Overview

Brief Summary

Overview

Ants, are classified in the family Formicidae. These insects are native to nearly all terrestrial habitats and all parts of the globe except for Antarctica, Iceland, Greenland, some parts of Polynesia, and a few remote Indian Ocean and Atlantic Ocean islands(3), and are often extremely abundant locally(2). Though there are over 8,800 species known (and perhaps over 11,000 more that have not been described)(3), ants generally have a distinctive body structure: while they have, like many insects, a head, thorax (the midsection), and abdomen (the rear section), their “waist” connecting their thorax to the main part of their abdomen is unusually thin and pinched (1,3). Most ants are also characterized by the presence of a metapleural gland, an organ that produces a chemical called phenylacetic acid that is used for fighting bacteria and fungi(2,3); this gland may have helped ants colonize the moist environments where most ant species now live(2). Like only a few other groups of insects, ants have evolved a complex system of social interaction that qualifies them as “eusocial” insects(2,3). They live and work together in multi-generational colonies that are generally organized in “castes” of queens and males (who reproduce) and worker females (who cannot reproduce)(2,3), communicating via a chemical communication system that may be more complicated than that of any other kind of animal(2). In addition to these extraordinary social structures, ants have complex and extremely important relationships with many other species, giving them a central role in ecosystems across the globe(2). Some ants have partnerships with fungi(2). Some ants defend plants from herbivores, help plants reproduce by pollinating their flowers and spreading their seeds, and help plants grow by turning over the soil (which keeps it rich and healthy)(1,2,3). In fact, many plants depend on ants for their survival(3). On the other hand, some ants are the primary plant-eaters in their environments(1,2), and in many cases ants are major predators of small animals(2). Although some ant species can be pests themselves(1,2), some are beneficial to humans by feeding on harmful crop pests(2)—and by serving as subjects for a wide range of scientific studies(2,3).

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Comprehensive Description

Key to Subfamilies of Malagasy male ants (alate) [[ Formicidae ]]

A few genera are divided into parts, reflecting distinct characters sets outlined in Table 1.

1. Two distinct, long, narrow spines or lobes present on apical portion of abdominal sternum IX (Fig. 3a) or, if absent, then mandibles extremely elongated, distinctly longer than head, and volsella massive, claw-shaped, directed dorsally. Pygostyles absent........................................................... Cerapachyinae HNS

- Spines or lobes absent on apical portion of abdominal sternum IX or the apical portion bilobed, with each lobe very wide (Fig. 3b). Mandibles not elongated, distinctly shorter than head. Volsella moderate, not claw-shaped, not directed dorsally. Pygostyles present or absent......................................................2

2. Abdominal segment III much smaller than segment IV in lateral view (Fig. 3c)....................................3

- Abdominal segment III nearly as large as segment IV in lateral view (Fig. 3d)......................................4

3. Hind tibia with two spurs (Fig. 3e)......................................................................... Pseudomyrmecinae HNS

- Hind tibia with one spur or without spurs (Fig. 3f) ............................................................. Myrmicinae HNS

4. Anal region of hind wing vestigial. Oblique mesopleural furrow reaching pronotum close to its posteroventral corner (Fig. 4a)...................................................................................................... Proceratiinae HNS

- Anal region of hind wing well developed. Oblique mesopleural furrow not reaching pronotum, its anterior termination well separated from the pronotum (Fig. 1a)..................................................................5

5. Petiole (abdominal segment II) broadly and dorsally attached to abdominal segment III; dorsal constriction between petiole and abdominal segment III very shallow or indistinct in lateral view (Fig. 4b......... ........................................................................................................................................ Amblyoponinae HNS

- Petiole (abdominal segment II) narrowly and ventrally attached to abdominal segment III; dorsal constriction between petiole and abdominal segment III deep in lateral view (Fig. 6a)...6

6. Scuto-scutellar suture usually longitudinally sculptured. Forewing clearly with cross vein 2rs-m (Fig. 1b); if vein weak then at least with vestigial branches on Radial sector and Media. Scape short, not reaching posterior margin of head in full-face view (Fig. 5c). Constriction between abdominal segments III and IV present in some cases.............................................................................................. Ponerinae HNS

- Scuto-scutellar suture not longitudinally sculptured. Forewing usually without any trace of cross vein 2rs-m (Fig. 4c). Scape short (Fig. 4d) or long (Fig. 2a). Constriction between abdominal segments III and IV absent...........................................................................................................................................7

7. Many minute, serrate teeth present on masticatory margin of mandible (Fig. 4d), or, if teeth absent, then scape not reaching posterior margin of head in full-face view........................................ Dolichoderinae HNS

- Several larger teeth present on masticatory margin of mandible (Fig. 2a). Scape long, distinctly exceeding posterior margin of head in full-face view (Fig. 2a)........................................................ Formicinae HNS

  • Yoshimura, M., Fisher, B. L. (2007): A revision of male ants of the Malagasy region (Hymenoptera: Formicidae): Key to subfamilies and treatment of the genera of Ponerinae. Zootaxa 1654, 21-40: 28-28, URL:http://www.mapress.com/zootaxa/2007f/zt01654p040.pdf
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KEY TO CALIFORNIA ANT GENERA BASED ON THE WORKER CASTE [[ Formicidae ]]

Morphological terminology follows Bolton (1994), on which much of this key is based. The term mesosoma is used in preference to alitrunk, to refer to the body part formed from fusion of the thorax and the first abdominal segment (i.e., thorax + propodeum). The promesonotum is that part of the mesosoma composed of the pronotum and the mesonotum. Metasoma refers to the apparent abdomen, comprising the segments posterior to the propodeum, i.e., abdominal segment 2 and succeeding segments. In ants abdominal segment2 forms a node- or scale-like petiole, which is separated by a constriction from the rest of the metasoma. In some species abdominal segment 3 is also node-like and in this case it is said to form a postpetiole.

The key has been designed to apply only to those ant species occurring in California but it should also work for most of western North America, excluding Arizona and New Mexico where additional genera occur.

1 Postpetiole present: abdominal segment 3 separated from segment 4 by a strong constriction and distinctly smaller in size, so that anteriorly the metasoma has two nodelike segments, the petiole and postpetiole .................................................................... 2

- Postpetiole absent: abdominal segment 3 separated from segment 4 by a weak to moderate constriction or by none at all, and when viewed in profile not distinctly smaller in size (height) than segment 4; metasoma anteriorly with a single, isolated node-like or scale-like segment ....................................................................................................... 26

2 Eye reduced to a single ommatidium or absent; antennal insertions fully exposed in a full-face view of head ( Ecitoninae HNS ) ........................................ Neivamyrmex Borgmeier HNS

- Eye very rarely reduced, usually consisting of multiple ommatidia; antennal insertions not fully exposed, covered partially by frontal lobes or medial extensions of the antennal sclerites, when the head is observed in full-face view ........................................... 3

3 Eye very large, eye length about one-half of head length (excluding mandibles); pronotum freely articulating with mesonotum ( Pseudomyrmecinae HNS ) ............................. ....................................................................................................... Pseudomyrmex Lund HNS

- Eye less than one-half head length; pronotum fused with mesonotum ( Myrmicinae HNS ) ... ....................................................................................................................................... 4

4 Antenna with 6 segments, including a 2-segmented club ............................................. 5

- Antenna with 10 segments, including a 2-segmented club ......... Solenopsis Westwood HNS

- Antenna with 11 segments; club variable ..................................................................... 6

- Antenna with 12 segments; club variable.. ................................................................ 14

5 Mandible elongate and linear, with an apical fork of two spiniform teeth ...................... ...................................................................................................... Strumigenys F. Smith HNS

- Mandible short and subtriangular, with a multi-denticulate masticatory margin ........... ............................................................................................................... Pyramica Roger HNS

6 Postpetiole attached to the dorsal surface of the following abdominal segment; petiole dorsoventrally flattened, not node-like in profile ........................... Crematogaster Mayr HNS

- Postpetiole attached to the anterior face of the following segment; petiole node-like in profile, not dorsoventrally flattened ............................................................................. 7

7 Head in lateral view with a diagonal carina running from above the eye down toward the mandibular insertion; promesonotum with conspicuous tubercles or spines ......... 8

- Head in lateral view lacking such a diagonal carina; promesonotum without conspicuous tubercles or spines .................................................................................................9

8 Frontal lobes expanded laterally and covering the sides of the head below the eyes, in full-face view; body lacking erect pilosity.................................... Cyphomyrmex Mayr HNS

- Frontal lobes not expanded laterally to cover the sides of the head; body with erect pilosity.............................................................................................. Acromyrmex Mayr HNS

9 Antenna with a distinct 2-segmented apical club............................... Wasmannia Forel HNS

- Antenna lacking a distinct 2-segmented apical club, either 3-segmented or indistinct.. .....................................................................................................................................10

10 Eye absent or rudimentary; propodeum unarmed, basal face rounding into declivitous face ................................................................................................ Solenopsidini HNS new genus

- Eye well developed, with multiple ommatidia; propodeum angulate or spinose.......11

11 Lateral portions of clypeus, in front of the antennal insertions, developed in the form of a raised ridge or shield-wall; frontal carinae extending almost to the posterior margin of the head......................................................................... Tetramorium Mayr HNS (part)

- Lateral portions of clypeus not developed as a raised ridge or shield-wall; frontal carinae very short or absent...............................................................................................12

12 Eye with short erect setae projecting between the ommatidia........ Formicoxenus Mayr HNS

- Eye lacking erect setae................................................................................................13

