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Observer Cards Ants Ants Ants www.eol.org

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Adam Lazarus, Consult Ants Edited by Jeff Holmes, EOL, Harvard University About Observer Cards EOL Observer Cards Observer cards are designed to foster the art and science of observing nature. Each set provides information about key traits and techniques necessary to make accurate and useful scientific observations. The cards are not designed to identify species but rather to encourage detailed observations. Take a journal or notebook along with you on your next nature walk and use these cards to guide your explorations. Observing Ants Ants are some of the most fascinating creatures in nature and can be observed almost everywhere! There are approximately 15,000 described species of ants (and perhaps 10,000 not yet described) with endless variations and adaptations to their environment. Use these cards to help you focus on the key traits and behaviors that make different ant species unique. Drawings and photographs are a great way to supplement your field notes as you explore the tiny world of these amazing animals. Icon Legend: = Observable by eye = Magnification necessary

Author: Adam Lazarus, Consult Ants. Editor: Jeff Holmes, EOL, Harvard University. Created by the Encyclopedia of Life - www.eol.org Content Licensed Under a Creative Commons Attribution 3.0 License. Table of Contents Anatomy Diet Associations Body Shape Carnivory Visible Symbiosis Body Color Herbivory Hidden Symbiosis Eyes Diet Specialists Mites Mandibles Mimicry Organization Antennae Inquilines Worker Castes Thorax Human Impact Communication Petiole Segments Nest Location Pilosity (Hair) Life Cycle Ants in Groups Tip of Gaster Development Stages Wings Behavior Techniques Body Specialists Behavior Specialists Photography Ants at a Glance Perception Collecting Scientific Records Body Shape Ants Ants

1 Anatomy Anatomy 2

3 4 Body Shape EOL Observer Cards Many ants have tell-tale body shapes. Guides and keys often begin with a few words about overall shape and include terms such as compact, cyclindrical, or flattened. You might also note out-standing physical features such as a flattened head or long spines.

Supplemental Notes An ant’s body shape may say a lot about its livelihood. For example slender, gracile (delicately built) ants tend to be fast. But cylindrical ants usually live in long, thin tunnels.

Images: 1. Orectognathus clarki © servitude via Flickr; 2. Cephalotes sp. © Ricardo Solar via Flickr; 3. Solenopsis invicta (Red Fire Ant) © AntWeb.org; 4. Pseudomyrmex apache © Jen Fogarty via AntWeb.org.

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8 9 Body Color EOL Observer Cards Record the overall body color but keep in mind that a single species may come in a variety of colors. Ants come in a huge range of colors but the most common are red, brown and black. If you see a white ant, chances are it has just completed its change (metamorphosis) from a wormlike larva to an adult ant. In some species, it takes a few days for the adult pigments to fully develop. Supplemental Notes Color has no bearing on how dangerous an ant species is. There are harmless red ants, and black ants with very powerful stings. Some ant species have names that reflect their color, such as Rhytidoponera metallica, or Tapinoma melanocephalum (not pictured). Images: 1. © Gary Alpert; 2. Messor angularis, © April Nobile via Antweb; 3. Rhytidoponera metallica, © April Nobile via Antweb; 5. Oecophylla smaragdina, © April Nobile via Antweb; 6.© Adam Lazarus & Christopher Rieken; 7. Myrmecia gulosa, John Tann via Flickr; 8 © Adam Lazarus & Christopher Rieken; 9. © April Nobile via Antweb.

3 4 Eyes EOL Observer Cards Note the presence, number and size of eyes, and whether the eyes extend outside the outer margin of a frontal view of the face. Ants can have two, five or zero eyes. On some ants you can observe microscopic hairs growing within the compound eye. Supplemental Notes Some ants have acute vision, while others are totally blind. Most have two compound eyes. These faceted organs are composed of fixed-angle, hexagonal eyelets called “ommatidia”. The ant’s brain assembles the visual inputs from each ommatidium into a single image. Many species also have three additional eyelets arranged in a triangle on the top of the head. These “ocelli” are believed to be especially light sensitive. Sometimes only queens & males have ocelli, while the worker caste does not.

Images: 1-3. © April Nobile via Antweb; 1. Camponotus anthrax showing two eyes; 2. Formica xerophila - the circle highlights three ocelli; 3. Dorylus affinis: a blind species; 4. Magnified view of a compound eye showing interfacetal hairs © David Graham & James Style, Marine Biological Lab.

