51-1057: Hyenas or hyaenas ( / h aɪ ˈ iː n ə z / hi- EE -nəz ; from Ancient Greek ὕαινα , hýaina ) are feliform carnivoran mammals belonging to the family Hyaenidae ( / h aɪ ˈ ɛ n ɪ d iː / ). With just four extant species (each in its own genus ), it is the fifth-smallest family in the order Carnivora and one of the smallest in the class Mammalia . Despite their low diversity, hyenas are unique and vital components of most African ecosystems. Although phylogenetically closer to felines and viverrids , hyenas are behaviourally and morphologically similar to canids in several elements due to convergent evolution : both hyenas and canines are non- arboreal , cursorial hunters that catch prey with their teeth rather than claws. Both eat food quickly and may store it, and their calloused feet with large, blunt, nonretractable claws are adapted for running and making sharp turns. However, hyenas' grooming, scent marking , defecation habits, mating and parental behavior are consistent with
102-543: 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
153-580: A change in climate, along with the arrival of canids into Eurasia. Of the dog-like hyena lineage, only the insectivorous aardwolf survived, while the bone-crushing hyenas (including the extant spotted, brown and striped hyenas) became the undisputed top scavengers of Eurasia and Africa. The descendants of Plioviverrops reached their peak 15 million years ago, with more than 30 species having been identified. Unlike most modern hyena species, which are specialised bone-crushers, these dog-like hyenas were nimble-bodied, wolfish animals; one species among them
204-477: 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
255-419: 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
306-550: 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
357-608: 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
408-630: 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
459-510: 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
510-570: 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
561-418: 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
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#1732780949210612-666: 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
663-448: 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
714-545: Is likely that its unrivaled ability to digest the terpene excretions from soldier termites is a modification of the strong digestive system its ancestors used to consume fetid carrion. The striped hyena may have evolved from Hyaenictitherium namaquensis of Pliocene Africa . Striped hyena fossils are common in Africa, with records going back as far as the Villafranchian . As fossil striped hyenas are absent from
765-1190: The Furninha Cave in Portugal and the Genista Caves in Gibraltar . The European form was similar in appearance to modern populations, but was larger, being comparable in size to the brown hyena . The spotted hyena ( Crocuta crocuta ) diverged from the striped and brown hyena 10 million years ago. Its direct ancestor was the Indian Crocuta sivalensis , which lived during the Villafranchian. Ancestral spotted hyenas probably developed social behaviours in response to increased pressure from rivals on carcasses, thus forcing them to operate in teams. Spotted hyenas evolved sharp carnassials behind their crushing premolars, therefore they did not need to wait for their prey to die, and thus became pack hunters as well as scavengers. They began forming increasingly larger territories , necessitated by
816-759: 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
867-861: The Mediterranean region, it is likely that the species is a relatively late invader to Eurasia, having likely spread outside Africa only after the extinction of spotted hyenas in Asia at the end of the Ice Age . The striped hyena occurred for some time in Europe during the Pleistocene, having been particularly widespread in France and Germany . It also occurred in Montmaurin , Hollabrunn in Austria ,
918-576: The Mid-Pleistocene transition . The four extant species are the striped hyena ( Hyaena hyaena ), the brown hyena ( Parahyaena brunnea ), the spotted hyena ( Crocuta crocuta ), and the aardwolf ( Proteles cristata ). The aardwolf can trace its lineage directly back to Plioviverrops 15 million years ago, and is the only survivor of the dog-like hyena lineage. Its success is partly attributed to its insectivorous diet, for which it faced no competition from canids crossing from North America. It
969-524: The Middle Miocene through the Pliocene , existing for about 8 million years . The first percrocutids are known from the middle Miocene of Europe and western Asia and belonged to the genus Percrocuta . Percrocuta already had large premolars , but did not carry such a massive bite as the later form Dinocrocuta , from the later Miocene. Originally, these carnivores were placed with
