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69-428: Perissodactyla ( / p ə ˌ r ɪ s oʊ ˈ d æ k t ɪ l ə / , from Ancient Greek περισσός , perissós  'odd' and δάκτυλος , dáktylos  'finger, toe') is an order of ungulates . The order includes about 17 living species divided into three families : Equidae ( horses , asses , and zebras ), Rhinocerotidae ( rhinoceroses ), and Tapiridae ( tapirs ). They typically have reduced

138-537: 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

207-637: A close family relationship with at least some of the so-called Meridiungulata , a very diverse group of mammals living from the Paleocene to the Pleistocene in South America, whose systematic unity is largely unexplained. Some of these were classified based on their paleogeographic distribution. However, a close relationship can be worked out to perissodactyls by protein sequencing and comparison with fossil collagen from remnants of phylogenetically young members of

276-584: A discredited alternative scenario, a close relationship exists between perissodactyls, carnivores, and bats, this assembly comprising the Pegasoferae .   Artiodactyla  (Even-toed ungulates and cetaceans)   Perissodactyla  (Odd-toed ungulates)   † Meridiungulata  (South American ungulates,     especially Notoungulata and Litopterna ) According to studies published in March 2015, odd-toed ungulates are in

345-472: 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

414-551: A large, paired, intramembranous bone behind the premaxilla called the septomaxilla . Because this bone is vestigial in Acristatherium (a Cretaceous eutherian ) this species is believed to be the oldest known therian mammal. Intriguingly the septomaxilla is still present in monotremes . However, embryonic and fossil studies in 2021 suggest that the incisive bone, which has been called "premaxilla" in therian mammals , has been largely replaced by septomaxilla; and that only

483-419: 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

552-551: A palatal part of the incisive bone remains a vestige of premaxilla. If this hypothesis is accurate, the bones that have been called "premaxilla" in therian mammals are not entirely homologous to the original premaxilla of other vertebrates. This homology is, however, contended. The differences in the size and composition in the premaxilla of various families of bats is used for classification. The premaxillae of squamates are fused; this feature can be used to distinguish fossil squamates from relatives. In 1573, Volcher Coiter

621-411: A part of the palate originates from the medial nasal prominence . This may be due to the replacement of most of the incisive bone with septomaxilla in the therian mammal , as following section. In any case, the development and evolution of this region is complex and needs to be considered carefully. In bilateral cleft lip and palate , the growth pattern of the premaxilla differs significantly from

690-543: 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

759-671: A separate historical stage, though its earliest form closely resembles Attic Greek , and its latest form approaches Medieval Greek , and Koine may be classified as Ancient Greek in a wider sense. 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

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828-582: A small fraction of their original range. Members of this group are now found only in Central and South America, eastern and southern Africa, and central, southern, and southeastern Asia. During the peak of odd-toed ungulate existence, from the Eocene to the Oligocene , perissodactyls were distributed over much of the globe, the major exceptions being Australia and Antarctica. Horses and tapirs arrived in South America after

897-609: 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

966-411: A suture is often observable up to five years of age. It is also common in non-mammals, such as chickens, that premaxilla is derived from medial nasal prominence . However, experiments using mice have shown a different result. The bone that has been called the "premaxilla" (incisive bone) in mice consists of two parts: most of the bone covering the face originates from the maxillary prominence , and only

1035-510: 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

1104-714: A worldwide distribution, and feral animals of both species are now also found in regions outside their original range, such as in Australia. Perissodactyls inhabit a number of different habitats, leading to different lifestyles. Tapirs are solitary and inhabit mainly tropical rainforests. Rhinos tend to live alone in rather dry savannas, and in Asia, wet marsh or forest areas. Horses inhabit open areas such as grasslands, steppes, or semi-deserts, and live together in groups. Odd-toed ungulates are exclusively herbivores that feed, to varying degrees, on grass, leaves, and other plant parts. A distinction

1173-556: 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

1242-405: Is a term used for mammals, and it has been generally thought to be homologous to premaxilla in non-mammalian animals. However, there are counterarguments. According to them, the incisive bone is a novel character first acquired in therian mammals as a composition of premaxilla derived from medial nasal prominence and septomaxilla derived from maxillary prominence . In the incisive bones, only

