114-542: Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees , belong to the order Hymenoptera . Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22,000 species have been classified. They are easily identified by their geniculate (elbowed) antennae and the distinctive node-like structure that forms their slender waists. Ants form colonies that range in size from
228-410: A Proto-Indo-European word *morwi- was the root for Sanskrit vamrah , Greek μύρμηξ mýrmēx , Old Church Slavonic mraviji , Old Irish moirb , Old Norse maurr , Dutch mier , Swedish myra , Danish myre , Middle Dutch miere , and Crimean Gothic miera . The family Formicidae belongs to the order Hymenoptera , which also includes sawflies , bees , and wasps . Ants evolved from
342-486: A cosmopolitan distribution . They 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. Ants occupy a wide range of ecological niches and exploit many different food resources as direct or indirect herbivores, predators and scavengers. Most ant species are omnivorous generalists , but
456-478: A multicellular organism ; castes fulfill a specific role that contributes to the functioning and survival of the whole colony, while being incapable of independent survival outside the colony. In 1969, Charles D. Michener further expanded Batra's classification with his comparative study of social behavior in bees. He observed multiple species of bees ( Apoidea ) in order to investigate the different levels of animal sociality, all of which are different stages that
570-543: A pitch accent . In Modern Greek, all vowels and consonants are short. Many vowels and diphthongs once pronounced distinctly are pronounced as /i/ ( iotacism ). Some of the stops and glides in diphthongs have become fricatives , and the pitch accent has changed to a stress accent . Many of the changes took place in the Koine Greek period. The writing system of Modern Greek, however, does not reflect all pronunciation changes. The examples below represent Attic Greek in
684-429: A colony may pass through. Eusociality, which is the highest level of animal sociality a species can attain, specifically had three characteristics that distinguished it from the other levels: E. O. Wilson extended the terminology to include other social insects, such as ants, wasps, and termites. Originally, it was defined to include organisms (only invertebrates) that had the following three features: Eusociality
798-407: A colony, there are both pleated fan-shaped "nest" fronds that collect and hold water, and gutter-shaped "strap" fronds that channel water: no solitary Platycerium species has both types. At the bottom of a colony, there are "nest" fronds that clasp the trunk of the tree supporting the fern, and drooping photosynthetic fronds. These are argued to be adapted to support the colony structurally, i.e. that
912-448: A conservative estimate of the ants at about 20 × 10 (20 quadrillion). Ants range in size from 0.75 to 52 millimetres (0.030–2.0 in), the largest species being the fossil Titanomyrma giganteum , the queen of which was 6 cm ( 2 + 1 ⁄ 2 in) long with a wingspan of 15 cm (6 in). Ants vary in colour; most ants are yellow to red or brown to black, but a few species are green and some tropical species have
1026-496: A few are specialist feeders. There is considerable variation in ant abundance across habitats, peaking in the moist tropics to nearly six times that found in less suitable habitats. Their ecological dominance has been examined primarily using estimates of their biomass : myrmecologist E. O. Wilson had estimated in 2009 that at any one time the total number of ants was between one and ten quadrillion ( short scale ) (i.e., between 10 and 10) and using this estimate he had suggested that
1140-596: A few dozen individuals often living in small natural cavities to highly organised colonies that may occupy large territories with sizeable nest that consist of millions of individuals or into the hundreds of millions in super colonies . Typical colonies consist of various castes of sterile, wingless females, most of which are workers (ergates), as well as soldiers (dinergates) and other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called " queens " ( gynes ). The colonies are described as superorganisms because
1254-406: A high resolution image. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization . Compared to vertebrates , ants tend to have blurrier eyesight, particularly in smaller species, and a few subterranean taxa are completely blind . However, some ants, such as Australia's bulldog ant , have excellent vision and are capable of discriminating
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#17327660799781368-419: A hive of half-sisters that share only 25% of their genes. The association between haplodiploidy and eusociality is below statistical significance. Haplodiploidy is thus neither necessary nor sufficient for eusociality to emerge. Relatedness does still play a part, as monogamy (queens mating singly) is the ancestral state for all eusocial species so far investigated. If kin selection is an important force driving
1482-477: A lack of contemporaneous evidence. Several theories exist about what Hellenic dialect groups may have existed between the divergence of early Greek-like speech from the common Proto-Indo-European language and the Classical period. They have the same general outline but differ in some of the detail. The only attested dialect from this period is Mycenaean Greek , but its relationship to the historical dialects and
1596-419: A lesser degree. Pamphylian Greek , spoken in a small area on the southwestern coast of Anatolia and little preserved in inscriptions, may be either a fifth major dialect group, or it is Mycenaean Greek overlaid by Doric, with a non-Greek native influence. Regarding the speech of the ancient Macedonians diverse theories have been put forward, but the epigraphic activity and the archaeological discoveries in
1710-517: A lineage within the stinging wasps , and a 2013 study suggests that they are a sister group of the Apoidea . However, since Apoidea is a superfamily, ants must be upgraded to the same rank. A more detailed basic taxonomy was proposed in 2020. Three species of the extinct mid- Cretaceous genera Camelomecia and Camelosphecia were placed outside of the Formicidae, in a separate clade within