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3 5 6 Mandibles EOL Observer Cards The mandibles (or jaws) come in many styles and their structure can say a lot about the lifestyle of an ant. There may also be teeth present on the mandibles that may require a microscope to be seen. Supplemental Notes Perhaps the most widespread mandible type is some variation of that on the Carpenter Ant at center. The short teeth and short mandible length are ideal for digging, cutting and prey capture, as well as manipulating delicate objects like eggs and larvae. However, many ant species have modified mandibles to suit specialized lifestyles. Jaws that are sharply hooked help some ants hold onto live, struggling victims. Blunt jaws attached to a large head may be characteristic of a seed crusher. Lengthy and serrated mandibles are probably used for sawing through leaves or prey. For catching fast prey, some ants even have mandibles that snap shut by reflex when special trigger hairs are touched. Images: 1-3,5-6 © April Nobile via AntWeb; 1. Odontomachus troglodytes; 2. Atta mexicana; 3. Gigantiops destructor; 4. Myrmecia rubripes. © Jean and Fred Hort via Flickr; 5. Amblyopone australis; 6. Eciton vagans Author: Adam Lazarus, Consult Ants. Editor: Jeff Holmes, EOL, Harvard University. Created by the Encyclopedia of Life - www.eol.org Content Licensed Under a Creative Commons Attribution 3.0 License. Antennae 1 2 3 Ants Ants Anatomy Anatomy

4 5 Antennae EOL Observer Cards Antennae vary widely across ant species in their shape, number of segments, and sensitivity. Some antennae end in a swollen “club”, while others can be drawn into specialized grooves called scrobes for protection. Supplemental Notes The antenna is an ant’s single most important tool for interacting with the environment. It is used for smell, touch, spatial perception, and even as an aid in manipulating small, delicate objects. If an ant is born without functional antennae or loses them in a fight, it is totally helpless. Colony members often reject or cannibalise such individuals. Some ants use their antennae with utmost care, tapping lightly, cleaning them fastidiously, and holding them protectively against the body at the slightest sign of danger. Others drag these appendages along the ground, and seem less perceptive of the surrounding landscape.

Images: 1) Scanning electron micrograph of Tetraponera aethiops face, highlighting antennal anatomy. © Roberto Keller Perez/AMNH; 2 - 5) A comparison of antennal variability in ants. © April Nobile & Eli M. Sarnat via Antweb.

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3 5 Thorax EOL Observer Cards Most accurately called the alitrunk in ants, the middle section of an ant’s body is a trove of identifying features. The metapleural gland is present in many species, but conspicuously absent from others. The presence and shape of spines is important, as are the grooves and other sculpturing on the integument. Sometimes it is easiest to identify a queen from a mass of seething workers by the hump and wing scars on her alitrunk. Supplemental Notes Mesosoma is a general term for the middle section of an insect. In ants, the first abdomenal segment is fused to the thoracic segments and is part of the mesosoma. Alitrunk is the term used for this special arrangement. Images: 1) Aphaenogaster © Kim Fleming via Flickr; 2) A scanning electron micrograph of Tapinoma simrothi shows the location of the metapleural gland. © Roberto Keller/AMNH; 3) Polyrhachis bihamata is adorned with elaborate spines. © April Nobile via AntWeb; 4) © Ratabago via Flickr; 5) Queen carpenter ant showing wing scar on thorax. © Silversyrpher via Flickr.

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3 Petiole Segments EOL Observer Cards The petiole (waist) is a pinched abdominal segment common to all ants, and is sometimes followed by another pinched segment called the postpetiole. Be aware that the petiole may be hidden by other abdominal segments or adorned with elaborate spines. The postpetiole is a major diagnostic tool for species determination. It is absent from some ant subfamilies, and fused to the rest of the abdomen (gaster) in others. Supplemental Notes The petiole imparts tremendous flexibility. In combat, ants thrust the abdomen forward to sting or spray defensive chemicals. Images: 1–4 © April Nobile and/or Antweb.org; 1) Camponotus thadeus has a petiole, but no postpetiole; 2) Pogonomyrmex (Harvester Ant) species have a distinct petiole & postpetiole; 3) The petiole of Tapinoma sessile (the Coconut, Stink or Odorous House Ant) is obscured by overhanging gastral segments; 4) A thick segment behind the petiole of Hypoponera punctatissima that is fused to the gaster is referred to as Abdominal Segment 3 and is part of the gaster; 5) The long waist of Tetraponera sp. gives it extra flexibility. © James Niland via Flickr.