1311-418: 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

1380-644: 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

1449-448: 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

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1518-427: Is formed from the fusion of a pair of small cranial bones at the very tip of the jaws of many animals, usually bearing teeth , but not always. They are connected to the maxilla and the nasals. While Johann Wolfgang von Goethe was not the first one to discover the incisive bone in humans, he was the first to prove its presence across mammals. Hence, the incisive bone is also known as Goethe's bone . Incisive bone

1587-455: Is high and the mandible is enlarged. Rhinos have one or two horns made of agglutinated keratin , unlike the horns of even-toed ungulates ( Bovidae and pronghorn ), which have a bony core. The number and form of the teeth vary according to diet. The incisors and canines can be very small or completely absent, as in the two African species of rhinoceros. In horses, usually only the males possess canines. The surface shape and height of

1656-445: Is of particular concern in equine veterinary practice , due to its frequent involvement in some serious infections. Aspergillosis (infection with Aspergillus mould) of the guttural pouch (also called guttural pouch mycosis ) can cause serious damage to the tissues of the pouch, as well as surrounding structures including important cranial nerves (nerves IX-XII: glossopharyngeal , vagus , accessory and hypoglossal nerves ) and

1725-447: Is often made between primarily grass feeders ( white rhinos , equines ) and leaf feeders (tapirs, other rhinos). Odd-toed ungulates are characterized by a long gestation period and a small litter size, usually delivering a single young. The gestation period is 330–500 days, being longest in rhinos. Newborn perissodactyls are precocial , meaning offspring are born already quite independent: for example, young horses can begin to follow

1794-420: Is relatively short, and the clavicle is absent. Odd-toed ungulates have a long upper jaw with an extended diastema between the front and cheek teeth , giving them an elongated head. The various forms of snout between families are due to differences in the form of the premaxilla . The lacrimal bone has projecting cusps in the eye sockets and a wide contact with the nasal bone. The temporomandibular joint

1863-404: Is very long, reaching up to 26 m (85 ft) in horses. Extraction of nutrients from food is relatively inefficient, which probably explains why no odd-toed ungulates are small; nutritional requirements per unit of body weight are lower for large animals, as their surface-area-to-volume ratio is smaller. Unlike artiodactyls, perissodactyls lack a carotid rete , a heat exchange that reduces

1932-745: 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

2001-600: 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

2070-411: The incisor teeth , and encompasses the anterior nasal spine and alar region. In the nasal cavity , the premaxillary element projects higher than the maxillary element behind. The palatal portion of the premaxilla is a bony plate with a generally transverse orientation. The incisive foramen is bound anteriorly and laterally by the premaxilla and posteriorly by the palatine process of the maxilla. It

2139-412: The internal carotid artery . Strangles ( Streptococcus equi equi infection) is a highly transmissible respiratory infection of horses that can cause pus to accumulate in the guttural pouch; horses with S. equi equi colonising their guttural pouch can continue to intermittently shed the bacteria for several months, and should be isolated from other horses during this time to prevent transmission. Due to

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2208-438: The molars is heavily dependent on whether soft leaves or hard grass make up the main component of their diets. Three or four cheek teeth are present on each jaw half, so the dental formula of odd-toed ungulates is: 0-3 . 0-1 . 2-4 . 3 1-3 . 1 . 2-4 . 3  × 2 = 30-44 The guttural pouch , a small outpocketing of the auditory tube that drains the middle ear , is a characteristic feature of Perissodactyla. The guttural pouch

2277-501: 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)

2346-453: The 19th century by the zoologist Richard Owen , who also coined the order's name. The largest odd-toed ungulates are rhinoceroses, and the extinct Paraceratherium , a hornless rhino from the Oligocene , is considered one of the largest land mammals of all time. At the other extreme, an early member of the order, the prehistoric horse Eohippus , had a withers height of only 30 to 60 cm (12 to 24 in). Apart from dwarf varieties of

2415-1031: 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"):

2484-490: 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

2553-563: 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,   Premaxilla In human anatomy , the premaxilla is referred to as the incisive bone ( os incisivum ) and is the part of the maxilla which bears

2622-711: The Meridiungulata (specifically Macrauchenia from the Litopterna and Toxodon from the Notoungulata ). Both kinship groups, the odd-toed ungulates and the Litopterna-Notoungulata, are now in the higher-level taxon of Panperissodactyla . This kinship group is included among the Euungulata, which also contains the even-toed ungulates and whales (Artiodactyla). The separation of the Litopterna-Notoungulata group from

