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Ichthyostega

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102-550: Ichthyostega (from Greek : ἰχθῦς ikhthûs , 'fish' and Greek : στέγη stégē , 'roof') is an extinct genus of limbed tetrapodomorphs from the Late Devonian of what is now Greenland . It was among the earliest four-limbed vertebrates ever in the fossil record and was one of the first with weight-bearing adaptations for terrestrial locomotion. Ichthyostega possessed lungs and limbs that helped it navigate through shallow water in swamps. Although Ichthyostega

204-577: A basis for coinages: anthropology , photography , telephony , isomer , biomechanics , cinematography , etc. Together with Latin words , they form the foundation of international scientific and technical vocabulary ; for example, all words ending in -logy ('discourse'). There are many English words of Greek origin . Greek is an independent branch of the Indo-European language family. The ancient language most closely related to it may be ancient Macedonian , which, by most accounts,

306-430: A different reason. Unlike dolphins, these fish do not feel the bubbles, because they have bony fins without nerve endings. Nevertheless, they cannot swim faster because the cavitation bubbles create a vapor film around their fins that limits their speed. Lesions have been found on tuna that are consistent with cavitation damage. Scombrid fishes (tuna, mackerel and bonito) are particularly high-performance swimmers. Along

408-558: A fairly stable set of consonantal contrasts . The main phonological changes occurred during the Hellenistic and Roman period (see Koine Greek phonology for details): In all its stages, the morphology of Greek shows an extensive set of productive derivational affixes , a limited but productive system of compounding and a rich inflectional system. Although its morphological categories have been fairly stable over time, morphological changes are present throughout, particularly in

510-560: A faster, more convenient cursive writing style with the use of ink and quill . The Greek alphabet consists of 24 letters, each with an uppercase ( majuscule ) and lowercase ( minuscule ) form. The letter sigma has an additional lowercase form (ς) used in the final position of a word: In addition to the letters, the Greek alphabet features a number of diacritical signs : three different accent marks ( acute , grave , and circumflex ), originally denoting different shapes of pitch accent on

612-540: A foreign language. It is also often stated that the historical changes have been relatively slight compared with some other languages. According to one estimation, " Homeric Greek is probably closer to Demotic than 12-century Middle English is to modern spoken English ". Greek is spoken today by at least 13 million people, principally in Greece and Cyprus along with a sizable Greek-speaking minority in Albania near

714-442: A homocercal tail. These come in a variety of shapes, and can appear: (D) - Diphycercal means the vertebrae extend to the tip of the tail and the tail is symmetrical and expanded (as in the bichir , lungfish , lamprey , coelacanths and † Tarrasiiformes ). Most Palaeozoic fishes had a diphycercal heterocercal tail. Finlets are small fins, generally behind the dorsal and anal fins (in bichirs , there are only finlets on

816-515: A liquid, which then promptly and violently collapse. It can cause significant damage and wear. Cavitation damage can occur to the tail fins of powerful swimming marine animals, such as dolphins and tuna. Cavitation is more likely to occur near the surface of the ocean, where the ambient water pressure is relatively low. Even if they have the power to swim faster, dolphins may have to restrict their speed because collapsing cavitation bubbles on their tail are too painful. Cavitation also slows tuna, but for

918-508: A modified fin to deliver sperm; thresher sharks use their caudal fin to whip and stun prey; reef stonefish have spines in their dorsal fins that inject venom as an anti-predator defense ; anglerfish use the first spine of their dorsal fin like a fishing rod to lure prey; and triggerfish avoid predators by squeezing into coral crevices and using spines in their fins to anchor themselves in place. Fins can either be paired or unpaired . The pectoral and pelvic fins are paired, whereas

1020-439: A more disproportionate way than other fins on female fish." There are two prevailing hypotheses that have been historically debated as models for the evolution of paired fins in fish: the gill arch theory and the lateral fin-fold theory. The former, commonly referred to as the " Gegenbaur hypothesis ," was posited in 1870 and proposes that the "paired fins are derived from gill structures". This fell out of popularity in favor of

1122-416: A neural network in the fin, indicating that it likely has a sensory function, but are still not sure exactly what the consequences of removing it are. A comparative study in 2013 indicates the adipose fin can develop in two different ways. One is the salmoniform-type way, where the adipose fin develops from the larval-fin fold at the same time and in the same direct manner as the other median fins. The other

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1224-705: A pancake, and will fit into fissures in rocks. Their pelvic and pectoral fins have evolved differently, so they act together with the flattened body to optimise manoeuvrability. Some fishes, such as puffer fish , filefish and trunkfish , rely on pectoral fins for swimming and hardly use tail fins at all. Male cartilaginous fishes (sharks and rays), as well as the males of some live-bearing ray finned fishes , have fins that have been modified to function as intromittent organs , reproductive appendages which allow internal fertilization . In ray finned fish, they are called gonopodia or andropodia , and in cartilaginous fish, they are called claspers . Gonopodia are found on

1326-549: A prominent dorsal fin. Like scombroids and other billfish , they streamline themselves by retracting their dorsal fins into a groove in their body when they swim. The huge dorsal fin, or sail, of the sailfish is kept retracted most of the time. Sailfish raise them if they want to herd a school of small fish, and also after periods of high activity, presumably to cool down. The oriental flying gurnard has large pectoral fins which it normally holds against its body, and expands when threatened to scare predators. Despite its name, it

