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76-467: The term coelom derives from the Ancient Greek word κοιλία ( koilía ) 'cavity'. The coelom is the mesodermally lined cavity between the gut and the outer body wall. During the development of the embryo , coelom formation begins in the gastrulation stage. The developing digestive tube of an embryo forms as a blind pouch called the archenteron . In protostomes , the coelom forms by

152-549: A backbone or spine, consisting of vertebrae and intervertebral discs . The vertebrae are irregular bones , and the intervertebral discs are of fibrocartilage . The vertebral column surrounds and protects the spinal cord . The other feature unique to vertebrates is the presence of a cranium , or skull. The vertebrates make up the subphylum Vertebrata with some 65,000 species in the phylum Chordata . The vertebrates include mammals , birds , amphibians , and various classes of reptiles and fish . Classes of fish include

228-427: A jaw , hyoid and/or the middle ear ossicles . An iodine-concentrating organ called the endostyle , which functions as a filter feeding organ in aquatic animals has evolved into the thyroid in most vertebrates. Vertebrates vary in body length ranging from the frog species Brachycephalus pulex , a Brazilian flea toad, with a minimum adult snout–vent length of 6.45 millimetres (0.254 in) to

304-452: A molecular phylogeny study mistakenly called this classification scheme the "traditional, morphology-based phylogeny". Coelomate animals or Coelomata (also known as eucoelomates – "true coelom") have a body cavity called a coelom with a complete lining called peritoneum derived from mesoderm (one of the three primary tissue layers ). The complete mesoderm lining allows organs to be attached to each other so that they can be suspended in

380-543: 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

456-417: A central nervous system arising from a single nerve cord dorsal to the gut tube , headed by a series of (typically paired) brain vesicles, is unique to vertebrates. This is in stark contrast to invertebrates with well-developed central nervous systems such as arthropods and cephalopods , which have an often ladder-like ventral nerve cord made of paired segmental ganglia on the opposite ( ventral ) side of

532-424: A coelom is correlated with a reduction in body size. Coelom is sometimes incorrectly used to refer to any developed digestive tract. Some organisms may not possess a coelom or may have a false coelom ( pseudocoelom ). Animals having coeloms are called coelomates , and those without are called acoelomates . There are also subtypes of coelom: According to Brusca and Brusca , the following bilaterian phyla possess

608-437: A coelom: In some protostomes , the embryonic blastocoele persists as a body cavity. These protostomes have a fluid filled main body cavity unlined or partially lined with tissue derived from mesoderm. This fluid-filled space surrounding the internal organs serves several functions like distribution of nutrients and removal of waste or supporting the body as a hydrostatic skeleton . A pseudocoelomate or blastocoelomate

684-476: A coelomate. All pseudocoelomates are protostomes ; however, not all protostomes are pseudocoelomates. An example of a pseudocoelomate is the roundworm. Pseudocoelomate animals are also referred to as blastocoelomate . Acoelomate animals, like flatworms , have no body cavity at all. Semi-solid mesodermal tissues between the gut and body wall hold their organs in place. Coeloms developed in triploblasts but were subsequently lost in several lineages. The lack of

760-433: A fluid-filled body cavity between the body wall and digestive tract. This can cause some serious disadvantages. Fluid compression is negligible, while the tissue surrounding the organs of these animals will compress. Therefore, acoelomate organs are not protected from crushing forces applied to the animal's outer surface. The coelom can be used for diffusion of gases and metabolites etc. These creatures do not have this need, as

836-525: A hollow neural tube ) running along the dorsal aspect of the notochord . Of particular importance and unique to vertebrates is the presence of neural crest cells, which are progenitor cells critical to coordinating the functions of cellular components. Neural crest cells migrate through the body from the dorsal nerve cord during development, initiate the formation of neuronal ganglia and various special sense organs. The peripheral nervous system forms when neural crest cells branch out laterally from

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912-477: 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

