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96-532: The region's rocks were laid down in the Devonian period and are known as Rhenish Facies . This is a fossil-bearing sedimentary rock type consisting mainly of slate . The rocks underwent considerable folding during the Carboniferous period. The gorge was carved out during a much more recent uplift to leave the river contained within steep walls 200 m (660 ft) high, the most famous feature being

192-565: A Mid-Devonian cooling of around 5 °C (9 °F). The Late Devonian warmed to levels equivalent to the Early Devonian; while there is no corresponding increase in CO 2 concentrations, continental weathering increases (as predicted by warmer temperatures); further, a range of evidence, such as plant distribution, points to a Late Devonian warming. The climate would have affected the dominant organisms in reefs ; microbes would have been

288-461: A bundle from a small chamber connected to the spiracle . This type of tracheal system has almost certainly evolved from the book lungs, and indicates that the tracheae of arachnids are not homologous with those of insects. Further adaptations to terrestrial life are appendages modified for more efficient locomotion on land, internal fertilisation, special sensory organs, and water conservation enhanced by efficient excretory structures as well as

384-444: A clade called Arachnopulmonata was also well supported. Pseudoscorpiones may also belong here, as all six orders share the same ancient whole genome duplication , and analyses support pseudoscorpions as the sister group of scorpions. Genetic analysis has not yet been done for Ricinulei, Palpigradi, or Solifugae, but horseshoe crabs have gone through two whole genome duplications, which gives them five Hox clusters with 34 Hox genes ,

480-464: A county in southwestern England, where a controversial argument in the 1830s over the age and structure of the rocks found throughout the county was resolved by adding the Devonian Period to the geological timescale. The Great Devonian Controversy was a lengthy debate between Roderick Murchison , Adam Sedgwick and Henry De la Beche over the naming of the period. Murchison and Sedgwick won

576-630: A diverse ecosystem of reefs and marine life. Devonian marine deposits are particularly prevalent in the midwestern and northeastern United States. Devonian reefs also extended along the southeast edge of Laurussia, a coastline now corresponding to southern England , Belgium , and other mid-latitude areas of Europe. In the Early and Middle Devonian, the west coast of Laurussia was a passive margin with broad coastal waters, deep silty embayments, river deltas and estuaries, found today in Idaho and Nevada . In

672-476: A fourth pair usually appears when they moult into nymphs . However, mites are variable: as well as eight, there are adult mites with six or, like in Eriophyoidea , even four legs. While the adult males in some members of Podapolipidae have six legs, the adult females have only a single pair. Arachnids are further distinguished from insects by the fact they do not have antennae or wings . Their body

768-763: A low-latitude archipelago to the north of Gondwana. They were separated from the southern continent by an oceanic basin: the Paleo-Tethys . Although the western Paleo-Tethys Ocean had existed since the Cambrian, the eastern part only began to rift apart as late as the Silurian. This process accelerated in the Devonian. The eastern branch of the Paleo-Tethys was fully opened when South China and Annamia (a terrane equivalent to most of Indochina ), together as

864-529: A massive extinction event . ( See Late Devonian extinction ). Primitive arthropods co-evolved with this diversified terrestrial vegetation structure. The evolving co-dependence of insects and seed plants that characterized a recognizably modern world had its genesis in the Late Devonian Epoch. The development of soils and plant root systems probably led to changes in the speed and pattern of erosion and sediment deposition. The rapid evolution of

960-543: A string of small settlements has grown up along the banks. Constrained in size, many of these old towns retain a historic feel today. With increasing wealth, roughly 40 hilltop castles were built and the valley became a core region of the Holy Roman Empire . It was at the centre of the Thirty Years' War , which left many of the castles in ruins , a particular attraction for today's cruise ships which follow

1056-664: A terrestrial ecosystem that contained copious animals opened the way for the first vertebrates to seek terrestrial living. By the end of the Devonian, arthropods were solidly established on the land. The Late Devonian extinction is not a single event, but rather is a series of pulsed extinctions at the Givetian-Frasnian boundary, the Frasnian-Famennian boundary, and the Devonian-Carboniferous boundary. Together, these are considered one of

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1152-413: A thin membrane. Inside the pit, a small hair touches the underside of the membrane, and detects its motion. Slit sense organs are believed to be involved in proprioception , and possibly also hearing. Arachnids may have one or two gonads , which are located in the abdomen. The genital opening is usually located on the underside of the second abdominal segment. In most species, the male transfers sperm to

1248-427: A transverse fold of the ectoderm . The ancestors of modern arachnids probably had both types, but modern ones often lack one type or the other. The cornea of the eye also acts as a lens, and is continuous with the cuticle of the body. Beneath this is a transparent vitreous body, and then the retina and, if present, the tapetum. In most arachnids, the retina probably does not have enough light sensitive cells to allow

