Crematogaster cerasi

Ge­o­graphic Range

Cre­mato­gaster cerasi, a species of ac­ro­bat ant, is na­tive to the Nearc­tic re­gion. It is found in south­ern Canada, through­out much of the United States, and into north­ern Mex­ico. In the United States, it ranges west to the Rocky Moun­tains and New Mex­ico, east to the east­ern coastal states, and as far south as Florida. It is par­tic­u­larly com­mon in the Mid­west and in the north­east­ern states. (John­son, 1988; Smith, 1965)

Habi­tat

Cre­mato­gaster cerasi builds its nests pri­mar­ily in trees such as under bark, in hol­low stems, or other cav­i­ties, but also lives in logs and stumps or under leaf lit­ter and stones. These ants also nest in­side houses and other build­ings, usu­ally in tight spaces such as be­tween shin­gles and in or around doors and win­dows. They are most com­mon in forests, but can also be found in grass­lands, pas­tures, bogs, and marshes. They are even oc­ca­sion­ally found in moun­tain re­gions, as high as 2,350 m. In the south­ern por­tion of their range, they tend to live in drier, ri­par­ian habi­tats, while in the north they are more fre­quent in wet­ter wood­lands and fields. (Ellis, et al., 2000; Jan­der, 1990; John­son, 1988; Kan­nowski, 1959; Mackay and Mackay, 2003; Rice, 2013; Smith, 1965; Thomp­son, 1990)

  • Range elevation
    2350 (high) m
    7709.97 (high) ft

Phys­i­cal De­scrip­tion

Work­ers of Cre­mato­gaster cerasi are typ­i­cally 2.6 to 4.0 mm in length. Queens are larger at 7.0 to 8.0 mm in length. These ants range from red­dish-brown to dark brown to black in color. Ants of genus Cre­mato­gaster have unique heart shaped gasters, with the gaster flat­tened dor­sally and con­vex ven­trally. This genus has 11 seg­mented an­ten­nae, with 3 seg­mented an­ten­nal clubs. The dor­sal face of the peti­ole is flat­tened, while the post-peti­ole has two lobes sep­a­rated by a crevice, the post-peti­ole at­taches to the dor­sum of the gaster. While genus Cre­mato­gaster is rea­son­ably easy to iden­tify, es­pe­cially due to the dis­tinct gaster, in­di­vid­ual species are more dif­fi­cult to dis­tin­guish. Cre­mato­gaster cerasi can be dis­tin­guished from other Cre­mato­gaster species by one or two long, erect hairs on each cor­ner of the prono­tum. Males and re­pro­duc­tive fe­males have wings, while work­ers do not. Queens lose their wings after mat­ing. Lar­vae are 1.0 to 1.1 mm in length when they have first hatched and can grow to 1.9 mm long. The lar­val body is curved and seg­mented and the in­tegu­ment grows smooth when the larva nears molt­ing. (El­li­son, et al., 2012; Mackay and Mackay, 2003; Rice, 2013; Smith, 1965; Thomp­son, 1990; Wheeler and Wheeler, 1973)

  • Range length
    2.6 to 4.0 mm
    0.10 to 0.16 in

De­vel­op­ment

Ants are holometabolous. Eggs hatch into lar­vae, which re­quire sig­nif­i­cant care from the adults to sur­vive. They then de­velop into pupae, which even­tu­ally de­velop into adults. For a short pe­riod time after the pupal stage, the adult ants are in a cal­low stage, where the ant is white in color, weak, and less ac­tive. Most off­spring de­velop into fe­male work­ers. Males and alate fe­males are pre­sent in the nest in late sum­mer. (Kan­nowski, 1959)

Re­pro­duc­tion

Mat­ing takes place in late sum­mer, after male and alate fe­male broods hatch and de­velop in the nest. Adult re­pro­duc­tives are pre­sent start­ing in July. Nup­tial flights occur from late July to early Sep­tem­ber. Dur­ing the nup­tial flights, swarms can occur, typ­i­cally with males seek­ing out fe­male mates. Fe­males mate only once in their lives, with males dying shortly after mat­ing. (Kan­nowski, 1959; Mar­shall, 2006)