13 Median portion of clypeus with a smooth, longitudinally excavate surface, and lacking carinae .............................................................................................. Leptothorax Mayr HNS

- Median portion of clypeus with several longitudinal carinae......................................... ................................................................................................. Temnothorax Mayr HNS (part)

14 Hind tibial spur finely pectinate (as seen at 50-100x magnification).........................15

- Hind tibial spur simple or absent.................................................................................17

15 Metanotal groove absent or very weakly impressed, not breaking the dorsal profile of the mesosoma; psammophore usually present............................ Pogonomyrmex Mayr HNS

- Metanotal groove present and interrupting the dorsal profile of the mesosoma; psammophore absent...........................................................................................................16

16 Propodeum unarmed; mandible with more than 12 teeth ....................... Manica Jurine HNS

- Propodeum armed with a pair of spines; mandible with 6-10 teeth .. Myrmica Latreille HNS

17 Lateral portions of clypeus, in front of the antennal insertions, developed in the form of a raised ridge or shield-wall; apex of sting with triangular lamellate appendage...... ................................................................................................. Tetramorium Mayr HNS (part)

- Lateral portions of clypeus not developed as a raised ridge or shield-wall; apex of sting without triangular lamellate appendage .....................................................................18

18 Petiole short and sessile, lacking well differentiated anterior peduncle and dorsal node; ventrolateral margin of head with sharp, longitudinal carina extending from mandibular base to posterolateral corner of head........................................... Myrmecina Curtis HNS

- Petiole with anterior peduncle and dorsal node; ventrolateral margin of head without sharp, longitudinal carina ...........................................................................................19

19 Dorsum of head and mesosoma without standing pilosity .......... Cardiocondyla Emery HNS

- Dorsum of head and mesosoma with standing pilosity ............................................... 20

20 Anteromedian portion of clypeus notably elevated and bounded by a pair of carinae that diverge anteriorly ................................................................................................ 21

- Anteromedian portion of clypeus not abruptly elevated and lacking a pair of anteriorly diverging carinae ........................................................................................................ 23

21 Propodeum unarmed ....................................................................... Monomorium Mayr HNS

- Propodeum armed with a pair of teeth or spines ........................................................ 22

22 Antennal club 3-segmented; propodeal spiracle large and located close to the declivitous face of the propodeum, separated from latter by no more than the diameter of the spiracle .................................................................................................. Rogeria Emery HNS

- Antennal club 4-segmented; propodeal spiracle relatively small and separated from the declivitous face of the propodeum by more than the spiracle diameter .......................... ....................................................................................................... Stenamma Westwood HNS

23 Antennal club 3- (rarely 4-) segmented ....................................................................... 24

- Antenna lacking a distinct club ................................................................................... 25

24 In profile promesonotum domed and distinctly elevated above the propodeal dorsum; workers dimorphic ......................................................................... Pheidole Westwood HNS

- In profile entire mesosoma dorsum flat to weakly convex, promesonotum not domed or markedly elevated above the level of the propodeum; workers monomorphic .......... ................................................................................................. Temnothorax Mayr HNS (part)

25 Head narrow, longer than broad; mandible slender and triangular, outer margin not strongly curving towards the midline; psammophore absent ...... Aphaenogaster Mayr HNS

- Head broad, subquadrate; mandible short and thick, outer margin strongly curving towards the midline; psammophore usually present ................................. Messor Forel HNS

26 Pygidium (last visible abdominal tergite) flattened and bordered laterally with a row of peg-like teeth or spines that converge distally ( Cerapachyinae HNS ).... Cerapachys F. Smith HNS

- Pygidium (last visible abdominal tergite) convex and rounded, lacking a row of teeth or spines ..................................................................................................................... 27

27 Distinct constriction between abdominal segments 3 and 4; terga and sterna of abdominal segments 3 and 4 laterally fused ......................................................................... 28

- No constriction between abdominal segments 3 and 4; terga of abdominal segments 3 and 4 overlapping the corresponding sterna, not laterally fused with them ............... 30

28 Articulation of petiole (second abdominal segment) to third abdominal segment very broad; petiole without a distinct posterior face ( Amblyoponinae HNS ) .................................. ..................................................................................................... Amblyopone Erichson HNS

- Articulation of petiole (second abdominal segment) to third abdominal segment narrow; petiole with a distinct posterior face .................................................................. 29

29 Pronotum freely articulating with the mesonotum; abdominal tergite 4 not strongly enlarged and not curved ventrally; apex of metasoma directed posteriorly ( Ponerinae HNS ) ....................................................................................................... Hypoponera Santschi HNS

- Pronotum fused immovably to the mesonotum; abdominal tergite 4 strongly enlarged and curved ventrally; apex of metasoma directed anteriorly ( Proceratiinae HNS ).................. .......................................................................................................... Proceratium Roger HNS

30 Apex of metasoma with a circular orifice, often fringed with short setae (the acidopore) ( Formicinae HNS )...................................................................................................31

- Apex of metasoma with a slit-shaped orifice ( Dolichoderinae HNS )..................................38

31 Antenna with 9 segments ............................................................. Brachymyrmex Mayr HNS

- Antenna with 11 segments ................................................................. Plagiolepis Mayr HNS

- Antenna with 12 segments .........................................................................................32

32 Metapleural gland absent; antennal insertions well separated from the posterior clypeal margin; in profile mesosoma dorsum usually evenly convex........................ Camponotus Mayr HNS

- Metapleural gland present; antennal insertions adjacent to the posterior clypeal margin; in profile promesonotum separated from the dorsal face of the propodeum by a distinct impression .....................................................................................................33

33 Maxillary palp segments 3 and 4 greatly elongated, segment 3 (counting from base) half the head length or more; psammophore present.............. Myrmecocystus Wesmael HNS

- Maxillary palp segments 3 and 4 not greatly elongated, segment 3 much less than half the head length; psammophore absent .......................................................................34

34 Ocelli present; propodeal spiracle elliptical to oval....................................................35

- Ocelli absent or indistinct; propodeal spiracle circular to subcircular.......................36

35 Mandible triangular, with seven or more distinct teeth on the masticatory margin........ ............................................................................................................ Formica Linnaeus HNS

- Mandible falcate (sickle-shaped) and lacking distinct teeth............ Polyergus Latreille HNS

36 Dorsum of head and mesosoma with coarse setae, arranged in distinct pairs; eye situated in relatively anterior position, at or in front of midlength of side of head ............. ............................................................................................ Paratrechina Motschoulsky HNS

- Pilosity on dorsum of head and mesosoma variable, but not arranged as coarse setae in pairs; eye situated in relatively posterior position, behind midlength of side of head ... .....................................................................................................................................37

37 Mandible with six teeth; antennal scape long, surpassing posterior margin of head by more than half its length; mesonotum in dorsal view strongly constricted behind pronotum .............................................................................................................. Prenolepis Mayr HNS

- Mandible with seven or more teeth; antennal scape shorter, surpassing posterior margin of head by less than a third its length; mesonotum in dorsal view not strongly constricted behind pronotum...................................................................... Lasius Fabricius HNS

38 Propodeum with a prominent conical tooth at the junction of the dorsal and declivitous faces; maxillary palp segment 3 elongate, subequal in length to segments 4-6; apical mandibular tooth much enlarged...................................................... Dorymyrmex Mayr HNS

- Propodeum rounded or subangulate at the junction of the dorsal and declivitous faces, but without a conical tooth; maxillary palp segment 3 short, subequal in length to segments4; apical mandibular tooth not notably enlarged .............................................. 39

39 Mesosoma dorsum lacking standing pilosity .............................................................. 40

- Mesosoma dorsum with standing pilosity .................................................................. 41

40 Petiole flattened, plate-like, and without a conspicuous, dorsally protruding scale (petiole often overhung by the succeeding abdominal segment); dorsal face of propodeum much shorter than the declivitous (posterior) face ............................ Tapinoma Foerster HNS

- Petiole with a well developed, dorsally protruding scale; dorsal face of propodeum subequal in length to declivitous face .............................................. Linepithema Mayr HNS

41 In profile mesosoma dorsum without an impressed metanotal groove, the promesonotum and propodeum forming a continuous surface; workers variable in size within a colony ........................................................................................................... Liometopum Mayr HNS

- In profile mesosoma dorsum interrupted by a well marked metanotal groove; workers showing little intra-colony size variation .................................................................... 42

42 Petiole lacking an erect scale; side of mesosoma with conspicuous microreticulate sculpture; dark brown-black, with contrastingly paler tarsi .......... Technomyrmex Mayr HNS

- Petiole with well developed erect scale; side of mesosoma without conspicuous microreticulate sculpture; varying in color from yellowish-orange to dark brown, but without contrastingly paler tarsi ............................................................ Forelius Emery HNS

  • Ward, P. S. (2005): A synoptic review of the ants of California (Hymenoptera: Formicidae). Zootaxa 936, 1-68: 20-25, URL:http://antbase.org/ants/publications/21008/21008.pdf
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Formicidae and Myrmicidae (Ants)
Ants are small to medium-sized insects that usually live in the ground, within cavities of trees, or rotting wood of buildings. They are various shades of black, brown, or red. Ants are highly social insects, consisting primarily of sterile workers without wings. Ants tend to be omnivorous scavengers, although some species cut leaves for their underground fungus farms, while others tend aphids for their honeydew. Ants sometimes visit flowering plants for nectar, particularly those with extra-floral nectaries, such as Trumpet Creeper, Wild Senna, and Partridge Pea. Species observed on such wildflowers include Acrobatic ants, Mound ants, and Carpenter ants.