1344-541: A unified continent, detached from the northeastern sector of Gondwana. Nevertheless, they remained close enough to Gondwana that their Devonian fossils were more closely related to Australian species than to north Asian species. Other Asian terranes remained attached to Gondwana, including Sibumasu (western Indochina), Tibet, and the rest of the Cimmerian blocks. While the South China-Annamia continent

1440-602: A volcanically active region during the Devonian, as it continued to assimilate smaller island arcs. The island arcs of the region, such as the Balkhash-West Junggar Arc, exhibited biological endemism as a consequence of their location. Siberia was located just north of the equator as the largest landmass in the Northern Hemisphere. At the beginning of the Devonian, Siberia was inverted (upside down) relative to its modern orientation. Later in

1536-435: A waxy layer covering the cuticle. The excretory glands of arachnids include up to four pairs of coxal glands along the side of the prosoma, and one or two pairs of Malpighian tubules , emptying into the gut. Many arachnids have only one or the other type of excretory gland, although several do have both. The primary nitrogenous waste product in arachnids is guanine . Arachnid blood is variable in composition, depending on

1632-509: Is derived from the Greek word ἀράχνη ( aráchnē , 'spider'), from the myth of the hubristic human weaver Arachne , who was turned into a spider. Almost all adult arachnids have eight legs, unlike adult insects which all have six legs. However, arachnids also have two further pairs of appendages that have become adapted for feeding, defense, and sensory perception. The first pair, the chelicerae , serve in feeding and defense. The next pair,

1728-695: Is formally broken into Early, Middle and Late subdivisions. The rocks corresponding to those epochs are referred to as belonging to the Lower, Middle and Upper parts of the Devonian System. The Early Devonian lasted from 419.2 to 393.3 Ma. It began with the Lochkovian Stage 419.2 to 410.8 Ma, which was followed by the Pragian from 410.8 to 407.6 Ma and then by the Emsian , which lasted until

1824-578: Is good evidence that Rheic oceanic crust experienced intense subduction and metamorphism under Mexico and Central America. The closure of the eastern part of the Rheic Ocean is associated with the assemblage of central and southern Europe. In the early Paleozoic, much of Europe was still attached to Gondwana, including the terranes of Iberia , Armorica (France), Palaeo-Adria (the western Mediterranean area), Bohemia , Franconia , and Saxothuringia . These continental blocks, collectively known as

1920-430: Is observed in many of those plants. Some of the early land plants such as Drepanophycus likely spread by vegetative growth and spores. The earliest land plants such as Cooksonia consisted of leafless, dichotomous axes with terminal sporangia and were generally very short-statured, and grew hardly more than a few centimetres tall. Fossils of Armoricaphyton chateaupannense , about 400 million years old, represent

2016-403: Is organized into two tagmata , called the prosoma and opisthosoma , also referred to as the cephalothorax and abdomen . However, there are questions about the validity of the latter terms. While the term cephalothorax implies a fused cephalon (head) and thorax , there is currently neither fossil nor embryological evidence that arachnids ever had a separate thorax-like division. Likewise,

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2112-507: The cephalothorax . In some species the frontmost pair of legs has converted to a sensory function, while in others, different appendages can grow large enough to take on the appearance of extra pairs of legs. Almost all extant arachnids are terrestrial , living mainly on land. However, some inhabit freshwater environments and, with the exception of the pelagic zone , marine environments as well. They comprise over 110,000 named species , of which 51,000 are species of spiders. The term

2208-669: The Hadley cell and the Ferrel cell . In these near-deserts, the Old Red Sandstone sedimentary beds formed, made red by the oxidised iron ( hematite ) characteristic of drought conditions. The abundance of red sandstone on continental land also lends Laurussia the name "the Old Red Continent". For much of the Devonian, the majority of western Laurussia (North America) was covered by subtropical inland seas which hosted

2304-666: The Loreley . The gorge produces its own microclimate and has acted as a corridor for species not otherwise found in the region. Its slopes have long been terraced for agriculture, in particular viticulture which has good conditions on south-facing slopes. Most of the vineyards belong to the wine region Mittelrhein , but the southernmost parts of the Rhine Gorge fall in Rheingau and Nahe . The river has been an important trade route into central Europe since prehistoric times and

2400-525: The Ordovician period. Fishes , especially jawed fish , reached substantial diversity during this time, leading the Devonian to often be dubbed the Age of Fishes . The armored placoderms began dominating almost every known aquatic environment. In the oceans, cartilaginous fishes such as primitive sharks became more numerous than in the Silurian and Late Ordovician . Tetrapodomorphs , which include