1824-648: A male is haploid, his daughters share 100% of his genes and 50% of their mother's. Therefore, they share 75% of their genes with each other. This mechanism of sex determination gives rise to what W. D. Hamilton first termed "supersisters", more closely related to their sisters than they would be to their own offspring. Even though workers often do not reproduce, they can pass on more of their genes by helping to raise their sisters than by having their own offspring (each of which would only have 50% of their genes). This unusual situation, where females may have greater fitness when they help rear sisters rather than producing offspring,
1938-477: A metallic lustre . More than 13,800 species are currently known (with upper estimates of the potential existence of about 22,000; see the article List of ant genera ), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntWeb and the Hymenoptera Name Server, help to keep track of
2052-508: A pheromone, helping their nest mates to find the food. Reproductive specialization generally involves the production of sterile members of the species, which carry out specialized tasks to care for the reproductive members. Individuals may have behavior and morphology modified for group defense, including self-sacrificing behavior . For example, members of the sterile caste of the honeypot ants such as Myrmecocystus fill their abdomens with liquid food until they become immobile and hang from
2166-466: A possible resolution to the paradox might lie in the close family relationship, which W.D. Hamilton quantified a century later with his 1964 inclusive fitness theory. After the gene-centered view of evolution was developed in the mid-1970s, non-reproductive individuals were seen as an extended phenotype of the genes, which are the primary beneficiaries of natural selection. According to inclusive fitness theory, organisms can gain fitness by increasing
2280-550: A prefix /e-/, called the augment . This was probably originally a separate word, meaning something like "then", added because tenses in PIE had primarily aspectual meaning. The augment is added to the indicative of the aorist, imperfect, and pluperfect, but not to any of the other forms of the aorist (no other forms of the imperfect and pluperfect exist). The two kinds of augment in Greek are syllabic and quantitative. The syllabic augment
2394-403: A protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates . Insects do not have lungs ; oxygen and other gases, such as carbon dioxide , pass through their exoskeleton via tiny valves called spiracles . Insects also lack closed blood vessels ; instead, they have a long, thin, perforated tube along the top of
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#17327660799782508-435: A segment fuse together, can occur partly or fully on the second, third and fourth abdominal segment and is used in identification. Fourth abdominal tergosternal fusion was formerly used as character that defined the poneromorph subfamilies, Ponerinae and relatives within their clade, but this is no longer considered a synapomorphic character. Like other arthropods, ants have an exoskeleton , an external covering that provides
2622-608: A separate historical stage, though its earliest form closely resembles Attic Greek , and its latest form approaches Medieval Greek . There were several regional dialects of Ancient Greek; Attic Greek developed into Koine. Ancient Greek was a pluricentric language , divided into many dialects. The main dialect groups are Attic and Ionic , Aeolic , Arcadocypriot , and Doric , many of them with several subdivisions. Some dialects are found in standardized literary forms in literature , while others are attested only in inscriptions. There are also several historical forms. Homeric Greek
2736-581: A set of pheromones that alter the behavior of specific castes in the colony. These pheromones may act across different species, as observed in Apis andreniformis (black dwarf honey bee), where worker bees responded to queen pheromone from the related Apis florea (red dwarf honey bee). Pheromones are sometimes used in these castes to assist with foraging. Workers of the Australian stingless bee Tetragonula carbonaria , for instance, mark food sources with
2850-427: A single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant ( Myrmecia pilosula ) has only a single pair of chromosomes (with the males having just one chromosome as they are haploid ), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects. Genome size
2964-416: A species. Multilevel selection has been heavily criticized for its conflict with the kin selection theory. A reversal to solitarity is an evolutionary phenomenon in which descendants of a eusocial group evolve solitary behavior once again. Bees have been model organisms for the study of reversal to solitarity, because of the diversity of their social systems. Each of the four origins of eusociality in bees
3078-630: A standard subject of study in educational institutions of the Western world since the Renaissance . This article primarily contains information about the Epic and Classical periods of the language, which are the best-attested periods and considered most typical of Ancient Greek. From the Hellenistic period ( c. 300 BC ), Ancient Greek was followed by Koine Greek , which is regarded as
3192-510: A vowel or /n s r/ ; final stops were lost, as in γάλα "milk", compared with γάλακτος "of milk" (genitive). Ancient Greek of the classical period also differed in both the inventory and distribution of original PIE phonemes due to numerous sound changes, notably the following: The pronunciation of Ancient Greek was very different from that of Modern Greek . Ancient Greek had long and short vowels ; many diphthongs ; double and single consonants; voiced, voiceless, and aspirated stops ; and
3306-488: A weak form of eusociality. It has been suggested that the colonial and epiphytic staghorn fern , too, may make use of a primitively eusocial division of labor. The term "eusocial" was introduced in 1966 by Suzanne Batra , who used it to describe nesting behavior in Halictid bees, on a scale of subsocial/solitary, colonial/communal, semisocial, and eusocial, where a colony is started by a single individual. Batra observed
3420-497: Is paraphyletic In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant ( Sphecomyrma ) that lived in the Cretaceous period. The specimen, trapped in amber dating back to around 92 million years ago, has features found in some wasps, but not found in modern ants. The oldest fossils of ants date to the mid-Cretaceous, around 100 million years ago, which belong to extinct stem-groups such as
3534-529: Is 50%. Therefore, helping behavior is only advantageous if it is biased to helping sisters, which would drive the population to a 1:3 sex ratio of males to females. At this ratio, males, as the rarer sex, increase in reproductive value, reducing the benefit of female-biased investment. Further, not all eusocial species are haplodiploid: termites, some snapping shrimps, and mole rats are not. Conversely, many non-eusocial bees are haplodiploid, and among eusocial species many queens mate with multiple males, resulting in