1428-433: A specialized ear, including a stapes with a unique morphology compared to other tetrapods or to any fish hyomandibula. The legs were large compared to contemporary relatives. It had seven digits on each hind leg, along with a peculiar, poorly ossified mass which lies anteriorly adjacent to the digits. The exact number of digits on the forelimb is not yet known, since fossils of the hand have not been found. While in water,

1530-454: A thin stretch of scaleless skin ; in lobe-finned fish ( Sarcopterygii ) such as coelacanths and lungfish , fins are short rays based around a muscular central bud supported by jointed bones ; in cartilaginous fish ( Chondrichthyes ) and jawless fish ( Agnatha ), fins are fleshy " flippers " supported by a cartilaginous skeleton. Fins at different locations of the fish body serve different purposes, and are divided into two groups:

1632-413: Is a demersal fish , not a flying fish, and uses its pelvic fins to walk along the bottom of the ocean. Fins can have an adaptive significance as sexual ornaments. During courtship, the female cichlid , Pelvicachromis taeniatus , displays a large and visually arresting purple pelvic fin . "The researchers found that males clearly preferred females with a larger pelvic fin and that pelvic fins grew in

1734-804: Is also found in Bulgaria near the Greek-Bulgarian border. Greek is also spoken worldwide by the sizable Greek diaspora which has notable communities in the United States , Australia , Canada , South Africa , Chile , Brazil , Argentina , Russia , Ukraine , the United Kingdom , and throughout the European Union , especially in Germany . Historically, significant Greek-speaking communities and regions were found throughout

1836-773: Is an Indo-European language, constituting an independent Hellenic branch within the Indo-European language family. It is native to Greece , Cyprus , Italy (in Calabria and Salento ), southern Albania , and other regions of the Balkans , Caucasus , the Black Sea coast, Asia Minor , and the Eastern Mediterranean . It has the longest documented history of any Indo-European language, spanning at least 3,400 years of written records. Its writing system

1938-479: Is an Indo-European language, but also includes a number of borrowings from the languages of the populations that inhabited Greece before the arrival of Proto-Greeks, some documented in Mycenaean texts ; they include a large number of Greek toponyms . The form and meaning of many words have changed. Loanwords (words of foreign origin) have entered the language, mainly from Latin, Venetian , and Turkish . During

2040-643: Is now East Greenland. Ichthyostega' s skull seems more fish-like than that of Acanthostega , but its pelvic girdle morphology seems stronger and better adapted to life on land. Ichthyostega also had more supportive ribs and stronger vertebrae with more developed zygapophyses . Whether or not these traits were independently evolved in Ichthyostega is debated. It does, however, show that Ichthyostega may have ventured onto land on occasions, unlike contemporaneous limbed vertebrates, such as Elginerpeton and Obruchevichthys . Traditionally, Ichthyostega

2142-400: Is often labelled a ' tetrapod ' because of its limbs and fingers, it evolved long before true crown group tetrapods and could more accurately be referred to as a stegocephalian or stem tetrapod . Likewise, while undoubtedly of amphibian build and habit, it is not a true member of the group in the narrow sense, as the first modern amphibians (members of the group Lissamphibia ) appeared in

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2244-628: Is one type of living lobe-finned fish. Both extant members of this group, the West Indian Ocean coelacanth ( Latimeria chalumnae ) and the Indonesian coelacanth ( Latimeria menadoensis ), are found in the genus Latimeria . Coelacanths are thought to have evolved roughly into their current form about 408 million years ago, during the early Devonian. Locomotion of the coelacanths is unique to their kind. To move around, coelacanths most commonly take advantage of up or downwellings of

2346-436: Is prohibited in many countries. Foil shaped fins generate thrust when moved, the lift of the fin sets water or air in motion and pushes the fin in the opposite direction. Aquatic animals get significant thrust by moving fins back and forth in water. Often the tail fin is used, but some aquatic animals generate thrust from pectoral fins . Cavitation occurs when negative pressure causes bubbles (cavities) to form in

2448-519: Is protected and promoted officially as a regional and minority language in Armenia, Hungary , Romania, and Ukraine. It is recognized as a minority language and protected in Turkey by the 1923 Treaty of Lausanne . The phonology , morphology , syntax , and vocabulary of the language show both conservative and innovative tendencies across the entire attestation of the language from the ancient to

2550-432: Is relatively conservative in lobe-finned fishes. However, there are a few examples from the fossil record that show aberrant morphologies , such as Allenypterus , Rebellatrix , Foreyia or the tetrapodomorphs . Ray-finned fishes form a class of bony fishes called Actinopterygii. Their fins contain spines or rays. A fin may contain only spiny rays, only soft rays, or a combination of both. If both are present,

2652-402: Is something of a mystery. It is frequently clipped off to mark hatchery-raised fish, though data from 2005 showed that trout with their adipose fin removed have an 8% higher tailbeat frequency. Additional information released in 2011 has suggested that the fin may be vital for the detection of, and response to, stimuli such as touch, sound and changes in pressure. Canadian researchers identified

