42-649: Agnatha ( / ˈ æ ɡ n ə θ ə , æ ɡ ˈ n eɪ θ ə / ; from Ancient Greek ἀ- ( a- ) 'without' and γνάθος ( gnáthos ) 'jaws') is a paraphyletic infraphylum of non- gnathostome vertebrates , or jawless fish , in the phylum Chordata , subphylum Vertebrata , consisting of both living ( cyclostomes ) and extinct ( conodonts , anaspids , and ostracoderms , among others). Among recent animals , cyclostomes are sister to all vertebrates with jaws , known as gnathostomes . Molecular data , both from rRNA and from mtDNA as well as embryological data, strongly supports
84-543: 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
126-445: A dorsal or a caudal fin . Some fossil agnathans, such as osteostracans and pituriaspids , did have paired fins, a trait inherited in their jawed descendants . Fertilization in lampreys is external. Mode of fertilization in hagfishes is not known. Development in both groups probably is external. There is no known parental care. Not much is known about the hagfish reproductive process. It is believed that hagfish only have 30 eggs over
168-828: A junior synonym of vertebrata. The new phylogeny from Miyashita et al . (2019) is considered compatible with both morphological and molecular evidence. † Myllokunmingiidae [REDACTED] † Euphanerida [REDACTED] † Jamoytiiformes [REDACTED] † Endeiolepis † Birkeniiformes [REDACTED] † Paraconodontida † Proconodontida † Protopanderodontida † Panderontida † Paracordylodus † Balognathidae [REDACTED] † Prioniodinida † Ozarkodinida Hagfish [REDACTED] Lampreys [REDACTED] † Anatolepis † Arandaspida [REDACTED] † Astraspidiformes [REDACTED] † Tesakoviaspidida † Eriptychiida † Tesseraspidiformes Ancient Greek language Ancient Greek ( Ἑλληνῐκή , Hellēnikḗ ; [hellɛːnikɛ́ː] ) includes
210-477: 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
252-419: 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
294-491: A lifetime. There is very little of the larval stage that characterizes the lamprey. Lamprey are only able to reproduce once. After external fertilization, the lamprey's cloacas remain open, allowing a fungus to enter their intestines, killing them. Lampreys reproduce in freshwater riverbeds, working in pairs to build a nest and burying their eggs about an inch beneath the sediment. The resulting hatchlings go through four years of larval development before becoming adults. Although
336-431: A long gut, more or less homogeneous throughout its length. Lampreys feed on other fish and mammals. Anticoagulant fluids preventing blood clotting are injected into the host, causing the host to yield more blood. Hagfish are scavengers, eating mostly dead animals. They use a row of sharp teeth to break down the animal. Because Agnathan teeth are unable to move up and down it limits their possible food types. In addition to
378-409: A minor element of modern marine fauna , agnathans were prominent among the early fish in the early Paleozoic . Two types of Early Cambrian animal apparently having fins, vertebrate musculature, and gills are known from the early Cambrian Maotianshan shales of China : Haikouichthys and Myllokunmingia . They have been tentatively assigned to Agnatha by Janvier. A third possible agnathid from
420-550: 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
462-608: 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
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#1732779933683504-630: 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
546-636: A thymus-like lympho-epithelial structure, termed thymoids. VLRB+ cells are B-cell-like, develop in hematopoietic organs, and differentiate into "VLRB antibody"-secreting plasma cells. Conodont (extinct) The taxon contains the subgroups Heterostraci , Astraspida , Arandaspida . Phylogeny based on the work of Mikko Haaramo and Delsuc et al. Hyperotreti / Myxini (hagfishes) Petromyzontomorpha (lampreys) ?† Conodonta † Pteraspidomorpha ?† Jamoytiiformes ?† Euphanerida † Anaspida † Thelodonti † Galeaspida † Pituriaspida † Osteostraci Gnathostomata (vertebrates with jaws) While
588-510: 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
630-570: 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
672-418: 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
714-666: 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
756-448: 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
798-735: Is no parental care in the Agnatha class. The Agnatha are ectothermic or cold, with a cartilaginous skeleton , and the heart contains 2 chambers. In modern agnathans, the body is covered in skin, with neither dermal or epidermal scales . The skin of hagfish has copious slime glands, the slime constituting their defense mechanism. The slime can sometimes clog up enemy fishes' gills, causing them to die. In direct contrast, many extinct agnathans sported extensive exoskeletons composed of either massive, heavy dermal armour or small mineralized scales. Almost all agnathans, including all extant agnathans , have no paired appendages, although most do have