Ant - Misplaced Pages Continue
3648-448: Is a fundamental characteristic of an organism. Ants have been found to have tiny genomes, with the evolution of genome size suggested to occur through loss and accumulation of non-coding regions, mainly transposable elements , and occasionally by whole genome duplication. This may be related to colonisation processes, but further studies are needed to verify this. Eusociality Eusociality ( Greek εὖ eu "good" and social )
3762-570: Is a literary form of Archaic Greek (derived primarily from Ionic and Aeolic) used in the epic poems , the Iliad and the Odyssey , and in later poems by other authors. Homeric Greek had significant differences in grammar and pronunciation from Classical Attic and other Classical-era dialects. The origins, early form and development of the Hellenic language family are not well understood because of
3876-446: Is a species of ambrosia beetle native to Australia, and is the first beetle (order Coleoptera ) to be recognized as eusocial. This species forms colonies in which a single female is fertilized, and is protected by many unfertilized females, which serve as workers excavating tunnels in trees. This species has cooperative brood care, in which individuals care for juveniles that are not their own. Some gall-inducing insects , including
3990-418: Is added to stems beginning with consonants, and simply prefixes e (stems beginning with r , however, add er ). The quantitative augment is added to stems beginning with vowels, and involves lengthening the vowel: Some verbs augment irregularly; the most common variation is e → ei . The irregularity can be explained diachronically by the loss of s between vowels, or that of the letter w , which affected
4104-427: Is an aggregation of relatives (because the shrimp do not have to disperse to find food), and much competition for those nesting sites. Being the target of attack promotes a good defense system (soldier caste); soldiers promote the fitness of the whole nest by ensuring safety and reproduction of the queen. Eusociality offers a competitive advantage in shrimp populations. Eusocial species are more abundant, occupy more of
4218-666: Is called 'East Greek'. Arcadocypriot apparently descended more closely from the Mycenaean Greek of the Bronze Age. Boeotian Greek had come under a strong Northwest Greek influence, and can in some respects be considered a transitional dialect, as exemplified in the poems of the Boeotian poet Pindar who wrote in Doric with a small Aeolic admixture. Thessalian likewise had come under Northwest Greek influence, though to
4332-448: Is considered by some linguists to have been closely related to Greek . Among Indo-European branches with living descendants, Greek is often argued to have the closest genetic ties with Armenian (see also Graeco-Armenian ) and Indo-Iranian languages (see Graeco-Aryan ). Ancient Greek differs from Proto-Indo-European (PIE) and other Indo-European languages in certain ways. In phonotactics , ancient Greek words could end only in
4446-501: Is correlated with the number of females in the nest, implying that having more females leads to more efficient building and provisioning of cells. In similar species with only one queen, such as Lasioglossum malachurum in Europe, the degree of eusociality depends on the clime in which the species is found. Termites (order Blattodea , infraorder Isoptera ) make up another large portion of highly advanced eusocial animals. The colony
4560-563: Is differentiated into various castes: the queen and king are the sole reproducing individuals; workers forage and maintain food and resources; and soldiers defend the colony against ant attacks. The latter two castes, which are sterile and perform highly specialized, complex social behaviors, are derived from different stages of pluripotent larvae produced by the reproductive caste. Some soldiers have jaws so enlarged (specialized for defense and attack) that they are unable to feed themselves and must be fed by workers. Austroplatypus incompertus
4674-615: Is distinguished from all other social systems because individuals of at least one caste usually lose the ability to perform behaviors characteristic of individuals in another caste. Eusocial colonies can be viewed as superorganisms . Eusociality has evolved among the insects , crustaceans , trematoda and mammals . It is most widespread in the Hymenoptera ( ants , bees , and wasps ) and in Blattodea ( termites ). A colony has caste differences: queens and reproductive males take
Ant - Misplaced Pages Continue
4788-542: Is not divided between castes. Though controversial, it has been suggested that male homosexuality and female menopause could have evolved through kin selection . This would mean that humans sometimes exhibit a type of alloparental behavior known as " helpers at the nest ", with juveniles and sexually mature adolescents helping their parents raise subsequent broods, as in some birds, some non-eusocial bees , and meerkats . These species are not eusocial: they do not have castes, and helpers reproduce on their own if given
4902-497: Is often invoked to explain the multiple independent evolutions of eusociality (at least nine separate times) within the Hymenoptera. Against the supposed benefits of haplodiploidy for eusociality, Robert Trivers notes that while females share 75% of genes with their sisters in haplodiploid populations, they only share 25% of their genes with their brothers. Accordingly, the average relatedness of an individual to their sibling
5016-687: Is often reliably predicted by ecological conditions, and switches in behavioral type have been experimentally induced by translocating offspring of solitary or social populations to warm and cool climates. In H. rubicundus , females produce a single brood in cooler regions and two or more broods in warmer regions, so the former populations are solitary while the latter are social. In another species of sweat bees, L. calceatum , social phenotype has been predicted by altitude and micro-habitat composition, with social nests found in warmer, sunnier sites, and solitary nests found in adjacent, cooler, shaded locations. Facultatively social bee species, however, which comprise
5130-430: Is the highest level of organization of sociality . It is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generations within a colony of adults , and a division of labor into reproductive and non-reproductive groups. The division of labor creates specialized behavioral groups within an animal society, sometimes called castes. Eusociality