2754-841: Is sometimes called aljamiado , as when Romance languages are written in the Arabic alphabet. Article 1 of the Universal Declaration of Human Rights in Greek: Transcription of the example text into Latin alphabet : Article 1 of the Universal Declaration of Human Rights in English: Proto-Greek Mycenaean Ancient Koine Medieval Modern Fin ray Fins are moving appendages protruding from

2856-526: Is spoken by at least 13.5 million people today in Greece, Cyprus, Italy, Albania, Turkey , and the many other countries of the Greek diaspora . Greek roots have been widely used for centuries and continue to be widely used to coin new words in other languages; Greek and Latin are the predominant sources of international scientific vocabulary . Greek has been spoken in the Balkan peninsula since around

2958-763: Is still used internationally for the writing of Ancient Greek . In Greek, the question mark is written as the English semicolon, while the functions of the colon and semicolon are performed by a raised point (•), known as the ano teleia ( άνω τελεία ). In Greek the comma also functions as a silent letter in a handful of Greek words, principally distinguishing ό,τι ( ó,ti , 'whatever') from ότι ( óti , 'that'). Ancient Greek texts often used scriptio continua ('continuous writing'), which means that ancient authors and scribes would write word after word with no spaces or punctuation between words to differentiate or mark boundaries. Boustrophedon , or bi-directional text,

3060-650: Is the Greek alphabet , which has been used for approximately 2,800 years; previously, Greek was recorded in writing systems such as Linear B and the Cypriot syllabary . The alphabet arose from the Phoenician script and was in turn the basis of the Latin , Cyrillic , Coptic , Gothic , and many other writing systems. The Greek language holds a very important place in the history of the Western world. Beginning with

3162-487: Is the characiform-type way, where the adipose fin develops late after the larval-fin fold has diminished and the other median fins have developed. They claim the existence of the characiform-type of development suggests the adipose fin is not "just a larval fin fold remainder" and is inconsistent with the view that the adipose fin lacks function. Research published in 2014 indicates that the adipose fin has evolved repeatedly in separate lineages . (A) - Heterocercal means

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3264-433: Is the heterocercal tail, which aids in locomotion. Most sharks have eight fins. Sharks can only drift away from objects directly in front of them because their fins do not allow them to move in the tail-first direction. Unlike modern cartilaginous fish, members of stem chondrichthyan lineages (e.g. the † climatiids and the † diplacanthids ) possessed pectoral dermal plates as well as dermal spines associated with

3366-400: Is the largest class of vertebrates in existence today, making up more than 50% of species. In the distant past, lobe-finned fish were abundant; however, there are currently only 8 species. Bony fish have fin spines called lepidotrichia or "rays" (due to how the spines spread open). They typically have swim bladders , which allow the fish to alter the relative density of its body and thus

3468-598: Is thought that their rostral organ helps give the coelacanth electroperception, which aids in their movement around obstacles. Lungfish are also living lobe-finned fish. They occur in Africa ( Protopterus ), Australia ( Neoceratodus ), and South America ( Lepidosiren ). Lungfish evolved during the Devonian Period. Genetic studies and paleontological data confirm that lungfish are the closest living relatives of land vertebrates . Fin arrangement and body shape

3570-575: The Danish East Greenland Expedition . Additional specimens were collected between 1933 and 1955. Ichthyostega was a fairly large animal for its time, as it was broadly built and about 1.5 m (4.9 ft) long. The skull was low, with dorsally placed eyes and large labyrinthodont teeth. The posterior margin of the skull formed an operculum covering the gills. The spiracle was situated in an otic notch behind each eye. Computed tomography has revealed that Ichthyostega had

3672-495: The Eastern Mediterranean , in what are today Southern Italy , Turkey , Cyprus , Syria , Lebanon , Israel , Palestine , Egypt , and Libya ; in the area of the Black Sea , in what are today Turkey, Bulgaria , Romania , Ukraine , Russia , Georgia , Armenia , and Azerbaijan ; and, to a lesser extent, in the Western Mediterranean in and around colonies such as Massalia , Monoikos , and Mainake . It

3774-584: The Humane Society International , approximately 100 million sharks are killed each year for their fins, in an act known as shark finning . After the fins are cut off, the mutilated sharks are thrown back in the water and left to die. In some countries of Asia , shark fins are a culinary delicacy, such as shark fin soup . Currently, international concerns over the sustainability and welfare of sharks have impacted consumption and availability of shark fin soup worldwide. Shark finning

3876-516: The Indo-Iranian languages (see Graeco-Aryan ), but little definitive evidence has been found. In addition, Albanian has also been considered somewhat related to Greek and Armenian, and it has been proposed that they all form a higher-order subgroup along with other extinct languages of the ancient Balkans; this higher-order subgroup is usually termed Palaeo-Balkan , and Greek has a central position in it. Linear B , attested as early as

3978-484: The Middle Triassic † Saurichthys , the oldest known example of viviparity in a ray-finned fish. Claspers are found on the males of cartilaginous fishes . They are the posterior part of the pelvic fins that have also been modified to function as intromittent organs, and are used to channel semen into the female's cloaca during copulation. The act of mating in sharks usually includes raising one of