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One side only 4 5 Pilosity (Hair) EOL Observer Cards Observe the pilosity or hair characteristics of the ants. Often slight variations in hair orientation, shape or density are the main characters distinguishing one species from another. Common terms to describe pilosity are: recumbent (flat against the surface), decumbent (flat against the surface with ends raising up), erect, suberect, flagellate (whip-like), spatulate (club-like), dense, and glabrous (smooth, devoid of hair). Supplemental Notes Hairs are often used as sensory organs. Some ants of the genus Basiceros stuff dirt between their spoon-shaped hairs as a way to camouflage themselves. Images: 1) Camponotus floridanus. © Bob Peterson via Flickr; 2) Sericomyrmex sp. © Ricardo Solar via Flickr; 3) The hair pattern on these Aphaenogaster femurs is useful to tell species apart , ©John Longino & Stefan Cover; 4) Polyrhachis illaudata, myrmex via Flickr; 5) Camponotus molossus, Jean and Fred Hort via Flickr.

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3 Tip of Gaster EOL Observer Cards Observe the apparatus at the tip of the gaster. Depending on the subfamily or genus, an ant may have an acidipore, a ventral slit that is either armed or unarmed with stiff hairs, or a sting. Supplemental Notes Though ants are famous for spraying formic acid, only some do this. As a group, they produce a huge range of defensive chemicals which may be sprayed, smeared, shot or injected, depending on the species. Though the sting is generally hypodermic, it may be modified for smearing instead of injecting. It may also be modified for laying trails. Some people have very strong reactions to ant stings. For treatment information, visit http://www.antallergy.org.

Images: 1. Photo: Myrmecocystus mexicanus ©Jen Fogarty, AntWeb.org; 2. Dolichoderus pustulatus © California Academy of Science, Michael Branstetter; 3. Hypoponera opaciceps © April Nobile AntWeb.org; Schematics: Redrawn from Sarnat, E. M., PIAkey: Identification guide to ants of the Pacific Islands, Edition 2.0, Lucid v. 3.4. USDA/APHIS/PPQ Center for Plant Health Science and Technology and University of California.

Anatomy Anatomy 3 Wings EOL Observer Cards Males and unmated queen ants are born with wings. At certain times of the year (rainy season is typical), huge swarms of ants emerge as the colony forces these sexual forms out of the nest. Exposure to the outside triggers these ants to fly, and flying triggers them to mate. After mating, the male dies - his only role in ant society has been fulfilled. The female, now a queen, tears off her wings and looks for a safe place to found a new colony. She carries with her a lifetime supply of sperm. Supplemental Notes Despite the number and ubiquity of ants, it’s still not clear what makes an ant colony force its sexuals outside, or what causes many colonies of the same species do this simultaneously. Mated queens face a gauntlet of dangers that puts their estimated success rate at less than 1%. Images: 1) Lasius niger (Black garden ant) mating. © Daniele Benucci via Flickr; 2) Queen ant sheds her wings. © John Pittman via Flickr; 3) Males take flight. © Erik bij de Vaate via Flickr.

Anatomy Anatomy 3 4 Body Specialists EOL Observer Cards Males and unmated queen ants are born with wings. At certain times of the year (rainy season is typical), huge swarms of ants emerge as the colony forces these sexual forms out of the nest. Exposure to the outside triggers these ants to fly, and flying triggers them to mate. After mating, the male dies - his only role in ant society has been fulfilled. The female, now a queen, tears off her wings and looks for a safe place to found a new colony. She carries with her a lifetime supply of sperm. Supplemental Notes Despite the number and ubiquity of ants, it’s still not clear what makes an ant colony force its breeding outside, or what causes many colonies of the same species do this simultaneously. Mated queens face a gauntlet of dangers that puts their estimated success rate at less than 1%. Images: 1) Majors of the Turtle ant or Cephalotes varians use their uniquely shaped head to seal the nest from the inside. © April Nobile via AntWeb.org; 2) When threatened, Camponotus saundersi contracts the giant venom glands running the length of its body, showering its enemy in poison and guts. © Ulrich Maschwitz; 3) Cephalotes atratus uses airfoils in its head to direct a fall from a tree back to the trunck. © Yonatan Munk via Flickr; 4) This Crematogaster species curls its gastor and chemical weaponry over its head. © Sean McCann via Flickr. Author: Adam Lazarus, Consult Ants. Editor: Jeff Holmes, EOL, Harvard University. Created by the Encyclopedia of Life - www.eol.org Content Licensed Under a Creative Commons Attribution 3.0 License. Ants at a Glance Ants Ants Anatomy Anatomy Ants at a Glance EOL Observer Cards Ants come in a number of different body plans. Some anatomical features are always present, such as mandibles and antennae. Other features are more unique, such as the postpetiole or acidipore. With practice, you’ll be able to tell much about an ant at just a glance. Knowing more of these details will increase your appreciation and possibly help you discover something yet unknown! Supplemental Notes Closely related species have similar body plans, but the same parts may be called by different names. An ant’s posterior is commonly defined as the abdomen, and scientifically defined as the gaster. Technically, the first of the seven abdominal segments resides on what is commonly called the thorax. The true thorax and the first abdominal segment is termed the mesosoma.