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Description of Formicidae

The Formicidae are the ants, social insects. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than 12,500 out of an estimated total of 22,000 species have been described. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist. Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of workers, soldiers, or other specialised types. Nearly all ant colonies also have some fertile males called drones and one or more fertile females called queens. Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and a few remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15–25% of the terrestrial animal biomass. Their success in so many environments has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study. Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. Ants come into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, aggressively establishing themselves in areas where they are accidentally introduced.
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Characteristics

Bolton (2003) gives the following list of general synapomorphies for the Formicidae (but note that several of these are not unique to ants and have evolved independently in other vespoids, and that several have been secondarily lost in certain ant lineages):
  • Eusocial, with perennial colonies
  • Wingless worker caste
  • Females with prognathous heads
  • Infrabuccal sac between labium and hypopharynx
  • Antennae elbowed (geniculate) between the funiculus and the elongated scape (scape short in the primitive subfamilies Armaniinae and Sphecomyrminae)
  • Metapleural gland in females
  • Abdominal segment II differentiated, forming a petiole (weakly differentiated in the primitive subfamily Armaniinae)
  • Wings of alate queens shed after mating
  • Mating performed in mass nuptial flights
  • Forewings always lacking cross-veins 3rs-m and 2m-cu

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Distribution

Geographic Range

Ants are very important insects all around the world, especially in tropical regions. There are over 11,000 species in the world, and at least 90 species in Michigan.

Biogeographic Regions: nearctic (Introduced , Native ); palearctic (Introduced , Native ); oriental (Introduced , Native ); ethiopian (Native ); neotropical (Native ); australian (Introduced , Native ); oceanic islands (Introduced , Native )

Other Geographic Terms: cosmopolitan

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Physical Description

Morphology

Physical Description

Ants are skinny insects. Like their relatives the wasps, they have a narrow connection between their abdomen and thorax. They have chewing mouthparts, and their antennae are bent in the middle. Some ant species are very tiny (1-2mm long), but some tropical species are very large (30 mm). Most species are 5-15 mm long. Some ant species can sting, and all can bite. Ants have lots of glands for producing chemicals. Most ant species are brown, but some are black, some are yellowish, and some are partly or entirely red.

Each ant colony has several different kinds of ants. They are all the same species, but they look different. Most of the ants you see are workers. They are female, but they cannot reproduce. They do all the work in the nest and protect it from enemies. Some ant species have different sizes of workers for different jobs: large ones with big jaws hunt or protect the nest, while smaller ones work inside, tending the young and digging. Inside the nest is the queen, she is a large female, and is the only one who can lay eggs. Some ant species have several queens in a nest, some have only one. At certain times in the summer there will be new queens and males in the nest as well. They have wings, and fly out to mate and start new nests. Males are usually smaller than females. Only males and queen ants have wings, but the queens remove their wings when they start a new nest. Worker ants never have wings.

Other Physical Features: ectothermic ; bilateral symmetry ; polymorphic

Sexual Dimorphism: female larger

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Diagnostic Description

Morphological terminology follows Bolton (1994), on which much of this key is based. The term mesosoma is used in preference to alitrunk, to refer to the body part formed from fusion of the thorax and the first abdominal segment (i.e., thorax + propodeum). The promesonotum is that part of the mesosoma composed of the pronotum and the mesonotum. Metasoma refers to the apparent abdomen, comprising the segments posterior to the propodeum, i.e., abdominal segment 2 and succeeding segments. In ants abdominal segment2 forms a node- or scale-like petiole, which is separated by a constriction from the rest of the metasoma. In some species abdominal segment 3 is also node-like and in this case it is said to form a postpetiole.

 

The key has been designed to apply only to those ant species occurring in California but it should also work for most of western North America, excluding Arizona and New Mexico where additional genera occur.

 

1 Postpetiole present: abdominal segment 3 separated from segment 4 by a strong constriction and distinctly smaller in size, so that anteriorly the metasoma has two nodelike segments, the petiole and postpetiole .................................................................... 2

 

- Postpetiole absent: abdominal segment 3 separated from segment 4 by a weak to moderate constriction or by none at all, and when viewed in profile not distinctly smaller in size (height) than segment 4; metasoma anteriorly with a single, isolated node-like or scale-like segment ....................................................................................................... 26

 

2 Eye reduced to a single ommatidium or absent; antennal insertions fully exposed in a full-face view of head ( Ecitoninae ) ........................................ Neivamyrmex Borgmeier

 

- Eye very rarely reduced, usually consisting of multiple ommatidia; antennal insertions not fully exposed, covered partially by frontal lobes or medial extensions of the antennal sclerites, when the head is observed in full-face view ........................................... 3

 

3 Eye very large, eye length about one-half of head length (excluding mandibles); pronotum freely articulating with mesonotum ( Pseudomyrmecinae ) ............................. ....................................................................................................... Pseudomyrmex Lund

 

- Eye less than one-half head length; pronotum fused with mesonotum ( Myrmicinae ) ... ....................................................................................................................................... 4

 

4 Antenna with 6 segments, including a 2-segmented club ............................................. 5

 

- Antenna with 10 segments, including a 2-segmented club ......... Solenopsis Westwood

 

- Antenna with 11 segments; club variable ..................................................................... 6

 

- Antenna with 12 segments; club variable.. ................................................................ 14

 

5 Mandible elongate and linear, with an apical fork of two spiniform teeth ...................... ...................................................................................................... Strumigenys F. Smith

 

- Mandible short and subtriangular, with a multi-denticulate masticatory margin ........... ............................................................................................................... Pyramica Roger

 

6 Postpetiole attached to the dorsal surface of the following abdominal segment; petiole dorsoventrally flattened, not node-like in profile ........................... Crematogaster Mayr

 

- Postpetiole attached to the anterior face of the following segment; petiole node-like in profile, not dorsoventrally flattened ............................................................................. 7

 

7 Head in lateral view with a diagonal carina running from above the eye down toward the mandibular insertion; promesonotum with conspicuous tubercles or spines ......... 8

 

- Head in lateral view lacking such a diagonal carina; promesonotum without conspicuous tubercles or spines .................................................................................................9

 

8 Frontal lobes expanded laterally and covering the sides of the head below the eyes, in full-face view; body lacking erect pilosity.................................... Cyphomyrmex Mayr

 

- Frontal lobes not expanded laterally to cover the sides of the head; body with erect pilosity.............................................................................................. Acromyrmex Mayr

 

9 Antenna with a distinct 2-segmented apical club............................... Wasmannia Forel

 

- Antenna lacking a distinct 2-segmented apical club, either 3-segmented or indistinct.. .....................................................................................................................................10

 

10 Eye absent or rudimentary; propodeum unarmed, basal face rounding into declivitous face ................................................................................................ Solenopsidininew genus

 

- Eye well developed, with multiple ommatidia; propodeum angulate or spinose.......11

 

11 Lateral portions of clypeus, in front of the antennal insertions, developed in the form of a raised ridge or shield-wall; frontal carinae extending almost to the posterior margin of the head......................................................................... Tetramorium Mayr (part)

 

- Lateral portions of clypeus not developed as a raised ridge or shield-wall; frontal carinae very short or absent...............................................................................................12

 

12 Eye with short erect setae projecting between the ommatidia........ Formicoxenus Mayr

 

- Eye lacking erect setae................................................................................................13

 

13 Median portion of clypeus with a smooth, longitudinally excavate surface, and lacking carinae .............................................................................................. Leptothorax Mayr

 

- Median portion of clypeus with several longitudinal carinae......................................... ................................................................................................. Temnothorax Mayr (part)

 

14 Hind tibial spur finely pectinate (as seen at 50-100x magnification).........................15

 

- Hind tibial spur simple or absent.................................................................................17

 

15 Metanotal groove absent or very weakly impressed, not breaking the dorsal profile of the mesosoma; psammophore usually present............................ Pogonomyrmex Mayr

 

- Metanotal groove present and interrupting the dorsal profile of the mesosoma; psammophore absent...........................................................................................................16

 

16 Propodeum unarmed; mandible with more than 12 teeth ....................... Manica Jurine

 

- Propodeum armed with a pair of spines; mandible with 6-10 teeth .. Myrmica Latreille

 

17 Lateral portions of clypeus, in front of the antennal insertions, developed in the form of a raised ridge or shield-wall; apex of sting with triangular lamellate appendage...... ................................................................................................. Tetramorium Mayr (part)

 

- Lateral portions of clypeus not developed as a raised ridge or shield-wall; apex of sting without triangular lamellate appendage .....................................................................18

 

18 Petiole short and sessile, lacking well differentiated anterior peduncle and dorsal node; ventrolateral margin of head with sharp, longitudinal carina extending from mandibular base to posterolateral corner of head........................................... Myrmecina Curtis

 

- Petiole with anterior peduncle and dorsal node; ventrolateral margin of head without sharp, longitudinal carina ...........................................................................................19

 

19 Dorsum of head and mesosoma without standing pilosity .......... Cardiocondyla Emery

 

- Dorsum of head and mesosoma with standing pilosity ............................................... 20

 

20 Anteromedian portion of clypeus notably elevated and bounded by a pair of carinae that diverge anteriorly ................................................................................................ 21

 

- Anteromedian portion of clypeus not abruptly elevated and lacking a pair of anteriorly diverging carinae ........................................................................................................ 23

 