4080-515: The Triassic Period. Until finds of other early stegocephalians and closely related fishes in the late 20th century, Ichthyostega stood alone as a transitional fossil between fish and tetrapods, combining fish and tetrapod features. Newer research has shown that it had an unusual anatomy, functioning more akin to a seal than a salamander , as previously assumed. In 1932 Gunnar Säve-Söderbergh described four Ichthyostega species from

4182-417: The buoyancy , so it can sink or float without having to use the fins to swim up and down. However, swim bladders are absent in many fish, most notably in lungfishes , who have evolved their swim bladders into primitive lungs , which may have a shared evolutionary origin with those of their terrestrial relatives, the tetrapods . Bony fishes also have a pair of opercula that function to draw water across

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4284-409: The extinct † Petalodontiformes (e.g. † Belantsea , † Janassa , † Menaspis ), which belong to Holocephali (ratfish and their fossil relatives), or in † Aquilolamna ( Selachimorpha ) and † Squatinactis (Squatinactiformes). Some cartilaginous fishes have an eel-like locomotion (e.g. Chlamydoselachus , † Thrinacoselache , † Phoebodus ) According to

4386-426: The gills , which help them breathe without needing to swim forward to force the water into the mouth across the gills. Lobe-finned fishes form a class of bony fishes called Sarcopterygii. They have fleshy, lobed , paired fins, which are joined to the body by a series of bones. The fins of lobe-finned fish differ from those of all other fish in that each is borne on a fleshy, lobe-like, scaly stalk extending from

4488-831: The midsagittal unpaired fins and the more laterally located paired fins . Unpaired fins are predominantly associated with generating linear acceleration via oscillating propulsion , as well as providing directional stability ; while paired fins are used for generating paddling acceleration , deceleration, and differential thrust or lift for turning , surfacing or diving and rolling . Fins can also be used for other locomotions other than swimming, for example, flying fish use pectoral fins for gliding flight above water surface, and frogfish and many amphibious fishes use pectoral and/or pelvic fins for crawling . Fins can also be used for other purposes: remoras and gobies have evolved sucker -like dorsal fins for attaching to surfaces and "hitchhiking"; male sharks and mosquitofish use

4590-492: The nominal and verbal systems. The major change in the nominal morphology since the classical stage was the disuse of the dative case (its functions being largely taken over by the genitive ). The verbal system has lost the infinitive , the synthetically -formed future, and perfect tenses and the optative mood . Many have been replaced by periphrastic ( analytical ) forms. Pronouns show distinctions in person (1st, 2nd, and 3rd), number (singular, dual , and plural in

4692-471: The ventral portion. This is because the shark's vertebral column extends into that dorsal portion, providing a greater surface area for muscle attachment. This allows more efficient locomotion among these negatively buoyant cartilaginous fish. By contrast, most bony fish possess a homocercal caudal fin. Tiger sharks have a large upper lobe , which allows for slow cruising and sudden bursts of speed. The tiger shark must be able to twist and turn in

4794-470: The 3rd millennium BC, or possibly earlier. The earliest written evidence is a Linear B clay tablet found in Messenia that dates to between 1450 and 1350 BC, making Greek the world's oldest recorded living language . Among the Indo-European languages, its date of earliest written attestation is matched only by the now-extinct Anatolian languages . The Greek language is conventionally divided into

4896-486: The Greek alphabet since approximately the 9th century BC. It was created by modifying the Phoenician alphabet , with the innovation of adopting certain letters to represent the vowels. The variant of the alphabet in use today is essentially the late Ionic variant, introduced for writing classical Attic in 403 BC. In classical Greek, as in classical Latin, only upper-case letters existed. The lower-case Greek letters were developed much later by medieval scribes to permit

4998-425: The Greek language are often emphasized. Although Greek has undergone morphological and phonological changes comparable to those seen in other languages, never since classical antiquity has its cultural, literary, and orthographic tradition been interrupted to the extent that one can speak of a new language emerging. Greek speakers today still tend to regard literary works of ancient Greek as part of their own rather than

5100-495: The Greek language was the Cypriot syllabary (also a descendant of Linear A via the intermediate Cypro-Minoan syllabary ), which is closely related to Linear B but uses somewhat different syllabic conventions to represent phoneme sequences. The Cypriot syllabary is attested in Cyprus from the 11th century BC until its gradual abandonment in the late Classical period, in favor of the standard Greek alphabet. Greek has been written in

5202-629: The Greek verb have likewise remained largely the same over the course of the language's history but with significant changes in the number of distinctions within each category and their morphological expression. Greek verbs have synthetic inflectional forms for: Many aspects of the syntax of Greek have remained constant: verbs agree with their subject only, the use of the surviving cases is largely intact (nominative for subjects and predicates, accusative for objects of most verbs and many prepositions, genitive for possessors), articles precede nouns, adpositions are largely prepositional, relative clauses follow

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5304-685: The Greek-Albanian border. A significant percentage of Albania's population has knowledge of the Greek language due in part to the Albanian wave of immigration to Greece in the 1980s and '90s and the Greek community in the country. Prior to the Greco-Turkish War and the resulting population exchange in 1923 a very large population of Greek-speakers also existed in Turkey , though very few remain today. A small Greek-speaking community

