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Scutigeromorpha

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68-1037: Scutigeromorpha is an order of centipedes also known as house centipedes . These centipedes are found in the temperate and tropical parts of every continent except Antarctica, with their distribution significantly expanded by the introduction of the Mediterranean species Scutigera coleoptrata throughout Europe, Asia, and North America. The common species S. coleoptrata is a typical representative of this order, lying in wait for other arthropods, then seizing prey using great speed, and all species in this order reflect adaptations for this mode of life. House centipedes are hemianamorphic , and adults in this order have 15 leg-bearing segments. Adults have bodies that measure 2 to 3.5 cm in length, but some can reach 8 cm in body length. Species in this order can be readily recognized by their long legs and antennae . The antennae end in annulated flagella that are each divided into two or three segments. The ultimate legs are much longer than

136-543: 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

204-476: A spermatophore for the female to take up. In temperate areas, egg laying occurs in spring and summer. A few parthenogenetic species are known. Females provide parental care, both by curling their bodies around eggs and young, and by grooming them, probably to remove fungi and bacteria. Centipedes are longer-lived than insects; the European Lithobius forficatus may live for 5 to 6 years, and

272-560: A broad range of prey. Common prey items include lumbricid earthworms, dipteran fly larvae, collembolans , and other centipedes. They are carnivorous; study of gut contents suggests that plant material is an unimportant part of their diets, although they eat vegetable matter when starved during laboratory experiments. Species of Scolopendromorph , noticeably members from the genera Scolopendra and Ethmostigmus , are able to hunt for substantial prey items, including large invertebrates and sizable vertebrates, which could be larger than

340-516: A food item, certain large centipedes are consumed in China , usually skewered and grilled or deep fried. They are often seen in street vendors’ stalls in large cities, including Donghuamen and Wangfujing markets in Beijing . Large centipedes are steeped in alcohol to make centipede vodka. Some species of centipedes can be hazardous to humans because of their bite. While a bite to an adult human

408-477: 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

476-419: 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

544-430: A moist microhabitat because they lack the waxy cuticle of insects and arachnids , therefore causing them to rapidly lose water. Accordingly, they avoid direct sunlight by staying under cover or by being active at night . Centipedes have a rounded or flattened head, bearing a pair of antennae at the forward margin. They have a pair of elongated mandibles , and two pairs of maxillae . The first pair of maxillae form

612-550: 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

680-451: A role in mating rituals. Because glandular pores occur more frequently on ultimate legs than on the "walking" legs, they may serve a sensory role. They are sometimes used in defensive postures, and some species use them to capture prey, defend themselves against predators, or suspend themselves from objects such as branches, using the legs as pincers. Several species use their ultimate legs upon encountering another centipede, trying to grab

748-608: A separate historical stage, though its earliest form closely resembles Attic Greek , and its latest form approaches Medieval Greek . There were several regional dialects of Ancient Greek; Attic Greek developed into Koine. Ancient Greek was a pluricentric language , divided into many dialects. The main dialect groups are Attic and Ionic , Aeolic , Arcadocypriot , and Doric , many of them with several subdivisions. Some dialects are found in standardized literary forms in literature , while others are attested only in inscriptions. There are also several historical forms. Homeric Greek

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816-628: A simple tube, with digestive glands attached to the mouthparts. Like insects, centipedes breathe through a tracheal system , typically with a single opening, or spiracle , on each body segment. They excrete waste through a single pair of malpighian tubules . Just as the first pair of legs are modified into forcipules, the back legs are modified into "ultimate legs", also called anal legs, caudal legs, and terminal legs. Their use varies between species, but does not include locomotion. The ultimate legs may be elongated and thin, thickened, or pincer-like. They are frequently sexually dimorphic , and may play

884-630: 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

952-415: A variable number of ocelli , sometimes clustered together to form true compound eyes . However, these eyes are only capable of discerning light from dark, and provide no true vision. In some species, the first pair of legs can function as sensory organs, similar to antennae; unlike the antennae of most other invertebrates, these point backwards. An unusual clustering of sensory organs found in some centipedes

1020-510: 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

1088-403: Is anamorphic : more segments and pairs of legs are grown between moults . For example, Scutigera coleoptrata , the house centipede, hatches with only four pairs of legs and in successive moults has 5, 7, 9, 11, 15, 15, 15 and 15 pairs respectively, before becoming a sexually mature adult. Life stages with fewer than 15 pairs of legs are called larval stadia (there are about five stages). After