5244-497: The Damaraland mole-rat ( Fukomys damarensis ), both of which are highly inbred . Usually living in harsh or limiting environments, these mole-rats aid in raising siblings and relatives born to a single reproductive queen. However, this classification is controversial owing to disputed definitions of 'eusociality'. To avoid inbreeding , mole rats sometimes outbreed and establish new colonies when resources are sufficient. Most of
5358-759: The Greek region of Macedonia during the last decades has brought to light documents, among which the first texts written in Macedonian , such as the Pella curse tablet , as Hatzopoulos and other scholars note. Based on the conclusions drawn by several studies and findings such as Pella curse tablet , Emilio Crespo and other scholars suggest that ancient Macedonian was a Northwest Doric dialect , which shares isoglosses with its neighboring Thessalian dialects spoken in northeastern Thessaly . Some have also suggested an Aeolic Greek classification. The Lesbian dialect
5472-694: The Haidomyrmecinae , Sphecomyrminae and Zigrasimeciinae , with modern ant subfamilies appearing towards the end of the Cretaceous around 80–70 million years ago. Ants diversified extensively during the Angiosperm Terrestrial Revolution and assumed ecological dominance around 60 million years ago. Some groups, such as the Leptanillinae and Martialinae , are suggested to have diversified from early primitive ants that were likely to have been predators underneath
5586-808: The Paleogene period . By the Oligocene and Miocene , ants had come to represent 20–40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, around one in 10 genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Termites live in colonies and are sometimes called "white ants", but termites are only distantly related to ants. They are
5700-471: The gall -forming aphid , Pemphigus spyrothecae (order Hemiptera ), and thrips such as Kladothrips (order Thysanoptera ), are described as eusocial. These species have very high relatedness among individuals due to their asexual reproduction (sterile soldier castes being clones produced by parthenogenesis ), but the gall-inhabiting behavior gives these species a defensible resource. They produce soldier castes for fortress defense and protection of
5814-519: The gene-centered view of evolution , eusociality was seen as paradoxical: if adaptive evolution unfolds by differential reproduction of individual organisms, the evolution of individuals incapable of passing on their genes presents a challenge. In On the Origin of Species , Darwin referred to the existence of sterile castes as the "one special difficulty, which at first appeared to me insuperable, and actually fatal to my theory". Darwin anticipated that
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#17327660799785928-437: 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 . For instance, these replete workers develop in
6042-482: The mandibles , used to carry food, manipulate objects, construct nests, and for defence. In some species, a small pocket (infrabuccal chamber) inside the mouth stores food, so it may be passed to other ants or their larvae. Both the legs and wings of the ant are attached to the mesosoma ("thorax"). The legs terminate in a hooked claw which allows them to hook on and climb surfaces. Only reproductive ants ( queens and males) have wings. Queens shed their wings after
6156-455: The nuptial flight , leaving visible stubs, a distinguishing feature of queens. In a few species, wingless queens ( ergatoids ) and males occur. 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. In
6270-501: The present , future , and imperfect are imperfective in aspect; the aorist , present perfect , pluperfect and future perfect are perfective in aspect. Most tenses display all four moods and three voices, although there is no future subjunctive or imperative. Also, there is no imperfect subjunctive, optative or imperative. The infinitives and participles correspond to the finite combinations of tense, aspect, and voice. The indicative of past tenses adds (conceptually, at least)
6384-604: The red imported fire ant ( Solenopsis invicta ) of South America, are regarded as invasive species in other parts of the world, establishing themselves in areas where they have been introduced accidentally. The word ant and the archaic word emmet are derived from ante , emete of Middle English , which come from ǣmette of Old English ; these are all related to Low Saxon e(e)mt , empe and varieties ( Old Saxon emeta ) and to German Ameise ( Old High German āmeiza ). All of these words come from West Germanic * ǣmaitjōn , and
6498-1031: The 5th century BC. Ancient pronunciation cannot be reconstructed with certainty, but Greek from the period is well documented, and there is little disagreement among linguists as to the general nature of the sounds that the letters represent. /oː/ raised to [uː] , probably by the 4th century BC. Greek, like all of the older Indo-European languages , is highly inflected. It is highly archaic in its preservation of Proto-Indo-European forms. In ancient Greek, nouns (including proper nouns) have five cases ( nominative , genitive , dative , accusative , and vocative ), three genders ( masculine , feminine , and neuter ), and three numbers (singular, dual , and plural ). Verbs have four moods ( indicative , imperative , subjunctive , and optative ) and three voices (active, middle, and passive ), as well as three persons (first, second, and third) and various other forms. Verbs are conjugated through seven combinations of tenses and aspect (generally simply called "tenses"):
6612-495: The Archaic period of ancient Greek (see Homeric Greek for more details): Μῆνιν ἄειδε, θεά, Πηληϊάδεω Ἀχιλῆος οὐλομένην, ἣ μυρί' Ἀχαιοῖς ἄλγε' ἔθηκε, πολλὰς δ' ἰφθίμους ψυχὰς Ἄϊδι προΐαψεν ἡρώων, αὐτοὺς δὲ ἑλώρια τεῦχε κύνεσσιν οἰωνοῖσί τε πᾶσι· Διὸς δ' ἐτελείετο βουλή· ἐξ οὗ δὴ τὰ πρῶτα διαστήτην ἐρίσαντε Ἀτρεΐδης τε ἄναξ ἀνδρῶν καὶ δῖος Ἀχιλλεύς. The beginning of Apology by Plato exemplifies Attic Greek from
6726-756: The North American honeypot ant Myrmecocystus mexicanus . Usually the largest workers in the colony develop into repletes; and, if repletes are removed from the colony, other workers become repletes, demonstrating the flexibility of this particular polymorphism . This polymorphism in morphology and behaviour of workers initially was thought to be determined by environmental factors such as nutrition and hormones that led to different developmental paths ; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in