After mat­ing, the queens go out in search of a suit­able lo­ca­tion to begin a new colony. Queens often take shel­ter in aban­doned bee­tle or ter­mite gal­leries in branches of trees, as well as other lo­ca­tions, and they re­move their wings. Some may over­win­ter be­fore pro­duc­ing a brood, while oth­ers begin egg pro­duc­tion soon after. Queens have a very high mor­tal­ity rate at this point in their lives, with only a small frac­tion of mated fe­males suc­cess­fully found­ing colonies. The first batch of eggs de­vel­ops into the first work­ers that tend the queen and fu­ture broods. When the colony is es­tab­lished, re­pro­duc­tive fe­males and males are pro­duced. Queens store sperm and lay eggs through­out their lives. Al­though the work­ers are fe­male, they do not mate. Cre­mato­gaster colonies can last 10 to 15 years. Some re­ports sug­gest this species may be polyg­y­nous, with mul­ti­ple queens per nest, as nests are often found with sev­eral dealate fe­males. (Kan­nowski, 1959; Mackay and Mackay, 2003; Rice, 2013)

  • Breeding interval
    Queens mate only once in their lifetime and males die shortly after mating.
  • Breeding season
    Mating takes place during late summer, but brood production continues throughout a queen's life.

Fe­male work­ers of Cre­mato­gaster cerasi in­vest sig­nif­i­cant care in the brood pro­duced by the queen. Since the lar­vae and pupae are help­less and con­fined to the nest, the work­ers must tend them and sup­ply food. Work­ers feed the brood via trophal­laxis, which is the mouth to mouth re­gur­gi­ta­tion of food. Work­ers also pro­tect the brood from po­ten­tial threats. The bee­tle Fustiger knausii often lives in the nests of Cre­mato­gaster cerasi, and if any bee­tles are pre­sent near the brood, the work­ers ants re­move them. (Leschen, 1991; Mar­shall, 2006)

  • Parental Investment
  • female parental care
  • pre-hatching/birth
    • provisioning
      • female
    • protecting
      • female
  • pre-weaning/fledging
    • provisioning
      • female
    • protecting
      • female
  • pre-independence
    • provisioning
      • female
    • protecting
      • female

Lifes­pan/Longevity

Males live only a few weeks to a month, and die shortly after mat­ing. While the spe­cific lifes­pan is not avail­able for Cre­mato­gaster cerasi, it is likely sim­i­lar to many other ant species, with work­ers liv­ing sev­eral months. Colonies of Cre­mato­gaster can per­sist for more than 10 years, so queens likely live for sev­eral years. (Mar­shall, 2006; Rice, 2013)

Be­hav­ior

Cre­mato­gaster cerasi for­ages for food both dur­ing the day and night. Work­ers mark trails with pheromones for their nest mates to fol­low. They move in lines, with one ant fol­low­ing an­other. This species is mainly ar­bo­real, con­struct­ing their nests and doing much of their for­ag­ing within trees. These ants are not strictly ar­bo­real and can nest in leaf lit­ter, under stones, or in build­ings. Their colonies can be enor­mous, with 10 thou­sand or more in­di­vid­u­als. Ants of genus Cre­mato­gaster per­form a dis­tinc­tive be­hav­ior when dis­turbed, which gives them the com­mon name 'ac­ro­bat ants'. They have sig­nif­i­cant flex­i­bil­ity in the con­nec­tion be­tween the tho­rax and gaster, al­low­ing them to hold their gaster up over their head and body, and wave it in the air when alarmed. As a eu­so­cial in­sect, Cre­mato­gaster cerasi has a di­vi­sion of labor be­tween the work­ers and re­pro­duc­tives. The role of alate fe­males and males is to mate, and then for the fe­males to found new colonies and pro­duce off­spring. Work­ers tend to the brood, for­age, and pro­tect the nest. The work­ers of most Cre­mato­gaster species are monomor­phic, though some are slightly poly­mor­phic. While Cre­mato­gaster cerasi is not specif­i­cally men­tioned in the lit­er­a­ture, work­ers are likely monomor­phic with no or lit­tle di­vi­sion of labor. (Ellis, et al., 2000; El­li­son, et al., 2012; Mar­shall, 2006; Rice, 2013; Thomp­son, 1990)

Home Range

There is cur­rently no in­for­ma­tion avail­able about the home range size of these ants.