Images: Ant schematics are adapted from The Ants (Holldobler & Wilson 1990, Belknap Press) and are previously adapted from (top to bottom): Brown, 1958 (Pachycondyla castanea), Bolton, 1981 (Baracidris meketra) & Wheeler, 1929 (Gesomyrmex kalshoveni).

3 4 Diet Carnivory EOL Observer Cards Almost all ants are occasional predators or scavengers. Look for dead insects being dragged back to the nest or check for ants on animal remains. Many ants are dedicated predators, and some are even specialized to hunt specific prey. Ants often attack insects attracted to lights at night.

Supplemental Notes Some ants hunt in coordinated groups while others are solitary hunters. It is a myth that large numbers of ants can kill a healthy human or other large animal but smaller prey such as frogs or snakes can be unfortunate victoms.

Images: 1. Weaver ants and Blue Banded bee © Troup Dresser via Flickr; 2. Eating another ant © gundy via Flickr; 3. Scavaging a dead catepillar © Glen Peterson via Flickr; 4. Harpegnathos sp. in Malaysia © Steve Shattuck via Flickr.

3 4 Diet Herbivory EOL Observer Cards Very few ants eat plants directly although almost all will drink the sweet secretions produced by plants or sap-drinking insects. No ants eat leaves, though some grow an edible fungus on leaves and flower petals that they have turned into mulch. Supplemental Notes Some plants produce seeds with arils - nutritious parts of a seed covering that entice an ant to carry the seed home, dispersing it in the process. This symbiosis is called myrmecochory. In some cases, specific types of ant and plant have co-evolved to be completely dependent on one another. The plant grows nutritionally balanced food and provides hollow cavities to house their ants in return for zealous protection from herbivores and the encroaching foliage of other plants.

Images: 1) Feeding at extrafloral nectaries on a Crotalaria species © Sean McCann via Flickr; 2) A Florida carpenter ant (Camponotus floridanus) on the bud of a hibiscus plant. © Bob Peterson via Flickr; 3) Ants consuming nectar from Milkweed (Asclepias physocarpa). © Leonora Enking via Flickr; 4) Dragging a seed back to the nest. © Steve Shattuck via Flickr.

Diet 4 Diet Specialists EOL Observer Cards Observe what the ants are using as food. Some ant genera specialize on a single food source such as seeds, millipedes, or fungi. Ant colonies that appear to favor one particular habitat may be food specialists. Supplemental Notes Some plants produce seeds with arils - nutritious parts of a seed covering that entice an ant to carry the seed Most ants are omnivores, though some are highly adapted to exploit a specific food source. For example, many species collect seeds, eat a specialized part of the husk, and leave rest - dispersing the plant in the process. But a few species get virtually all their sustenance by grinding up entire seeds. Many specialist ants will, in a pinch, eat some other food. But fungus growing ants are locked into their farming lifestyle - neither the fungus nor the ant has any hope of existing alone. home, dispersing it in the process. This symbiosis is called myrmecochory. In some cases, specific types of ant and plant have co-evolved to be completely dependent on one another. The plant grows nutritionally balanced food and provides hollow cavities to house their ants in return for zealous protection from herbivores and the encroaching foliage of other plants. Images: 1) Leafcutter ants (Atta sp.) chop leaves up as a substrate for growing fungus (see photo 4) that they can eat. © Yonatan Munk via Flickr; 2) Ants on a corn stalk. © LadyDragonflyCC via Flickr; 3) Honey ants (Prenolepis imparis) tending aphids on White Snakeroot (Ageratina altissima). © Dendroica cerulea via Flickr; 4) Atta colombica tending fungus that grows on a leaf-based substrate. © Christian R. Linder via Wikipedia. Author: Adam Lazarus, Consult Ants. Editor: Jeff Holmes, EOL, Harvard University. Created by the Encyclopedia of Life - www.eol.org Content Licensed Under a Creative Commons Attribution 3.0 License. Worker Castes 1 2 Ants Ants Organization

3 4 Worker Castes EOL Observer Cards Most ants you encounter are sterile females of the worker caste. Look for variations in size. The workers of some ant species are monomorphic - all one size. Others have a range of worker sizes (polymorphic). Still others have a largely dimorphic worker force - members are either one size or another. Supplemental Notes When multiple worker sizes are present, they typically look similar but for dimension. However, some species have highly specialized worker castes that are adapted to specific jobs. Large workers are often called soldiers, but “major” is a more accurate term - in most cases, colony defense is just one of many duties for these bigger ants. Even monomorphic species usually have a division of labor among the work force. Often this is a factor of age - new workers engage in safe chores like caring for the young or nest maintenance, while older workers take on increasingly risky jobs like foraging and colony defense. A worker may change jobs, but never shape or size- she will look the same her entire adult life.