2496-481: The Paleo-Tethys Ocean and Rheic Ocean . By the early Devonian, the continent Laurussia (also known as Euramerica ) was fully formed through the collision of the continents Laurentia (modern day North America) and Baltica (modern day northern and eastern Europe). The tectonic effects of this collision continued into the Devonian, producing a string of mountain ranges along the southeastern coast of

2592-509: The cladogram below. Including fossil taxa does not fundamentally alter this view, although it introduces some additional basal groups. Chelicerata (sea spiders, horseshoe crabs and arachnids ) [REDACTED] [REDACTED] [REDACTED] Myriapoda (centipedes, millipedes, and allies) [REDACTED] [REDACTED] Pancrustacea (crustaceans and hexapods) [REDACTED] [REDACTED] The extant chelicerates comprise two marine groups: Sea spiders and horseshoe crabs, and

2688-475: The cladoxylopsids and progymnosperm Archaeopteris . These tracheophytes were able to grow to large size on dry land because they had evolved the ability to biosynthesize lignin , which gave them physical rigidity and improved the effectiveness of their vascular system while giving them resistance to pathogens and herbivores. In Eifelian age, cladoxylopsid trees formed the first forests in Earth history. By

2784-424: The house dust mite , are also the only arachnids able to ingest solid food, which exposes them to internal parasites, although it is not unusual for spiders to eat their own silk. And one species of spider is mostly herbivorous. Scorpions, spiders and pseudoscorpions secrete venom from specialized glands to kill prey or defend themselves. Their venom also contains pre-digestive enzymes that helps breaking down

2880-453: The pedipalps , have been adapted for feeding, locomotion, and/or reproductive functions. In scorpions, pseudoscorpions, and ricinuleids the pedipalps end in a pair of pinchers, while in whip scorpions, Schizomida , Amblypygi , and most harvestmen, they are raptorial and used for prey capture. In Solifugae , the palps are quite leg-like, so that these animals appear to have ten legs. The larvae of mites and Ricinulei have only six legs;

2976-452: The " Big Five " mass extinctions in Earth's history. The Devonian extinction crisis primarily affected the marine community, and selectively affected shallow warm-water organisms rather than cool-water organisms. The most important group to be affected by this extinction event were the reef-builders of the great Devonian reef systems. Amongst the severely affected marine groups were the brachiopods, trilobites, ammonites, and acritarchs , and

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3072-453: The 'abdomen' of many arachnids contains organs atypical of an abdomen, such as a heart and respiratory organs. The cephalothorax is usually covered by a single, unsegmented carapace. The abdomen is segmented in the more primitive forms, but varying degrees of fusion between the segments occur in many groups. It is typically divided into a preabdomen and postabdomen, although this is only clearly visible in scorpions, and in some orders, such as

3168-646: The Armorican Terrane Assemblage, split away from Gondwana in the Silurian and drifted towards Laurussia through the Devonian. Their collision with Laurussia leads to the beginning of the Variscan Orogeny , a major mountain-building event which would escalate further in the Late Paleozoic. Franconia and Saxothuringia collided with Laurussia near the end of the Early Devonian, pinching out the easternmost Rheic Ocean. The rest of

3264-536: The Armorican terranes followed, and by the end of the Devonian they were fully connected with Laurussia. This sequence of rifting and collision events led to the successive creation and destruction of several small seaways, including the Rheno-Hercynian, Saxo-Thuringian, and Galicia-Moldanubian oceans. Their sediments were eventually compressed and completely buried as Gondwana fully collided with Laurussia in

3360-574: The Carboniferous. Sea levels in the Devonian were generally high. Marine faunas continued to be dominated by conodonts, bryozoans , diverse and abundant brachiopods , the enigmatic hederellids , microconchids , and corals . Lily-like crinoids (animals, their resemblance to flowers notwithstanding) were abundant, and trilobites were still fairly common. Bivalves became commonplace in deep water and outer shelf environments. The first ammonites also appeared during or slightly before

3456-612: The Carboniferous. In 19th-century texts the Devonian has been called the "Old Red Age", after the red and brown terrestrial deposits known in the United Kingdom as the Old Red Sandstone in which early fossil discoveries were found. Another common term is "Age of the Fishes", referring to the evolution of several major groups of fish that took place during the period. Older literature on the Anglo-Welsh basin divides it into

3552-639: The Devonian extinction events were caused by an asteroid impact. However, while there were Late Devonian collision events (see the Alamo bolide impact ), little evidence supports the existence of a large enough Devonian crater. Arachnid Arachnids are arthropods in the class Arachnida ( / ə ˈ r æ k n ɪ d ə / ) of the subphylum Chelicerata . Arachnida includes, among others, spiders , scorpions , ticks , mites , pseudoscorpions , harvestmen , camel spiders , whip spiders and vinegaroons . Adult arachnids have eight legs attached to