21 Propodeum unarmed ....................................................................... Monomorium Mayr

 

- Propodeum armed with a pair of teeth or spines ........................................................ 22

 

22 Antennal club 3-segmented; propodeal spiracle large and located close to the declivitous face of the propodeum, separated from latter by no more than the diameter of the spiracle .................................................................................................. Rogeria Emery

 

- Antennal club 4-segmented; propodeal spiracle relatively small and separated from the declivitous face of the propodeum by more than the spiracle diameter .......................... ....................................................................................................... Stenamma Westwood

 

23 Antennal club 3- (rarely 4-) segmented ....................................................................... 24

 

- Antenna lacking a distinct club ................................................................................... 25

 

24 In profile promesonotum domed and distinctly elevated above the propodeal dorsum; workers dimorphic ......................................................................... Pheidole Westwood

 

- In profile entire mesosoma dorsum flat to weakly convex, promesonotum not domed or markedly elevated above the level of the propodeum; workers monomorphic .......... ................................................................................................. Temnothorax Mayr (part)

 

25 Head narrow, longer than broad; mandible slender and triangular, outer margin not strongly curving towards the midline; psammophore absent ...... Aphaenogaster Mayr

 

- Head broad, subquadrate; mandible short and thick, outer margin strongly curving towards the midline; psammophore usually present ................................. Messor Forel

 

26 Pygidium (last visible abdominal tergite) flattened and bordered laterally with a row of peg-like teeth or spines that converge distally ( Cerapachyinae ).... Cerapachys F. Smith

 

- Pygidium (last visible abdominal tergite) convex and rounded, lacking a row of teeth or spines ..................................................................................................................... 27

 

27 Distinct constriction between abdominal segments 3 and 4; terga and sterna of abdominal segments 3 and 4 laterally fused ......................................................................... 28

 

- No constriction between abdominal segments 3 and 4; terga of abdominal segments 3 and 4 overlapping the corresponding sterna, not laterally fused with them ............... 30

 

28 Articulation of petiole (second abdominal segment) to third abdominal segment very broad; petiole without a distinct posterior face ( Amblyoponinae ) .................................. ..................................................................................................... Amblyopone Erichson

 

- Articulation of petiole (second abdominal segment) to third abdominal segment narrow; petiole with a distinct posterior face .................................................................. 29

 

29 Pronotum freely articulating with the mesonotum; abdominal tergite 4 not strongly enlarged and not curved ventrally; apex of metasoma directed posteriorly ( Ponerinae ) ....................................................................................................... Hypoponera Santschi

 

- Pronotum fused immovably to the mesonotum; abdominal tergite 4 strongly enlarged and curved ventrally; apex of metasoma directed anteriorly ( Proceratiinae ).................. .......................................................................................................... Proceratium Roger

 

30 Apex of metasoma with a circular orifice, often fringed with short setae (the acidopore) ( Formicinae )...................................................................................................31

 

- Apex of metasoma with a slit-shaped orifice ( Dolichoderinae )..................................38

 

31 Antenna with 9 segments ............................................................. Brachymyrmex Mayr

 

- Antenna with 11 segments ................................................................. Plagiolepis Mayr

 

- Antenna with 12 segments .........................................................................................32

 

32 Metapleural gland absent; antennal insertions well separated from the posterior clypeal margin; in profile mesosoma dorsum usually evenly convex........................ Camponotus Mayr

 

- Metapleural gland present; antennal insertions adjacent to the posterior clypeal margin; in profile promesonotum separated from the dorsal face of the propodeum by a distinct impression .....................................................................................................33

 

33 Maxillary palp segments 3 and 4 greatly elongated, segment 3 (counting from base) half the head length or more; psammophore present.............. Myrmecocystus Wesmael

 

- Maxillary palp segments 3 and 4 not greatly elongated, segment 3 much less than half the head length; psammophore absent .......................................................................34

 

34 Ocelli present; propodeal spiracle elliptical to oval....................................................35

 

- Ocelli absent or indistinct; propodeal spiracle circular to subcircular.......................36

 

35 Mandible triangular, with seven or more distinct teeth on the masticatory margin........ ............................................................................................................ Formica Linnaeus

 

- Mandible falcate (sickle-shaped) and lacking distinct teeth............ Polyergus Latreille

 

36 Dorsum of head and mesosoma with coarse setae, arranged in distinct pairs; eye situated in relatively anterior position, at or in front of midlength of side of head ............. ............................................................................................ Paratrechina Motschoulsky

 

- Pilosity on dorsum of head and mesosoma variable, but not arranged as coarse setae in pairs; eye situated in relatively posterior position, behind midlength of side of head ... .....................................................................................................................................37

 

37 Mandible with six teeth; antennal scape long, surpassing posterior margin of head by more than half its length; mesonotum in dorsal view strongly constricted behind pronotum .............................................................................................................. Prenolepis Mayr

 

- Mandible with seven or more teeth; antennal scape shorter, surpassing posterior margin of head by less than a third its length; mesonotum in dorsal view not strongly constricted behind pronotum...................................................................... Lasius Fabricius

 

38 Propodeum with a prominent conical tooth at the junction of the dorsal and declivitous faces; maxillary palp segment 3 elongate, subequal in length to segments 4-6; apical mandibular tooth much enlarged...................................................... Dorymyrmex Mayr

 

- Propodeum rounded or subangulate at the junction of the dorsal and declivitous faces, but without a conical tooth; maxillary palp segment 3 short, subequal in length to segments4; apical mandibular tooth not notably enlarged .............................................. 39

 

39 Mesosoma dorsum lacking standing pilosity .............................................................. 40

 

- Mesosoma dorsum with standing pilosity .................................................................. 41

 

40 Petiole flattened, plate-like, and without a conspicuous, dorsally protruding scale (petiole often overhung by the succeeding abdominal segment); dorsal face of propodeum much shorter than the declivitous (posterior) face ............................ Tapinoma Foerster

 

- Petiole with a well developed, dorsally protruding scale; dorsal face of propodeum subequal in length to declivitous face .............................................. Linepithema Mayr

 

41 In profile mesosoma dorsum without an impressed metanotal groove, the promesonotum and propodeum forming a continuous surface; workers variable in size within a colony ........................................................................................................... Liometopum Mayr

 

- In profile mesosoma dorsum interrupted by a well marked metanotal groove; workers showing little intra-colony size variation .................................................................... 42

 

42 Petiole lacking an erect scale; side of mesosoma with conspicuous microreticulate sculpture; dark brown-black, with contrastingly paler tarsi .......... Technomyrmex Mayr

 

- Petiole with well developed erect scale; side of mesosoma without conspicuous microreticulate sculpture; varying in color from yellowish-orange to dark brown, but without contrastingly paler tarsi ............................................................ Forelius Emery

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A few genera are divided into parts, reflecting distinct characters sets outlined in Table 1.

 

1. Two distinct, long, narrow spines or lobes present on apical portion of abdominal sternum IX (Fig. 3a) or, if absent, then mandibles extremely elongated, distinctly longer than head, and volsella massive, claw-shaped, directed dorsally. Pygostyles absent........................................................... Cerapachyinae

 

- Spines or lobes absent on apical portion of abdominal sternum IX or the apical portion bilobed, with each lobe very wide (Fig. 3b). Mandibles not elongated, distinctly shorter than head. Volsella moderate, not claw-shaped, not directed dorsally. Pygostyles present or absent......................................................2

 

2. Abdominal segment III much smaller than segment IV in lateral view (Fig. 3c)....................................3

 

- Abdominal segment III nearly as large as segment IV in lateral view (Fig. 3d)......................................4

 

3. Hind tibia with two spurs (Fig. 3e)......................................................................... Pseudomyrmecinae

 

- Hind tibia with one spur or without spurs (Fig. 3f) ............................................................. Myrmicinae

 

4. Anal region of hind wing vestigial. Oblique mesopleural furrow reaching pronotum close to its posteroventral corner (Fig. 4a)...................................................................................................... Proceratiinae

 

- Anal region of hind wing well developed. Oblique mesopleural furrow not reaching pronotum, its anterior termination well separated from the pronotum (Fig. 1a)..................................................................5

 

5. Petiole (abdominal segment II) broadly and dorsally attached to abdominal segment III; dorsal constriction between petiole and abdominal segment III very shallow or indistinct in lateral view (Fig. 4b......... ........................................................................................................................................ Amblyoponinae

 

- Petiole (abdominal segment II) narrowly and ventrally attached to abdominal segment III; dorsal constriction between petiole and abdominal segment III deep in lateral view (Fig. 6a)...6

 

6. Scuto-scutellar suture usually longitudinally sculptured. Forewing clearly with cross vein 2rs-m (Fig. 1b); if vein weak then at least with vestigial branches on Radial sector and Media. Scape short, not reaching posterior margin of head in full-face view (Fig. 5c). Constriction between abdominal segments III and IV present in some cases.............................................................................................. Ponerinae

 

- Scuto-scutellar suture not longitudinally sculptured. Forewing usually without any trace of cross vein 2rs-m (Fig. 4c). Scape short (Fig. 4d) or long (Fig. 2a). Constriction between abdominal segments III and IV absent...........................................................................................................................................7

 

7. Many minute, serrate teeth present on masticatory margin of mandible (Fig. 4d), or, if teeth absent, then scape not reaching posterior margin of head in full-face view........................................ Dolichoderinae

 

- Several larger teeth present on masticatory margin of mandible (Fig. 2a). Scape long, distinctly exceeding posterior margin of head in full-face view (Fig. 2a)........................................................ Formicinae

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Ecology

Habitat

Ants are found just about every habitat on land except the very coldest. All ant species need sheltered places to nest and take care of their offspring. Most species nest underground, but some nest in trees. Some very small ant species can make nests inside acorns and other small hiding places.