6840-424: The anatomy of the worker caste, which is sterile and provides support for the reproductive caste. Most eusocial societies exist in arthropods , while a few are found in mammals . Some ferns may exhibit a primitive form of eusocial behavior. Eusociality has evolved multiple times in different insect orders, including hymenopterans, termites, thrips, aphids, and beetles. The order Hymenoptera contains
6954-1167: The ants appear to operate as a unified entity, collectively working together to support the colony. Ants have colonised almost every landmass on Earth . The only places lacking indigenous ants are Antarctica and a few remote or inhospitable islands. Ants thrive in moist tropical ecosystems and may exceed the combined biomass of wild birds and mammals. Their success in so many environments has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic , commensal , parasitic , and mutualistic relationships. Ant societies have division of labour , communication between individuals, and an ability to solve complex problems . These parallels with human societies have long been an inspiration and subject of study. Many human cultures make use of ants in cuisine, medication, and rites. Some species are valued in their role as biological pest control agents. Their ability to exploit resources may bring ants into conflict with humans, however, as they can damage crops and invade buildings. Some species, such as
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#17327660799787068-550: The aorist. Following Homer 's practice, the augment is sometimes not made in poetry , especially epic poetry. The augment sometimes substitutes for reduplication; see below. Almost all forms of the perfect, pluperfect, and future perfect reduplicate the initial syllable of the verb stem. (A few irregular forms of perfect do not reduplicate, whereas a handful of irregular aorists reduplicate.) The three types of reduplication are: Irregular duplication can be understood diachronically. For example, lambanō (root lab ) has
7182-419: The augment when it was word-initial. In verbs with a preposition as a prefix, the augment is placed not at the start of the word, but between the preposition and the original verb. For example, προσ(-)βάλλω (I attack) goes to προσ έ βαλoν in the aorist. However compound verbs consisting of a prefix that is not a preposition retain the augment at the start of the word: αὐτο(-)μολῶ goes to ηὐ τομόλησα in
7296-555: The body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph toward the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. An ant's head contains many sensory organs . Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ant eyes are good for acute movement detection, but do not offer
7410-585: The ceilings of the underground nests, acting as food storage for the rest of the colony. Not all social insects have distinct morphological differences between castes. For example, in the Neotropical social wasp Synoeca surinama , caste ranks are determined by social displays in the developing brood. These castes are sometimes further specialized in their behavior based on age, as in Scaptotrigona postica workers. Between approximately 0–40 days old,
7524-438: The center of Greek scholarship, this division of people and language is quite similar to the results of modern archaeological-linguistic investigation. One standard formulation for the dialects is: West vs. non-West Greek is the strongest-marked and earliest division, with non-West in subsets of Ionic-Attic (or Attic-Ionic) and Aeolic vs. Arcadocypriot, or Aeolic and Arcado-Cypriot vs. Ionic-Attic. Often non-West
7638-515: The colonies of a few ant species, there are physical castes—workers in distinct size-classes, called minor (micrergates), median, and major ergates (macrergates). Often, the larger ants have disproportionately larger heads, and correspondingly stronger mandibles . Although formally known as dinergates, such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they still are workers and their "duties" typically do not vary greatly from
7752-421: The colony against predators, kleptoparasites , and competitors. In these groups, eusociality is produced by both high relatedness and by living in a restricted, shared area. Eusociality has evolved in three different lineages in the colonial crustacean genus Synalphaeus . S. regalis , S. microneptunus , S. filidigitus , S. elizabethae , S. chacei , S. riosi , S. duffyi , and S. cayoneptunus are
7866-523: The colony, it is protected by a specialized caste of sterile soldier trematodes. Soldiers are smaller, more mobile, and develop along a different pathway than sexually mature reproductives. One difference is that a soldier's mouthparts (pharynx) is five times as big as those of the reproductives. They make up nearly a quarter of the volume of the soldier. These soldiers do not have a germinal mass, never metamorphose to be reproductive, and are, therefore, obligately sterile. Soldiers are readily distinguished from
7980-432: The cooperative behavior of the bees, males and females alike, as they took responsibility for at least one duty (i.e., burrowing, cell construction, oviposition ) within the colony. The cooperativeness was essential as the activity of one labor division greatly influenced the activity of another. Eusocial colonies can be viewed as superorganisms , with individual castes being analogous to different tissue or cell types in
8094-422: The cost of investing in non-reproductive offspring, and an increased risk of disease. All reversals to solitarity have occurred among primitively eusocial groups; none have followed the emergence of advanced eusociality. The "point of no return" hypothesis posits that the morphological differentiation of reproductive and non-reproductive castes prevents highly eusocial species such as the honeybee from reverting to
8208-615: The dialect of Sparta ), and Northern Peloponnesus Doric (including Corinthian ). All the groups were represented by colonies beyond Greece proper as well, and these colonies generally developed local characteristics, often under the influence of settlers or neighbors speaking different Greek dialects. After the conquests of Alexander the Great in the late 4th century BC, a new international dialect known as Koine or Common Greek developed, largely based on Attic Greek , but with influence from other dialects. This dialect slowly replaced most of
8322-530: The distance and size of objects moving nearly a meter away. Based on experiments conducted to test their ability to differentiate between selected wavelengths of light, some ant species such as Camponotus blandus, Solenopsis invicta, and Formica cunicularia are thought to possess a degree of colour vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents , and vibrations ; they also are used to transmit and receive signals through touch. The head has two strong jaws,