840-759: 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
882-501: 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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#1732779933683924-412: The "agnatha" Conodonta was indeed jawless, if it would have continued to live, its descendants would still be closer related to e.g. humans than to lampreys, and also contempory it was closer related to the ancestor of humans. Due to such considerations, Agnatha can not be consolidated into a coherent grouping without either removing any non-cyclostomata, or by including all vertebrata thus rendering it into
966-1031: 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"):
1008-495: 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
1050-428: The absence of jaws , modern agnathans are characterised by absence of paired fins ; the presence of a notochord both in larvae and adults; and seven or more paired gill pouches. Lampreys have a light sensitive pineal eye (homologous to the pineal gland in mammals ). All living and most extinct Agnatha do not have an identifiable stomach or any appendages . Fertilization and development are both external. There
1092-550: 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
1134-419: 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
1176-438: 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
1218-615: 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
1260-673: 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
1302-590: The gnathostomes than to the surviving agnathans, known as cyclostomes. Cyclostomes apparently split from other agnathans before the evolution of dentine and bone, which are present in many fossil agnathans, including conodonts . Agnathans declined in the Devonian and never recovered. Approximately 500 million years ago, two types of recombinatorial adaptive immune systems (AISs) arose in vertebrates. The jawed vertebrates diversify their repertoire of immunoglobulin domain-based T and B cell antigen receptors mainly through
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1344-561: 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
1386-476: 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
1428-473: The hypothesis that both groups of living agnathans, hagfishes and lampreys , are more closely related to each other than to jawed fish , forming the clade Cyclostomi . The oldest fossil agnathans appeared in the Cambrian . Living jawless fish comprise about 120 species in total. Hagfish are considered members of the subphylum Vertebrata , because they secondarily lost vertebrae; before this event
1470-676: The late Cambrian to the Triassic. Many Ordovician, Silurian, and Devonian agnathans were armored with heavy bony-spiky plates. The first armored agnathans—the Ostracoderms , precursors to the bony fish and hence to the tetrapods (including humans )—are known from the middle Ordovician , and by the Late Silurian the agnathans had reached the high point of their evolution. Most of the ostracoderms, such as thelodonts , osteostracans , and galeaspids , were more closely related to
1512-508: 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
1554-487: 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
1596-644: The rearrangement of V(D)J gene segments and somatic hypermutation, but none of the fundamental AIS recognition elements in jawed vertebrates have been found in jawless vertebrates. Instead, the AIS of jawless vertebrates is based on variable lymphocyte receptors (VLRs) that are generated through recombinatorial usage of a large panel of highly diverse leucine-rich-repeat (LRR) sequences. Three VLR genes (VLRA, VLRB, and VLRC) have been identified in lampreys and hagfish, and are expressed on three distinct lymphocytes lineages. VLRA+ cells and VLRC+ cells are T-cell-like and develop in
1638-565: The same region is Haikouella . A possible agnathid that has not been formally described was reported by Simonetti from the Middle Cambrian Burgess Shale of British Columbia . Conodonts , a class of agnathans which arose in the early Cambrian, remained common enough until their extinction in the Triassic that their teeth (the only parts of them that were usually fossilized) are often used as index fossils from
1680-517: 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
1722-480: 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 ,
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1764-475: Was inferred from molecular and developmental data, the Craniata hypothesis was accepted (and is still sometimes used as a strictly morphological descriptor) to reference hagfish plus vertebrates. Agnathans are ectothermic , meaning they do not regulate their own body temperature. Agnathan metabolism is slow in cold water, and therefore they do not have to eat very much. They have no distinct stomach, but rather
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