5406-470: The Late Devonian of East Greenland and one species belonging to the genus Ichthyostegopsis , I. wimani . These species could be synonymous (in which case only I. stensioei would remain), because their morphological differences are not very pronounced. The species differ in skull proportions, skull punctuation and skull bone patterns. The comparisons were done on 14 specimens collected in 1931 by

5508-434: The ability to lock their spines outwards. Triggerfish also use spines to lock themselves in crevices to prevent them being pulled out. Lepidotrichia are usually composed of bone , but those of early osteichthyans - such as Cheirolepis - also had dentine and enamel . They are segmented and appear as a series of disks stacked one on top of another. They may have been derived from dermal scales. The genetic basis for

5610-404: The acute during the late 20th century, and it has only been retained in typography . After the writing reform of 1982, most diacritics are no longer used. Since then, Greek has been written mostly in the simplified monotonic orthography (or monotonic system), which employs only the acute accent and the diaeresis. The traditional system, now called the polytonic orthography (or polytonic system),

5712-402: The ancient language; singular and plural alone in later stages), and gender (masculine, feminine, and neuter), and decline for case (from six cases in the earliest forms attested to four in the modern language). Nouns, articles, and adjectives show all the distinctions except for a person. Both attributive and predicative adjectives agree with the noun. The inflectional categories of

5814-427: The body of fish that interact with water to generate thrust and help the fish swim . Apart from the tail or caudal fin , fish fins have no direct connection with the back bone and are supported only by muscles . Fish fins are distinctive anatomical features with varying structures among different clades : in ray-finned fish ( Actinopterygii ), fins are mainly composed of bony spines or rays covered by

5916-521: The body. Pectoral and pelvic fins have articulations resembling those of tetrapod limbs. These fins evolved into legs of the first tetrapod land vertebrates ( amphibians ) in the Devonian Period . Sarcopterygians also possess two dorsal fins with separate bases, as opposed to the single dorsal fin of most ray-finned fish (except some teleosts ). The caudal fin is either heterocercal (only fossil taxa ) or diphycercal. The coelacanth

6018-548: The claspers to allow water into a siphon through a specific orifice . The clasper is then inserted into the cloaca, where it opens like an umbrella to anchor its position. The siphon then begins to contract expelling water and sperm. Other uses of fins include walking and perching on the sea floor, gliding over water, cooling of body temperature, stunning of prey, display (scaring of predators, courtship), defence (venomous fin spines, locking between corals), luring of prey, and attachment structures. The Indo-Pacific sailfish has

6120-465: The current and drift. They use their paired fins to stabilize their movement through the water. While on the ocean floor their paired fins are not used for any kind of movement. Coelacanths can create thrust for quick starts by using their caudal fins. Due to the high number of fins they possess, coelacanths have high maneuverability and can orient their bodies in almost any direction in the water. They have been seen doing headstands and swimming belly up. It

6222-738: The dorsal surface and no dorsal fin). In some fish such as tuna or sauries , they are rayless, non-retractable, and found between the last dorsal and/or anal fin and the caudal fin. Bony fishes ( Actinopterygii and Sarcopterygii ) form a taxonomic group called Osteichthyes (or Euteleostomi , which includes also land vertebrates ); they have skeletons made of bone mostly, and can be contrasted with cartilaginous fishes (see below), which have skeletons made mainly of cartilage (except for their teeth , fin spines , and denticles ). Bony fishes are divided into ray-finned and lobe-finned fish . Most living fish are ray-finned, an extremely diverse and abundant group consisting of over 30,000 species . It

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6324-525: The dorsal, anal and caudal fins are unpaired and situated along the midline of the body. For every type of fin, there are a number of fish species in which this particular fin has been lost during evolution (e.g. pelvic fins in † Bobasatrania , caudal fin in ocean sunfish ). In some clades , additional unpaired fins were acquired during evolution (e.g. additional dorsal fins, adipose fin). In some † Acanthodii ("spiny sharks"), one or more pairs of "intermediate" or "prepelvic" spines are present between

6426-404: The early 19th century that was used for literary and official purposes in the newly formed Greek state. In 1976, Dimotiki was declared the official language of Greece, after having incorporated features of Katharevousa and thus giving birth to Standard Modern Greek , used today for all official purposes and in education . The historical unity and continuing identity between the various stages of

6528-618: The epics of Homer , ancient Greek literature includes many works of lasting importance in the European canon . Greek is also the language in which many of the foundational texts in science and philosophy were originally composed. The New Testament of the Christian Bible was also originally written in Greek. Together with the Latin texts and traditions of the Roman world , the Greek texts and Greek societies of antiquity constitute

6630-442: The female. The male shortly inserts the organ into the sex opening of the female, with hook-like adaptations that allow the fish to grip onto the female to ensure impregnation. If a female remains stationary and her partner contacts her vent with his gonopodium, she is fertilized. The sperm is preserved in the female's oviduct. This allows females to fertilize themselves at any time without further assistance from males. In some species,