Adult ants can live in drier conditions than many other invertebrates, but ant eggs and young need humid conditions to survive.

Habitat Regions: temperate ; tropical ; polar ; terrestrial

Terrestrial Biomes: tundra ; taiga ; desert or dune ; savanna or grassland ; chaparral ; forest ; rainforest ; scrub forest ; mountains

Wetlands: marsh ; swamp ; bog

Other Habitat Features: urban ; suburban ; agricultural

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Dispersal

Ant Mating Behavior

Most ants are females. In fact only the princes are males. Nearing mating season the queen produces large amounts of male unfertilized egg to cope up with the short supply of males and ensure successful breeding. During mating seaon a number of ants swarm outside their colony to take part or root for for the "marriage fligt" wherein prince and princess ants take off from their colony , mate in the air and fly off to a new place to start a new colony. The only role of male ants is to mate with the queen and they die alone shortly after mating.  

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Trophic Strategy

Food Habits

Ant species eat many different foods. Some specialize in sugary liquids like nectar and the "honeydew" produced by aphids and other insects (see Aphididae). Many eat other Insecta and other small animals, and scavenge dead meat. Some others specialize in eating seeds or fungus. Ants drink from dew, rain drops, and puddles, and sometimes they get their moisture from their food (like nectar).

Many ant species store food in their nests, especially the seed-eating ants. Others eat fungus that they grow in their nest. Ants that find a big food source leave a chemical trail, so that their nestmates can find the food too. Pretty soon there is a busy column of ants going back and forth from the nest to the food source.

Leaf-cutter ants live in warm climates, they cut up leaves and carry them into their nests underground. They eat the fungus that grows on the leaves. Army ants and driver ants roam through jungles and tropical habitats eating any animals they can find. They are big ants with sharp jaws, and there are many thousands of them in a group. They will eat any animals, even large ones, that they can catch.

Foraging Behavior: stores or caches food

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Associations

Ecosystem Roles

Ants are very important in lots of roles. Some species disperse seeds, some are important predators of insects, some tropical species are important herbivores. Their digging often improves the soil for plant growth.

Ecosystem Impact: disperses seeds; creates habitat; biodegradation ; soil aeration

Mutualist Species:

  • Aphids
  • Some caterpillars

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Predation

Worker ants attack predators, they will die to protect their nest. Some ant species can sting, and all can bite (though the little ones can't hurt a large animal). Many ants also have toxic chemicals they can spray on their enemies.

Known Predators:

  • Aves
  • Ursidae (break open nests and eat the immature ants)
  • Eumeces fasciatus
  • Anura
  • Araneae
  • other Formicidae 
  • Carabidae

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Animal / parasite / endoparasite
cyst of Dicrocoelium lanceolatum endoparasitises brain of Formicoidea

Animal / parasitoid
larva of Eucharis adscendens is parasitoid of larva of Formicoidea

Animal / predator
adult of Systellonotus triguttatus is predator of pupa of Formicoidea
Remarks: season: late 5-mid 8

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Animal / guest
nymph of Alydus calcaratus is a guest in nest of Formicidae

Animal / predator
larva of Chrysotoxum is predator of brood of Formicidae
Remarks: Other: uncertain

Animal / predator
larva of Doros conopseus is predator of brood of Formicidae
Remarks: Other: uncertain

Animal / honeydew feeder
Formicidae feeds on honeydew Maculolachnus submacula

Animal / honeydew feeder
Formicidae feeds on honeydew Aphis sambuci

Animal / slave maker
Formicidae makes a slave of Aphidoidea

Animal / slave maker
Formicidae makes a slave of Pseudococcidae

Animal / slave maker
Formicidae makes a slave of Coccoidea

In Great Britain and/or Ireland:
Animal / associate
colony of anamorph of Hormiscium pithyophilum var. myrmecophilum is associated with nest of Formicidae

Animal / associate
larva of Microdon analis is associated with nest of Formicidae

Animal / associate
larva of Microdon mutabilis is associated with nest of Formicidae

Animal / predator
leaf of Pinguicula vulgaris is predator of adult of Formicidae
Other: minor host/prey

Animal / inquiline
larva of Smaragdina affinis is inquiline in nest of Formicidae
Remarks: Other: uncertain

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Known predators

  • Waide RB, Reagan WB (eds) (1996) The food web of a tropical rainforest. University of Chicago Press, Chicago
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Known prey organisms

Formicidae preys on:
Eleutherodactylus coqui
Collembola
Auchenorrhyncha
Sternorrhyncha
fungi
fruit
seeds
dead leaves
detritus
Acari
fruit and seeds
nectar and floral
Zenaida asiatica
Vanessa cardui
Misumena vatia

Based on studies in:
Puerto Rico, El Verde (Rainforest)

This list may not be complete but is based on published studies.
  • Waide RB, Reagan WB (eds) (1996) The food web of a tropical rainforest. University of Chicago Press, Chicago
  • Myers, P., R. Espinosa, C. S. Parr, T. Jones, G. S. Hammond, and T. A. Dewey. 2006. The Animal Diversity Web (online). Accessed February 16, 2011 at http://animaldiversity.org. http://www.animaldiversity.org
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Life History and Behavior

Behavior

Communication and Perception

Ants can see, but not very well. They mainly communicate with scent and touch. They have complicated chemical signals that allow them to work together on different tasks. They often spread information by touching each other's antennae or head. Some ants also make noises by rubbing their legs against their body.

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Life Cycle

Development

Ants have complete metamorphosis. Queen ants lay eggs. The baby ant that hatches from the egg is a larva, with no legs, just a soft white body like a worm and a small head. The larvae are fed by the queen (in the first generation) and then by workers. The amount and kind of food an ant larva gets helps determine how big it will be as an adult, and whether it will be a worker or a queen. Each larva grows and molts, and eventually spins a small cocoon of silk, and inside the cocoon it transforms into a pupa. The pupa is a resting stage, it doesn't move or eat, but just completes the transformation into an adult ant. The new adult emerges from cocoon to join start working for the nest.

Development - Life Cycle: metamorphosis

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Life Expectancy

Lifespan/Longevity

Queen ants can live for several years. Workers may live for a year but many only live for a few months. Males die as soon as they mate, so they only live for a few weeks. Sometimes a nest has several queens, and they can keep a large colony going for many years.

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Reproduction

Mating System: monogamous ; eusocial

Ants live in colonies where one or a few females, called queens, lay all the eggs. Most of the queens' offspring become worker ants that do not reproduce. A few are males, and some become new queens. Each queen ant can lay thousands of eggs per year.

Breeding interval: In most species, queen ants only mate once in their life.

Breeding season: Summer

Key Reproductive Features: iteroparous ; year-round breeding ; sexual ; fertilization (Internal ); oviparous ; sperm-storing

When a new queen finds a good place for a nest, she builds a small chamber and lays some eggs. When the eggs hatch, she finds food, and feeds and takes care of them until they mature. They become workers, and they take over all the work in the nest. The queen does nothing but lay eggs.

Parental Investment: female parental care

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Evolution and Systematics

Functional Adaptations

Functional adaptation

Air scoops provide cooling: ants
 

Air scoops on the sides of ants cool them through evaporation.

     
  "Another reason ants succeed so well is that they're superb lawn-traversing machines. When this first one backs away from the shadow of the giant human and reenters the main part of the sunny, hot lawn, little air-scoops on its side automatically switch on. A mist of cooling water vapor puffs upward from them. That keeps the ant's temperature down, but it could also mean that the ant's nitrogen--the equivalent of our urine substances--would become overconcentrated." (Bodanis 1992: 39)
  Learn more about this functional adaptation.
  • Bodanis, D. 1992. The Secret Garden: Dawn to Dusk in the Astonishing Hidden World of the Garden. Simon & Schuster. 187 p.
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Functional adaptation

Tongue sticks out elastically: ants
 

The tongue of ants protracts using elastic mechanisms.

   
  "The mouthparts are very important tools for almost any task performed by ants. In particular, the labiomaxillary complex is essential for food intake. In the present study we investigated the anatomical design of the labiomaxillary complex in various ant species, focusing on movement mechanisms. Six labial and six maxillary muscles with different functions control the several joints and ensure the proper performance of the labiomaxillary complex…the labial and maxillary muscles feature rather slow than fast muscle characteristics and do not seem to be specialized for specific tasks. Since glossa [tongue] protractor muscles are absent, the protraction of the glossa, the distal end of the labium, is a nonmuscular movement. By histological measurements of hemolymph volumes we could exclude a pressure-driven mechanism. Additional experiments showed that, upon relaxation of the glossa retractor muscles, the glossa protracts elastically. This elastic mechanism possibly sets an upper limit to licking frequency, thus influencing food intake rates and ultimately foraging behavior. In contrast to many other elastic mechanisms among arthropods, glossa protraction in ants is based on a mechanism where elasticity works as an actual antagonist to muscles. We compared the design of the labiomaxillary complex of ants with that of the honeybee and suggest an elastic mechanism for glossa protraction in honeybees as well." (Paul et al. 2002:39)

  Learn more about this functional adaptation.
  • Paul J; Roces F, Hölldobler. 2002. How do ants stick out their tongues?. Journal of Morphology. 254(1): 39-52.
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Functional adaptation

Communication is resilient: Temnothorax ants
 

Communication behaviors in Temnothorax ants are resilient because they have evolved as anytime algorithms.