988-419: 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

1064-458: A non-taxonomic, utilitarian way, as the Acoelomata, Pseudocoelomata, and Coelomata. These groups were never intended to represent related animals, or a sequence of evolutionary traits. However, although this scheme was followed by a number of college textbooks and some general classifications, it is now almost totally abandoned as a formal classification. Indeed, as late as 2010, one author of

1140-438: A particular order while still being able to move freely within the cavity. Most bilateral animals, including all the vertebrates , are coelomates. Pseudocoelomate animals have a pseudocoelom (literally "false cavity"), which is a fluid filled body cavity. Tissue derived from mesoderm partly lines the fluid filled body cavity of these animals. Thus, although organs are held in place loosely, they are not as well organized as in

1216-550: 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

1292-404: A process known as schizocoely . The archenteron initially forms, and the mesoderm splits into two layers: the first attaches to the body wall or ectoderm , forming the parietal layer and the second surrounds the endoderm or alimentary canal forming the visceral layer . The space between the parietal layer and the visceral layer is known as the coelom or body cavity. In deuterostomes ,

1368-589: A protein related to ubiquitin carboxyl-terminal hydrolase are exclusively shared by all vertebrates and reliably distinguish them from all other animals. A specific relationship between vertebrates and tunicates is strongly supported by two CSIs found in the proteins Rrp44 (associated with the exosome complex ) and serine palmitoyltransferase . These are exclusively shared by species from these two subphyla, but not cephalochordates . This indicates hat vertebrates are more closely related to tunicates than cephalochordates. The "Notochordata hypothesis" suggested that

1444-608: 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

1520-606: A series of crescentic openings from the pharynx to the outside. Each gill is supported by a cartilaginous or bony gill arch , which develop embryonically from pharyngeal arches . Bony fish have three pairs of gill arches, cartilaginous fish have five to seven pairs, while the primitive jawless fish have seven pairs. The ancestral vertebrates likely had more arches than seven, as some of their chordate relatives have more than 50 pairs of gill opens, although most, if not all, of these openings are actually involved in filter feeding rather than respiration . In jawed vertebrates ,

1596-630: 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

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1672-510: 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

1748-570: 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

1824-418: 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

1900-520: Is any invertebrate animal with a three-layered body and a pseudocoel . The coelom was apparently lost or reduced as a result of mutations in certain types of genes that affected early development. Thus, pseudocoelomates evolved from coelomates. "Pseudocoelomate" is no longer considered a valid taxonomic group , since it is not monophyletic . However, it is still used as a descriptive term. Important characteristics: Bilaterian pseudocoelomate phyla according to Brusca and Brusca,: Acoelomates lack

1976-666: 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

2052-448: 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

2128-440: Is found in invertebrate chordates such as lancelets (a sister subphylum known as the cephalochordates ), though it lacks eyes and other complex special sense organs comparable to those of vertebrates. Other chordates do not show any trends towards cephalization. The rostral end of the neural tube is expanded by a thickening of the walls and expansion of the central canal of spinal cord into three primary brain vesicles :

2204-635: The Cephalochordata is the sister taxon to Craniata (Vertebrata). This group, called the Notochordata, was placed as sister group to the Tunicata (Urochordata). Studies since 2006 analyzing large sequencing datasets however strongly support Olfactores (tunicates + vertebrates) as a clade, and hence the placement of Cephalochordata as sister-group to Olfactores (known as the " Olfactores hypothesis "). The following cladogram summarizes

2280-559: The Chengjiang biota and lived about 518 million years ago. These include Haikouichthys , Myllokunmingia , Zhongjianichthys , and probably Haikouella . Unlike the other fauna that dominated the Cambrian, these groups had the basic vertebrate body plan: a notochord, rudimentary vertebrae, and a well-defined head and tail. All of these early vertebrates lacked jaws in the common sense and relied on filter feeding close to

