation (Pharaoh-Rid®) specifically targeting pharaoh ants. Other pesticide manufacturers tested and marketed carbohydrate-and-protein-based ant baits infused with various toxicants and growth regulators.
Additionally, a bewildering array of novel, mostly exotic, pesticide molecules arrived on the scene during this same period, making it difficult for practitioners in the field to keep abreast of modern technology. Some amongst us (myself included), in the midst of the perplexity that so many “advances” produced, questioned aloud whether we should be considered beneficiaries or victims of this flurry of progress.
Answers, for those who sought them, were not long in coming.
The carbamate growth regulator with the chemical name fenoxycarb — ethyl N-[2-(4-phenoxyphenoxy)ethyl]carbamate — was marketed extensively in the late 1980’s as a somewhat exotic substitute for methoprene and a close cousin, hydroprene; this novel molecule, we were told, had an excellent toxicity profile, touted to be “similar to that of common table salt,” making it as safe around humans and pets as terpenoids like methoprene and hydroprene. Plus, unlike these terpenoids, fenoxycarb was effective against all insects, while methoprene was only effective against those with complete metamorphosis, and and hydroprene only worked with insects with incomplete metamorphosis. Those like me who got excited about low-toxicity products tried fenoxycarb out right away. We liked it’s apparent effectiveness, and worked it into our arthropod management protocols. When it appeared to work at least as well as methoprene and hydroprene, it made sense to switch between these IGRs each month, as an aid to lessening the risk of creating resistant pest strains.
Several years later the EPA announced their latest toxicological findings. Fenoxycarb, far from being “as safe as table salt,” was now considered a class-B2-carcinogen…
This was a bitter pill to swallow. Worse, similar examples soon surfaced, reminding the observant practitioner that unheralded risks sometimes attended the use of “new and safe” chemicals that came into the pesticide marketplace. It bears pointing out that few of these cases presented clear-cut risks that absolutely must be avoided at all costs. Controversy swirled about every negative finding, enough to fill a good-sized book, placing the practicing pest manager squarely on the horns of a dilemma.
One more example before moving on: The fluoridated organic compound commonly known as sulfluramid — N-ethyl-1,1,2,2,3,3,4,4,5,5,6,6,7,7,8,8,8-heptadecafluoro-1-octanesulfonamide — was and still is (until 2014 or 2016, according to various sources) the active ingredient in a variety of ant and termite baits that were first marketed in the 1980’s. Again, sulfluramid, though an exotic molecule, was initially promoted for having an unusually safe toxicity profile.
Many years later EPA judged this highly persistent compound to be potentially teterogenic, and toxic to developing human fetuses, human reproductive organs, and human kidneys…
Ingestion of the contents of a single ant bait packet by a human child has, according to some investigators, the potential of causing permanent damage to the child’s reproductive organs. Absorption of sulfluramid through the skin of adult humans who come into direct contact with the active ingredients of baits containing sulfluramid has the potential of producing (1) injury to developing fetuses carried by pregnant women, (2) damage to the adult’s reproductive organs, and (3) damage to the adult’s renal (urinary) organs. Again, the studies that led to such judgments are controversial and subject to various interpretations. However, the EPA considered them significant enough to require phasing out all sulfluramid-based products, but not significant enough to force a halt to the manufacture and sale of those products before existing stockpiles of baits and technical grade materials are exhausted.
My point in discussing these conundrums here is to demonstrate the range and depth of the difficulties faced — then and now — by those in the pest management community who must decide the best way to deal with pharaoh ant and similar types of arthropod infestations. One has to weigh the potential risks posed by the remedy against the potential risks represented by the target pest, and as the above should make clear neither risk is so adequately defined as to enable even a careful judge to view the scales with absolute clarity of vision. Ultimately the question distills down to which way one should err… A. On behalf of a new and exotic molecule’s proponent, and their promises of greater profits? Or B. With humanity’s and one’s own health concerns, avoiding the newly announced exotics until their safety is firmly established, while exhausting every known, firmly safe measure at one’s disposal? For me the answer was clear and unequivocal.