1156-570: 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

1224-418: 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

1292-666: 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

1360-448: 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

1428-501: Is epimorphic, meaning that all pairs of legs are developed in the embryonic stages, and offspring do not develop more legs between moults. This clade contains the longest centipedes. In the Geophilomorpha, the number of thoracic segments usually varies within species, often on a geographical basis, and in most cases, females bear more legs than males. The number of leg-bearing segments varies within each order (usually 21 or 23 in

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1496-556: 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

1564-415: Is the organ of Tömösváry . The organs, at the base of the antennae, consist of a disc-like structure and a central pore, with an encircling of sensitive cells. They are likely used for sensing vibrations, and may provide a weak form of hearing. Forcipules are unique to centipedes. The forcipules are modifications of the first pair of legs (the maxillipeds), forming a pincer -like appendage , just behind

1632-483: Is usually very painful and may cause severe swelling, chills, fever , and weakness, it is unlikely to be fatal. Bites can be dangerous to small children and those with allergies to bee stings. The venomous bite of larger centipedes can induce anaphylactic shock in such people. Smaller centipedes are generally incapable of piercing human skin. Even small centipedes that cannot pierce human skin are considered frightening by some humans due to their dozens of legs moving at

1700-517: The Cretaceous . The following cladogram shows the position of the Chilopoda within the arthropods as of 2019: inc. horseshoe crabs and spiders [REDACTED] [REDACTED] Chilopoda [REDACTED] millipedes and allies [REDACTED] inc. crustaceans and insects [REDACTED] [REDACTED] Within the myriapods, centipedes are believed to be the first of

1768-759: 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

1836-629: The IUCN Red List , there are one vulnerable , six endangered , and three critically endangered species of centipede. For example, the Serpent Island centipede ( Scolopendra abnormis) is vulnerable, and Turk's earth centipede (Nothogeophilus turki) and the Seychelles long-legged centipede ( Seychellonema gerlachi) are both endangered. The fossil record of centipedes extends back to 430  million years ago , during

1904-676: The Late Silurian ( Crussolum ), though they are rare throughout the Paleozoic . The Devonian Panther Mountain Formation contains two species of centipede. One is a species of the scutigeromorph Crussolum . The other is Devonobius , which is included in the extinct group Devonobiomorpha. Another Devonian site, the Rhynie chert , also bears Crussolum fossils, and possible scutigeromorph head material. Rhyniognatha , which

1972-501: 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)

2040-1031: 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"):

2108-495: 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

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2176-531: The Scolopendromorpha; from 27 to 191 in the Geophilomorpha), but the total number of pairs is always odd, so there are never exactly 100 legs or 100 pairs, despite the group's common name. Centipede segments are developed in two phases. Firstly, the head gives rise to a fixed but odd number of segments, driven by Hox genes as in all arthropods. Secondly, pairs of segments are added at

2244-550: 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

2312-419: 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

2380-415: The body of the other centipede. Members of the genus Alipes can stridulate their leaf-like ultimate legs to distract or threaten predators. Rhysida immarginata togoensis makes a faint creaking sound when it swings its ultimate legs. There are many differences between millipedes and centipedes. Both groups of myriapods have long, multi-segmented bodies, many legs, a single pair of antennae, and

2448-438: 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

2516-575: The class Chilopoda ( Ancient Greek χεῖλος , kheilos , "lip", and Neo-Latin suffix -poda , "foot", describing the forcipules ) of the subphylum Myriapoda , an arthropod group which includes millipedes and other multi-legged animals. Centipedes are elongated segmented ( metameric ) creatures with one pair of legs per body segment. All centipedes are venomous and can inflict painful stings , injecting their venom through pincer-like appendages known as forcipules or toxicognaths, which are actually modified legs instead of fangs . Despite

2584-549: The crown group of Scutigeromorpha to at least the Early Cretaceous . There are 88 species and 27 genera of Scutigeromorph centipedes distributed among three families: Pselliodidae , Scutigeridae , and Scutigerinidae . Most scutigeromorph species belong to the largest family, Scutigeridae, with a cosmopolitan distribution. The family Scutigeridae includes two subfamilies: Scutigerinae and Thereuoneminae. The small family Pselliodidae contains several species and includes

2652-615: 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

2720-618: The extant classes to branch from the last common ancestor . The five orders of centipedes are: Craterostigmomorpha , Geophilomorpha , Lithobiomorpha , Scolopendromorpha , and Scutigeromorpha . These orders are united into the clade Chilopoda by the following synapomorphies : The Chilopoda are then split into two clades: the Notostigmophora including the Scutigeromorpha and the Pleurostigmophora including