2691-550: 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

2760-419: 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

2829-449: The brain. All perissodactyls are hindgut fermenters. In contrast to ruminants , hindgut fermenters store digested food that has left the stomach in an enlarged cecum , where the food begins digestion by microbes, with the fermentation continuing in the large colon . No gallbladder is present. The stomach of perissodactyls is simply built, while the cecum accommodates up to 90 L (24 US gal) in horses. The small intestine

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2898-438: 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

2967-494: The dependence of the temperature of the brain on that of the body. As a result, perissodactyls have limited thermoregulatory flexibility compared to artiodactyls which has restricted them to habitats of low seasonality and rich in food and water, such as tropical forests. In contrast, artiodactyls occupy a wide range of habits ranging from the Arctic Circle to deserts and tropical savannahs. Most extant perissodactyl species occupy

3036-611: 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

3105-423: The domestic horse and donkey, living perissodactyls reach a body length of 180–420 cm (71–165 in) and a weight of 150 to 4,500 kg (330 to 9,920 lb). While rhinos have only sparse hair and exhibit a thick epidermis , tapirs and horses have dense, short coats. Most species are grey or brown, although zebras and young tapirs are striped. The main axes of both the front and rear feet pass through

3174-992: The exact count is still controversial. Rhinos and tapirs are more closely related to each other than to horses. The separation of horses from other perissodactyls took place according to molecular genetic analysis in the Paleocene some 56 million years ago, while the rhinos and tapirs split off in the lower-middle Eocene , about 47 million years ago.   Equus ferus   Equus asinus   Equus hemionus   Equus kiang   Equus zebra   Equus grevyi   Equus quagga   Tapirus indicus   Tapirus bairdii   Tapirus kabomani   Tapirus pinchaque   Tapirus terrestris   Ceratotherium simum   Diceros bicornis   Dicerorhinus sumatrensis   Rhinoceros sondaicus   Rhinoceros unicornis There are many perissodactyl fossils of multivariant form. The major lines of development include

3243-401: The face during the fourth week of gestation. A pair of symmetrical nasal placodes (thickenings in the epithelium ) are each divided into medial and lateral processes by the nasal pits. The medial processes become the septum , philtrum , and premaxilla. The first ossification centers in the area of the future premaxilla appear during the seventh week above the germ of the second incisor on

3312-400: The following groups:   Palaeotheriidae (†)   Equidae   Lambdotheriidae (†)   Brontotheriidae (†)   Isectolophidae (†)   Lophiodontidae (†)   Chalicotheriidae (†) Ancient Greek language Ancient Greek ( Ἑλληνῐκή , Hellēnikḗ ; [hellɛːnikɛ́ː] ) includes the forms of the Greek language used in ancient Greece and

3381-568: The formation of the Isthmus of Panama around 3 million years ago in the Pliocene . Their North American counterparts died out around 10,000 years ago, leaving only Baird's tapir with a range extending to what is now southern Mexico. The tarpans were pushed to extinction in 19th century Europe. Hunting and habitat destruction have reduced the surviving perissodactyl species to fragmented populations. In contrast, domesticated horses and donkeys have gained

3450-556: 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

3519-472: 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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3588-410: The intermittent nature of S. equi equi shedding, prematurely reintroducing an infected horse may risk exposing other horses to the infection, even though the shedding horse appears well and may have previously returned negative samples. The function of the guttural pouch has been difficult to determine, but it is now believed to play a role in cooling blood in the internal carotid artery before it enters

3657-541: The mother after a few hours. The young are nursed for a relatively long time, often into their second year, with rhinos reaching sexual maturity around eight or ten years old, but horses and tapirs maturing around two to four years old. Perissodactyls are long-lived, with several species, such as rhinos, reaching an age of almost 50 years in captivity. Traditionally, the odd-toed ungulates were classified with other mammals such as artiodactyls , hyraxes , elephants and other "ungulates". A close family relationship with hyraxes

3726-424: The normal case; in utero growth is excessive and directed more horizontally, resulting in a protrusive premaxilla at birth. Forming the oral edge of the upper jaw in most jawed vertebrates , the premaxillary bones comprise only the central part in more primitive forms. They are fused in blowfishes and absent in cartilaginous fishes such as sturgeons . Reptiles and most non-mammalian therapsids have