6732-400: The first dorsal fin spine was modified, forming a spine-brush complex. As with most fish, the tails of sharks provide thrust, making speed and acceleration dependent on tail shape. Caudal fin shapes vary considerably between shark species, due to their evolution in separate environments. Sharks possess a heterocercal caudal fin in which the dorsal portion is usually noticeably larger than

6834-452: The following periods: In the modern era, the Greek language entered a state of diglossia : the coexistence of vernacular and archaizing written forms of the language. What came to be known as the Greek language question was a polarization between two competing varieties of Modern Greek: Dimotiki , the vernacular form of Modern Greek proper, and Katharevousa , meaning 'purified', a compromise between Dimotiki and Ancient Greek developed in

6936-430: The foot would have functioned like a fleshy paddle more than a fin. The vertebral column and ribcage of Ichthyostega was unusual and highly specialized relative to both its contemporaries and later tetrapods. The thoracic vertebrae at the front of the trunk and the short neck have tall neural spines that lean backwards. They attach to pointed ribs which increase in size and acquire prominent overlapping flanges . Past

7038-501: The formation of a linked chain of vortex rings" and that "the dorsal and anal fin wakes are rapidly entrained by the caudal fin wake, approximately within the timeframe of a subsequent tail beat". Once motion has been established, the motion itself can be controlled with the use of other fins. The bodies of reef fishes are often shaped differently from open water fishes . Open water fishes are usually built for speed, streamlined like torpedoes to minimise friction as they move through

7140-553: The formation of the fin rays is thought to be genes coded for the production of certain proteins. It has been suggested that the evolution of the tetrapod limb from lobe-finned fishes is related to the loss of these proteins. Cartilaginous fishes form a class of fishes called Chondrichthyes. They have skeletons made of cartilage rather than bone . The class includes sharks , rays and chimaeras . Shark fin skeletons are elongated and supported with soft and unsegmented rays named ceratotrichia, filaments of elastic protein resembling

7242-522: The gonopodium may be half the total body length. Occasionally the fin is too long to be used, as in the "lyretail" breeds of Xiphophorus helleri . Hormone treated females may develop gonopodia. These are useless for breeding. Similar organs with similar characteristics are found in other fishes, for example the andropodium in the Hemirhamphodon or in the Goodeidae or the gonopodium in

7344-466: The hips have unusually large ribs similar to those of the thoracic region. The caudal vertebrae have slender spines that lean backwards. The tail of Ichthyostega retained a low fin supported by bony lepidotrichia ( fin rays ). The tail fin was not as deep as in Acanthostega, and would have been less useful for swimming. Ichthyostega is related to Acanthostega gunnari , which is also from what

7446-410: The horny keratin in hair and feathers. Originally the pectoral and pelvic girdles, which do not contain any dermal elements, did not connect. In later forms, each pair of fins became ventrally connected in the middle when scapulocoracoid and puboischiadic bars evolved. In rays , the pectoral fins have connected to the head and are very flexible. One of the primary characteristics present in most sharks

7548-439: The infinitive entirely (employing a raft of new periphrastic constructions instead) and uses participles more restrictively. The loss of the dative led to a rise of prepositional indirect objects (and the use of the genitive to directly mark these as well). Ancient Greek tended to be verb-final, but neutral word order in the modern language is VSO or SVO. Modern Greek inherits most of its vocabulary from Ancient Greek, which in turn

7650-467: The late 15th century BC, was the first script used to write Greek. It is basically a syllabary , which was finally deciphered by Michael Ventris and John Chadwick in the 1950s (its precursor, Linear A , has not been deciphered and most likely encodes a non-Greek language). The language of the Linear B texts, Mycenaean Greek , is the earliest known form of Greek. Another similar system used to write

7752-443: The lateral fin-fold theory, first suggested in 1877, which proposes that paired fins budded from longitudinal, lateral folds along the epidermis just behind the gills. There is weak support for both hypotheses in the fossil record and in embryology. However, recent insights from developmental patterning have prompted reconsideration of both theories in order to better elucidate the origins of paired fins. Carl Gegenbaur 's concept of

7854-499: The males of some species in the Anablepidae and Poeciliidae families. They are anal fins that have been modified to function as movable intromittent organs and are used to impregnate females with milt during mating. The third, fourth and fifth rays of the male's anal fin are formed into a tube-like structure in which the sperm of the fish is ejected. When ready for mating, the gonopodium becomes erect and points forward towards

7956-458: The margin at the rear of their bodies is a line of small rayless, non-retractable fins, known as finlets . There has been much speculation about the function of these finlets. Research done in 2000 and 2001 by Nauen and Lauder indicated that "the finlets have a hydrodynamic effect on local flow during steady swimming" and that "the most posterior finlet is oriented to redirect flow into the developing tail vortex, which may increase thrust produced by

8058-640: The membership of Greece and Cyprus in the European Union, Greek is one of the organization's 24 official languages . Greek is recognized as a minority language in Albania, and used co-officially in some of its municipalities, in the districts of Gjirokastër and Sarandë . It is also an official minority language in the regions of Apulia and Calabria in Italy. In the framework of the European Charter for Regional or Minority Languages , Greek

