52-494: Gnathostomata ( / ˌ n æ θ oʊ ˈ s t ɒ m ə t ə / ; from Ancient Greek : γνάθος ( gnathos ) 'jaw' + στόμα ( stoma ) 'mouth') are the jawed vertebrates . Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all living vertebrates, including humans. Most gnathostomes have retained ancestral traits like true teeth , a stomach , and paired appendages (pectoral and pelvic fins, arms, legs, wings, etc.). Other traits are elastin ,
104-561: A cartilaginous skeleton , but their fins had a wide, bony base and were reinforced on their anterior margin with a dentine spine. As a result, fossilized spines and scales are often all that remains of these fishes in ancient sedimentary rocks . The earliest acanthodians were marine, but during the Devonian , freshwater species became predominant. Acanthodians have been divided into four orders: Acanthodiformes , Climatiiformes , Diplacanthiformes , and Ischnacanthiformes . "Climatiiformes"
156-400: A paraphyletic grade of various fish lineages basal to extant Chondrichthyes , which includes living sharks , rays , and chimaeras . Acanthodians possess a mosaic of features shared with both osteichthyans (bony fish) and chondrichthyans (cartilaginous fish). In general body shape, they were similar to modern sharks, but their epidermis was covered with tiny rhomboid platelets like
208-537: 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
260-620: A branch of Placoderms was most likely the ancestor of present-day gnathostomes. A 419-million-year-old fossil of a placoderm named Entelognathus had a bony skeleton and anatomical details associated with cartilaginous and bony fish, demonstrating that the absence of a bony skeleton in Chondrichthyes is a derived trait. The fossil findings of primitive bony fishes such as Guiyu oneiros and Psarolepis , which lived contemporaneously with Entelognathus and had pelvic girdles more in common with placoderms than with other bony fish, show that it
312-411: A horizontal semicircular canal of the inner ear, myelin sheaths of neurons , and an adaptive immune system which has discrete lymphoid organs ( spleen and thymus ), and uses V(D)J recombination to create antigen recognition sites, rather than using genetic recombination in the variable lymphocyte receptor gene. It is now assumed that Gnathostomata evolved from ancestors that already possessed
364-472: 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
416-419: 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
468-418: A neck, and a flat or slightly curved diamond-shaped crown. Despite being called "spiny sharks", acanthodians predate sharks. Scales that have been tentatively identified as belonging to acanthodians, or "shark-like fishes" have been found in various Ordovician strata, though, they are ambiguous, and may actually belong to jawless fishes such as thelodonts . The earliest unequivocal acanthodian fossils date from
520-471: A pair of both pectoral and pelvic fins . Until recently these ancestors, known as antiarchs , were thought to have lacked pectoral or pelvic fins. In addition to this, some placoderms (extinct fish with bony plates) were shown to have a third pair of paired appendages, that had been modified to claspers in males and basal plates in females—a pattern not seen in any other vertebrate group. The Osteostraci (bony armored jawless fish) are generally considered
572-1219: A paraphyletic assemblage leading to cartilaginous fish, while bony fish evolved from placoderm ancestors. Burrow et al. 2016 provides vindication by finding chondrichthyans to be nested among Acanthodii, most closely related to Doliodus and Tamiobatis . A 2017 study of Doliodus morphology points out that it appears to display a mosaic of shark and acanthodian features, making it a transitional fossil and further reinforcing this idea. Phylogeny after Galeaspida Osteostraci " Placodermi " Osteichthyes Tetanopsyrus Nerepisacanthus Ischnacanthus Poracanthodes Culmacanthus Uraniacanthus Diplacanthus Rhadinacanthus Cassidiceps Mesacanthus Lodeacanthus Triazeugacanthus Promesacanthus Acanthodes Cheiracanthus Homalacanthus Euthacanthus Ptomacanthus Brachyacanthus Climatius Parexus Vernicomacanthus Lupopsyrus Obtusacanthus Kathemacanthus Brochoadmones Gyracanthides Chondrichthyes (conventionally defined) The oldest remains attributed acanthodian-grade chondrichthyans are Fanjingshania and Qianodus from
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#1732844137076624-543: 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
676-608: 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
728-450: A single dorsal fin, toothless jaws, and long gill rakers . They were the last and most specialized off the traditional acanthodians, as they survived up until the Permian period. The scales of Acanthodii have distinctive ornamentation peculiar to each order. Because of this, the scales are often used in determining relative age of sedimentary rock. The scales are tiny, with a bulbous base,
780-609: 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
832-617: A study of early jawed vertebrate relationships, Davis et al. (2012) found acanthodians to be split among the two major clades Osteichthyes (bony fish) and Chondrichthyes (cartilaginous fish). The well-known acanthodian Acanthodes was placed within Osteichthyes, despite the presence of many chondrichthyan characteristics in its braincase. However, a newly described Silurian placoderm , Entelognathus , which has jaw anatomy shared with bony fish and tetrapods , has led to revisions of this phylogeny: acanthodians were then considered to be
884-510: 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
936-556: 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
988-474: Is a paraphyletic assemblage of early acanthodians such as climatiids , gyracanthids , and diplacanthids ; they had robust bony shoulder girdles and many small sharp spines ("intermediate" or "prepelvic" spines) between the pectoral and pelvic fins. The climatiiform subgroup Diplacanthida has subsequently been elevated to its own order, Diplacanthiformes. Ischnacanthiforms were predators with tooth plates fused to their jaws. Acanthodiforms were filter feeders with
1040-418: 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
1092-644: 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
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#17328441370761144-448: 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
1196-405: Is that jaws are homologous to the gill arches . In jawless fishes a series of gills opened behind the mouth, and these gills became supported by cartilaginous elements. The first set of these elements surrounded the mouth to form the jaw. The upper portion of the second embryonic arch supporting the gill became the hyomandibular bone of jawed fish, which supports the skull and therefore links
1248-745: 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