Com­mu­ni­ca­tion and Per­cep­tion

An­ten­nae are the main sen­sory or­gans of ants. They are used to de­tect chem­i­cals and pheromones, to iden­tify nest mates and re­pro­duc­tive mates, and also to feel the en­vi­ron­ment around them. Ants reg­u­larly groom their an­ten­nae to pre­vent buildup of chem­i­cals and par­ti­cles. Pheromones are an im­por­tant method of com­mu­ni­ca­tion be­tween in­di­vid­u­als. Cre­mato­gaster cerasi emits an alarm pheromone to alert its nest mates of a threat. Cre­mato­gaster ants also use trail pheromones to mark paths to food sources and other lo­ca­tions. Ants in this genus are unique, as the gland for trail chem­i­cals is in their hind legs, not in their gaster. Most ants use the tip of their gaster to lay trail pheromones, but be­cause of the unique shape of the Cre­mato­gaster gaster, they can­not put the tip of their gaster in con­tact with the ground. So in­stead, the gland is lo­cated in the hind legs, and the ants adopt a strange-look­ing run when mark­ing the trail. (Boroczky, et al., 2013; Crewe, et al., 1972; Mor­gan, et al., 2004)

Food Habits

Cre­mato­gaster cerasi is om­niv­o­rous. Hon­ey­dew from in­sects such as aphids makes up a large por­tion of its diet, as well as nec­tar from extra flo­ral nec­taries. They also eat the ed­i­ble parts of seeds. Cre­mato­gaster cerasi is also a preda­tor. They prey on live in­sects and also scav­enge dead in­sects. Ants that find their way in­side homes and build­ings read­ily eat what­ever human foods they come across. (Beat­tie and Cul­ver, 1981; Rice, 2013; Smith, 1965; Stephen­son, 1981; Thomp­son, 1990)

  • Animal Foods
  • insects
  • Plant Foods
  • seeds, grains, and nuts
  • nectar

Pre­da­tion

Cre­mato­gaster cerasi is preyed on by spi­ders, bee­tles, and other in­sects, as well as birds, small mam­mals such as mice, and oc­ca­sion­ally black bears. (Noyce, et al., 1997; Rice, 2013)

Ecosys­tem Roles

In some for­est habi­tats, Cre­mato­gaster cerasi can be one of the dom­i­nant ant species. It is a preda­tor of other in­sects, and can serve as prey to a va­ri­ety of other an­i­mals in­clud­ing spi­ders, birds, bears, and other in­sects. These ants likely have a min­i­mal im­pact on seed dis­per­sal. The ants trans­port seeds and eat the ed­i­ble part, the elaio­some, and leave the rest, al­low­ing the seed to ger­mi­nate in a new lo­ca­tion. These ants also form mu­tu­al­is­tic re­la­tion­ships with hon­ey­dew ex­cret­ing in­sects such as aphids and scale in­sects. In turn for eat­ing the hon­ey­dew these in­sects pro­duce, the ants pro­vide care and pro­tec­tion from preda­tors and par­a­sitoids. Some of the aphid species they tend in­clude Sipha flava, Aphis pomi, Aphis spi­raecola, and Neo­ceru­raphis vibur­ni­cola, as well as scale in­sects such as Eu­le­ca­nium cera­so­rum and Ne­ole­ca­nium cor­nu­parvum. Nests of ac­ro­bat ants are often in­hab­ited by other or­gan­isms, form­ing com­men­sal re­la­tion­ships, in which the ants are largely un­af­fected. The bee­tle Fustiger knausii is one species that can be found in the nests. The ants do not seem both­ered by the bee­tles, though the work­ers will re­move bee­tles if they are near the brood. Work­ers will groom the bee­tles, and vice versa. The bee­tles will also ride on the ants; the ants do not re­spond and ap­pear un­af­fected. There has also been some ev­i­dence of trophal­laxis be­tween the species. Crick­ets of genus Myrme­cophilus also often in­habit the nest, with no ef­fect on the ants. There are also sev­eral species of phoretic mites that at­tach them­selves to work­ers of C. cerasi and es­sen­tially hitch a ride to an­other lo­ca­tion. The mites are not par­a­sitic and the ants are not both­ered. Cre­mato­gaster cerasi does have one recorded par­a­sitoid species, phorid flies, the lar­vae of which de­velop in­side the ant's head, even­tu­ally killing it. (Beat­tie and Cul­ver, 1981; Brad­shaw, et al., 2010; Camp­bell, et al., 2013; Ellis, et al., 2000; El­li­son, et al., 2012; Leonard, 1971; Leschen, 1991; Rice, 2013; Smith, 1965; Steyskal, 1944; Vanek and Pot­ter, 2010)

