Overview
Brief Summary
Overview
- 1. “Ant Information.” Center for Insect Science Education Outreach, University of Arizona. 1997. 29 Jul. 2011. http://insected.arizona.edu/antinfo.htm
- 2. Hölldobler, Bert and Edward O. Wilson. The Ants. Cambridge: Harvard University Press, 1990.
- 3. Roof, Jennifer. “Family Formicidae: Ants.” Animal Diversity Web. University of Michigan Museum of Zoology. 2001. 1 Sept. 2011. http://animaldiversity.ummz.umich.edu/site/accounts/information/Formicidae.html
Unreviewed
Comprehensive Description
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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Hilty, J. Editor. 2010. Insect Visitors of Illinois Wildflowers. World Wide Web electronic publication. flowervisitors.info, version (09/2010).
See: Abbreviations for Insect Activities, Abbreviations for Scientific Observers, References for behavioral observations H
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Description of Formicidae
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Characteristics
- 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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Physical Description
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
Associations
Known predators
Amphisbaena caeca
Eleutherodactylus coqui
Eleutherodactylus richmondi
Eleutherodactylus portoricensis
Eleutherodactylus wightmanae
Eleutherodactylus eneidae
Melanerpes portoricensis
Todus mexicanus
Mimocichla plumbea
Margarops fuscatus
Anolis cuvieri
Anolis evermanni
Anolis stratulus
Anolis gundlachi
Leptodactylus albilabris
Myiarchus antillarum
Nesospingus speculiferus
Icterus dominicensis
Vireo altiloquus
Seiurus aurocapillus
Sphaerodactylus klauberi
Sphaerodactylus macrolepis
Diploglossus pleei
Anthracothorax viridis
Parula americana
Dendroica caerulescens
Dendroica discolor
Typhlops rostellatus
Eptesicus fuscus
Lasiurus borealis
Dendroica petechia
Loxigilla noctis
Trochilidae
Anolis gingivinus
Anolis pogus
Orthoptera
Araneae
Chilopoda
Based on studies in:
Puerto Rico, El Verde (Rainforest)
This list may not be complete but is based on published studies.
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Known prey organisms
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.
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Associations
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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Evolution and Systematics
Functional Adaptations
Functional adaptation
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
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 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 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
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
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
Barcode
Locations of barcode samples
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Statistics of barcoding coverage
| Specimen Records: | 48,357 |
| Specimens with Sequences: | 35,037 |
| Specimens with Barcodes: | 31,063 |
| Public Records: | 4,439 |
| Species: | 4,210 |
| Species With Barcodes: | 3,546 |
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Relevance to Humans and Ecosystems
Benefits
Pollinator
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
A phylogeny of the extant ant subfamilies.[1][2]
*Cerapachyinae is paraphyletic
Ants are social insects of the family Formicidae (pronounced /fɔrˈmɪsəˌdiː/), and along with the related wasps and bees, they 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.[3] More than 12,500 species are classified and the upper estimates of species is about 22,000.[4][5][6] 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 groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.[7]
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15–25% of the terrestrial animal biomass.[8] Their success 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.[9]
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems.[10] 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.[11] However, their ability to exploit resources brings ants 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, since they have established themselves in new areas where they have been accidentally introduced.[12]
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Etymology
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German āmeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut").[13][14] The family name Formicidae is derived from the Latin formīca ("ant")[15] from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnică and French fourmi are derived.
Taxonomy and evolution
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.[16][17][18] In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps.[19] Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.[2]
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During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary 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, approximately one in ten 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).[16][21]
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution.[22] Velvet ants look like large ants, but are wingless female wasps.[23][24]
Distribution and diversity
| Region | Number of species [25] |
|---|---|
| Neotropics | 2162 |
| Nearctic | 580 |
| Europe | 180 |
| Africa | 2500 |
| Asia | 2080 |
| Melanesia | 275 |
| Australia | 985 |
| Polynesia | 42 |
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.[26][27] 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.[8]
Ants range in size from 0.75 to 52 millimetres (0.030–2.0 in).[28][29] Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), 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.[30] The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.[31][32]
Morphology
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 or gaster 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 can be formed by one or two nodes (the second alone, or the second and third abdominal segments).[33]
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 like carbon dioxide pass through their exoskeleton through 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 towards 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.[34]
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization.[35] 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 exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also 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.[34] In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.[36]
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.[34]
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.[34]
Polymorphism
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 are still 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.[37] Weaver ants, for example, have a distinct bimodal size distribution.[38] [39] 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.[40] 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.[41] This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp.[42] 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.[43] The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have 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.[44][45]
Development and reproduction
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 larval 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", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa.[46] The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony.[47]
A new worker spends the first few days of its adult life caring for the queen and young. It 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 that are older and are likely to die soon of natural causes.[48][49]
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 can 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.[50] The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year.[51] During the species specific breeding period, new reproductives, winged males and females 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 some others they may mate with anywhere from one to ten or more different males.[9] 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 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,[52] a process akin to swarming in honeybees.
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[53] and one species, Mycocepurus smithii is known to be all-female.[54]
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, and survive only a few weeks.[55] Ant queens are estimated to live 100 times longer than solitary insects of a similar size.[56]
Ants are active all year long in the tropics but, in cooler regions, 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.[57]
Behaviour and ecology
Communication
Ants communicate with each other using pheromones.[58] These chemical signals are more developed in ants 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 can 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 finding the best path.[59]
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 further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves.[60] 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.[56] Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony.[61] This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to.[62] In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone.[63]
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.[64][65]
Defence
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. 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,[66] and an antivenom has been developed.[67] Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids.[68] Their stings are painful and can be dangerous to hypersensitive people.[69]
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.[70] One study of
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