53-472: The Lyginopteridales are an extinct group of seed plants known from the Paleozoic . They were the first plant fossils to be described as pteridosperms (a polyphyletic group sometimes referred to as "seed ferns") and, thus, the group on which the concept of pteridosperms was first developed; they are the stratigraphically oldest-known pteridosperms, occurring first in late Devonian strata; and they have
106-406: A clade of gymnosperms , with the gnetophytes in or near the conifers. For example, one common proposed set of relationships is known as the gne-pine hypothesis and looks like: (flowering plants) [REDACTED] Cycads [REDACTED] Ginkgo [REDACTED] Pinaceae (the pine family) [REDACTED] Gnetophytes [REDACTED] other conifers [REDACTED] However,
159-495: A phanerogam (taxon Phanerogamae ) or a phaenogam (taxon Phaenogamae ), is any plant that produces seeds . It is a category of embryophyte (i.e. land plant) that includes most of the familiar land plants, including the flowering plants and the gymnosperms , but not ferns , mosses , or algae . The term phanerogam or phanerogamae is derived from the Greek φανερός ( phanerós ), meaning "visible", in contrast to
212-543: 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
265-405: A cupule (there are some lyginopteridalean ovules that have been reported without a cupule, but these may simply have been shed from the cupule before being fossilised). In Late Devonian and early Mississippian species, the cupule contained several ovules, but by Pennsylvanian times there was normally just a single ovule per cupule. A number of cases have been found of cupules occurring in clusters at
318-477: 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
371-531: A lagenostome, despite its function in pollen capture having been replaced by the micropyle. As with all seed-plants, the lyginopteridalean ovules had just one functional seed megaspore within the nucellus. In some, however, three other, aborted megaspores were still present that together with the functional megaspore represented the remains of what would have been the tetrad of megaspores in the ancestral pre-seed plants. Most if not all lyginopteridalean ovules were borne in an outer protective sheath of tissue known as
424-419: 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
477-441: A much less elongate, almost radial pad. The Mississippian-aged lyginopteridaleans tended to have a simple protostele usually surrounded by secondary wood, but in later forms there was a eustele with a central core of pith or mixed-pith. In most cases the amount of secondary wood was limited suggesting they were stems of scrambling or climbing plants, but some Mississippian-aged forms (e.g., Pitys ) had substantial secondary wood and
530-431: A pinnate manner similar to that seen in fern fronds. The ultimate segments (pinnules) of the fronds are mostly lobate or digitate. Various fossil genera are recognised for these fronds, distinguished on whether the frond was bipartite or quadripartite, whether there were pinnae attached to the main rachis below the main dichotomy, the general shape of the pinnules, and the surface markings on the rachises and stem reflecting
583-550: 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
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#1732790867072636-608: 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
689-630: 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
742-558: A system to guide the pollen to the seed. Runcaria was followed shortly after by plants with a more condensed cupule, such as Spermasporites and Moresnetia . Seed-bearing plants had diversified substantially by the Famennian , the last stage of the Devonian. Examples include Elkinsia , Xenotheca , Archaeosperma , " Hydrasperma ", Aglosperma , and Warsteinia . Some of these Devonian seeds are now classified within
795-510: 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
848-570: 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
901-418: 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
954-422: Is an integumented megasporangium surrounded by a cupule. The megasporangium bears an unopened distal extension protruding above the mutlilobed integument . It is suspected that the extension was involved in anemophilous (wind) pollination . Runcaria sheds new light on the sequence of character acquisition leading to the seed. Runcaria has all of the qualities of seed plants except for a solid seed coat and
1007-666: 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
1060-448: 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
1113-481: Is the flowering plants , also known as angiosperms or magnoliophytes, the largest and most diverse group of spermatophytes: In addition to the five living taxa listed above, the fossil record contains evidence of many extinct taxa of seed plants, among those: By the Triassic period, seed ferns had declined in ecological importance, and representatives of modern gymnosperm groups were abundant and dominant through
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#17327908670721166-708: 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
1219-709: The Late Permian , but subsequently became extinct. Most evidence of the Lyginopteridales suggests that they grew in tropical latitudes of the time, in North America, Europe and China. As the Lyginopteridales are the earliest-known gymnosperms, and the development of ovules was one of the key innovations that enabled seed plants eventually to dominate land vegetation, the evolution of lyginopteridalean ovules has attracted considerable interest from palaeobotanists. The most important work on these early ovules
1272-791: The Medullosales took over as the more important of the larger pteridosperms but the Lyginopteridales continued to flourish as climbing (lianescent) and scrambling plants. However, later in Middle Pennsylvanian times the Lyginopteridales went into serious decline, probably being out-competed by the Callistophytales that occupied similar ecological niches but had more sophisticated reproductive strategies. A few species continued into Late Pennsylvanian times, and in Cathaysia and east equatorial Gondwana they persisted into
1325-501: 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)
1378-1031: 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"):
1431-495: 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
1484-550: 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
1537-419: 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
1590-438: 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
1643-615: 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
Lyginopteridales - Misplaced Pages Continue
1696-552: The end of the Cretaceous , when the angiosperms radiated. A whole genome duplication event in the ancestor of seed plants occurred about 319 million years ago . This gave rise to a series of evolutionary changes that resulted in the origin of modern seed plants. A middle Devonian (385-million-year-old) precursor to seed plants from Belgium has been identified predating the earliest seed plants by about 20 million years. Runcaria , small and radially symmetrical,