2788-472: The final two segments are small and legless. Each pair of legs is slightly longer than the pair preceding them, ensuring that they do not overlap, which reduces the chance that they will collide and trip the animal. The last pair of legs may be as much as twice the length of the first pair. The final segment bears a telson , and includes the openings of the reproductive organs. Centipedes mainly use their antennae to seek out their prey. The digestive tract forms

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2856-409: The first 50 million years of the clade's evolutionary history, centipede venoms appear to have consisted of a simple cocktail of about four different components, and differentiation into specific venom types appears to have only occurred after the currently recognized five orders had developed. The evolution of the venom includes horizontal gene transfer , involving bacteria, fungi and oomycetes . As

2924-510: The fourth covers segments 7, 8, and 9, and the remaining three each cover two segments per tergite. An especially striking feature that distinguishes this order from all other centipedes is the arrangement of spiracles . In this order, spiracles are arranged in a series down the middle of the centipede's back, whereas spiracles in all other centipedes are on the sides of their bodies. This distinction places this order in its own subclass, Notostigmomorpha , whereas all other centipedes are placed in

2992-431: The full complement of legs is achieved, the now postlarval stadia (about five more stages) develop gonopods, sensory pores, more antennal segments, and more ocelli. All mature lithobiomorph centipedes have 15 leg-bearing segments. The Craterostigmomorpha only have one phase of anamorphosis, with embryos having 12 pairs, and adults 15. The clade Epimorpha, consisting of the orders Geophilomorpha and Scolopendromorpha ,

3060-518: The genus Sphendononema , found in the Neotropics and tropical Africa. The smallest family, Scutigerinidae, contains only a few species and includes the genera Scutigerina and Madagassophora , found in South Africa and Madagascar. Centipede Centipedes (from Neo-Latin centi- , "hundred", and Latin pes, pedis , " foot ") are predatory arthropods belonging to

3128-402: The head. Forcipules are not oral mouthparts , though they are used to subdue prey by injecting venom and gripping the prey animal. Venom glands run through a tube, from inside the head to the tip of each forcipule. Behind the head, the body consists of at least fifteen segments. Most of the segments bear a single pair of legs; the maxillipeds project forward from the first body segment, while

3196-561: 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

3264-476: 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

3332-454: The lower lip, and bear short palps . The first pair of limbs stretch forward from the body over the mouth. These limbs, or forcipules, end in sharp claws and include venom glands that help the animal to kill or paralyze its prey. Their size ranges from a few millimetres in the smaller lithobiomorphs and geophilomorphs to about 30 cm (12 in) in the largest scolopendromorphs . Many species of centipedes lack eyes, but some possess

3400-623: The myriapod itself. For instance, Scolopendra gigantea (the Amazonian giant centipede) preys on tarantulas , scorpions , lizards , frogs , birds , mice , snakes , and even bats , catching them in midflight. Three species ( Scolopendra cataracta , S. paradoxa , and S. alcyona ) are amphibious, believed to hunt aquatic or amphibious invertebrates. Many larger animals prey upon centipedes, such as mongooses , mice , salamanders , beetles and some specialist snake species. They form an important item of diet for many species and

3468-425: The name, no species of centipede has exactly 100 legs; the number of pairs of legs is an odd number that ranges from 15 pairs to 191 pairs. Centipedes are predominantly generalist carnivorous , hunting for a variety of prey items that can be overpowered. They have a wide geographical range, which can be found in terrestrial habitats from tropical rainforests to deserts . Within these habitats, centipedes require

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3536-508: 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

3604-460: The order Scutigeromorpha based on the morphology of their legs and their maxillipeds . These Paleozoic fossils date the crown group of Chilopoda to at least the late Silurian , 418 million years ago. These fossils place scutigeromorphs among the earliest terrestrial arthropods. These older fossils differ enough from extant species to be assigned to the scutigeromorph stem group , but younger fossils may be similar enough to extant species to date

3672-641: The other four orders. The following physical and developmental traits can be used to separate members of the Pleurostigmomorpha from Notostigmomorpha: It was previously believed that Chilopoda was split into Anamorpha (Lithobiomorpha and Scutigeromorpha) and Epimorpha (Geophilomorpha and Scolopendromorpha), based on developmental modes, with the relationship of the Craterostigmomorpha being uncertain. Recent phylogenetic analyses using combined molecular and morphological characters supports