8160-441: The modern period. The division into conventional periods is, as with all such periodizations, relatively arbitrary, especially because, in all periods, Ancient Greek has enjoyed high prestige, and the literate borrowed heavily from it. Across its history, the syllabic structure of Greek has varied little: Greek shows a mixed syllable structure, permitting complex syllabic onsets but very restricted codas. It has only oral vowels and

8262-399: The noun they modify and relative pronouns are clause-initial. However, the morphological changes also have their counterparts in the syntax, and there are also significant differences between the syntax of the ancient and that of the modern form of the language . Ancient Greek made great use of participial constructions and of constructions involving the infinitive, and the modern variety lacks

8364-536: The objects of study of the discipline of Classics . During antiquity , Greek was by far the most widely spoken lingua franca in the Mediterranean world . It eventually became the official language of the Byzantine Empire and developed into Medieval Greek . In its modern form , Greek is the official language of Greece and Cyprus and one of the 24 official languages of the European Union . It

8466-706: The older periods of Greek, loanwords into Greek acquired Greek inflections, thus leaving only a foreign root word. Modern borrowings (from the 20th century on), especially from French and English, are typically not inflected; other modern borrowings are derived from Albanian , South Slavic ( Macedonian / Bulgarian ) and Eastern Romance languages ( Aromanian and Megleno-Romanian ). Greek words have been widely borrowed into other languages, including English. Example words include: mathematics , physics , astronomy , democracy , philosophy , athletics , theatre, rhetoric , baptism , evangelist , etc. Moreover, Greek words and word elements continue to be productive as

8568-696: The paired fins. The oldest species demonstrating these features is the † acanthodian † Fanjingshania renovata from the lower Silurian ( Aeronian ) of China. Fanjingshania possess compound pectoral plates composed of dermal scales fused to a bony plate and fin spines formed entirely of bone. Fin spines associated with the dorsal fins are rare among extant cartilaginous fishes, but are present, for instance, in Heterodontus or Squalus . Dorsal fin spines are typically developed in many fossil groups, such as in † Hybodontiformes , † Ctenacanthiformes or † Xenacanthida . In † Stethacanthus ,

8670-403: The pectoral and pelvic fins, but these are not associated with fins. The pelvic fin assists the fish in going up or down through the water, turning sharply, and stopping quickly. The dorsal fins are located on the back. A fish can have up to three dorsal fins. The dorsal fins serve to protect the fish against rolling, and assist it in sudden turns and stops. The function of the adipose fin

8772-637: The required range of movement to push the body up and forward, probably allowing the animal to drag itself across flat land by synchronous (rather than alternate) "crutching" movements, much like that of a mudskipper or a seal . It was incapable of typical quadrupedal gaits as the forelimbs lacked the necessary rotary motion range. [REDACTED] [REDACTED] [REDACTED] [REDACTED] [REDACTED] [REDACTED] Greek language Greek ( Modern Greek : Ελληνικά , romanized :  Elliniká , [eliniˈka] ; Ancient Greek : Ἑλληνική , romanized :  Hellēnikḗ )

8874-425: The sixth or seventh flanged rib, the ribs abruptly decrease in size and lose their flanges. The lumbar vertebrae at the back of the trunk have strong muscle scars and neural spines which are bent forwards and decrease in size towards the hips. The sacral vertebrae above the hips have fan-shaped neural spines that transition from forward-leaning to backward-leaning as they approach the tail. The vertebrae right behind

8976-402: The spiny rays are always anterior . Spines are generally stiff and sharp. Rays are generally soft, flexible, segmented, and may be branched. This segmentation of rays is the main difference that separates them from spines; spines may be flexible in certain species, but they will never be segmented. Spines have a variety of uses. In catfish , they are used as a form of defense; many catfish have

9078-437: The stressed vowel; the so-called breathing marks ( rough and smooth breathing ), originally used to signal presence or absence of word-initial /h/; and the diaeresis , used to mark the full syllabic value of a vowel that would otherwise be read as part of a diphthong. These marks were introduced during the course of the Hellenistic period. Actual usage of the grave in handwriting saw a rapid decline in favor of uniform usage of

9180-552: The tail of swimming mackerel". Fish use multiple fins, so it is possible that a given fin can have a hydrodynamic interaction with another fin. In particular, the fins immediately upstream of the caudal (tail) fin may be proximate fins that can directly affect the flow dynamics at the caudal fin. In 2011, researchers using volumetric imaging techniques were able to generate "the first instantaneous three-dimensional views of wake structures as they are produced by freely swimming fishes". They found that "continuous tail beats resulted in

9282-450: The tip of the tail and the tail is symmetrical but not expanded (as in the first fishes and the cyclostomes , and a more primitive precursor in lancelets ) (C) - Homocercal where the fin usually appears superficially symmetric but in fact the vertebrae extend for a very short distance into the upper lobe of the fin. Homocercal caudal fins can, however, also appear asymmetric (e.g. blue flying fish ). Most modern fishes ( teleosts ) have

9384-419: The vertebrae extend into the upper lobe of the tail, often making it longer than the lower lobe (as in sharks , † Placodermi , most stem Actinopterygii , and sturgeons and paddlefish ). However, the external shape of heterocercal tail fins can also appear symmetric (e.g. † Birgeria , † Bobasatrania ). Heterocercal is the opposite of hypocercal (B) - Protocercal means the vertebrae extend to