2356-759: 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

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2432-474: The Izu–Ogasawara Trench at a depth of 8,336 metres (27,349 feet). Many fish varieties are the main predators in most of the world's freshwater and marine water bodies . The rest of the vertebrate species are tetrapods, a single lineage that includes amphibians (with roughly 7,000 species); mammals (with approximately 5,500 species); and reptiles and birds (with about 20,000 species divided evenly between

2508-633: The Jurassic . The Cenozoic world saw great diversification of bony fishes, amphibians, reptiles, birds and mammals. Over half of all living vertebrate species (about 32,000 species) are fish (non-tetrapod craniates), a diverse set of lineages that inhabit all the world's aquatic ecosystems, from the Tibetan stone loach ( Triplophysa stolickai ) in western Tibetan hot springs near Longmu Lake at an elevation of 5,200 metres (17,100 feet) to an unknown species of snailfish (genus Pseudoliparis ) in

2584-464: The Latin vertebratus ("jointed"), from vertebra meaning "joint", in turn from Latin vertere to turn. All vertebrates are built along the basic chordate body plan of five synapomorphies . These are a rigid axial skeleton that includes a vertebral column developed around an elastic notochord . The notochord becomes the intervertebral discs , and runs dorsally to the gut tube along

2660-574: The Placodermi and the Acanthodii , both considered paraphyletic . Other ways of classifying the vertebrates have been devised, particularly with emphasis on the phylogeny of early amphibians and reptiles. An example based on Janvier (1981, 1997), Shu et al. (2003), and Benton (2004) is given here († = extinct ): While this traditional classification is orderly, most of the groups are paraphyletic , i.e. do not contain all descendants of

2736-399: The blue whale , at up to 33 m (108 ft). With only one exception, the defining characteristic of all vertebrates is the vertebral column , in which the embryonic notochord found in all chordates is replaced by a segmented series of mineralized elements called vertebrae separated by fibrocartilaginous intervertebral discs , which are embryonic and evolutionary remnants of

2812-565: The genetics of organisms. Phylogenetic classification is based solely on phylogeny . Evolutionary systematics gives an overview; phylogenetic systematics gives detail. The two systems are thus complementary rather than opposed. Conventional classification has extant vertebrates grouped into seven classes based on traditional interpretations of gross anatomical and physiological traits. The commonly held classification lists three classes of fish and four of tetrapods . In addition to these, there are two classes of extinct armoured fishes,

2888-501: 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)

2964-410: The prosencephalon ( forebrain ), mesencephalon ( midbrain ) and rhombencephalon ( hindbrain ), which are further differentiated in the various vertebrate groups. Two laterally placed retinas and optical nerves form around outgrowths from the midbrain, except in hagfish which may have secondarily lost the structures. The forebrain is more well-developed in most tetrapods and subdivided into

3040-432: The telencephalon and diencephalon , while the midbrain dominates in fish and some salamanders . In vertebrates with paired appendages, especially tetrapods, a pair of secondary enlargements of the hindbrain become the cerebella , which modulate complex motor coordinations . The brain vesicles are usually bilaterally symmetrical , giving rise to the paired cerebral hemispheres in mammals . The resultant anatomy of

3116-1031: 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"):

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3192-495: 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

3268-564: 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,   Vertebrate Ossea Batsch, 1788 Vertebrates ( / ˈ v ɜːr t ə b r ɪ t s , - ˌ b r eɪ t s / ) are animals with

3344-550: 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

3420-419: 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

3496-418: The axial endoskeleton (enclosed by protective skeletal extensions known as neural arches ), with a fore-end enlargement that is contained within a distinct skeletonized braincase (hence the alternative name for vertebrates, the craniates ). All vertebrate embryos develop transient pharyngeal arches , which in fish develop into the branchial arches that support the gills . Other vertebrate features are

3572-531: The body wall — this feature can be seen in the digestive tract of tardigrades (water bears) which is suspended within the body in the mesentery derived from a mesoderm-lined coelom. The fluid inside the coelom is known as coelomic fluid. This is circulated by mesothelial cilia or by contraction of muscles in the body wall. The coelomic fluid serves several functions: it acts as a hydroskeleton; it allows free movement and growth of internal organs; it serves for transport of gases, nutrients and waste products around