Images: 1) Major worker of Pheidole sp. surrounded by smaller minors. © Steve Shattuck via Flickr; 2) Pheidologeton diversus majors are several times larger than the minor worker caste. © Nicholas Kanakis via Flickr; 3) Anoplolepis gracilipes (Yellow Crazy ant) have monomorphic workers. © John Tann via Flickr; 4) Workers from rival colonies battle for territory. © Mike Kelley via Flickr.

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6 7 Communication EOL Observer Cards Watch how ants respond to one another and to you. An ant’s language is a combination of chemical and physical cues that affect what you might observe. Examples include trail-laying (ants following specific routes), defensive or offensive posturing, sharing food, and a varity of odors and sounds. Supplemental Notes Although totally foreign to us, ant language rivals ours in complexity. However, not all the ants communicate to the same extent, nor with the same tactics. The particular style of communicating can be excellent for identifying species.

Images: 1) Ants lay pheromone trails that can be followed by other ants. © James Cridland via Flickr; 2) Ants trophallax (exchange food) to feed other nestmates and communicate what food they have found. © Ajay Narendra via Flickr; 3) Pachycondyla villosa rubs her armor plates against the ridges of this “stridulatory” organ to send vibration signals. © Roberto Keller/AMNH; 4) Myrmecia rubripes patrol outside their nest. © Jean and Fred Hort via Flickr; 5) A Formica worker’s defensive stance. © Sedeer via Flickr; 6) Weaver ant carrying a dead comrade. © Ihsan Khairir via Flickr; 7) Antennae can transmit and receive signals to aid in communication. © Matt Stanford via Flickr.

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5 6 Nest Location EOL Observer Cards Most ant species have a preferred place in which to nest so the location can aid identification. But not all ants are choosy when it comes to housing a colony - some are as much at home in an electrical outlet as they are under a rock. Others, like some Army Ants, make temporary nests out of their own bodies! Supplemental Notes In many cases, an entire colony will relocate if living conditions become unfavorable or if a better spot presents itself. Many temperate ants change their nest seasonally. They expand their living quarters & even establish large satellite nests in the summer, then re-group into a relatively small space as winter approaches.

Images: 1) Camponotus texanus lives in wood. © Adam Lazarus; 2) A Turneria bidentata nest found in a small twig in northern Queensland. The nest was broken open to show the size of the inner chamber. © Steve Shattuck via Flickr; 3) A Camponotus americanus nest extends from the soil into crystal-lined channels in a large piece of quartz. © Adam Lazarus; 4) A small Lasius sp. colony living in the stem of a morel mushroom. © Adam Lazarus; 5) Ant nest in Common whelk. © Bas Kers via Flickr; 6) Weaver ants building a nest. © Troup Dresser via Flickr.

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3 Ants in Groups EOL Observer Cards Ants form large groups for many different reasons. Because many of these behaviors are extremely species specific, the nature of a group can be a fast way to correctly identify an ant. Look for evidence of group activities such as attacks on other colonies, hunting for food, or migration. Supplemental Notes Though all ants work together, the ways in which they do so vary a great deal. People tend to think of ants as excellent recruiters or enthusiastic warmongers, but there is no generalizing- some species forage alone, and some shy away from the first sign of danger.

Images: 1) Army ants attacking a crab. © Axel Rouvin via Flickr; 2) Species such as Tetramorium caespitum (pavement or sidewalk ant) is famous for engaging in epic battles against rival colonies. © nick chapman via Flickr; 3) A problematic invasive species, yellow crazy ants (Anoplolepis gracilipes) form super-colonies that can devastate local environments. © henna lion via Flickr; 4) Army ants (Eciton burchellii) in Costa Rica with larvae of a raided wasp nest. © Geoff Gallice via Flickr.

Behavior 4 Behavior Specialists EOL Observer Cards With about 15,000 described species of ants, it is not surprising that many interact with their surroundings in striking and unusual ways. Such specialized behavior is often unique to one taxonomic group of ants. Look for ways that ants are changing their environment, building with own bodies, and adapting to specific conditions, such as frequent flooding.

Supplemental Notes A worker’s behavior may seem counterintuitive when it puts itself at extreme disadvantage or even commits outright suicide. But in a way, ants are just like cells in a body - taken as a whole, the colony acts just like many other animals.