3740-432: The others and resemble the antennae. These centipedes also feature compound eyes , which are divided into ommatidia with crystalline cones. The trunk of these centipedes features eight tergites : The first trunk segment corresponds to a short first tergite, then seven large tergites follow. The first large tergite covers the second trunk segment, but the second covers segments 3 and 4, the third covers segments 5 and 6,

3808-487: 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

3876-450: The presence of postantennal organs . Centipedes have one pair of legs per segment, while millipedes have two. Their heads differ in that millipedes have short, elbowed antennae , a pair of robust mandibles and a single pair of maxillae fused into a lip; centipedes have long, threadlike antennae, a pair of small mandibles, two pairs of maxillae and a pair of large venom claws. Centipede reproduction does not involve copulation. Males deposit

3944-456: The previous phylogeny. The Epimorpha still exist as a monophyletic group within the Pleurostigmophora, but the Anamorpha are paraphyletic , as shown in the cladogram: Scutigeromorpha [REDACTED] Lithobiomorpha [REDACTED] Craterostigmomorpha [REDACTED] Scolopendromorpha [REDACTED] Geophilomorpha [REDACTED] All centipedes are venomous . Over

4012-445: The same time and their tendency to dart swiftly out of the darkness towards one's feet. A 19th-century Tibetan poet warned his fellow Buddhists , "if you enjoy frightening others, you will be reborn as a centipede." Ancient Greek Ancient Greek ( Ἑλληνῐκή , Hellēnikḗ ; [hellɛːnikɛ́ː] ) includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It

4080-447: The secretions of Geophilus vittatus are sticky and odorous, and contain hydrogen cyanide. The giant desert centipede of Arizona, Scolopendra polymorpha , has a black head and tail, and an orange body; this conspicuous pattern may be aposematic , an honest signal of the animal's toxicity. Many species raise and splay their ultimate legs and display the spines found on the legs in a defensive threat posture. Because centipedes lack

4148-576: The staple diet of some such as the African ant Amblyopone pluto , which feeds solely on geophilomorph centipedes, and the South African Cape black-headed snake Aparallactus capensis . Some Geophilomorph, Lithobiomorph, and Scolopendromorph centipedes produce sticky, toxic secretions to defend themselves. The various secretions ward off or entangle predators. Scolopendromorph secretions contain hydrogen cyanide . Among Geophilomorphs,

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4216-401: The subclass Pleurostigmomorpha . Centipedes in this order have a single spiracle opening through a slit at the posterior of each of the seven large tergites. The dorsal placement of spiracles reflects the adaptation of species in this order to life in the open rather than under stones or bark, where other centipedes live. The earliest known fossil centipedes have been identified as belonging to

4284-517: 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

4352-636: The tail (posterior) end by the creation of a prepattern unit, a double segment, which is then always divided into two. The repeated creation of these prepattern units is driven by an oscillator clock, implemented with the Notch signalling pathway . The segments are homologous with the legs of other arthropods such as trilobites ; it would be sufficient for the Notch clock to run faster , as it does in snakes , to create more legs. Centipedes are predominantly generalist predators, which means they are adapted to eat

4420-510: The waxy water-resistant cuticle of other arthropods, they are more susceptible to water loss via evaporation. Thus, centipedes are most commonly found in high-humidity environments to avoid dehydration, and are mostly nocturnal . Centipedes live in many different habitats including in soil and leaf litter; they are found in environments as varied as tropical rain forests, deserts, and caves. Some geophilomorphs are adapted to littoral habitats, where they feed on barnacles . According to

4488-451: The wide-ranging Scolopendra subspinipes can live for over 10 years. The combination of a small number of eggs laid, long gestation period, and long time of development to reproduction has led authors to label lithobiomorph centipedes as K-selected . Centipedes grow their legs at different points in their development. In the primitive condition, seen in the orders Lithobiomorpha , Scutigeromorpha , and Craterostigmomorpha , development

4556-480: 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 ,

4624-582: Was once thought to be the oldest insect fossil, is also found in the Rhynie Chert. Three species, one scutigeromorph ( Latzelia ) and two scolopendromorphs ( Mazoscolopendra and the poorly known Palenarthrus ), have been described from the Mazon Creek fossil beds , which are Carboniferous , 309–307 mya. More species appear in the Mesozoic , including scolopendromorphs and scutigeromorphs in

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