3648-431: The body; it allows storage of sperm and eggs during maturation; and it acts as a reservoir for waste. In the past, some zoologists grouped bilaterian animal phyla based on characteristics related to the coelom for practical purposes, knowing, and explicitly stating, that these groups were not phylogenetically related. Animals were classified in three informal groups according to the type of body cavity they possess, in

3724-427: The brain (which itself is a fused cluster of segmental ganglia from the rostral metameres ). Molecular markers known as conserved signature indels (CSIs) in protein sequences have been identified and provide distinguishing criteria for the vertebrate subphylum. Specifically, 5 CSIs in the following proteins: protein synthesis elongation factor-2 , eukaryotic translation initiation factor 3 , adenosine kinase and

3800-438: 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

3876-409: The class's common ancestor. For instance, descendants of the first reptiles include modern reptiles, mammals and birds; the agnathans have given rise to the jawed vertebrates ; the bony fishes have given rise to the land vertebrates ; the traditional " amphibians " have given rise to the reptiles (traditionally including the synapsids or mammal-like "reptiles"), which in turn have given rise to

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3952-487: The coelom forms by enterocoely . The archenteron wall produces buds of mesoderm , and these mesodermal diverticula hollow to become the coelomic cavities. Deuterostomes are therefore known as enterocoelomates . Examples of deuterostome coelomates belong to three major clades: chordates ( vertebrates , tunicates , and lancelets ), echinoderms ( starfish , sea urchins , sea cucumbers ), and hemichordates ( acorn worms and graptolites ). The evolutionary origin of

4028-464: The coelom is uncertain. The oldest known animal to have had a body cavity was the Vernanimalcula . Current hypothesis include: A coelom can absorb shock or provide a hydrostatic skeleton . It can also support an immune system in the form of coelomocytes that may either be attached to the wall of the coelom or may float about in it freely. The coelom allows muscles to grow independently of

4104-556: The demise of virtually all jawless fishes save for lampreys and hagfishes, as well as the Placodermi , a group of armoured fish that dominated the entirety of that period since the late Silurian as well as the eurypterids , dominant animals of the preceding Silurian , and the anomalocarids . By the middle of the Devonian, several droughts, anoxic events and oceanic competition led a lineage of sarcopterygii to leave water, eventually establishing themselves as terrestrial tetrapods in

4180-615: 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

4256-408: The dorsal nerve cord and migrate together with the mesodermal somites to innervate the various different structures that develop in the body. The vertebrates are the only chordate group with neural cephalization , and their neural functions are centralized towards a series of enlarged clusters in the head , which give rise to a brain . A slight swelling of the anterior end of the nerve cord

4332-483: The first gill arch pair evolved into the jointed jaws and form an additional oral cavity ahead of the pharynx . Research also suggests that the sixth branchial arch contributed to the formation of the vertebrate shoulder, which separated the head as a distinct part of the body. In amphibians and some primitive bony fishes, the larvae bear external gills , branching off from the gill arches. These are reduced in adulthood, their respiratory function taken over by

4408-673: 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

4484-421: The gill arches form during fetal development , and form the basis of essential structures such as jaws , the thyroid gland , the larynx , the columella (corresponding to the stapes in mammals ) and, in mammals, the malleus and incus . The central nervous system of vertebrates is based on the embryonic dorsal nerve cord (which then flattens into a neural plate before folding and fusing over into

4560-400: The gut tube, with a split brain stem circumventing the foregut around each side to form a brain on the dorsal side of the mouth . The higher functions of the vertebrate CNS are highly centralized towards the brain (particularly the forebrain), while the invertebrate CNS is significantly more decentralized with the segmental ganglia having substantial neural autonomy independent of

4636-561: 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

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4712-476: 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