3648-408: The Devonian seas. The first abundant genus of cartilaginous fish, Cladoselache , appeared in the oceans during the Devonian Period. The great diversity of fish around at the time has led to the Devonian being given the name "The Age of Fishes" in popular culture. The Devonian saw significant expansion in the diversity of nektonic marine life driven by the abundance of planktonic microorganisms in

3744-650: The Downtonian, Dittonian, Breconian, and Farlovian stages, the latter three of which are placed in the Devonian. The Devonian has also erroneously been characterised as a "greenhouse age", due to sampling bias : most of the early Devonian-age discoveries came from the strata of western Europe and eastern North America , which at the time straddled the Equator as part of the supercontinent of Euramerica where fossil signatures of widespread reefs indicate tropical climates that were warm and moderately humid. In fact

3840-506: The Early Devonian and the existence of fossils such as Protichnites suggest that amphibious arthropods may have appeared as early as the Cambrian ). By far the largest land organism at the beginning of this period was the enigmatic Prototaxites , which was possibly the fruiting body of an enormous fungus, rolled liverwort mat, or another organism of uncertain affinities that stood more than 8 metres (26 ft) tall, and towered over

3936-700: The Late Devonian the tectonic situation had relaxed and much of South America was covered by shallow seas. These south polar seas hosted a distinctive brachiopod fauna, the Malvinokaffric Realm, which extended eastward to marginal areas now equivalent to South Africa and Antarctica. Malvinokaffric faunas even managed to approach the South Pole via a tongue of Panthalassa which extended into the Paraná Basin . The northern rim of Gondwana

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4032-491: The Late Devonian, an approaching volcanic island arc reached the steep slope of the continental shelf and began to uplift deep water deposits. This minor collision sparked the start of a mountain-building episode called the Antler orogeny , which extended into the Carboniferous. Mountain building could also be found in the far northeastern extent of the continent, as minor tropical island arcs and detached Baltic terranes re-join

4128-682: The Late Devonian. The Altai-Sayan region was shaken by volcanism in the Early and Middle Devonian, while Late Devonian magmatism was magnified further to produce the Vilyuy Traps , flood basalts which may have contributed to the Late Devonian Mass Extinction. The last major round of volcanism, the Yakutsk Large Igneous Province, continued into the Carboniferous to produce extensive kimberlite deposits. Similar volcanic activity also affected

4224-497: The Middle Devonian began, 393.3 Ma. During this time, the first ammonoids appeared, descending from bactritoid nautiloids . Ammonoids during this time period were simple and differed little from their nautiloid counterparts. These ammonoids belong to the order Agoniatitida , which in later epochs evolved to new ammonoid orders, for example Goniatitida and Clymeniida . This class of cephalopod molluscs would dominate

4320-640: The Silurian and Devonian, it decreased across the span of the Devonian, particularly during the Pragian, and that the overall diversity of nektonic taxa did not increase significantly during the Devonian compared to during other geologic periods, and was in fact higher during the intervals spanning from the Wenlock to the Lochkovian and from the Carboniferous to the Permian. The study's authors instead attribute

4416-436: The abdomen has no appendages. Like all arthropods, arachnids have an exoskeleton , and they also have an internal structure of cartilage -like tissue, called the endosternite , to which certain muscle groups are attached. The endosternite is even calcified in some Opiliones . Most arachnids lack extensor muscles in the distal joints of their appendages. Spiders and whip scorpions extend their limbs hydraulically using

4512-496: The ancestors of all four- limbed vertebrates (i.e. tetrapods ), began diverging from freshwater lobe-finned fish as their more robust and muscled pectoral and pelvic fins gradually evolved into forelimbs and hindlimbs , though they were not fully established for life on land until the Late Carboniferous . The first ammonites , a subclass of cephalopod molluscs , appeared. Trilobites , brachiopods and

4608-409: The arachnids has proven difficult as of March 2016 , with successive studies producing different results. A study in 2014, based on the largest set of molecular data to date, concluded that there were systematic conflicts in the phylogenetic information, particularly affecting the orders Acariformes , Parasitiformes and Pseudoscorpiones , which have had much faster evolutionary rates. Analyses of

4704-406: The beginning and end of which are marked with extinction events. This lasted until the end of the Devonian, 358.9 Ma. The Devonian was a relatively warm period, although significant glaciers may have existed during the Early and Middle Devonian. The temperature gradient from the equator to the poles was not as large as it is today. The weather was also very arid, mostly along the equator where it