The diagram at left is a generalized anatomical sketch of an ant, courtesy of the author, Mary Villarreal, Hamburg, Germany, who released it into the public domain for all to use. Thank you, Ms. Villarreal. We will be referring to this diagram in the anatomical descriptions of the little black ant, below. As with all images 0n this website, this image can be enlarged for more detailed viewing by moving your cursor to it and clicking.
Suffice it to say that, over a period of about fifteen years, enough excellent, good, and fair remedies reached the market to make a solid dent in the pharaoh ant populations throughout the United States. As a result that species, though not completely decimated, has today lost much of its “ferocity” as a pest of habitations and medical facilities.
Which brings us back to the little black ant (Monomorium minimum).
Unlike the pharaoh ant, the little black ant (or LBA, as some are wont to call it) historically was not accustomed to living its entire life cycle within the walls of man-made structures. Nor did it have a reputation for producing polygyne colonies (with more than one queen) or for “budding” when the population of a given colony dropped precipitously low (as may occur when pesticides wipe out large numbers of the colony’s worker ants).
Budding is an adaptational defense to predation that leads the colony’s brood workers to split the nest into several satellite “buds” that — along with at least one queen — march off in different directions to found new colonies elsewhere.
When pesticides are used against ant colonies prone to budding, the end result does not reduce the scope of the original infestation, but rather enlarges it. Overnight single colonies become fragmented into a multitude of budded colonies that, though initially small, speedily become as large as, or even larger than, the original colony they spring from. But, as previously noted, the little black ant was not known for exhibiting this behavior, at least as long as the pharaoh ant remained its superior. When that changed, toward the end of the 1990’s, the little black ant began to take over the territories — and at least some of the habits — of the pharaoh ant.
Today this tiny ant has become nearly as important a pest of homes and medical facilities as the pharaoh ant of old. Evidence is now beginning to surface that suggests the little black ant is adapting to its new ascendancy in much the same way as its predecessor, the pharaoh ant, with polygyne colonies capable of living entirely within man-made structures, prone to budding when threatened. However, its dietary requirements and behavior differ from that of the pharaoh ant. Those differences are significant enough that measures successful against the pharaoh ant are not effective against the little black ant. Thus, curbing its infestations in homes and medical facilities require new approaches.
EntomoBiotics Inc. is currently studying a number of LBA infestations in Round Rock, Texas, as well as in several other locations throughout Texas. Unique protocols, using non-toxic habitat modifiers and insect growth regulators (IGRs) are being developed in the process, and will be tested in each of these locations to determine the best way to entirely resolve LBA infestations without exposing humans or their pets to pesticide toxicants in the process.
This posting is a work in progress, and will be fleshed out as time permits. Details of the little black ant’s anatomical characters, and the exhaustive measures we are taking to deal with its infestations — focusing on the safest products presently on the market — will soon be added.
General Anatomy: Samuel Botsford Buckley’s 1866 analysis of this and 66 other species of ants was carried out while he and a group of associate geologists were conducting a geological survey of Texas. The ants, he wrote, secured his interest because of “their wonderful works, which, in Texas, evince so much intelligence, industry, and perseverance as to attract the attention of the most ordinary observer. Finally, although pressed with other duties, we gave a large portion of our leisure to collecting them and observing their habits, which we continued to do up to the present time, with much pleasure and satisfaction.” Regarding the species before us, he wrote the following words (pp. 338-339 of Buckley, S. B. 1866-7. Descriptions of new species of North American Formicidae; Proc. Entomol. Soc. Phila. 6: 152-350):
43. Myrmica (Monomarium) minima, n. sp.
Female. Length 0.22 inch. — Color jet-black, the whole body smooth and shining; legs and margins of the segments of the abdomen brownish-black, or sub-hyaline; head triangular, rounded above; occiput not emarginate, the posterior angles rounded and of about the same width as the thorax; mandibles small, curved, acute and triangular; eyes small, circular, lateral, and placed about midway of the head; antennae long, filiform, and enlarged anteriorly; thorax with its sides compressed, the divisions but slightly marked; mesothorax the highest; scales large, the front one slightly compressed, inclined forwards, and subacute, the other rounded; abdomen large, ovate, oblong, upper surface of the body and head sprinkled with a few gray hairs; slender in outline.