   
  "Tandem runs are a form of recruitment in ants. During a tandem run,  a single leader teaches one follower the route to important resources  such as sources of food or better nest sites. In the present  study, we investigate what tandem leaders and followers do,  in the context of nest emigration, if their partner goes missing.  Our experiments involved removing either leaders or followers  at set points during tandem runs. Former leaders first stand  still and wait for their missing follower but then most often  proceed alone to the new nest site. By contrast, former followers  often first engage in a Brownian search, for almost exactly  the time that their former leader should have waited for  them, and then former followers switch to a superdiffusive search.  In this way, former followers first search their immediate neighbourhood  for their lost leader before becoming ever more wide ranging  so that in the absence of their former leader they can often  find the new nest, re-encounter the old one or meet a new  leader. We also show that followers gain useful information even  from incomplete tandem runs. These observations point to the  important principle that sophisticated communication behaviours may  have evolved as anytime algorithms, i.e. procedures that are  beneficial even if they do not run to completion." (Franks et al. 2010:1697)

  Learn more about this functional adaptation.
  • Franks NR; Richardson TO; Keir S; Inge SJ; Bartumeus F; Sendova-Franks AB. 2010. Ant search strategies after interrupted tandem runs. Journal of Experimental Biology. 213: 1697-1708.
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Functional adaptation

Colonies distribute food after famine: Temnothorax ants
 

Colonies of Temnothorax ants distribute food effectively and safely after famine using multiple techniques, including food dilution, strategic food location within community, and living 'silos'.

       
  "Resource distribution is fundamental to social organization, but it  poses a dilemma. How to facilitate the spread of useful resources but  restrict harmful substances? This dilemma reaches a zenith in famine  relief. Survival depends on distributing food fast but that could  increase vulnerability to poisons. We tested how Temnothorax  albipennis ants solve this dilemma in the distribution of honey  solution after 48 h of starvation in four colonies with individually  marked workers. We constructed the complete network of liquid food  transmission (trophallaxis) between individuals. Within the first 30 min  of famine relief, 95% of the workers received food and the distribution  rate was an order of magnitude faster compared to the controls. We  tested the assumptions of a simple analytical model that best fitted our  data. Good mixing during famine relief was facilitated by the movement  of internal workers away from the brood pile and the movement of  foragers with food away from the nest entrance. This is intriguing  because T. albipennis workers have spatial fidelity zones and in  the controls internal and external workers were segregated. We  discovered that colony vulnerability to poisons during famine relief  might be mitigated by: (1) the dilution of food from the same source  through mixing, (2) the concentration of food in workers positioned  midway between the colony centre and its periphery and (3) the existence  of living 'silos'. The latter are expendable foragers, who stay inside  the nest and store food during famine relief, thus acting as potential  disposable testers for food toxicity." (Sendova-Franks et al. 2010:473)

  Learn more about this functional adaptation.
  • Sendova-Franks AB; Hayward RK; Wulf B; Klimek T; James R; Planqué R; Britton NF; Franks NR. 2010. Emergency networking: famine relief in ant colonies. Animal Behaviour. 79(2): 473-485.
  • Viegas J. 2010. Poison-taster ants help save colonies. Discovery News [Internet],
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Functional adaptation

Collaborating for group decisions: honeybees
 

Honeybees collaborate when foraging, selecting a new hive through knowledge sharing.

     
  "Researchers at the Univ. of Illinois at Urbana-Champaign, led by principal researcher Feniosky Pena-Mora, are looking at ways to improve human collaboration during disaster relief efforts. They are attempting to draw inspiration from the collaboration patterns that honeybees use in their decision-making process when selecting a new hive or foraging, ants' behavior when they are under threat, and how infectious diseases spread among human populations. The team includes biological, computer, and social scientists, and civil engineers. The team believes that civil engineers should be a fourth group of first-responders at disaster relief efforts involving critical physical infrastructures. The researchers will develop ad hoc communication networks to spread critical information among first responders, similar to how a virus spreads. Models of collaboration based on study of ants and bees may be useful in understanding the basic principles and best practices when developing strategies to coordinate knowledge sharing in chaotic social settings." (Courtesy of the Biomimicry Guild)
  Learn more about this functional adaptation.
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Functional adaptation

Relationship provides nutrients, protection: ant-plants and ants
 

Ant-plants and their ant lodgers gain nutrients and protection thanks to their mutualistic relationship.

         
  "One group of plants, the ant-plants, provide even more lavish accomodation for their ant-lodgers. They are epiphytes, and are very common growing on the branches of mangroves. In such a position, without roots in the ground, they are in particular need of mineral nutrients. Their guests provide it. The ant-plant's stem is swollen into a globe the size of a football and armoured on the outside by prickles. Ants swarm all over it, scurrying in and out of holes on the surface. Within, there are a number of large interconnected chambers. Some are the ants' living quarters. There the queen sits, steadily producing her eggs, and there too are the nurseries where the young larvae are kept and reared. These apartments have smooth light-coloured walls. But other chambers are different. These have darker walls which are covered with small warty outgrowths. Here the ants deposit the remains of their insect meals and their droppings. Both are rich in phosphates and nitrates, exactly the nutrients that the plant badly needs since, hanging on the branches of a mangrove tree in a brackish swamp, it is cut off from the soil. It absorbs them through the walls of these compartments and so is able to flourish in one of the most difficult and impoverished of habitats for a plant. But it can only do so because its insect lodgers pay rent by feeding it." (Attenborough 1995:209-211)
  Learn more about this functional adaptation.
  • Attenborough, D. 1995. The Private Life of Plants: A Natural History of Plant Behavior. London: BBC Books. 320 p.
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Molecular Biology and Genetics

Molecular Biology

Statistics of barcoding coverage

Barcode of Life Data Systems (BOLD) Stats
Specimen Records: 84292
Specimens with Sequences: 63455
Specimens with Barcodes: 55355
Species: 6384
Species With Barcodes: 4762
Public Records: 33345
Public Species: 1628
Public BINs: 6791
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Barcode data

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Conservation

Conservation Status

Several ant species are considered threatened or in danger of extinction. This is because they live in special habitats that are very rare and may be destroyed by human construction or environmental change.

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Relevance to Humans and Ecosystems

Benefits

Economic Importance for Humans: Negative

Ants can be major pests. Carpenter ants make their nests in wood, including houses, and several ant species come into houses looking for food. In the tropics leaf-cutter ants attack crops in the fields. Some stinging ants can be dangerous to people. Recently an ant species from South America was accidentally brought to the southern United States. It is called the Imported Fire Ant, and it makes large nests with thousands of stinging defenders.

Negative Impacts: injures humans (bites or stings); crop pest; household pest

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Economic Importance for Humans: Positive

Ants are important predators of insects, including flies, caterpillars, and other pests.

Positive Impacts: controls pest population

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Pollinator

Ants (Order: Hymenoptera; Family: Formicidae) are highly social insects and are often associated in one way or another with plants. Ants sometimes form mutualistic relationships with plants, which may benefit from ant predation on plant herbivores or seed dispersal by ants. However, there are relatively few examples of pollination by ants.

In some cases, ants actually appear to interfere with pollination, sometimes reducing plant reproductive output: they may consume nectar without providing the plant with any reproductive benefit; they are aggressive toward other insects, including pollinators; they can destroy floral tissue; and their secretions may reduce pollen viability. Some plants appear to have evolved means of minimizing ant visitation to their flowers. In one example of ant interference with pollination, the ant, Crematogaster scutellaris, is a major predator of the fig wasp, which forms an obligate pollination mutualism with the Mediterranean fig tree (Genus: Ficus L.)

  • Complex interactions on fig trees: ants capturing parasitic wasps as possible indirect mutualists of the fig - fig wasp interaction, B. Schatz, M. Proffit, B. V. Rakhi, R. M. Borges, and M. Hossaert-McKey, In OIKOS, Volume 113, pages 344-352, 2006
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Wikipedia

Ant

For other uses, see Ant (disambiguation).


Myrmicinae



Formicinae






A phylogeny of the extant ant subfamilies.[1][2]
*Cerapachyinae is paraphyletic
‡ The previous dorylomorph subfamilies were synonymized under Dorylinae by Brady et al. in 2014[3]

Ants are eusocial insects of the family Formicidae /fɔrˈmɪsɨd/ and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than 12,500 of an estimated total of 22,000 species have been classified.[4][5] They are easily identified by their elbowed antennae and the distinctive node-like structure that forms their slender waists.

Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies that may occupy large territories and consist of millions of individuals. Larger colonies consist mostly of sterile, wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies sometimes are described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.[6]

Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and a few remote or inhospitable islands. Ants thrive in most ecosystems and may form 15–25% of the terrestrial animal biomass.[7] Their success in so many environments has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships.[8]

Ant societies have division of labour, communication between individuals, and an ability to solve complex problems.[9] These parallels with human societies have long been an inspiration and subject of study. Many human cultures make use of ants in cuisine, medication, and rituals. Some species are valued in their role as biological pest control agents.[10] Their ability to exploit resources may bring ants into conflict with humans, however, as they can damage crops and invade buildings. Some species, such as the red imported fire ant (Solenopsis invicta), are regarded as invasive species, establishing themselves in areas where they have been introduced accidentally.[11]

Etymology[edit]

The word "ant" is derived from ante, emete of Middle English which are derived from ǣmette of Old English, and is related to the dialectal Dutch emt and the Old High German āmeiza, hence the modern German Ameise. All of these words come from West Germanic *ēmaitijǭ, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut").[12][13] The family name Formicidae is derived from the Latin formīca ("ant")[14] from which the words in other Romance languages, such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnică, and French fourmi are derived. It has been hypothesised that a Proto-Indo-European word *morwi- was used, cf. Sanskrit vamrah, Latin formīca, Greek μύρμηξ mýrmēx, Old Church Slavonic mraviji, Old Irish moirb, Old Norse maurr, Dutch mier.[15]

Taxonomy and evolution[edit]

Ants fossilised in Baltic amber
Aculeata

Chrysidoidea


 
 

Vespidae



Rhopalosomatidae



 
 

Pompilidae



Tiphiidae



 

Scolioidea


 

Apoidea



Formicidae







Phylogenetic position of the Formicidae.[16]

The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees, and wasps. Ants evolved from a lineage within the aculeate wasps, and a 2013 study suggests they are a sister group of the Apoidea.[16] In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma) that lived in the Cretaceous period. The specimen, trapped in amber dating back to around 92 million years ago, has features found in some wasps, but not found in modern ants.[17] Sphecomyrma possibly was a ground forager, while Haidomyrmex and Haidomyrmodes, related genera in subfamily Sphecomyrminae, are reconstructed as active arboreal predators.[18] After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.[19][20][21][22] Some groups, such as the Leptanillinae and Martialinae, are suggested to have diversified from early primitive ants that were likely to have been predators underneath the surface of the soil.[2][23]

During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian supercontinent (the Northern Hemisphere). They were scarce in comparison to the populations of other insects, representing only about 1% of the entire insect population. Ants became dominant after adaptive radiation at the beginning of the Paleogene period. By the Oligocene and Miocene, ants had come to represent 20–40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, around one in 10 genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene).[19][24]

Termites, although sometimes called 'white ants', are not ants. They belong to the order Isoptera. Termites are more closely related to cockroaches and mantids. Termites are eusocial, but differ greatly in the genetics of reproduction. The similarity of their social structure to that of ants is attributed to convergent evolution.[25] Velvet ants look like large ants, but are wingless female wasps.[26][27]

Distribution and diversity[edit]

RegionNumber of
species [28]
Neotropics2162
Nearctic580
Europe180
Africa2500
Asia2080
Melanesia275
Australia985
Polynesia42

Ants are found on all continents except Antarctica, and only a few large islands, such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species.[29][30] Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators, and scavengers. Most species are omnivorous generalists, but a few are specialist feeders. Their ecological dominance may be measured by their biomass and estimates in different environments suggest that they contribute 15–20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.[7]

Ants range in size from 0.75 to 52 millimetres (0.030–2.0 in),[31][32] the largest species being the fossil Titanomyrma giganteum, the queen of which was 6 centimetres (2.4 in) long with a wingspan of 15 centimetres (5.9 in).[33] Ants vary in colour; most ants are red or black, but a few species are green and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of the potential existence of about 22,000) (see the article List of ant genera), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species.[34] The relative ease with which ants may be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.[35][36]

Morphology[edit]

Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma, and metasoma are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole may be formed by one or two nodes (the second alone, or the second and third abdominal segments).[37]

Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision

Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases such as carbon dioxide pass through their exoskeleton via tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph toward the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages.[38]

Diagram of a worker ant (Pachycondyla verenae)

Head[edit]

An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ant eyes are good for acute movement detection, but do not offer a high resolution image. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization.[39] Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have excellent vision and are capable of discriminating the distance and size of objects moving nearly a metre away.[40]

Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents, and vibrations; they also are used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence.[38] In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it may be passed to other ants or their larvae.[41]

Legs[edit]

All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and to hang onto surfaces.[citation needed]

Wings[edit]

Only reproductive ants, queens and males, have wings. Queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. Wingless queens (ergatoids) and males occur in a few species, however.[38]

Metasoma[edit]

The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae), and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.[38]

Polymorphism[edit]

Seven Leafcutter ant workers of various castes (left) and two Queens (right)

In the colonies of a few ant species, there are physical castes—workers in distinct size-classes, called minor, median, and major workers. Often, the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they still are workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species, the median workers are absent, creating a sharp divide between the minors and majors.[42] Weaver ants, for example, have a distinct bimodal size distribution.[43][44] Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights.[45] Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile, haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes.[46] For instance, these replete workers develop in the North American honeypot ant Myrmecocystus mexicanus. Rissing found that usually the largest workers in the colony develop into repletes, and if repletes are removed from the colony other workers become repletes, demonstrating the flexibility of this particular polymorphism.[47] This polymorphism in morphology and behaviour of workers initially was thought to be determined by environmental factors such as nutrition and hormones that led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp.[48] These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens.[49] The Australian jack jumper ant (Myrmecia pilosula) has only a single pair of chromosomes (with the males having just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.[50][51]

Development and reproduction[edit]

Meat eater ant nest during swarming

The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larva stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach". Larvae, especially in the later stages, may also be provided solid food such as trophic eggs, pieces of prey, and seeds brought by workers.

The larvae grow through a series of four or five moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa.[52] The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is influenced in some species by the nutrition the larvae obtain. Genetic influences and the control of gene expression by the developmental environment are complex and the determination of caste continues to be a subject of research.[53] Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so often, are moved around among the various brood chambers within the colony.[54]

A new worker spends the first few days of its adult life caring for the queen and young. She then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants who are older and are likely to die soon of natural causes.[55][56]

Fertilised meat-eater ant queen beginning to dig a new colony

Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens, while others may exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right.[57] The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, such as army ants, have wingless queens), and do nothing in life except eat and mate.

Most ants are univoltine, producing a new generation each year.[58] During the species-specific breeding period, new reproductives, females and winged males leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in others they may mate with as many as ten or more different males.[8]

Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food, and care for the other eggs. This is how new colonies start in most ant species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site,[59] a process akin to swarming in honeybees.

Ants mating

A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis. [60]

Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, being quite short-lived and surviving for only a few weeks.[61] Ant queens are estimated to live 100 times longer than solitary insects of a similar size.[62]

Ants are active all year long in the tropics, but, in cooler regions, they survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state, (diapause), while in others, the adults alone pass the winter in a state of reduced activity.[63]

Behaviour and ecology[edit]

Communication[edit]

An ant trail
A Plectroctena sp. attacks another of its kind to protect its territory

Ants communicate with each other using pheromones, sounds, and touch.[64] The use of pheromones as chemical signals is more developed in ants, such as the red harvester ant, than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin, and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that may be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually identifying the best path.[65]

Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from farther away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves.[66] Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum, and hind tibia.[62] Pheromones also are exchanged, mixed with food, and passed by trophallaxis, transferring information within the colony.[67] This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to.[68] In ant species with queen castes, when the dominant queen stops producing a specific pheromone, workers begin to raise new queens in the colony.[69]

Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.[70][71]

Defence[edit]

See also Insect defences

Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals, such as formic acid in the case of formicine ants, alkaloids and piperidines in fire ants, and a variety of protein components in other ants. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index.

The sting of jack jumper ants can be fatal,[72] and an antivenom has been developed for it.[73]

Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids.[74] Their stings are painful and can be dangerous to hypersensitive people.[75]

A weaver ant in fighting position, mandibles wide open

Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom.[76] One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 – 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backward to escape a threat.[76] Before striking, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory organs resembling hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws also are seen in the following genera: Anochetus, Orectognathus, and Strumigenys,[76] plus some members of the Dacetini tribe,[77] which are viewed as examples of convergent evolution.

A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies.[78]

Suicidal defences by workers are also noted in a Brazilian ant, Forelius pusillus, where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.[79]

Ant mound holes prevent water from entering the nest during rain

In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates.[80] Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana[81] while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates to trigger similar behaviour.[82]

Nests may be protected from physical threats such as flooding and overheating by elaborate nest architecture.[83][84] Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside.[85] Camponotus anderseni, which nests in the cavities of wood in mangrove habitats, deals with submergence under water by switching to anaerobic respiration.[86]

Learning[edit]

Many animals can learn behaviours by imitation, but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis will lead a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. The leader is acutely sensitive to the progress of the follower and slows down when the follower lags and speeds up when the follower gets too close.[87]

Controlled experiments with colonies of Cerapachys biroi suggest that an individual may choose nest roles based on her previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out fewer and fewer times. A month later, the successful foragers continued in their role while the others had moved to specialise in brood care.[88]

Nest construction[edit]

Main article: Ant colony

Complex nests are built by many ant species, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests may be found in the ground, under stones or logs, inside logs, hollow stems, or even acorns. The materials used for construction include soil and plant matter,[59] and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.[89]

The army ants of South America, such as the Eciton burchellii species, and the driver ants of Africa do not build permanent nests, but instead, alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together.[90]

Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.[91]

Formica polyctena, among other ant species, constructs nests that maintain a relatively constant interior temperature that aids in the development of larvae. The ants maintain the nest temperature by choosing the location, nest materials, controlling ventilation and maintaining the heat from solar radiation, worker activity and metabolism, and in some moist nests, microbial activity in the nest materials.[92]

Some ant species, such as those that use natural cavities, can be opportunistic and make use of the controlled micro-climate provided inside human dwellings and other artificial structures to house their colonies and nest structures.[93][94]

Cultivation of food[edit]