1020-520: The middle ear and dentition. The lineage of Plioviverrops prospered, and gave rise to descendants with longer legs and more pointed jaws, a direction similar to that taken by canids in North America . Hyenas then diversified into two distinct types: lightly built dog-like hyenas and robust bone-crushing hyenas. Although the dog-like hyenas thrived 15 million years ago (with one taxon having colonised North America), they became extinct after
1071-501: 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)
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#17327809492101122-1031: 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"):
1173-495: 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
1224-568: The Classical period of ancient Greek. (The second line is the IPA , the third is transliterated into the Latin alphabet using a modern version of the Erasmian scheme .) Ὅτι [hóti Hóti μὲν men mèn ὑμεῖς, hyːmêːs hūmeîs, Percrocutidae Percrocutidae is an extinct family of hyaenid feliform carnivores endemic to Asia , Africa , and Southern Europe from
1275-607: The World for extant genera. The percrocutids are, in contrast to McKenna and Bell's classification, not included as a subfamily into the Hyaenidae, but as the separate family Percrocutidae (though they are generally grouped as sister-taxa to hyenas). Furthermore, the living brown hyena and its closest extinct relatives are not included in the genus Pachycrocuta , but in the genus Parahyaena . However, some research has suggested Parahyaena may be synonymous with Pachycrocuta , making
1326-421: The ancestral bone-crushing hyenas coincided with the decline of the similarly built family Percrocutidae . The bone-crushing hyenas survived the changes in climate and the arrival of canids, which wiped out the dog-like hyenas, though they never crossed into North America, as their niche there had already been taken by the dog subfamily Borophaginae . By 5 million years ago, the bone-crushing hyenas had become
1377-550: 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
1428-419: 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
1479-435: The behavior of other feliforms. Hyenas feature prominently in the folklore and mythology of human cultures that live alongside them. Hyenas are commonly viewed as frightening and worthy of contempt. In some cultures, hyenas are thought to influence people's spirits, rob graves, and steal livestock and children. Other cultures associate them with witchcraft, using their body parts in traditional medicine . Hyenas originated in
1530-418: The broader context of late-Quaternary extinctions , as the late Pleistocene and early Holocene saw the disappearance of many primarily large mammals from Europe and the world. Expansion or duplication of the olfatory receptor gene family has been found in all 4 extant species, which would have led to the evolution of the more specialised feeding habits of hyenas. Expansion in immune-related gene families
1581-1368: The brown hyena the only extant member of this genus. The following cladogram illustrates the phylogenetic relationships between extant and extinct hyaenids based on the morphological analysis by Werdelin & Solounias (1991), as updated by Turner et al. (2008). Protictitherium crassum "Protictitherium" cingulatum "Protictitherium" intermedium "Protictitherium" llopisi "Protictitherium" punicum " Protictitherium" gaillardi "Protictitherium" sumegense "Protictitherium" csakvarense Plioviverrops gervaisi Plioviverrops orbignyi Plioviverrops guerini Plioviverrops faventinus Plioviverrops gaudryi Tungurictis spocki Thalassictis robusta "Thalassictis" certa "Thalassictis" montadai "Thalassictis" proava "Thalassictis" sarmatica "Thalassictis" spelaea Tongxinictis primordialis Proteles cristatus (aardwolf) [REDACTED] Proteles amplidentus Ictitherium viverrinum Ictitherium ebu Ictitherium tauricum Ictitherium ibericum Ictitherium kurteni Ictitherium intuberculatum Ictitherium pannonicum Miohyaenotherium bessarabicum Hyaenotherium wongii Hyaenictitherium hyaenoides "Hyaenictitherium" pilgrimi Ancient Greek Ancient Greek ( Ἑλληνῐκή , Hellēnikḗ ; [hellɛːnikɛ́ː] ) includes
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1632-438: 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
1683-615: 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
1734-416: The dominant scavengers of Eurasia, primarily feeding on large herbivore carcasses felled by sabre-toothed cats . One genus, Pachycrocuta , was a 110 kg (240 lb) mega-scavenger that could splinter the bones of elephants . Starting in the early Middle Pleistocene Pachycrocuta was replaced by the smaller Crocuta and Hyena , which corresponds to a general faunal change, perhaps in connection to