Images: 1) Fire ants in the southern USA build a raft during floods. © nile red via Flickr; 2) Army Ants build temporary bridges and shelters by linking their bodies together. © Sean McCann via Flickr; 3) Some species of ants harvest honeydew from mealybugs. “Devil’s gardens” are clearings in tropical forests where Myrmelachista sp. has used formic acid to kill off foliage except for the plant that supports their favorite mealybug. © John Tann via Flickr; 4) Weaver ants cooperate to extend their reach. © Budzlife via Flickr.

3 4 Behavior Perception EOL Observer Cards Ants have a sophisticated array of sensory equipment for interacting with their world. Observing local environment conditions may reveal how some ant species have developed extraordinary perceptual capabilities. Such conditions foster some sensory abilities but render others useless. For example, subterranean ants often lack eyes. Other ants may be seen glancing skyward, navigating by the position of the sun. Supplemental Notes Though perceptive capability varies between species, as a group, ants have the ability to see depth, movement & polarized light. They memorize, learn, and detect chemical stimuli at concentrations as low as a few trillionths of a gram. In the desert, shifting sands prevent the use of chemical trails so some ants have developed the ability to count their steps. Images: 1) By extending the normal leg length of Cataglyphis workers, researchers found they count the number of steps they take from the nest. © Harald Wolf; 2) Pachycondyla villosa has acute vision to hunt prey. This unfortunate individual is infected with fruiting Cordyceps, a parasitic fungus. © Yonatan Munk via Flickr; 3) Dorylus nigricans is completely blind, yet perceives a rich chemical landscape. © April Nobile via AntWeb; 4) Species of the drought tolerant Cataglyphis genus use olfaction and the position of the sun to pinpoint their nest location. © Jonathan Bliss via Flickr. Author: Adam Lazarus, Consult Ants. Editor: Jeff Holmes, EOL, Harvard University. Created by the Encyclopedia of Life - www.eol.org Content Licensed Under a Creative Commons Attribution 3.0 License. Visible Symbioses 1 Ants Ants

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4 Visible Symbioses EOL Observer Cards Record visible symbiotic relationships between ants and other species. Some ant/arthropod symbioses are quite specific. For example, certain parasitoid flies are only associated with one species. However, most of these relationships are facultative (optional). For example, any number of ants will tend an insect that offers an attractive edible reward. Supplemental Notes Aphids, plant hoppers and other insects that drink sap extract what nutrition they can, and give off concentrated sugar as waste. Ants maintain herds of such insects, and guard them vigorously as they can become major colony assets. Other insects, like Lycaenid caterpillars, give off chemicals which are specifically designed to attract and satiate ants in return for protection or other services. But symbiosis isn’t always for the benefit of both parties - parasitoid Phorid flies detect the odor of certain ant species and lay their eggs on the ant’s exoskeleton. The grubs burrow into the ant and eat it to death. Images: 1) Myrmica ruginodis tending aphids. © rowe_becky via Flickr; 2) Fungal growth can cause odd behavior before the ant finally succumbs to the parasite. © ron_n_beth via Flickr; 3) Myrmecodia, or Ant plant, creates nesting space for ants within its structure in return for mineral nourishment and protection. © BlueRidgeKitties via Flickr; 4) Lasius fuliginosus tending aphids in Austria. © Jacobo Martin via Flickr. Author: Adam Lazarus, Consult Ants. Editor: Jeff Holmes, EOL, Harvard University. Created by the Encyclopedia of Life - www.eol.org Content Licensed Under a Creative Commons Attribution 3.0 License. Hidden Symbioses 1 2 Ants Ants Associations

3 4 Hidden Symbioses EOL Observer Cards Many symbiotic relationships occur underground. Sometimes the symbiont, such as a beetle, spider, wasp, or fly larva, is found with a variety of ants. Other times, the relationship is more specific - only certain ants tend subterranean herds of sap-sucking insects, such as aphids or mealybugs. Making note of a hidden symbiosis can help in species identification. Supplemental Notes Hidden underground are ants that never see the sun, tending pale white aphids on root systems. Note that not all symbioses are mutually beneficial. Some invaders of other colonies are masters of surprise and chemical subterfuge, and can eat ant young with impunity. Others steal food directly from a passing ant’s mouth! Images: 1) The Yellow Meadow Ant (Lasius flavus) lives mostly underground and feeds on the honeydew from root aphids. © Steve Begin via Flickr; 2) The beetle Martineziana dutertrei, seems to function primarily as a parasite of its host ant colony, stealing prey and other nutrients, and even eating ant larvae. © Alison Bockoven, 6legs2many; 3) A phorid fly, Pseudacteon cultellatus and a red imported fire ant, Solenopsis invicta. The fly can lay eggs inside the ant’s head. Fly maggots that hatch will then slowly decapitate the ant. Photo by Sanford Porter, USDA-ARS. The inset shows a closeup view of the fly’s ovipositor (photo via Wikimedia Commons).