8436-680: The dwarf mongoose, the breeding pair receives food priority and protection from subordinates and rarely has to defend against predators. Scientists have debated whether humans are prosocial or eusocial. Edward O. Wilson called humans eusocial apes, arguing for similarities to ants, and observing that early hominins cooperated to rear their children while other members of the same group hunted and foraged. Wilson and others argued that through cooperation and teamwork, ants and humans form superorganisms. Wilson's claims were vigorously rejected by critics of group selection theory, which grounded Wilson's argument, and because human reproductive labor
8550-438: The early phase of infection. This strategic positioning allows them to effectively defend against invaders, similar to how soldier distribution patterns are seen in other animals with defensive castes. They "appear to be an obligately sterile physical caste, akin to that of the most advanced social insects". Among mammals, two species in the rodent group Phiomorpha are eusocial, the naked mole-rat ( Heterocephalus glaber ) and
8664-509: The eight recorded species of parasitic shrimp that rely on fortress defense and live in groups of closely related individuals in tropical reefs and sponges. They live eusocially with a single breeding female, and a large number of male defenders armed with enlarged snapping claws. There is a single shared living space for the colony members, and the non-breeding members act to defend it. The fortress defense hypothesis additionally points out that because sponges provide both food and shelter, there
8778-524: The eusocial phenotype, with one queen supported by sterile workers and sexually active males ( drones ). In queenless colonies, the lack of queen pheromones causes winged females to quickly shed their wings, develop ovaries and lay eggs. These virgin replacement queens assume the role of the queen and start to produce queen pheromones. Similarly, queen weaver ants Oecophylla longinoda have exocrine glands that produce pheromones which prevent workers from laying reproductive eggs. Similar mechanisms exist in
8892-426: The eusocial wasp Vespula vulgaris . For a queen to dominate all the workers, usually numbering more than 3000 in a colony, she signals her dominance with pheromones. The workers regularly lick the queen while feeding her, and the air-borne pheromone from the queen's body alerts those workers of her dominance. Ancient Greek Ancient Greek ( Ἑλληνῐκή , Hellēnikḗ ; [hellɛːnikɛ́ː] ) includes
9006-722: The evolution of eusociality is supported by evidence such as experimentally induced reproductive division of labor, for example when normally solitary queens are forced together. Conversely, female Damaraland mole-rats undergo hormonal changes that promote dispersal after periods of high rainfall. Climate too appears to be a selective agent driving social complexity; across bee lineages and Hymenoptera in general, higher forms of sociality are more likely to occur in tropical than temperate environments. Similarly, social transitions within halictid bees , where eusociality has been gained and lost multiple times, are correlated with periods of climatic warming. Social behavior in facultative social bees
9120-437: The evolution of eusociality, monogamy should be the ancestral state, because it maximizes the relatedness of colony members. Increased parasitism and predation rates are the primary ecological drivers of social organization. Group living affords colony members defense against enemies, specifically predators, parasites, and competitors, and allows them to gain advantage from superior foraging methods. The importance of ecology in
9234-673: The forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek ( c. 1400–1200 BC ), Dark Ages ( c. 1200–800 BC ), the Archaic or Epic period ( c. 800–500 BC ), and the Classical period ( c. 500–300 BC ). Ancient Greek was the language of Homer and of fifth-century Athenian historians, playwrights, and philosophers . It has contributed many words to English vocabulary and has been
9348-872: The general superfamily Formicoidea , which, together with Apoidea, forms the higher-ranking group Formicapoidina . Fernández et al. (2021) suggest that the common ancestors of ants and apoids within the Formicapoidina probably existed as early as in the end of the Jurassic period, before divergence in the Cretaceous. Chrysidoidea Vespidae Rhopalosomatidae Pompilidae Mutillidae Tiphiidae Chyphotidae Scolioidea Apoidea Formicidae Myrmicinae Ectatomminae Heteroponerinae Formicinae Dolichoderinae Aneuretinae Pseudomyrmecinae Myrmeciinae Dorylinae‡ Ponerinae Agroecomyrmecinae Paraponerinae Proceratiinae Amblyoponinae Apomyrminae Leptanillinae Martialinae *Cerapachyinae
9462-449: The habitat, and use more of the available resources than non-eusocial species. The trematodes are a class of parasitic flatworm, also known as flukes. One species, Haplorchis pumilio , has evolved eusociality involving a colony creating a class of sterile soldiers. One fluke invades a host and establishes a colony of dozens to thousands of clones that work together to take it over. Since rival trematode species can invade and replace
9576-561: The historical Dorians . The invasion is known to have displaced population to the later Attic-Ionic regions, who regarded themselves as descendants of the population displaced by or contending with the Dorians. The Greeks of this period believed there were three major divisions of all Greek people – Dorians, Aeolians, and Ionians (including Athenians), each with their own defining and distinctive dialects. Allowing for their oversight of Arcadian, an obscure mountain dialect, and Cypriot, far from
9690-476: The historical circumstances of the times imply that the overall groups already existed in some form. Scholars assume that major Ancient Greek period dialect groups developed not later than 1120 BC, at the time of the Dorian invasions —and that their first appearances as precise alphabetic writing began in the 8th century BC. The invasion would not be "Dorian" unless the invaders had some cultural relationship to
9804-429: The hormonally based behavioral development of workers and suppress their ovarian development. Both behavioral effects mediated by the nervous system often leading to recognition of queens ( releaser ) and physiological effects on the reproductive and endocrine system ( primer ) are attributed to the same pheromones. These pheromones volatilize or are deactivated within thirty minutes, allowing workers to respond rapidly to