1785-408: The end of the last glacial period and a subsequent displacement of open grassland by closed forests, which favoured wolves and humans instead. However, analyses have shown that climate change alone is insufficient to explain the spotted hyena's disappearance from Europe, suggesting that other factors – such as human pressure – must have played a role. This suggests that the events must be seen within
1836-480: The evolution of the feeding of termites Trinervitermes in this species. Mutations and variants in genes related to craniofacial shape were also found ( GARS , GMPR , STIP1 , SMO and PAPSS2 ). Another gene is related to protective epidermis function ( DSC1 ). The list follows McKenna and Bell's Classification of Mammals for prehistoric genera (1997) and Wozencraft (2005) in Wilson and Reeders Mammal Species of
1887-478: The fact that their prey was often migratory, and long chases in a small territory would have caused them to encroach into another clan's turf. Spotted hyenas spread from their original homeland during the Middle Pleistocene , and quickly colonised a very wide area from Europe, to southern Africa and China . The eventual disappearance of the spotted hyena from Europe has traditionally been attributed to
1938-673: 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
1989-488: The genera Percrocuta , Dinocrocuta , Belbus and Allohyaena . Later studies have suggested that Belbus and Allohyaena are true hyaenids and not percrocutids. The list follows McKenna and Bell's Classification of Mammals for prehistoric genera (1997). In contrast to McKenna and Bell's classification, they are not included as a subfamily into the Hyaenidae but as a separate family Percrocutidae. This article related to prehistoric animals from order Carnivora
2040-561: 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
2091-476: 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
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2142-491: The hyenas in the family Hyaenidae . As of 2013 , most scientists considered the Percrocutidae to be a distinct family - although usually as sister-taxa/immediate outgroup to Hyaenidae. Sometimes it was placed with the family Stenoplesictidae into the superfamily Stenoplesictoidea . However, studies in the 2020s placed Dinocrocuta and Percrocuta as true hyaenids, invalidating the family Percrocutidae. Percrocuta
2193-402: The jungles of Miocene Eurasia 22 million years ago, when most early feliform species were still largely arboreal . The first ancestral hyenas were likely similar to the modern African civet ; one of the earliest hyena species described, Plioviverrops , was a lithe, civet-like animal that inhabited Eurasia 20–22 million years ago, and is identifiable as a hyaenid by the structure of
2244-549: The land bridge into North America, being the only hyena to do so. Chasmaporthetes managed to survive for some time in North America by deviating from the endurance-running and bone-crushing niches monopolized by canids, and developing into a cheetah -like sprinter. Most of the dog-like hyenas had died off by 1.5 million years ago. By 10–14 million years ago, the hyena family had split into two distinct groups: dog-like hyenas and bone-crushing hyenas. The arrival of
2295-508: 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
2346-487: 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
2397-517: 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
2448-521: Was Ictitherium viverrinum , which was similar to a jackal . The dog-like hyenas were numerous; in some Miocene fossil sites, the remains of Ictitherium and other dog-like hyenas outnumber those of all other carnivores combined. The decline of the dog-like hyenas began 5–7 million years ago during a period of climate change, exacerbated by canids crossing the Bering land bridge to Eurasia. One species, Chasmaporthetes ossifragus , managed to cross
2499-480: 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 ,
2550-583: Was also found in the spotted hyena, striped hyena and brown hyena, which would have led to the evolution of the scavenging in these species. Mutations and variants were also found in digestion-related genes ( ASH1L , PTPN5 , PKP3 , AQP10 ). One of these digestion-related genes has variants also related to enhanced bone mineralisation ( PTPN5 ), while other have also a role in inflammatory skin responses ( PKP3 ). In aardwolves, expansion of genes related to toxin response were found ( Lipocalin and UDP Glucuronosyltransferase gene families), which would have led to
2601-404: Was first considered as a side-branch outside of Hyaenidae by Thenius in 1966. It was later named as a different subfamily, Percrocutinae, of Hyaenidae in 1976, and at that time was proposed to include Percrocuta , Adcrocuta eximia , and Allohyaena kadici . Dinocrocuta was elevated from a subgenus to a full genus in 1988. The family Percrocutidae was formally elevated in 1991, to include
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