Worker. Length 0.06 inch. — Black; head wider than the thorax; pedicle long; abdomen but little larger than the head, ovate and acute; otherwise like the female.
Lives under stones and beneath the bark of decaying trees. It also excavates into the ground, but two or three inches below the surface. Common in Central Texas, where it is often seen going in ranks on the ground or trees. It is rather slow in its movements.
When, in 1901, William Morton Wheeler, Professor of Zoology at the University of Texas, published a lengthy analysis of Mr. Buckley’s 1966 paper, he characterized Buckley’s descriptive notes as “fearfully and wonderfully made.” By this he alluded, as the preamble to his analysis made clear, to the taxonomic fiasco that Buckley’s 1866 paper foisted upon the scientific community of that day. Buckley’s training in entomology was, after all, practically nonexistent. Furthermore, Wheeler points out, “there are scarcely any insects more difficult of analysis and description than the Formicidae.” It annoyed Wheeler to have to sift through Buckley’s amateurish scribblings (he often mistook one sex for another, and would go to great pains to describe worthless characters while ignoring features of greater import) and winnow the paltry good from the chaff. But Wheeler did so with a serious eye to objectivity, and in the end — though finding many of Buckley’s “new species” to be of ants already well-known at the time, and others described so poorly as to be impossible to recognize in the extant fauna — he succeeded in crediting a number of the old geologist’s descriptions as new: “In the following pages I have seen fit to cite Buckley’s species seriatim with their conjectural identifications. I am not aware of having made any effort to strain a point in favor of Buckley, — for I hold that no zoologist deserves a particle of credit for writing a worse than useless description, — but if I have succeeded in throwing a little light on some of his species, I would wish this to be regarded as a tribute to a pioneer naturalist who long ago searched the woods and hill-slopes of Texas, collecting ants and observing their ways ‘with much pleasure and satisfaction‘”.
A detailed anatomical analysis of this species is in preparation.
- Kingdom Animalia (ahn-uh-MAYHL-yuh) — first described in 1758 by the Swedish taxonomist Carolus Linnaeus [23 May 1707 – 10 January 1778], using the Latin word animal = “a living being,” from the Latin word anima = “vital breath”, to refer to multicellular, eukaryotic organisms whose body plans become fixed during development, some of which undergo additional processes of metamorphosis later in their lives; most of which are motile, and thus exhibit spontaneous and independent movements; and all of whom are heterotrophs that feed by ingesting other organisms or their products;
- Phylum Arthropoda (ahr-THROPP-uh-duh) — first described in 1829 by the French zoologist Pierre André Latreille [November 20, 1762 – February 6, 1833], using the two Greek roots αρθρον (AR-thrawn) = jointed + ποδ (pawd) = foot, in an obvious reference to animals with jointed feet, but in the more narrow context of the invertebrates, which have segmented bodies as well as jointed appendages;
- Class Insecta (Linnaeus, 1758) — named using the Latin word insectum, a calque of the Greek word ἔντομον ( EN-toh-mawn) = “(that which is) cut into sections”; comprised of arthropods with chitinous external (exo-) skeletons, a three part body composed of a distinct head, thorax, and abdomen, the midmost part having three pairs of jointed legs, and the foremost part having a pair of compound eyes and antennae;
- Order Hymenoptera (hye-muhn-OPP-turr-uh) — first described in 1758 by the Swedish taxonomist Carl Linnaeus (1707 – 1778), who combined the Greek words ὑμήν (pron. humēn) = “membrane” + πτερόν (TARE-awn) = “wing”, thus ὑμενόπτερος (hew-men-OPP-tehr-ose) = “membrane-winged” to refer to insects with membranous wings, specifically the sawflies, wasps, bees, and ants; this is one of the largest orders of insects, and includes over 130,000 species;