4800-514: The best view being from one of a convoy of boats. Devonian The Devonian ( / d ə ˈ v oʊ n i . ən , d ɛ -/ də- VOH -nee-ən, deh- ) is a geologic period and system of the Paleozoic era during the Phanerozoic eon , spanning 60.3 million years from the end of the preceding Silurian period at 419.2 million years ago ( Ma ), to the beginning of

4896-466: The climate in the Devonian differed greatly during its epochs and between geographic regions. For example, during the Early Devonian, arid conditions were prevalent through much of the world including Siberia, Australia, North America, and China, but Africa and South America had a warm temperate climate . In the Late Devonian, by contrast, arid conditions were less prevalent across the world and temperate climates were more common. The Devonian Period

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4992-520: The continent. Deformed remnants of these mountains can still be found on Ellesmere Island and Svalbard . Many of the Devonian collisions in Laurussia produce both mountain chains and foreland basins , which are frequently fossiliferous. Gondwana was by far the largest continent on the planet. It was completely south of the equator, although the northeastern sector (now Australia) did reach tropical latitudes. The southwestern sector (now South America)

5088-741: The continent. In present-day eastern North America, the Acadian Orogeny continued to raise the Appalachian Mountains . Further east, the collision also extended the rise of the Caledonian Mountains of Great Britain and Scandinavia . As the Caledonian Orogeny wound down in the later part of the period, orogenic collapse facilitated a cluster of granite intrusions in Scotland. Most of Laurussia

5184-673: The continents. By the middle of the Devonian, several groups of vascular plants had evolved leaves and true roots , and by the end of the period the first seed -bearing plants ( pteridospermatophytes ) appeared. This rapid evolution and colonization process, which had begun during the Silurian, is known as the Silurian-Devonian Terrestrial Revolution . The earliest land animals , predominantly arthropods such as myriapods , arachnids and hexapods , also became well-established early in this period, after beginning their colonization of land at least from

5280-920: The data using sets of genes with different evolutionary rates produced mutually incompatible phylogenetic trees . The authors favoured relationships shown by more slowly evolving genes, which demonstrated the monophyly of Chelicerata, Euchelicerata and Arachnida, as well as of some clades within the arachnids. The diagram below summarizes their conclusions, based largely on the 200 most slowly evolving genes; dashed lines represent uncertain placements. Acariformes [REDACTED] Opiliones [REDACTED] Ricinulei [REDACTED] Solifugae [REDACTED] Parasitiformes [REDACTED] Pseudoscorpiones [REDACTED] Scorpiones [REDACTED] Araneae [REDACTED] Amblypygi [REDACTED] Uropygi (Thelyphonida s.s. ) [REDACTED] Tetrapulmonata , here consisting of Araneae , Amblypygi and Uropygi (Thelyphonida s.s. ) ( Schizomida

5376-622: The debate and named it the Devonian System. While the rock beds that define the start and end of the Devonian Period are well identified, the exact dates are uncertain. According to the International Commission on Stratigraphy , the Devonian extends from the end of the Silurian 419.2 Ma, to the beginning of the Carboniferous 358.9 Ma – in North America , at the beginning of the Mississippian subperiod of

5472-405: The early Devonian Period around 400  Ma. Bactritoids make their first appearance in the Early Devonian as well; their radiation, along with that of ammonoids, has been attributed by some authors to increased environmental stress resulting from decreasing oxygen levels in the deeper parts of the water column. Among vertebrates, jawless armored fish ( ostracoderms ) declined in diversity, while

5568-465: The ecosystems and completed the extinction of all calcite sponge reefs and placoderms. Devonian palaeogeography was dominated by the supercontinent Gondwana to the south, the small continent of Siberia to the north, and the medium-sized continent of Laurussia to the east. Major tectonic events include the closure of the Rheic Ocean , the separation of South China from Gondwana, and

5664-569: The end of the Devonian, the first seed-forming plants had appeared. This rapid appearance of many plant groups and growth forms has been referred to as the Devonian Explosion or the Silurian-Devonian Terrestrial Revolution. The 'greening' of the continents acted as a carbon sink , and atmospheric concentrations of carbon dioxide may have dropped. This may have cooled the climate and led to

5760-475: The environment necessary for certain early fish to develop such essential characteristics as well developed lungs and the ability to crawl out of the water and onto the land for short periods of time. Finally, the Late Devonian started with the Frasnian , 382.7 to 372.2 Ma, during which the first forests took shape on land. The first tetrapods appeared in the fossil record in the ensuing Famennian subdivision,

5856-419: The eyes to form a proper image. In addition to the eyes, almost all arachnids have two other types of sensory organs. The most important to most arachnids are the fine sensory hairs that cover the body and give the animal its sense of touch. These can be relatively simple, but many arachnids also possess more complex structures, called trichobothria . Finally, slit sense organs are slit-like pits covered with