Mu­tu­al­ist Species
Com­men­sal/Par­a­sitic Species

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

There are no known pos­i­tive ef­fects of Cre­mato­gaster cerasi on hu­mans.

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

Cre­mato­gaster cerasi can be a house­hold pest, and is es­pe­cially com­mon as a home in­vader in the north­east­ern United States. These ants have been known to live in­side tight spaces within house­holds and just out­side build­ings, ven­tur­ing in­side to feed on house­hold foods. It does not ap­pear to cause much dam­age, but can be a nui­sance. They are de­scribed as strangely dif­fi­cult to squish; those that are stepped on or hit seem merely stunned, be­fore straight­en­ing them­selves out and going back to work. Cre­mato­gaster cerasi is said to in­flict a painful bite. It may also nest in wood that has been dam­aged by other species, and often en­larges the cav­i­ties a bit, caus­ing more dam­age. Since it tends hon­ey­dew pro­duc­ing in­sects, many of which are crop pests, Cre­mato­gaster cerasi is often in­di­rectly re­spon­si­ble for crop dam­age. Since it pro­tects aphid pop­u­la­tions and other pests from preda­tors, it al­lows the pests to con­tinue feed­ing. This species may be an in­ter­me­di­ate host for the poul­try tape­worm, Rail­li­etina tetrag­ona, as it has been ob­served car­ry­ing seg­ments of tape­worm into its nest, which would then in­fect any bird that eats an in­fected ant. There has been lit­tle re­search con­ducted about this method of in­fec­tion and it may not be a huge con­cern, but it has the pos­si­bil­ity of caus­ing do­mes­tic an­i­mal losses. (Brad­shaw, et al., 2010; Mar­shall, 2006; Rice, 2013; Smith, 1965; Thomp­son, 1990)

  • Negative Impacts
  • household pest

Con­ser­va­tion Sta­tus

Cre­mato­gaster cerasi has no spe­cial con­ser­va­tion sta­tus.

Other Com­ments

Cre­mato­gaster cerasi was for­merly known as Cre­mato­gaster li­ne­o­lata var. cerasi. It may also be re­ferred to as the cherry ant, since it was first found tend­ing aphids on a cherry tree. How­ever, this name is not used with enough reg­u­larly to be con­sid­ered this species' com­mon name. (El­li­son, et al., 2012; Gaddy, 1986)

Con­trib­u­tors

An­gela Miner (au­thor), An­i­mal Di­ver­sity Web Staff, Leila Si­cil­iano Mar­tina (ed­i­tor), An­i­mal Di­ver­sity Web Staff.

Glossary

Nearctic

living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.