3795-499: 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

3864-404: The outer surface of the nasal capsule. After eleven weeks an accessory ossification center develops into the alar region of the premaxilla. Then a premaxillary process grow upwards to fuse with the frontal process of the maxilla; and later expands posteriorly to fuse with the alveolar process of the maxilla. The boundary between the premaxilla and the maxilla remains discernible after birth and

3933-469: The palatine process corresponds to the premaxilla, while the other parts are the septomaxilla . Based on this, the incisive bone is not completely homologous to the non-mammalian premaxilla. This was hypothesized by Ernst Gaupp in 1905 and demonstrated by developmental biological- and paleontological experiments in 2021. This issue is still under debate. In the embryo, the nasal region develops from neural crest cells which start their migration down to

4002-487: 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

4071-543: The perissodactyls probably took place before the Cretaceous–Paleogene extinction event . "Condylarths" can probably be considered the starting point for the development of the two groups, as they represent a heterogeneous group of primitive ungulates that mainly inhabited the northern hemisphere in the Paleogene . Odd-toed ungulates (Perissodactyla) comprise three living families with around 17 species—in horses, however,

4140-646: The perissodactyls. These in turn are in the Laurasiatheria , a superorder that had its origin in the former supercontinent Laurasia . Molecular genetic findings suggest that the cloven Artiodactyla (containing the cetaceans as a deeply nested subclade) are the sister taxon of the Perissodactyla; together, the two groups form the Euungulata . More distant are the bats (Chiroptera) and Ferae (a common taxon of carnivorans, Carnivora , and pangolins, Pholidota ). In

4209-404: The shape of the humerus . For example, often shorter, thicker, bones belong to the largest and heaviest ungulates like the rhinoceros . The ulnae and fibulae are reduced in horses. A common feature that clearly distinguishes this group from other mammals is the articulation between the astragalus , the scaphoid and the cuboid , which greatly restricts the mobility of the foot. The thigh

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4278-517: 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

4347-406: The third toe, which is always the largest. The remaining toes have been reduced in size to varying degrees. Tapirs, which are adapted to walking on soft ground, have four toes on their fore feet and three on their hind feet. Living rhinos have three toes on both the front and hind feet. Modern equines possess only a single toe; however, their feet are equipped with hooves, which almost completely cover

4416-442: The toe. Rhinos and tapirs, by contrast, have hooves covering only the leading edge of the toes, with the bottom being soft. Ungulates have stances that require them to stand on the tips of their toes. Equine ungulates with only one digit or hoof have decreased mobility in their limbs, which allows for faster running speeds and agility. Differences in limb structure and physiology between ungulates and other mammals can be seen in

4485-523: The two is that perissodactyls digest plant cellulose in their intestines , rather than in one or more stomach chambers as artiodactyls, with the exception of Suina , do. The order was considerably more diverse in the past, with notable extinct groups including the brontotheres , palaeotheres , chalicotheres , and the paraceratheres , with the paraceratheres including the largest known land mammals to have ever existed. Despite their very different appearances, they were recognized as related families in

4554-403: The weight-bearing toes to three or one of the five original toes, though tapirs retain four toes on their front feet. The nonweight-bearing toes are either present, absent, vestigial , or positioned posteriorly. By contrast, artiodactyls (even-toed ungulates) bear most of their weight equally on four or two (an even number) of the five toes: their third and fourth toes. Another difference between

4623-475: 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 ,

4692-405: Was suspected based on similarities in the construction of the ear and the course of the carotid artery . Molecular genetic studies, however, have shown the ungulates to be polyphyletic , meaning that in some cases the similarities are the result of convergent evolution rather than common ancestry . Elephants and hyraxes are now considered to belong to Afrotheria , so are not closely related to

4761-436: Was the first to illustrate the incisive suture in humans. Pierre Marie Auguste Broussonet and Félix Vicq-d'Azyr were the first to describe the incisive bone as a separate bone within the skull in 1779 and 1780, respectively. In the 1790s, Johann Wolfgang von Goethe began studying zoology , and formed the impression that all animals are similar , being bodies composed of vertebrae and their permutations. The human skull

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