1300-403: The operculum in later bony fishes . However, most of these characteristics are considered homologous characteristics derived from common placoderm ancestors , and present also in basal cartilaginous fish . Overall, the acanthodians' jaws are presumed to have evolved from the first gill arch of some ancestral jawless fishes that had a gill skeleton made of pieces of jointed cartilage. In
1352-501: 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)
1404-404: The sister taxon of Gnathostomata. Jaw development in vertebrates is likely a product of the supporting gill arches. This development would help push water into the mouth by the movement of the jaw, so that it would pass over the gills for gas exchange. The repetitive use of the newly formed jaw bones would eventually lead to the ability to bite in some gnathostomes. Newer research suggests that
1456-1031: 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"):
1508-490: 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
1560-555: 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, Acanthodians Acanthodii or acanthodians is an extinct class of gnathostomes (jawed fishes ). They are currently considered to represent
1612-694: The Early Silurian of China, dating to around 439 million years ago. Compared to other contemporary groups of fish, acanthodians were relatively morphologically and ecologically conservative. Acanthodians rose in diversity during the Late Silurian, reaching their apex of diversity during the Lochkovian stage of the Early Devonian, declining during the Pragian but rising again during the following Emsian , which
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1664-423: The acanthodians close to the ancestors of the bony fishes. Although their interior skeletons were made of cartilage , a bonelike material had developed in the skins of these fishes, in the form of closely fitting scales (see above) . Some scales were greatly enlarged and formed a bony covering on top of the head and over the lower shoulder girdle . Others developed a bony flap over the gill openings analogous to
1716-550: 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
1768-419: 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
1820-583: The beginning of the Silurian Period, some 50 million years before the first sharks appeared. Later, the acanthodians colonized fresh waters, and thrived in the rivers and lakes during the Devonian and in the coal swamps of Carboniferous . By this time bony fishes were already showing their potential to dominate the waters of the world, and their competition proved too much for the spiny sharks, which died out in Permian times (approximately 250 million years ago). Many palaeontologists originally considered
1872-438: 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
1924-483: The deep remodelling of the vertebrate skull that must have taken place as early jaws evolved. The ancestor of all jawed vertebrates have gone through two rounds of whole genome duplication. The first happened before the gnathostome and cyclostome split, and appears to have been an autopolyploidy event (happened within the same species). The second occurred after the split, and was an allopolyploidy event (the result of hybridization between two lineages). The customary view
1976-611: 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
2028-473: The early vertebrate jaw has been described as "a crucial innovation" and "perhaps the most profound and radical evolutionary step in the vertebrate history". Fish without jaws had more difficulty surviving than fish with jaws, and most jawless fish became extinct during the Triassic period. However studies of the cyclostomes , the jawless hagfishes and lampreys that did survive, have yielded little insight into
2080-663: 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
2132-556: 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
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2184-472: 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
2236-727: The jaw to the cranium. The hyomandibula is a set of bones found in the hyoid region in most fishes. It usually plays a role in suspending the jaws or the operculum in the case of teleosts . While potentially older Ordovician records are known, the oldest unambigious evidence of jawed vertebrates are Qianodus and Fanjingshania from the early Silurian ( Aeronian ) of Guizhou , China around 439 million years ago, which are placed as acanthodian -grade stem -chondrichthyans. [REDACTED] [REDACTED] [REDACTED] [REDACTED] [REDACTED] [REDACTED] [REDACTED] Ancient Greek Ancient Greek ( Ἑλληνῐκή , Hellēnikḗ ; [hellɛːnikɛ́ː] ) includes
2288-419: The jawless craniates Agnatha . † Placodermi [REDACTED] Acanthodians , incl. Chondrichthyes (cartilaginous fishes) [REDACTED] Actinopterygii [REDACTED] Amphibia [REDACTED] Sauria [REDACTED] Mammalia [REDACTED] The appearance of
2340-499: 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
2392-401: The origin of teeth along with, or soon after, the evolution of jaws. Late Ordovician -aged microfossils of what have been identified as scales of either acanthodians or "shark-like fishes", may mark Gnathostomata's first appearance in the fossil record. Undeniably unambiguous gnathostome fossils, mostly of primitive acanthodians, begin appearing by the early Silurian , and become abundant by
2444-487: 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
2496-462: The scales of holosteians ( gars , bowfins ). The popular name " spiny sharks " is because they were superficially shark-shaped, with a streamlined body, paired fins , a strongly upturned tail, and stout, largely immovable bony spines supporting all the fins except the tail—hence, "spiny sharks". However, acanthodians are not true sharks; their close relation to modern cartilaginous fish can lead them to be considered " stem -sharks". Acanthodians had
2548-415: The start of the Devonian . Gnathostomata is traditionally a infraphylum , broken into three top-level groupings: Chondrichthyes , or the cartilaginous fish; Placodermi , an extinct grade of armored fish; and Teleostomi , which includes the familiar classes of bony fish , birds , mammals , reptiles , and amphibians . Some classification systems have used the term Amphirhina . It is a sister group of
2600-517: 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
2652-475: 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 ,
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#17328441370762704-456: Was a relative rather than a direct ancestor of the extant gnathostomes. It also indicates that spiny sharks and Chondrichthyes represent a single sister group to the bony fishes. Fossil findings of juvenile placoderms, which had true teeth that grew on the surface of the jawbone and had no roots, making them impossible to replace or regrow as they broke or wore down as they grew older, proves the common ancestor of all gnathostomes had teeth and place
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