World Map

agricultural

living in landscapes dominated by human agriculture.

arboreal

Referring to an animal that lives in trees; tree-climbing.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

bog

a wetland area rich in accumulated plant material and with acidic soils surrounding a body of open water. Bogs have a flora dominated by sedges, heaths, and sphagnum.

chemical

uses smells or other chemicals to communicate

colonial

used loosely to describe any group of organisms living together or in close proximity to each other - for example nesting shorebirds that live in large colonies. More specifically refers to a group of organisms in which members act as specialized subunits (a continuous, modular society) - as in clonal organisms.

diurnal
  1. active during the day, 2. lasting for one day.
ectothermic

animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature

eusocial

the condition in which individuals in a group display each of the following three traits: cooperative care of young; some individuals in the group give up reproduction and specialize in care of young; overlap of at least two generations of life stages capable of contributing to colony labor

female parental care

parental care is carried out by females

fertilization

union of egg and spermatozoan

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

heterothermic

having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.

internal fertilization

fertilization takes place within the female's body

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

marsh

marshes are wetland areas often dominated by grasses and reeds.

metamorphosis

A large change in the shape or structure of an animal that happens as the animal grows. In insects, "incomplete metamorphosis" is when young animals are similar to adults and change gradually into the adult form, and "complete metamorphosis" is when there is a profound change between larval and adult forms. Butterflies have complete metamorphosis, grasshoppers have incomplete metamorphosis.

motile

having the capacity to move from one place to another.

native range

the area in which the animal is naturally found, the region in which it is endemic.

nocturnal

active during the night

omnivore

an animal that mainly eats all kinds of things, including plants and animals

oviparous

reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.

pheromones

chemicals released into air or water that are detected by and responded to by other animals of the same species

riparian

Referring to something living or located adjacent to a waterbody (usually, but not always, a river or stream).

seasonal breeding

breeding is confined to a particular season

sedentary

remains in the same area

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

sperm-storing

mature spermatozoa are stored by females following copulation. Male sperm storage also occurs, as sperm are retained in the male epididymes (in mammals) for a period that can, in some cases, extend over several weeks or more, but here we use the term to refer only to sperm storage by females.

suburban

living in residential areas on the outskirts of large cities or towns.

tactile

uses touch to communicate

temperate

that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).

terrestrial

Living on the ground.

tropical savanna and grassland

A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.

savanna

A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.

temperate grassland

A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.

visual

uses sight to communicate

Ref­er­ences

Beat­tie, A., D. Cul­ver. 1981. The Guild of Myrme­co­chores in the Herba­ceous Flora of West Vir­ginia Forests. Ecol­ogy, 62/1: 107-115.

Boroczky, K., A. Wada-Kat­sumata, D. Batch­e­lor, M. Zhukovskaya, C. Schal. 2013. In­sects groom their an­ten­nae to en­hance ol­fac­tory acu­ity. Pro­ceed­ings of the Na­tional Acad­emy of Sci­ences of the United States of Amer­ica, 110/9: 3615-3620.

Brad­shaw, J., J. Prasifka, K. Stef­fey, M. Gray. 2010. First Re­port of Field Pop­u­la­tions of Two Po­ten­tial Aphid Pests of the Bioen­ergy Crop Mis­cant­hus × Gi­gan­teus. Florida En­to­mol­o­gist, 93/1: 135-137.

Camp­bell, K., H. Klom­pen, T. Crist. 2013. The di­ver­sity and host speci­ficity of mites as­so­ci­ated with ants: The roles of eco­log­i­cal and life his­tory traits of ant hosts. In­sectes So­ci­aux, 60/1: 31-41.

Crewe, R., M. Blum, C. Colling­wood. 1972. Com­par­a­tive analy­sis of alarm pheromones in the ant genus Cre­mato­gaster. Com­par­a­tive Bio­chem­istry and Phys­i­ol­ogy Part B: Com­par­a­tive Bio­chem­istry, 43/3: 703-716.

Ellis, L., M. Molles, Jr., C. Craw­ford, F. Heinzel­mann. 2000. Sur­face-Ac­tive Arthro­pod Com­mu­ni­ties in Na­tive and Ex­otic Ri­par­ian Veg­e­ta­tion in the Mid­dle Rio Grande Val­ley, New Mex­ico. The South­west­ern Nat­u­ral­ist, 45/4: 456-471.