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5952-554: The female in a package, or spermatophore . The males in harvestmen and some mites have a penis. Complex courtship rituals have evolved in many arachnids to ensure the safe delivery of the sperm to the female. Members of many orders exhibit sexual dimorphism. Arachnids usually lay yolky eggs , which hatch into immatures that resemble adults. Scorpions, however, are either ovoviviparous or viviparous , depending on species, and bear live young. Also some mites are ovoviviparous and viviparous, even if most lay eggs. In most arachnids only

6048-640: The females provide parental care, with harvestmen being one of the few exceptions. The phylogenetic relationships among the main subdivisions of arthropods have been the subject of considerable research and dispute for many years. A consensus emerged from about 2010 onwards, based on both morphological and molecular evidence; extant (living) arthropods are a monophyletic group and are divided into three main clades: chelicerates (including arachnids), pancrustaceans (the paraphyletic crustaceans plus insects and their allies), and myriapods (centipedes, millipedes and allies). The three groups are related as shown in

6144-489: The food. It extends through most of the body, and connects to a short sclerotised intestine and anus in the hind part of the abdomen. Arachnids have two kinds of eyes: the lateral and median ocelli . The lateral ocelli evolved from compound eyes and may have a tapetum , which enhances the ability to collect light. With the exception of scorpions, which can have up to five pairs of lateral ocelli, there are never more than three pairs present. The median ocelli develop from

6240-492: The free water column as well as high ecological competition in benthic habitats, which were extremely saturated; this diversification has been labeled the Devonian Nekton Revolution by many researchers. However, other researchers have questioned whether this revolution existed at all; a 2018 study found that although the proportion of biodiversity constituted by nekton increased across the boundary between

6336-418: The great coral reefs were still common during the Devonian. The Late Devonian extinction , which started about 375 Ma, severely affected marine life, killing off most of the reef systems, most of the jawless fish, half of all placoderms, and nearly all trilobites save for a few species of the order Proetida . The subsequent end-Devonian extinction , which occurred at around 359 Ma, further impacted

6432-744: The increased overall diversity of nekton in the Devonian to a broader, gradual trend of nektonic diversification across the entire Palaeozoic. A now-dry barrier reef, located in present-day Kimberley Basin of northwest Australia , once extended 350 km (220 mi), fringing a Devonian continent. Reefs are generally built by various carbonate -secreting organisms that can erect wave-resistant structures near sea level. Although modern reefs are constructed mainly by corals and calcareous algae , Devonian reefs were either microbial reefs built up mostly by autotrophic cyanobacteria or coral-stromatoporoid reefs built up by coral-like stromatoporoids and tabulate and rugose corals . Microbial reefs dominated under

6528-403: The jawed fish (gnathostomes) simultaneously increased in both the sea and fresh water . Armored placoderms were numerous during the early ages of the Devonian Period and became extinct in the Late Devonian, perhaps because of competition for food against the other fish species. Early cartilaginous ( Chondrichthyes ) and bony fishes ( Osteichthyes ) also become diverse and played a large role within

6624-575: The low, carpet-like vegetation during the early part of the Devonian. Also, the first possible fossils of insects appeared around 416  Ma, in the Early Devonian. Evidence for the earliest tetrapods takes the form of trace fossils in shallow lagoon environments within a marine carbonate platform/shelf during the Middle Devonian, although these traces have been questioned and an interpretation as fish feeding traces ( Piscichnus ) has been advanced. Many Early Devonian plants did not have true roots or leaves like extant plants, although vascular tissue

6720-549: The main reef-forming organisms in warm periods, with corals and stromatoporoid sponges taking the dominant role in cooler times. The warming at the end of the Devonian may even have contributed to the extinction of the stromatoporoids. At the terminus of the Devonian, Earth rapidly cooled into an icehouse , marking the beginning of the Late Paleozoic icehouse . The Devonian world involved many continents and ocean basins of various sizes. The largest continent, Gondwana ,

6816-709: The marine fauna until the beginning of the Mesozoic Era. The Middle Devonian comprised two subdivisions: first the Eifelian , which then gave way to the Givetian 387.7 Ma. During this time the jawless agnathan fishes began to decline in diversity in freshwater and marine environments partly due to drastic environmental changes and partly due to the increasing competition, predation, and diversity of jawed fishes . The shallow, warm, oxygen-depleted waters of Devonian inland lakes, surrounded by primitive plants, provided