Author: Adam Lazarus, Consult Ants. Editor: Jeff Holmes, EOL, Harvard University. Created by the Encyclopedia of Life - www.eol.org Content Licensed Under a Creative Commons Attribution 3.0 License. Mites 1 2 Ants Ants Associations Mites EOL Observer Cards Look for mites on the ant exoskeleton. While some mites parasitize a range of ant species, others are shockingly adapted to the lifestyle of a specific ant host. For this reason, mites can be a useful tool for species identification.

Supplemental Notes Mites are tiny parasites that prey on the skin, secretions, or food stores of almost every terrestrial animal, including ants. Some mites will attach to the face of one worker, and force it to ask for food from another. When the unwitting nestmate obliges, the mite steals the offering! Other mites fit onto the feet of their ant hosts, like a living shoe. Mites tend to occur at higher densities in captive colonies. In fact, they are one of the only biological threats to ant farms.

Images: 1) Camponotus texanus with mites. © Dmitry Mozzherin; 2) Scanning electron micrograph of mite on Camponotus texanus. © David Graham & James Style.

3 4 Mimicry EOL Observer Cards Are you sure that’s an ant? Spiders, flies, true bugs and crickets are but a few of the many creatures that imitate ants. Watch for legs or wings being waved in front like antennae, count the number of bodyparts and eyes, and watch for movements that are somehow un-antlike. Usually mimics look much like a local ant, and so indicate what species are around. Supplemental Notes One possible reason for mimicry is to fool predators - ants tend to be aggressive, foul-tasting, and nutritionally poor. Insect eaters often avoid them in favor of more palatable fare. Some suppose ant mimics use their disguise to infiltrate and prey on ants, but ants primarily rely on smell to identify kin - not vision.

Images: 1) Appearing remarkably like an ant, Synemosyna formica, is really a spider. © cotinis via Flickr; 2) A mimic fly. © Scott Justis, INSET: Formica sp. © Christy Beal; 3) The Kerengga Ant-like Jumper (Myrmarachne plataleoides) male. © Vishal Bhave via EOL; 4) A true bug that looks like several ant species. © Brian Malow. INSET (from top) Camponotus americanus, © Adam Lazarus; Camponotus castaneus, © Thomas of Baltimore; Camponotus chromaiodes, © Adam Lazarus.

3 4 Inquilines EOL Observer Cards Inquiline ants avoid colony founding and other tasks by exploiting the labor of neighboring species. With less need to build resources, these species may begin immediate production of new queens - a process that can take years for more typical ants. Some inquilines look nothing like their hosts, while others are nearly impossible to distinguish. Supplemental Notes Parasites benefit from fast reproduction, plentiful food and safety from predators. But such a lifestyle is tenuous, and success can spell disaster if the host ants are overworked. Superparasitic ants have no worker caste - just queens who invade a host colony, kill the resident queen and then use the motherless workers to produce more superparasites. Images: 1) The saberlike jaws of the red slave-maker Polyergus breviceps are ideal for combat and stealing cocoons from Formica argentea colonies. © Tomasz Kucza via Flickr; 2) The frail superparasite Anergates atratulus moves into orphaned nests of Tetramorium species to take advantage of their workers. April Nobile, © California Academy of Sciences; 3) Megalomyrmex symmetochus lives in and feeds on fungus gardens of Trachymrmex. April Nobile, © California Academy of Sciences; 4) Pogonomyrmex barbatus workers. The worker- less queen from another species in this genus (P. anergismus), invades these related Harvester ant nests to lay her eggs. © mkarco via Flickr. Author: Adam Lazarus, Consult Ants. Editor: Jeff Holmes, EOL, Harvard University. Created by the Encyclopedia of Life - www.eol.org Content Licensed Under a Creative Commons Attribution 3.0 License. Human Impact 1 2 Ants Ants Associations

3 4 Human Impact EOL Observer Cards Since the dawn of humanity, ants have both inspired & annoyed us. They have played roles in religion, philosophy, law, mathematics and technology. We use them as food, for entertainment, and as models for parallel processing and testing efficiency. Both races build, grow and store food, live in organised societies, compete for resources, and go to great lengths to keep each other out of our spaces. Supplemental Notes There is scientific evidence demonstrating that human activity can have an effect on the introduction and spread of invasive species. One example of this is the imported red fire ant (Solenopsis invicta) which has had a negative impact on native species and human agriculture in several countries.