- Family Formicidae (fohr-MISS-uh-dee) — first described in 1809 by the French zoologist Pierre André Latreille (1762 – 1833), from the Latin formica = “ant” to refer to hymenopteran insects that have elbowed antennae and a narrow waist that separates the thorax from the abdomen with a node-like petiole; at present, 20 distinct subfamilies of ants are recognized:
- Subfamily Aenictogitoninae: a subfamily comprising a single genus, Aenictogiton, with seven known species of rarely collected ants found in Central Africa with morphological and phylogenetic affinities to the army ant genus Dorylus; only males have been collected, and nothing is known about their workers, queens or behavior;
- Subfamily Agroecomyrmecinae: characterized by the following derived traits (see Bolton 2003): mandibular masticatory margins oppose at full closure but do not overlap; eye at extreme posterior apex of deep antennal scrobe; antennal sockets and frontal lobes strongly migrated laterally, far apart and close to lateral margins of head; abdominal segment IV with complete tergosternal fusion; sternite of abdominal segment IV reduced, tergite much larger than sternite and strongly vaulted;
- Subfamily Amblyoponinae (including the subfamily Apomyrminae): mostly specialized subterranean predators, comprised of a single genus of two species native to California; characterized by the following traits (see Bolton 2003): workers of this ant subfamily — which was formerly considered a tribe within the subfamily Ponerinae — exhibit the following characters: eyes small or absent, if present situated behind the mid-length of side of head; anterior margin of clypeus with specialized dentiform setae; the promesonotal suture is flexible; the petiole is broadly attached to abdominal segment 3 and is absent a distinct posterior face; the postpetiole is absent; a sting is present and is well developed.
- Subfamily Aneuretinae: this subfamily is comprised of a single extant tribe containing a single extant genus and a single extant species (several extinct tribes, genera, and species have also been described), namely the Sri Lankan relict ant (Aneuretus simoni); this is one of the few ant species considered endangered;
- Subfamily Cerapachyinae: a subfamily of 5 genera and 217 recognized species, distributed throughout the tropics; they possess spines on the pygidium; their antennae are short and thick; and they lack dorsal thoracic structures; they prey on other ant species;
- Subfamily Dolichoderinae: presently not divided into tribes, but comprised of 24 genera, including the Argentine ant (Linepithema humile), the erratic ant (Tapinoma erraticum), the odorous house ant (Tapinoma sessile), and cone ants in the genus Dorymyrmex; these ants are distinguished by having a single petiole, absent a post-petiole, and lacking a sting but possessing an apical slit-like orifice at the posterior abdomen instead of the round acidopore encircled by hairs typical of the Formicinae subfamily;
- Subfamily Ecitoninae (incl. “Dorylinae” and “Aenictinae”): New World and Old World army ants; in the New World, these ants are found in the tribes Cheliomyrmecini (containing the single genus Cheliomyrmex) and Ecitonini (containing the four genera Neivamyrmex, Nomamyrmex, Labidus, and Eciton); the genus Neivamyrmex — the largest of all army ant genera — contains more than 120 species, all native to the United States; the predominant species of the genus Eciton, E. burchellii, has been given the common name “army ant” and is considered the archetypal species; Old World army ants are usually divided into two tribes, Aenictini and Dorylini, but are often treated as a single tribe, Dorylini, alone; each contains a single genus; the genus Aenictus contains over 100 species, and the genus Doryus contains the aggressive “driver ants”, of which 70 species are known;
- Subfamily Ectatomminae: In North America a single genus, Gnamptogenys, is represented; that genus is not native to North America but has been introduced;