6912-481: The mites, the abdominal sections are completely fused. A telson is present in scorpions, where it has been modified to a stinger, and into a flagellum in the Palpigradi , Schizomida (very short) and whip scorpions . At the base of the flagellum in the two latter groups there are glands which produce acetic acid as a chemical defense. Except for a pair of pectines in scorpions, and the spinnerets in spiders,

7008-486: The mode of respiration. Arachnids with an efficient tracheal system do not need to transport oxygen in the blood, and may have a reduced circulatory system. In scorpions and some spiders, however, the blood contains haemocyanin , a copper-based pigment with a similar function to haemoglobin in vertebrates. The heart is located in the forward part of the abdomen, and may or may not be segmented. Some mites have no heart at all. Arachnids are mostly carnivorous , feeding on

7104-416: The mouth. Behind the mouth is a muscular, sclerotised pharynx , which acts as a pump, sucking the food through the mouth and on into the oesophagus and stomach . In some arachnids, the oesophagus also acts as an additional pump. The stomach is tubular in shape, with multiple diverticula extending throughout the body. The stomach and its diverticula both produce digestive enzymes and absorb nutrients from

7200-411: The nearby microcontinent of Amuria (now Manchuria , Mongolia and their vicinities). Though certainly close to Siberia in the Devonian, the precise location of Amuria is uncertain due to contradictory paleomagnetic data. The Rheic Ocean, which separated Laurussia from Gondwana, was wide at the start of the Devonian, having formed after the drift of Avalonia away from Gondwana. It steadily shrunk as

7296-401: The oldest known plants with woody tissue. By the Middle Devonian, shrub-like forests of primitive plants existed: lycophytes , horsetails , ferns , and progymnosperms evolved. Most of these plants had true roots and leaves, and many were quite tall. The earliest-known trees appeared in the Middle Devonian. These included a lineage of lycopods and another arborescent, woody vascular plant,

7392-403: The period continued, as the two major continents approached near the equator in the early stages of the assembly of Pangaea . The closure of the Rheic Ocean began in the Devonian and continued into the Carboniferous. As the ocean narrowed, endemic marine faunas of Gondwana and Laurussia combined into a single tropical fauna. The history of the western Rheic Ocean is a subject of debate, but there

7488-472: The period it moved northwards and began to twist clockwise, though it was not near its modern location. Siberia approached the eastern edge of Laurussia as the Devonian progressed, but it was still separated by a seaway, the Ural Ocean . Although Siberia's margins were generally tectonically stable and ecologically productive, rifting and deep mantle plumes impacted the continent with flood basalts during

7584-457: The pre-digested bodies of insects and other small animals. But ticks, and many mites, are parasites, some of which are carriers of disease. The diet of mites also include tiny animals, fungi, plant juices and decomposing matter. Almost as varied is the diet of harvestmen , where we will find predators, decomposers and omnivores feeding on decaying plant and animal matter, droppings, animals and mushrooms. The harvestmen and some mites, such as

7680-460: The pressure of their hemolymph . Solifuges and some harvestmen extend their knees by the use of highly elastic thickenings in the joint cuticle. Scorpions, pseudoscorpions and some harvestmen have evolved muscles that extend two leg joints (the femur-patella and patella-tibia joints) at once. The equivalent joints of the pedipalps of scorpions though, are extended by elastic recoil. There are characteristics that are particularly important for

7776-407: The prey. The saliva of ticks contains anticoagulants and anticomplements, and several species produce a neurotoxin . Arachnids produce digestive enzymes in their stomachs, and use their pedipalps and chelicerae to pour them over their dead prey. The digestive juices rapidly turn the prey into a broth of nutrients, which the arachnid sucks into a pre-buccal cavity located immediately in front of

7872-523: The resulting expansion of the Paleo-Tethys Ocean . The Devonian experienced several major mountain-building events as Laurussia and Gondwana approached; these include the Acadian Orogeny in North America and the beginning of the Variscan Orogeny in Europe. These early collisions preceded the formation of the single supercontinent Pangaea in the Late Paleozoic. The period is named after Devon ,

7968-669: The river. At one time forming a border of France , in the 19th century the valley became part of Prussia and its landscape became the quintessential image of Germany . This part of the Rhine features strongly in folklore , such as a legendary castle on the Rhine being the setting for the opera Götterdämmerung . The annual Rhine in Flames festivals include spectacular firework displays at Sankt Goar in September and Koblenz in August,

8064-562: The succeeding Carboniferous period at 358.9 Ma. It is the fourth period of both the Paleozoic and the Phanerozoic. It is named after Devon , South West England , where rocks from this period were first studied. The first significant evolutionary radiation of life on land occurred during the Devonian, as free- sporing land plants ( pteridophytes ) began to spread across dry land , forming extensive coal forests which covered