Main article: Ant-fungus mutualism
Myrmecocystus, honeypot ants, store food to prevent colony famine

Most ants are generalist predators, scavengers, and indirect herbivores,[21] but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in related tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is found to be toxic to the fungus, the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria introduced into the nest that may harm the fungi.[95]

Navigation[edit]

Foraging ants travel distances of up to 200 metres (700 ft) from their nest [96] and scent trails allow them to find their way back even in the dark. In hot and arid regions, day-foraging ants face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants of the genus Cataglyphis such as the Sahara desert ant navigate by keeping track of direction as well as distance travelled. Distances travelled are measured using an internal pedometer that keeps count of the steps taken [97] and also by evaluating the movement of objects in their visual field (optical flow).[98] Directions are measured using the position of the sun.[99] They integrate this information to find the shortest route back to their nest.[100] Like all ants, they can also make use of visual landmarks when available [101] as well as olfactory and tactile cues to navigate.[102][103] Some species of ant are able to use the Earth's magnetic field for navigation.[104] The compound eyes of ants have specialised cells that detect polarised light from the Sun, which is used to determine direction.[105][106] These polarization detectors are sensitive in the ultraviolet region of the light spectrum.[107] In some army ant species, a group of foragers who become separated from the main column sometimes may turn back on themselves and form a circular ant mill. The workers may then run around continuously until they die of exhaustion.[108]

Locomotion[edit]

The female worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs.[109] There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.[110]

Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands.[111] Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they go to trapped pockets of air in the submerged nests to breathe.[112]

Cooperation and competition[edit]

Meat-eater ants feeding on a cicada, social ants cooperate and collectively gather food

Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal of ants. Like virtually all ants, they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using her large eyes instead of chemical senses to find prey.[113]

Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist, but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers or slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive.[114] Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults).[115]

A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen

Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.[116] Also, the reason why two separate colonies of ants will attack each other even if they are of the same species is because the genes responsible for pheromone production are different between them. The Argentine ant, however, does not have this characteristic, due to lack of genetic diversity, and has become a global pest because of it.

Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species such as Strumigenys xenos are entirely parasitic and do not have workers, but instead, rely on the food gathered by their Strumigenys perplexa hosts.[117][118] This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen may enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest.[119]

A conflict between the sexes of a species is seen in some species of ants with these reproducers apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process whereby a diploid egg loses its maternal contribution to produce haploid males who are clones of the father.[120]

Relationships with other organisms[edit]

The spider Myrmarachne plataleoides (female shown) mimics weaver ants to avoid predators.

Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They also are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae, and eggs, consuming the food stores of the ants, or avoiding predators. These inquilines may bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.[121][122]

Ant takes nectar
An ant collects honeydew from an aphid

Aphids and other hemipteran insects secrete a sweet liquid called honeydew, when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect.[123] In some cases, the aphids secrete the honeydew in response to ants tapping them with their antennae. The ants in turn keep predators away from the aphids and will move them from one feeding location to another. When migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs may become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.[124]

Myrmecophilous (ant-loving) caterpillars of the butterfly family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants.[125] Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ant nest where it feeds on the ant larvae.[126] Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant, Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.[127]

Ants may obtain nectar from flowers such as the dandelion but are only rarely known to pollinate flowers.

Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees, (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees.[128] Although some ants obtain nectar from flowers, pollination by ants is somewhat rare.[129] Some plants have special nectar exuding structures, extrafloral nectaries that provide food for ants, which in turn protect the plant from more damaging herbivorous insects.[130] Species such as the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) who defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants.[131] In return, the ants obtain food from protein- and lipid-rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.

Many tropical tree species have seeds that are dispersed by ants.[132] Seed dispersal by ants or myrmecochory is widespread and new estimates suggest that nearly 9% of all plant species may have such ant associations.[133][134] Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal as the seeds are transported to safety below the ground. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food.[135]

A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.[136]

A meat ant tending a common leafhopper nymph

Most ants are predatory and some prey on and obtain food from other social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants.[96] Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away predatory ant species.[137] The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellent chemical.[138] It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.[96]

Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants.[139] Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.[139]

Fungi in the genera Cordyceps and Ophiocordyceps infect ants. Ants react to their infection by climbing up plants and sinking their mandibles into plant tissue. The fungus kills the ants, grows on their remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores [140] in a microhabitat that best suits the fungus.[141] Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates.[142]

A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black-coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, causing them to carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits, such as Hyeronima alchorneoides, and eaten. The droppings of the bird are collected by other ants and fed to their young, leading to further spread of the nematode.[143]

Spiders sometimes feed on ants

South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants.[144]

Army ants forage in a wide roving column, attacking any animals in that path that are unable to escape. In Central and South America, Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers.[145][146] This behaviour was once considered mutualistic, but later studies found the birds to be parasitic. Although direct kleptoparasitism (birds stealing food from the ants' grasp) is rare, the birds eat many prey insects that the ants would otherwise eat and thus decrease their foraging success.[147] Birds indulge in a peculiar behaviour called anting that, as yet, is not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may be a means to remove ectoparasites from the birds.

Anteaters, aardvarks, pangolins, echidnas and numbats have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants. About 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.[148]

Relationship with humans[edit]

Weaver ants are used as a biological control for citrus cultivation in southern China

Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control.[10] On the other hand, ants may become nuisances when they invade buildings, or cause economic losses.

In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.[149][150][151]

Some ants have toxic venom and are of medical importance. The species include Paraponera clavata (tocandira) and Dinoponera spp. (false tocandiras) of South America [152] and the Myrmecia ants of Australia.[153]

In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds may be collected from one ant-heap.[154][155]

Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These tend to be island species that have evolved specialized traits and risk being displaced by introduced ant species. Examples include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.[156]

It has been estimated by E.O. Wilson that the total number of individual ants alive in the world at any one time is between one and ten quadrillion (short scale) (i.e. between 1015 and 1016). According to this estimate, the total biomass of all the ants in the world is approximately equal to the total biomass of the entire human race.[157] Also, according to this estimate, there are approximately 1 million ants for every human on Earth.[158]

As food[edit]

See also: Entomophagy
Roasted ants in Colombia
Ant larvae for sale in Isaan, Thailand

Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are used in Mexican escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.[159]

In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry.[160] Weaver ant eggs and larvae, as well as the ants, may be used in a Thai salad, yam (Thai: ยำ), in a dish called yam khai mot daeng (Thai: ยำไข่มดแดง) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".[161]

In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickling, acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).[161]

As pests[edit]

The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper

Some ant species are considered as pests. The presence of ants can be undesirable in places meant to be sterile. They can also come in the way of humans by their habit of raiding stored food, damaging indoor structures, causing damage to agricultural crops either directly or by aiding sucking pests or because of their stings and bites.[11] The adaptive nature of ant colonies make it nearly impossible to eliminate entire colonies and most pest management practices aim to control local populations and tend to be temporary solutions.

Some of the ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant, and European fire ant. Ant populations are managed by a combination of approaches that make use of chemical, biological and physical methods. Chemical methods include the use of insecticidal bait which is gathered by ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Management is based on the species and techniques can vary according to the location and circumstance.[11]

In science and technology[edit]

Observed by humans since the dawn of history, the behaviour of ants has been documented and the subject of early writings and fables passed from one century to another. Those using scientific methods, myrmecologists, study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbock in 1881.[162] Studies on ants have tested hypotheses in ecology and sociobiology, and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies.[163] Ant colonies may be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures.[164] Individuals may be tracked for study by marking them with dots of colours.[165]

The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems, for example Ant colony optimization and Ant robotics. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage, and networking algorithms.[9]

In culture[edit]

Aesop's ants: picture by Milo Winter, 1888–1956

Anthropomorphised ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They also are mentioned in religious texts.[166][167] In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army.[Quran 27:18][168] In parts of Africa, ants are considered to be the messengers of the deities. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief.[169] Ant bites are used in the initiation ceremonies of some Amazon Indian cultures as a test of endurance.[170][171]

Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his 1880 book A Tramp Abroad.[172] Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. H.G. Wells wrote about intelligent ants destroying human settlements in Brazil and threatening human civilization in his 1905 science-fiction short story, The Empire of the Ants. In more recent times, animated cartoons and 3-D animated films featuring ants have been produced including Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, and Atom Ant. Renowned myrmecologist E. O. Wilson wrote a short story, "Trailhead" in 2010 for The New Yorker magazine, which describes the life and death of an ant-queen and the rise and fall of her colony, from an ants' point of view.[173]

From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil, allowing greater visibility.[174] In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".[175]

Ants also are quite popular inspiration for many science-fiction insectoids, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the films Them! and Empire of the Ants, Marvel Comics' super hero Ant-Man, and ants mutated into super-intelligence in Phase IV. In computer strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.[176]

See also[edit]

Main article: Outline of ants

References[edit]

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Cited texts[edit]

Further reading[edit]

  • Bolton, Barry (1995). A New General Catalogue of the Ants of the World. Harvard University Press. ISBN 978-0-674-61514-4. 
  • Hölldobler B, Wilson EO (1998). Journey to the Ants: A Story of Scientific Exploration. Belknap Press. ISBN 0-674-48526-2. 
  • Hölldobler B, Wilson EO (1990). The Ants. Harvard University Press. ISBN 978-3-540-52092-4. 
  • Hölldobler B, Wilson EO (2009). The Superorganism: The Beauty, Elegance and Strangeness of Insect Societies. Norton & Co. ISBN 978-0-393-06704-0. 
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