- Subfamily Formicinae: (fohr-mih-SEE-nee) — first described in 1836 by the French entomologist Amédée Louis Michel le Peletier, comte de Saint-Fargeau (1770 – 1845), usu. referred to as Lepeletier, from the Latin formica = “ant” to refer to a subfamily of ants whose evolutionary development is not as robust as most other subfamilies, e.g., they generally retain such primitive features as pupal cocoons, ocelli in workers, and a lesser tendency toward reduced palpal or antennal segmentation; all formicines have reduced stings and enlarged venom reservoirs, with a venom gland that is uniquely specialized to produce formic acid, and a one-segmented petiole having the form of a vertical scale;;
- Subfamily Heteroponerinae:
- Subfamily Leptanillinae: comprised of two tribes, the Anomalomyrmini (two genera, seven species) and Leptanillini (three genera, 41 species); within the tribe Leptanillini the larva provide their hemolymph as food to the queen through specialized processes on their prothorax and third abdominal segment; this behavior resembles that of the unrelated Adetomyrma, also called Dracula ants, which actually pierce their larvae to get at the body fluids; ants in the genera Leptanilla and Phaulomyrma are minute, yellow, blind, and subterranean;
- Subfamily Leptanilloidinae: 1 tribe, 3 genera, 15 species;
- Subfamily Martialinae: 1 genus containing a single species, Martialis heureka, discovered in 2000 from the Amazon rainforest near Manaus, Brazil, and placed as the sole member of a new subfamily (Martialinae); the generic name, which means “from Mars,” refers to its unusual “out-of-this-world” morphology; the species epithet heureka honors the surprise that accompanied its discovery; it is the oldest known extant species of ants;
- Subfamily Myrmeciinae (incl. “Nothomyrmeciinae”): once distributed worldwide but now restricted to Australia and New Caledonia; one of several ant subfamilies which possess gamergates, i.e., female worker ants which are able to mate and reproduce, thus sustaining the colony after the loss of the queen; formerly composed of a single genus, Myrmecia, but revised (Ward & Brady 2003) to include two tribes and four genera; three additional genera, one form genus, and nine species were later described (Archibald, Cover and Moreau 2006) from the Early Eocene of Denmark, Canada, and Washington;
- Subfamily Myrmicinae: approximately 130 genera in 23 tribes, and 10 additional genera not assigned to specific tribes, all cosmopolitan; the pupae lack cocoons; some species retain a functional sting; the petioles have two nodes; nests are permanent, in soil, rotting wood, under stones or in trees; the subfamily includes leaf cutters (tribe Attini), acrobat (tribe Crematogasterini), harvester (tribe Myrmicini), big-headed (tribe Pheidolini), and fire (tribe Solenopsidini) ants;
- Subfamily Paraponerinae: comprised of a single genus, Paraponera, containing a single species (Paraponera clavata), known as the lesser giant hunting ant, the conga ant, or the bullet ant (so named for its powerful sting); this ant inhabits lowland rainforest, from Nicaragua and eastern Honduras, and south to Paraguay; the ant is called “hormiga veinticuatro” by locals to refer to the 24 hours of pain following each sting;
- Subfamily Ponerinae: about 1,600 species in 28 extant genera, including Dinoponera gigantea, which is one of the largest species of ant found in the world; distinguished from other formicine subfamilies by their constricted abdomens;
- Subfamily Proceratiinae: similar to Ponerinae but the promesonotal suture is fused and the frontal lobes, elevated rather than transverse, are frequently reduced; antennal sockets are exposed in frontal view; in most species abdominal tergite 4 is much enlarged and vaulted, while abdominal sternite 4 is reduced; these are specialized predatory ants that are represented in California by a single species;
- Subfamily Pseudomyrmecinae: three genera of slender, wasp-like ants that forage alone and readily sting when molested;