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Gaddy, L. 1986. Twelve New Ant-Dis­persed Species from the South­ern Ap­palachi­ans. Bul­letin of the Tor­rey Botan­i­cal Club, 113/3: 247-251.

Jan­der, R. 1990. Ar­bo­real search in ants- search on branches (Hy­menoptera, Formi­ci­dae). Jour­nal of In­sect Be­hav­ior, 3/4: 515-527.

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Kan­nowski, P. 1959. The flight ac­tiv­i­ties and colony-found­ing be­hav­ior of bog ants in south­east­ern Michi­gan. In­sectes So­ci­aux, 6/2: 115-162.

Leonard, M. 1971. More Records of New Jer­sey Aphids (Ho­moptera: Aphi­di­dae). Jour­nal of the New York En­to­mo­log­i­cal So­ci­ety, 79/2: 62-83.

Leschen, R. 1991. Be­hav­ioral ob­ser­va­tions on the myrme­cophile Fustiger knausii (Coleoptera: Pse­laphi­dae: Clav­igeri­nae) with a dis­cus­sion of grasp­ing notches in myrme­cophiles. En­to­mo­log­i­cal News, 102/5: 215-222.

Mackay, W., E. Mackay. 2003. The Ants of New Mex­ico (Hy­menoptera: Formi­ci­dae). Lewis­ton, New York: The Edwin Mellen Press.

Mar­shall, S. 2006. In­sects - Their Nat­ural His­tory and Di­ver­sity. Buf­falo, New York: Fire­fly Books Inc.

Mor­gan, E., J. Brand, K. Mori, S. Kee­gans. 2004. The trail pheromone of the ant Cre­mato­gaster cas­tanea. Chemoe­col­ogy, 14/2: 119-120.

Noyce, K., P. Kan­nowski, M. Riggs. 1997. Black bears as ant-eaters: sea­sonal as­so­ci­a­tions be­tween bear myrme­cophagy and ant ecol­ogy in north-cen­tral Min­nesota. Cana­dian Jour­nal of Zo­ol­ogy, 75/10: 1671-1686.

Rice, E. 2013. "Dr. Eleanor's Book of Com­mon Ants" (On-line pdf). Your Wild Life. Ac­cessed De­cem­ber 28, 2013 at http://​www.​yourwildlife.​org/​wp-content/​uploads/​2013/​04/​BCA_​demo.​pdf.

Smith, M. 1965. House-in­fest­ing ants of the east­ern United States: their recog­ni­tion, bi­ol­ogy, and eco­nomic im­por­tance. Wash­ing­ton, D.C.: U.S. De­part­ment of Agri­cul­ture.

Stephen­son, A. 1981. Toxic Nec­tar De­ters Nec­tar Thieves of Catalpa spe­ciosa. Amer­i­can Mid­land Nat­u­ral­ist, 105/2: 381-383.

Steyskal, G. 1944. A new ant-at­tack­ing fly of the genus Pseudacteon, with a key to the fe­males of the North Amer­i­can species (Diptera, Phori­dae). Oc­ca­sional Pa­pers of the Mu­seum of Zo­ol­ogy, Uni­ver­sity of Michi­gan, 489: 1-4.

Thomp­son, C. 1990. Ants that have pest sta­tus in the United States. Pp. 51-70 in R Van­der Meer, K Jaffe, A Ce­deno, eds. Ap­plied Myrme­col­ogy. Boul­der, Col­orado: West­view Press, Inc.

Vanek, S., D. Pot­ter. 2010. Ant-Ex­clu­sion to Pro­mote Bi­o­log­i­cal Con­trol of Soft Scales (Hemiptera: Coc­ci­dae) on Woody Land­scape Plants. En­vi­ron­men­tal En­to­mol­ogy, 39/6: 1829-1837.

Wheeler, G., J. Wheeler. 1973. The ant lar­vae of six tribes: sec­ond sup­ple­ment (Hy­menoptera: Formi­ci­dae: Myr­mic­i­nae). Jour­nal of the Geor­gia En­to­mo­log­i­cal So­ci­ety, 8/1: 27-39.