9486-511: The water easily when hunting to support its varied diet, whereas the porbeagle shark , which hunts schooling fish such as mackerel and herring , has a large lower lobe to help it keep pace with its fast-swimming prey. Other tail adaptations help sharks catch prey more directly, such as the thresher shark 's usage of its powerful, elongated upper lobe to stun fish and squid. On the other hand, rays rely on their enlarged pectoral fins for propulsion. Similarly enlarged pectoral fins can be found in

9588-697: The water. Reef fish operate in the relatively confined spaces and complex underwater landscapes of coral reefs . For this manoeuvrability is more important than straight line speed, so coral reef fish have developed bodies which optimize their ability to dart and change direction. They outwit predators by dodging into fissures in the reef or playing hide and seek around coral heads. The pectoral and pelvic fins of many reef fish, such as butterflyfish , damselfish and angelfish , have evolved so they can act as brakes and allow complex manoeuvres. Many reef fish, such as butterflyfish , damselfish and angelfish , have evolved bodies which are deep and laterally compressed like

9690-529: The “Archipterygium” was introduced in 1876. It was described as a gill ray, or "joined cartilaginous stem," that extended from the gill arch. Additional rays arose from along the arch and from the central gill ray. Gegenbaur suggested a model of transformative homology – that all vertebrate paired fins and limbs were transformations of the Archipterygium. Based on this theory, paired appendages such as pectoral and pelvic fins would have differentiated from

9792-574: Was a distinct dialect of Greek itself. Aside from the Macedonian question, current consensus regards Phrygian as the closest relative of Greek, since they share a number of phonological, morphological and lexical isoglosses , with some being exclusive between them. Scholars have proposed a Graeco-Phrygian subgroup out of which Greek and Phrygian originated. Among living languages, some Indo-Europeanists suggest that Greek may be most closely related to Armenian (see Graeco-Armenian ) or

9894-568: Was also used as the official language of government and religion in the Christian Nubian kingdoms , for most of their history. Greek, in its modern form, is the official language of Greece, where it is spoken by almost the entire population. It is also the official language of Cyprus (nominally alongside Turkish ) and the British Overseas Territory of Akrotiri and Dhekelia (alongside English ). Because of

9996-557: Was also used in Ancient Greek. Greek has occasionally been written in the Latin script , especially in areas under Venetian rule or by Greek Catholics . The term Frankolevantinika / Φραγκολεβαντίνικα applies when the Latin script is used to write Greek in the cultural ambit of Catholicism (because Frankos / Φράγκος is an older Greek term for West-European dating to when most of (Roman Catholic Christian) West Europe

10098-1396: Was considered part of an order named for it, the " Ichthyostegalia ". However, this group represents a paraphyletic grade of primitive stem-tetrapods and is not used by many modern researchers. Phylogenetic analysis has shown Ichthyostega is intermediate between other primitive stegocephalian stem-tetrapods. The evolutionary tree of early stegocephalians below follows the results of one such analysis performed by Swartz in 2012. Panderichthys [REDACTED] Tiktaalik [REDACTED] Elpistostege Elginerpeton [REDACTED] Ventastega [REDACTED] Acanthostega [REDACTED] Ichthyostega [REDACTED] Whatcheeriidae [REDACTED] Colosteidae [REDACTED] Crassigyrinus [REDACTED] Baphetidae [REDACTED] Crown group Tetrapoda [REDACTED] Early limbed vertebrates like Ichthyostega and Acanthostega differed from earlier tetrapodomorphs such as Eusthenopteron or Panderichthys in their increased adaptations for life on land. Though tetrapodomorphs possessed lungs, they used gills as their primary means of discharging carbon dioxide . Tetrapodomorphs used their bodies and tails for locomotion and their fins for steering and braking; Ichthyostega may have used its forelimbs for locomotion on land and its tail for swimming. Its massive ribcage

10200-482: Was made up of overlapping ribs and the animal possessed a stronger skeletal structure, a largely fishlike spine, and forelimbs apparently powerful enough to pull the body from the water. These anatomical modifications may have evolved to handle the lack of buoyancy experienced on land. The hindlimbs were smaller than the forelimbs and unlikely to have borne full weight in an adult, while the broad, overlapping ribs would have inhibited side-to-side movements. The forelimbs had

10302-580: Was under the control of the Frankish Empire ). Frankochiotika / Φραγκοχιώτικα (meaning 'Catholic Chiot') alludes to the significant presence of Catholic missionaries based on the island of Chios . Additionally, the term Greeklish is often used when the Greek language is written in a Latin script in online communications. The Latin script is nowadays used by the Greek-speaking communities of Southern Italy . The Yevanic dialect

10404-673: Was written by Romaniote and Constantinopolitan Karaite Jews using the Hebrew Alphabet . In a tradition, that in modern time, has come to be known as Greek Aljamiado , some Greek Muslims from Crete wrote their Cretan Greek in the Arabic alphabet . The same happened among Epirote Muslims in Ioannina . This also happened among Arabic-speaking Byzantine rite Christians in the Levant ( Lebanon , Palestine, and Syria ). This usage

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