- Subfamily Myrmicinae (murr-mih-SEE-nee) — approximately 130 genera in 23 tribes, and 10 additional genera not assigned to specific tribes, all cosmopolitan; the pupae lack cocoons; some species retain a functional sting; the petioles have two nodes; nests are permanent, in soil, rotting wood, under stones or in trees; the subfamily includes leaf cutters (tribe Attini), acrobat (tribe Crematogasterini), harvester (tribe Myrmicini), big-headed (tribe Pheidolini), and fire (tribe Solenopsidini) ants;
- Tribe Solenopsidini (soh-luhn-opp-suh-DEE-nee);
- Genus Monomorium (mohn-oh-MOHR-ee-uhm) — first described in 1855 by the Austrian entomologist Gustav Mayr (1830-1908), possibly using the Greek roots μονος (MOHN-os) = single + μορφη (MOHR-fee) = shape, form, in conjunction with the Greek diminutive suffix -ιυμ (ee-uhm) = denoting the quality or nature of, to refer to the fact that most ants in this genus, contrary to many others in the tribe Solenopsidini, are monomorphic (though some are decidedly polymorphic);
- Species minimum (MEHN-uh-muhm) — first described by the American naturalist and geologist Samuel Botsford Buckley (1809-1884) in 1866, using the epithet Monomarium minima in reference to the diminutive size of the workers of this species; Buckley, a curious student of nature whose even more curious descriptions of ant species became the source of much scientific consternation after his death, was born in New York in 1809 and died in Austin, Texas on 18 February, 1884; during his lifetime he made botanical collections in Virginia and Illinois, traveled extensively through the south discovering and describing twenty-four new species of plants, including a new genus, which was named Buckleya; he discovered and obtained in Alabama a nearly complete skeleton of a zeuglodon (an extinct carnivorous marine mammal related to whales, porpoises, and dolphins); in 1843 he studied at the College of physicians and surgeons, New York, and in the same year, in an expedition to Florida, discovered thirteen new species of shells; in 1858 he determined barometrically the height of several mountains in Tennessee and North Carolina; one of them, Mount Buckley, North Carolina, still bears his name; in 1859-1860 he collected materials for a supplement to Michaux and Nuttall’s Sylva; he was assistant geologist and naturalist of the Texas geological survey in 1860-1861, and from 1862 till 1865 was connected with the United States sanitary commission; he served as state geologist of Texas from 1866 till 1867, during which time he collected and described 67 species of ants, many of which had not been described previously; he assumed the post of Texas state geologist again from 1874 till 1877, and prepared two geological maps of the state, showing by his investigations that Texas had deposits of iron and coal of much greater extent than had been supposed; in 1871-18722 he was scientific editor of the “State Gazette,” Austin, and from 1877 till 1881 prepared a work on the geology and natural history of the state.
- Blair, W. F. 1949. The biotic provinces of Texas. Texas Journal of Science 2(I):93-117.
- Bolten, Barry. 1994. Identification Guide to the Ant Genera of the World. Harvard University Press.
- Bolten, Barry. 2003. Synopsis and Classification of Formicidae. Memoirs of the American Entomological Institute, Vol. 71.
- Borror, Donald J., and Richard E. White. 1970. A Field Guide to Insects, America north of Mexico. Houghton Mifflin Company
- Hansen, Laurel D., and John H. Klotz. 2005. Carpenter Ants of the United States and Canada. Comstock Publishing Associates
- Hölldobler, Bert, and Edward O. Wilson. 1990. The Ants. Belknap Press of Harvard University Press
- Hölldobler, Bert, and Edward O. Wilson. 1995. Journey to the Ants. Belknap Press of Harvard University Press
- Howard, Leland O. 1914. The Insect Book. Doubleday, Page and Company.
- Mallis, Arnold, Stoy Hedges (Ed.) et al. 2011. The Mallis Handbook of Pest Control, 10th Ed.
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