9918-416: The immature and mature reproductive worms. Soldiers are more aggressive than reproductives, attacking heterospecific trematodes that infect their host in vitro . H. pumilio soldiers do not attack conspecifics from other colonies. The soldiers are not evenly distributed throughout the host body. They are found in the highest numbers in the basal visceral mass, where competing trematodes tend to multiply during
10032-558: The individuals cooperatively care for the brood of a single reproductive female (the queen) to which they are most likely related. Thus, it is uncertain whether mole rats are truly eusocial, since their social behavior depends largely on their resources and environment. Some mammals in the Carnivora and Primates have eusocial tendencies, especially meerkats ( Suricata suricatta ) and dwarf mongooses ( Helogale parvula ). These show cooperative breeding and marked reproductive skews. In
10146-399: The individuals in the colony are to some degree specialized for tasks, a division of labor . Eusociality is a rare but widespread phenomenon in species in at least seven orders in the animal kingdom , as shown in the phylogenetic tree (non-eusocial groups not shown). All species of termites are eusocial, and it is believed that they were the first eusocial animals to evolve, sometime in
10260-443: The known and newly described species. The relative ease with which ants may be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies. Ants are distinct in their morphology from other insects in having geniculate (elbowed) antennae , metapleural glands , and a strong constriction of their second abdominal segment into a node-like petiole . The head, mesosoma , and metasoma are
10374-417: The largest group of eusocial insects, including ants , bees , and wasps —divided into castes: reproductive queens , drones , more or less sterile workers, and sometimes also soldiers that perform specialized tasks. In the well-studied social wasp Polistes versicolor , dominant females perform tasks such as building new cells and ovipositing, while subordinate females tend to perform tasks like feeding
10488-561: The larvae and foraging. The task differentiation between castes can be seen in the fact that subordinates complete 81.4% of the total foraging activity, while dominants only complete 18.6% of the total foraging. Eusocial species with a sterile caste are sometimes called hypersocial. While only a moderate percentage of species in bees (families Apidae and Halictidae ) and wasps ( Crabronidae and Vespidae ) are eusocial, nearly all species of ants ( Formicidae ) are eusocial. Some major lineages of wasps are mostly or entirely eusocial, including
10602-561: The loss of their queen. The levels of two of the aliphatic compounds increase rapidly in virgin queens within the first week after emergence from the pupa , consistent with their roles as sex attractants during the mating flight. Once a queen is mated and begins laying eggs, she starts producing the full blend of compounds. In several ant species, reproductive activity is associated with pheromone production by queens. Mated egg-laying queens are attractive to workers, whereas young winged virgin queens elicit little or no response. Among ants,
10716-610: The majority of social bee diversity, have their lowest diversity in the tropics, being largely limited to temperate regions. Once pre-adaptations such as group formation, nest building, high cost of dispersal, and morphological variation are present, between-group competition has been suggested as a driver of the transition to advanced eusociality. M. A. Nowak, C. E. Tarnita, and E. O. Wilson proposed in 2010 that since eusociality produces an extremely altruistic society, eusocial groups should out-reproduce their less cooperative competitors, eventually eliminating all non-eusocial groups from
10830-484: The minor or median workers. In a few species, the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants , for example, have a distinct bimodal size distribution. Some other species show continuous variation in the size of workers. The smallest and largest workers in Carebara diversa show nearly a 500-fold difference in their dry weights. Workers cannot mate; however, because of
10944-508: The older dialects, although the Doric dialect has survived in the Tsakonian language , which is spoken in the region of modern Sparta. Doric has also passed down its aorist terminations into most verbs of Demotic Greek . By about the 6th century AD, the Koine had slowly metamorphosed into Medieval Greek . Phrygian is an extinct Indo-European language of West and Central Anatolia , which
11058-401: The opportunity. One plant, the epiphytic staghorn fern, Platycerium bifurcatum ( Polypodiaceae ), may exhibit a primitive form of eusocial behavior amongst clones. The evidence for this is that individuals live in colonies, where they are structured in different ways, with fronds of differing size and shape, to collect and store water and nutrients for the colony to use. At the top of
11172-551: The original meaning of the word was "the biter" (from Proto-Germanic * ai- , "off, away" + * mait- "cut"). The family name Formicidae is derived from the Latin formīca ("ant") from which the words in other Romance languages , such as the Portuguese formiga , Italian formica , Spanish hormiga , Romanian furnică , and French fourmi are derived. It has been hypothesised that
11286-487: The perfect stem eilēpha (not * lelēpha ) because it was originally slambanō , with perfect seslēpha , becoming eilēpha through compensatory lengthening. Reduplication is also visible in the present tense stems of certain verbs. These stems add a syllable consisting of the root's initial consonant followed by i . A nasal stop appears after the reduplication in some verbs. The earliest extant examples of ancient Greek writing ( c. 1450 BC ) are in
11400-627: The queen pheromone system of the fire ant Solenopsis invicta includes both releaser and primer pheromones. A queen recognition (releaser) pheromone is stored in the poison sac along with three other compounds. These compounds elicit a behavioral response from workers. Several primer effects have also been demonstrated. Pheromones initiate reproductive development in new winged females, called female sexuals. These chemicals inhibit workers from rearing male and female sexuals, suppress egg production in other queens of multiple queen colonies, and cause workers to execute excess queens. These pheromones maintain