Images: 1) Satere-Mawe of the Brazilian Amazon weave Bullet Ants (Paraponera clavata) into a sleeve to be placed on a young warrior’s hand for a painful initiation ceremony. From “Bullet Ant Glove” otithelis.com on 12 April, 2011; 2) Human disturbance of the environment can favor the establishment of invasive species such as fire ants (Solenopsis invicta). © J. Stephen Conn via Flickr; 3) Tapinoma sp. ants live in an electrical outlet. © Wally Hartshorn via Flickr; 4) Toasted leafcutter ants on sale. © Songkran via Flickr.

2 4 Life Cycle Development Stages EOL Observer Cards Look for ants at the egg, larva, pupa, and adult stages. Some larvae spin silken cocoons in which to develop so pupae are not visible unless the silk is torn. Note that certain species produce new ants in discreet waves, while others have a constant turnover, with all life stages present at any one time so presence or absence can be a useful taxonomic indicator. Supplemental Notes An egg hatches into a helpless grublike larva and is diligently tended by adults. Some need regurgitated food, while others have well developed mandibles and eat insects that are placed within reach. All growth happens during the larval stage - adult ants do not change size. When the larva is ready to become an adult it turns into a pupa that looks like a white ant in the fetal position. Shortly before it becomes an adult, the pupa develops pigmentation. Ants that become active before their pigments fully develop (known as “callows”), look like white or pale colony members and are easily damaged until their exoskeleton hardens. Images: 1) Transporting some newly hatched eggs. © NZ Alex via Flickr; 2) Young larval forms Camponotus sp. © Arthur Chapman; 3) Pupae stage. © Steve Begin; 4) Late stage pupae. © Sean McCann.

3 4 Photography EOL Observer Cards Digital photography has revolutionized how we record, study, and share information about ants. Macrophotography captures ants life-size or bigger but requires special macro extension tubes or dedicated lenses and usually an SLR camera. However, good results can be gained using simple point-and-shoot cameras that have a “macro” setting. Depth of field (how much of the subject is in focus) is affected by the aperture setting. To blur the background, use a small “f” number (e.g. f2.8-4). A larger “f” number (e.g. f32) increases the depth of field but usually means additional lighting (a flash) is required. Using a minimal flash setting or a diffuser (tissue paper can work) can help reduce glare and unnatural lighting. Supplemental Notes Learn from an expert! Check out Alex Wild’s excellent article on photographing ants on his website at: http://myrmecos.net/2008/03/30/photo-technique-working-with-ants/. Images: 1) A tripod can help reduce image blurring by minimizing camera shake. © Shirley Buxton via Flickr; 2) Extension tube for close-up images. © Boris Mitendorfer via Flickr; 3) A homemade flash diffuser to reduce glare. © Danny Chapman via Flickr; 4) Macro lens mounted on a small camera. © Martha Gunawan via Flickr. Author: Adam Lazarus, Consult Ants. Editor: Jeff Holmes, EOL, Harvard University. Created by the Encyclopedia of Life - www.eol.org Content Licensed Under a Creative Commons Attribution 3.0 License. Collecting

Ants Ants 1

Techniques 2 4 5

3 Collecting EOL Observer Cards Although collectors develop their own suite of collecting implements, some deserve special mention: 1) Small ants can be sucked up with an aspirator 2) Forceps are essential - soft-grip or “featherweight” forceps reduce excess hand pressure while stiff, pointed forceps permit the most detailed manipulations; 3) Baiting with sugar-water; 4) Berlese traps or Tullgren funnels drive arthropods out of a soil sample. 5) The pitfall trap is a form of passive collection.

Supplemental Notes Collecting is great fun but it should be done with care to protect your samples and their environment. Rough handling can injure live ants, or destroy key taxonomic features on preserved ones. Collecting should also be done with a conscience - know your reason for removing something from its home.

2 4 Scientific Records EOL Observer Cards Your observations can be great memories or they can add to our collective scientific knowledge. The most useful field data include the date and time of day, location, GPS coordinates, habitat, observations, sample ants, and a personal reference number unique to the event. Voucher specimens serve as a hard copy of your ant observations. Supplemental Notes Include a personal reference number (e.g. “ccast464”) in your summary to link the voucher to more detailed information in your field notebook. Voucher specimens enable your work to be vetted (reviewed) - a critical part of the scientific process. When deposited in a well-regarded collection or museum, the voucher ID numbers they receive will be included in all relevant science papers.

Images: 1) Detailed field notes. © Michael Branstetter; 2) Pinned & labeled voucher specimens for deposit in the CASENT voucher system www.antweb.org/documentation.do. © Adam Lazarus; 3) Voucher label. © Adam Lazarus; 4. Samples of professional ant work. © Adam Lazarus.

Author: Adam Lazarus, Consult Ants. Editor: Jeff Holmes, EOL, Harvard University. Created by the Encyclopedia of Life - www.eol.org Content Licensed Under a Creative Commons Attribution 3.0 License.