1749-458: The ends of branching axes. Many palaeobotanists now interpret these clusters of cupulate organs as fertile fronds, in which the cupulate tissue was derived from the laminate part of the frond that surrounded the ovule and thereby provided added protection for it. Most lyginopteridalean ovules were radiospermic. The only notable exceptions were a distinctive group of Mississippian-age lyginopteridaleans that had platyspermic ovules and are referred to
1802-481: The foliage are the most commonly found fossilised remains of the Lyginopteridales. When found complete, the fronds always seem to have a main rachis that dichotomises in the lower (proximal) part. In some cases, the two branches each underwent a second dichotomy, resulting in what is termed a quadripartite frond, but in others there is just the main proximal dichotomy, resulting in a bipartite frond. The branches produced by these dichotomies then undergo further divisions in
1855-673: 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
1908-509: The fossil family the Eospermaceae. The Lyginopteridales produced small trilete pre-pollen that superficially resemble the spores of non-seed plants but with a fundamentally gymnosperm-like wall-structure. The pre-pollen was produced by sporangia that formed regular clusters (synangia). The stratigraphically older lyginopteridaleans had trusses of synangia borne on slender axes, which were attached to vegetative fronds; these are referred to
1961-418: The fossil genera Telangium if they are anatomically preserved or Telangiopsis if they occur as adpressions. The more primitive forms of Telangium had sporangial walls that were essentially uniform in thickness. In stratigraphically younger Telangium species, however, the side of each sporangium that formed the outer surface of the synangium tended to be thicker than the inwards-facing wall, suggesting that
2014-561: 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
2067-476: 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
2120-483: The most primitive features, most notably in the structure of their ovules. They probably evolved from a group of Late Devonian progymnosperms known as the Aneurophytales , which had large, compound frond-like leaves. The Lyginopteridales became the most abundant group of pteridosperms during Mississippian times, and included both trees and smaller plants. During early and most of middle Pennsylvanian times
2173-508: 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
Lyginopteridales - Misplaced Pages Continue
2226-540: The order Lyginopteridales . Seed-bearing plants are a clade within the vascular plants (tracheophytes). The spermatophytes were traditionally divided into angiosperms , or flowering plants, and gymnosperms , which includes the gnetophytes, cycads, ginkgo, and conifers. Older morphological studies believed in a close relationship between the gnetophytes and the angiosperms, in particular based on vessel elements . However, molecular studies (and some more recent morphological and fossil papers) have generally shown
2279-487: 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
2332-592: The relationships between these groups should not be considered settled. Other classifications group all the seed plants in a single division , with classes for the five groups: A more modern classification ranks these groups as separate divisions (sometimes under the Superdivision Spermatophyta ): Unassigned extinct spermatophyte orders, some of which qualify as "seed ferns": Ancient Greek language Ancient Greek ( Ἑλληνῐκή , Hellēnikḗ ; [hellɛːnikɛ́ː] ) includes
2385-812: The sclerotic tissue in the cortex. There is no consensus on the division of the Lyginopteridales into families, either in terms of whole organisms or as fossil families of particular plant organs. The most recent scheme, by Anderson et al. (2007) recognized 5 families based on ovulate structures. Some authors also recognise the Mariopteridaceae for the distinctive group of Pennsylvanian-aged lianescent plants with Mariopteris fronds, although details of their reproductive structures are unknown. Seed plants A seed plant or spermatophyte ( lit. ' seed plant ' ; from Ancient Greek σπέρματος ( spérmatos ) 'seed' and φυτόν (phytón) 'plant'), also known as
2438-450: The sporangia split to release the pre-pollen along this inner wall. In Pennsylvanian-age lyginopteridalean synangia, the sporangia were usually attached to a pad of tissue that was probably homologous to a pinnule on the vegetative fronds. For instance, Crossotheca synangia bore elongate, "epaulet"-shaped pads, that were arranged in a pinnate pattern in a frond that was wholly or partially fertile. The Feraxotheca synangium in contrast had
2491-517: 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
2544-484: The term "cryptogam" or " cryptogamae " (from Ancient Greek κρυπτός (kruptós) 'hidden'), together with the suffix γαμέω ( gaméō ), meaning "to marry". These terms distinguish those plants with hidden sexual organs (cryptogamae) from those with visible ones (phanerogamae). The extant spermatophytes form five divisions, the first four of which are classified as gymnosperms , plants that have unenclosed, "naked seeds": The fifth extant division
2597-480: 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 ,
2650-464: Was a cluster of sporangium-bearing axes, but where only one eventually retained its megasporangium, the others forming the surrounding pre-integument. Progressively, this sheath of axes became fused to form a continuous integument that surrounded and protected the nucellus, such as in Eurystoma and then Stamnostoma . These earliest ovules had the apical part of the nucellus exposed, from which there
2703-476: Was a projection known as a lagenostome (sometimes also called a salpinx) that facilitated capture of the pollen and directed it down to the pollen chamber above the megagametophyte. In later, Pennsylvanian-age ovules such as Lagenostoma the nucellus became almost entirely encased by and fused to the integument, leaving just the small distal opening in the integument known as the micropyle through which pollen passed. Nevertheless, most lyginopteridalean ovules retained
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#17327908670722756-468: Was by the British palaeobotanist Albert Long, based mainly on early Mississippian, anatomically preserved fossils from Scotland (UK). Some of the earliest ovules (e.g., Genomosperma ) consisted of a nucellus (the equivalent of the sporangium wall) surrounded by a sheath of slender axes known as a pre-integument. It is widely thought that this arrangement was derived from an ancestral condition where there
2809-489: Was probably the trunk of a large tree. The stele is surrounded by a zone of cortex, which in many genera contains bands of fibrous tissue. This fibrous tissue often results in distinctive markings on the surface of the stems even when preserved as adpressions and can help with their generic identification: Lyginopteris for instance shows a mesh-shaped patterning on the surface of the stems, whereas Heterangium has mainly transverse bars. As with most pteridosperms, fragments of
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