4788-443: The internal gills proper in fishes and by cutaneous respiration in most amphibians. While some amphibians such as axolotl retain the external gills into adulthood, the complex internal gill system as seen in fish apparently being irrevocably lost very early in the evolution of tetrapods , who evolved lungs (which are homologous to swim bladders ) to breathe air. While the more specialized terrestrial vertebrates lack gills,

4864-407: The jawless Agnatha , and the jawed Gnathostomata . The jawed fish include both the cartilaginous fish and the bony fish . Bony fish include the lobe-finned fish , which gave rise to the tetrapods , the animals with four limbs. Vertebrates make up less than five percent of all described animal species ; the rest are all invertebrates , that lack a backbone. The word 'vertebrate' derives from

4940-432: The length of an animal, hence the common name of backbone. The axial endoskeleton typically continues beyond the anus / cloaca to form an elongated tail . Some vertebrates evolved to become tailless with only a vestigial coccyx . A dorsal nerve cord , which folds and fuses into a hollow neural tube during embryonic development and eventually gives rise to the brain and spinal cord , runs more dorsally to

5016-460: The mammals and birds. Most scientists working with vertebrates use a classification based purely on phylogeny , organized by their known evolutionary history and sometimes disregarding the conventional interpretations of their anatomy and physiology. In phylogenetics , the relationships between animals are illustrated as a hierarchy known as a phylogenetic tree . The cladogram below is based on studies compiled by Philippe Janvier and others for

5092-411: The notochord. Hagfish are the only extant vertebrate whose notochord persists and is not integrated/ replaced by the vertebral column. A few vertebrates have secondarily lost this feature and retain the notochord into adulthood, such as the sturgeon . Most vertebrates are aquatic and carry out gas exchange via gills . The gills are carried right behind the head, bordering the posterior margins of

5168-508: 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

5244-487: 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

5320-609: The relationships between the Olfactores (vertebrates and tunicates) and the Cephalochordata.   Amphioxiformes (lancelets)  [REDACTED]   Tunicata /Urochordata ( sea squirts , salps , larvaceans )  [REDACTED]   Vertebrata   [REDACTED] Vertebrates originated during the Cambrian explosion , which saw a rise in organism diversity. The earliest known vertebrates belongs to

5396-661: The seabed. A vertebrate group of uncertain phylogeny, small eel-like conodonts , are known from microfossils of their paired tooth segments from the late Cambrian to the end of the Triassic. The first jawed vertebrates may have appeared in the late Ordovician (~445 mya) and became common in the Devonian period , often known as the "Age of Fishes". The two groups of bony fishes , the Actinopterygii and Sarcopterygii , evolved and became common. The Devonian also saw

5472-650: The succeeding Carboniferous . Amniotes branched from amphibious tetrapods early in the Carboniferous period. The synapsid amniotes were dominant during the late Paleozoic, the Permian , while diapsid amniotes became dominant during the Mesozoic . In the sea, the teleosts and sharks became dominant. Mesothermic synapsids called cynodonts gave rise to endothermic mammals and diapsids called dinosaurs eventually gave rise to endothermic birds , both in

5548-452: The surface area to volume ratio is large enough to allow absorption of nutrients and gas exchange by diffusion alone, due to dorso-ventral flattening. A body cavity is also absent in placozoans , cnidarians ( jellyfish and allies) and the ctenophores (comb jellies), but these animals are neither bilaterians or triploblastic . Ancient Greek Ancient Greek ( Ἑλληνῐκή , Hellēnikḗ ; [hellɛːnikɛ́ː] ) includes

5624-517: 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

5700-402: The two classes). Tetrapods comprise the dominant megafauna of most terrestrial environments and also include many partially or fully aquatic groups (e.g., sea snakes , penguins , cetaceans). There are several ways of classifying animals. Evolutionary systematics relies on anatomy , physiology and evolutionary history, which is determined through similarities in anatomy and, if possible,

5776-480: 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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