11514-626: The reproductive output of other individuals that share their genes, especially their close relatives. Natural selection favors individuals to help their relatives when the cost of helping is less than the benefit gained by their relative multiplied by the fraction of genes that they share, i.e. when Cost < relatedness * Benefit . W. D. Hamilton suggested in 1964 that eusociality could evolve more easily among haplodiploid species such as Hymenoptera, because of their unusual relatedness structure. In haplodiploid species, females develop from fertilized eggs and males develop from unfertilized eggs. Because
11628-535: The roles of the sole reproducers, while soldiers and workers work together to create and maintain a living situation favorable for the brood. Queens produce multiple queen pheromones to create and maintain the eusocial state in their colonies; they may also eat eggs laid by other females or exert dominance by fighting. There are two eusocial rodents : the naked mole-rat and the Damaraland mole-rat . Some shrimps , such as Synalpheus regalis , are eusocial. E. O. Wilson and others have claimed that humans have evolved
11742-574: The solitary state. Pheromones play an important role in the physiological mechanisms of eusociality. Enzymes involved in the production and perception of pheromones were important for the emergence of eusociality within both termites and hymenopterans. The best-studied queen pheromone system in social insects is that of the honey bee Apis mellifera . Queen mandibular glands produce a mixture of five compounds, three aliphatic and two aromatic , which control workers. Mandibular gland extracts inhibit workers from constructing queen cells, which can delay
11856-506: The sub-order Isoptera , and together with cockroaches , they form the order Blattodea . Blattodeans are related to mantids , crickets , and other winged insects that do not undergo complete metamorphosis . Like ants, termites are eusocial , with sterile workers, but they differ greatly in the genetics of reproduction. The similarity of their social structure to that of ants is attributed to convergent evolution . Velvet ants look like large ants, but are wingless female wasps . Ants have
11970-402: The subfamilies Polistinae and Vespinae . The corbiculate bees (subfamily Apinae of family Apidae ) contain four tribes of varying degrees of sociality: the highly eusocial Apini (honey bees) and Meliponini (stingless bees), primitively eusocial Bombini (bumble bees), and the mostly solitary or weakly social Euglossini (orchid bees). Eusociality in these families is sometimes managed by
12084-530: The surface of the soil. During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian supercontinent (the Northern Hemisphere ). Their representation in the fossil record is poor, in comparison to the populations of other insects, representing only about 1% of fossil evidence of insects in the era. Ants became dominant after adaptive radiation at the beginning of
12198-517: The syllabic script Linear B . Beginning in the 8th century BC, however, the Greek alphabet became standard, albeit with some variation among dialects. Early texts are written in boustrophedon style, but left-to-right became standard during the classic period. Modern editions of ancient Greek texts are usually written with accents and breathing marks , interword spacing , modern punctuation , and sometimes mixed case , but these were all introduced later. The beginning of Homer 's Iliad exemplifies
12312-409: The three distinct body segments (formally tagmata ). The petiole forms a narrow waist between their mesosoma ( thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole may be formed by one or two nodes (the second alone, or the second and third abdominal segments). Tergosternal fusion, when the tergite and sternite of
12426-399: The total biomass of all the ants in the world was approximately equal to the total biomass of the entire human race. More careful estimates made in 2022 which take into account regional variations puts the global ant contribution at 12 megatons of dry carbon, which is about 20% of the total human contribution, but greater than that of the wild birds and mammals combined. This study also puts
12540-749: The upper Jurassic period (~150 million years ago). The other orders shown contain both eusocial and non-eusocial species, including many lineages where eusociality is inferred to be the ancestral state. Thus the number of independent evolutions of eusociality ( clades ) is not known. The major eusocial groups are shown in boldface in the phylogenetic tree. Mole-rats [REDACTED] Synalpheus spp. [REDACTED] [REDACTED] Kladothrips spp. [REDACTED] various Aphids [REDACTED] Austroplatypus incompertus [REDACTED] many Vespidae (wasps) [REDACTED] [REDACTED] many Bees [REDACTED] Staghorn fern Platycerium bifurcatum [REDACTED] Prior to
12654-419: The workers perform tasks within the nest such as provisioning cell broods, colony cleaning, and nectar reception and dehydration. Once older than 40 days, S. postica workers move outside the nest for colony defense and foraging. In Lasioglossum aeneiventre , a halictid bee from Central America, nests may be headed by more than one female; such nests have more cells, and the number of active cells per female
12768-480: Was Aeolic. For example, fragments of the works of the poet Sappho from the island of Lesbos are in Aeolian. Most of the dialect sub-groups listed above had further subdivisions, generally equivalent to a city-state and its surrounding territory, or to an island. Doric notably had several intermediate divisions as well, into Island Doric (including Cretan Doric ), Southern Peloponnesus Doric (including Laconian ,
12882-422: Was followed by at least one reversal to solitarity, giving a total of at least nine reversals. In a few species, solitary and eusocial colonies appear simultaneously in the same population, and different populations of the same species may be fully solitary or eusocial. This suggests that eusociality is costly to maintain, and can only persist when ecological variables favor it. Disadvantages of eusociality include
12996-511: Was then discovered in a group of chordates , the mole-rats. Further research distinguished another possibly important criterion for eusociality, "the point of no return". This is characterized by having individuals fixed into one behavioral group, usually before reproductive maturity. This prevents them from transitioning between behavioral groups, and creates a society with individuals truly dependent on each other for survival and reproductive success. For many insects, this irreversibility has changed
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