8160-492: The terrestrial arachnids. These have been thought to be related as shown below. (Pycnogonida (sea spiders) may be excluded from the chelicerates, which are then identified as the group labelled "Euchelicerata". ) A 2019 analysis nests Xiphosura deeply within Arachnida. Pycnogonida (sea spiders) [REDACTED] Xiphosura (horseshoe crabs) [REDACTED] Arachnida [REDACTED] Discovering relationships within

8256-432: The terrestrial lifestyle of arachnids, such as internal respiratory surfaces in the form of tracheae , or modification of the book gill into a book lung , an internal series of vascular lamellae used for gas exchange with the air. While the tracheae are often individual systems of tubes, similar to those in insects, ricinuleids, pseudoscorpions, and some spiders possess sieve tracheae, in which several tubes arise in

8352-521: The warmer conditions of the early and late Devonian, while coral-stromatoporoid reefs dominated during the cooler middle Devonian. By the Devonian Period, life was well underway in its colonization of the land. The moss forests and bacterial and algal mats of the Silurian were joined early in the period by primitive rooted plants that created the first stable soils and harbored arthropods like mites , scorpions , trigonotarbids and myriapods (although arthropods appeared on land much earlier than in

8448-555: The world saw the disappearance of an estimated 96% of vertebrates like conodonts and bony fishes , and all of the ostracoderms and placoderms. Land plants as well as freshwater species, such as our tetrapod ancestors, were relatively unaffected by the Late Devonian extinction event (there is a counterargument that the Devonian extinctions nearly wiped out the tetrapods ). The reasons for the Late Devonian extinctions are still unknown, and all explanations remain speculative. Canadian paleontologist Digby McLaren suggested in 1969 that

8544-480: Was a time of great tectonic activity, as the major continents of Laurussia and Gondwana drew closer together. Sea levels were high worldwide, and much of the land lay under shallow seas, where tropical reef organisms lived. The enormous "world ocean", Panthalassa , occupied much of the Northern Hemisphere as well as wide swathes east of Gondwana and west of Laurussia. Other minor oceans were

8640-594: Was located entirely within the Southern Hemisphere . It corresponds to modern day South America , Africa , Australia , Antarctica , and India , as well as minor components of North America and Asia . The second-largest continent, Laurussia, was northwest of Gondwana, and corresponds to much of modern-day North America and Europe . Various smaller continents, microcontinents , and terranes were present east of Laurussia and north of Gondwana, corresponding to parts of Europe and Asia. The Devonian Period

8736-487: Was located south of the equator, but in the Devonian it moved northwards and began to rotate counterclockwise towards its modern position. While the most northern parts of the continent (such as Greenland and Ellesmere Island ) established tropical conditions, most of the continent was located within the natural dry zone along the Tropic of Capricorn , which (as nowadays) is a result of the convergence of two great air-masses,

8832-469: Was located to the far south, with Brazil situated near the South Pole . The northwestern edge of Gondwana was an active margin for much of the Devonian, and saw the accretion of many smaller land masses and island arcs. These include Chilenia , Cuyania , and Chaitenia , which now form much of Chile and Patagonia . These collisions were associated with volcanic activity and plutons , but by

8928-442: Was mostly a passive margin, hosting extensive marine deposits in areas such as northwest Africa and Tibet . The eastern margin, though warmer than the west, was equally active. Numerous mountain building events and granite and kimberlite intrusions affected areas equivalent to modern day eastern Australia , Tasmania , and Antarctica. Several island microcontinents (which would later coalesce into modern day Asia) stretched over

9024-498: Was not included in the study), received strong support. Somewhat unexpectedly, there was support for a clade comprising Opiliones , Ricinulei and Solifugae , a combination not found in most other studies. In early 2019, a molecular phylogenetic analysis placed the horseshoe crabs, Xiphosura , as the sister group to Ricinulei. It also grouped pseudoscorpions with mites and ticks, which the authors considered may be due to long branch attraction . The addition of Scorpiones to produce

9120-434: Was the driest. Reconstruction of tropical sea surface temperature from conodont apatite implies an average value of 30 °C (86 °F) in the Early Devonian. Early Devonian mean annual surface temperatures were approximately 16 °C. CO 2 levels dropped steeply throughout the Devonian Period. The newly evolved forests drew carbon out of the atmosphere, which were then buried into sediments. This may be reflected by

9216-577: Was the newest addition to the Asian microcontinents, it was not the first. North China and the Tarim Block (now northwesternmost China) were located westward and continued to drift northwards, powering over older oceanic crust in the process. Further west was a small ocean (the Turkestan Ocean), followed by the larger microcontinents of Kazakhstania , Siberia , and Amuria . Kazakhstania was

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