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146-423: Approximately 220 million years ago, during the late Triassic period, the supercontinent Pangaea began to break apart. The focus of the rifting began somewhere between where present-day eastern North America and north-western Africa were joined. As the rift began to separate from mainland North America, volcanic activity occurred, forming volcanoes and flood basalts . These flood basalts poured out over

292-630: A bolide impact, for which an impact crater containing Manicouagan Reservoir in Quebec , Canada , has been singled out. However, the Manicouagan impact melt has been dated to 214±1 Mya. The date of the Triassic-Jurassic boundary has also been more accurately fixed recently, at 201.4 Mya. Both dates are gaining accuracy by using more accurate forms of radiometric dating, in particular the decay of uranium to lead in zircons formed at time of

438-626: A cosmopolitan distribution . Coelacanths show their highest post- Devonian diversity in the Early Triassic . Ray-finned fishes (actinopterygians) went through a remarkable diversification in the beginning of the Triassic, leading to peak diversity during the Middle Triassic; however, the pattern of this diversification is still not well understood due to a taphonomic megabias . The first stem-group teleosts appeared during

584-569: A basin is any area that collects sediments. These "aborted rifts" (rifts that are tectonically inactive and no longer collecting sediments) extend from Alabama to Newfoundland . Along certain basins, rifting was not partial. Where full rifting occurred, the Atlantic Ocean was created. Along these rifts, magmatic activity never stopped, as shown by the ongoing eruption of lava along the Mid-Atlantic Ridge . The Fundy Basin

730-555: A chain of mountain ranges stretching from Turkey to Malaysia . Pangaea was fractured by widespread faulting and rift basins during the Triassic—especially late in that period—but had not yet separated. The first nonmarine sediments in the rift that marks the initial break-up of Pangaea, which separated eastern North America from Morocco , are of Late Triassic age; in the United States , these thick sediments comprise

876-402: A cusp covering the rear belly, between the pelvis and the belly ribs. The vertical mobility of this element suggests a function in breathing, compensating the relative rigidity of the chest cavity. The hindlimbs of pterosaurs were strongly built, yet relative to their wingspans smaller than those of birds. They were long in comparison to the torso length. The thighbone was rather straight, with

1022-474: A few exposures in the west. During the Triassic peneplains are thought to have formed in what is now Norway and southern Sweden. Remnants of this peneplain can be traced as a tilted summit accordance in the Swedish West Coast . In northern Norway Triassic peneplains may have been buried in sediments to be then re-exposed as coastal plains called strandflats . Dating of illite clay from

1168-458: A few millimetres thin transversely. The bony crest base would typically be extended by keratinous or other soft tissue. Since the 1990s, new discoveries and a more thorough study of old specimens have shown that crests are far more widespread among pterosaurs than previously assumed. That they were extended by or composed completely of keratin, which does not fossilize easily, had misled earlier research. For Pterorhynchus and Pterodactylus ,

1314-411: A flying creature in a letter to Georges Cuvier . Cuvier agreed in 1801, understanding it was an extinct flying reptile. In 1809, he coined the name Ptéro-Dactyle , "wing-finger". This was in 1815 Latinised to Pterodactylus . At first most species were assigned to this genus and ultimately "pterodactyl" was popularly and incorrectly applied to all members of Pterosauria. Today, paleontologists limit

1460-445: A limited mobility. These toes were clawed but the claws were smaller than the hand claws. The rare conditions that allowed for the fossilisation of pterosaur remains, sometimes also preserved soft tissues. Modern synchrotron or ultraviolet light photography has revealed many traces not visible to the naked eye. These are often imprecisely called "impressions" but mostly consist of petrifications , natural casts and transformations of

1606-471: A long beak-like snout), and Shringasaurus (a horned herbivore which reached a body length of 3–4 metres (9.8–13.1 ft)). One group of archosauromorphs, the archosauriforms , were distinguished by their active predatory lifestyle, with serrated teeth and upright limb postures. Archosauriforms were diverse in the Triassic, including various terrestrial and semiaquatic predators of all shapes and sizes. The large-headed and robust erythrosuchids were among

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1752-400: A membrane of skin, muscle, and other tissues stretching from the ankles to a dramatically lengthened fourth finger. There were two major types of pterosaurs. Basal pterosaurs (also called 'non-pterodactyloid pterosaurs' or ' rhamphorhynchoids ') were smaller animals with fully toothed jaws and, typically, long tails. Their wide wing membranes probably included and connected the hind legs. On

1898-516: A membrane that stretched between the legs, possibly connecting to or incorporating the tail, called the uropatagium ; the extent of this membrane is not certain, as studies on Sordes seem to suggest that it simply connected the legs but did not involve the tail (rendering it a cruropatagium ). A common interpretation is that non-pterodactyloid pterosaurs had a broader uro/cruropatagium stretched between their long fifth toes, with pterodactyloids, lacking such toes, only having membranes running along

2044-425: A new fossil of Tupandactylus cf. imperator was found to have melanosomes in forms that signal an earlier-than-anticipated development of patterns found in extant feathers. The new specimen suggested that pterosaur integumentary melanosomes exhibited a more complex organization than those previously known from other pterosaurs. This indicates the presence of a unique form of melanosomes within pterosaur integument at

2190-617: A pseudosuchian. Pseudosuchians were far more ecologically dominant in the Triassic, including large herbivores (such as aetosaurs ), large carnivores (" rauisuchians "), and the first crocodylomorphs (" sphenosuchians "). Aetosaurs were heavily-armored reptiles that were common during the last 30 million years of the Late Triassic until they died out at the Triassic-Jurassic extinction. Most aetosaurs were herbivorous and fed on low-growing plants, but some may have eaten meat. " rauisuchians " (formally known as paracrocodylomorphs ) were

2336-411: A rotation could be caused by an abduction of the thighbone, meaning that the legs would be spread. This would also turn the feet into a vertical position. They then could act as rudders to control yaw. Some specimens show membranes between the toes, allowing them to function as flight control surfaces. The uropatagium or cruropatagium would control pitch. When walking the toes could flex upwards to lift

2482-458: A short period of time, becoming extinct about 220 million years ago. They were exceptionally abundant in the middle of the Triassic, as the primary large herbivores in many Carnian-age ecosystems. They sheared plants with premaxillary beaks and plates along the upper jaw with multiple rows of teeth. Allokotosaurs were iguana-like reptiles, including Trilophosaurus (a common Late Triassic reptile with three-crowned teeth), Teraterpeton (which had

2628-567: A specialized subgroup of cynodonts, appeared during the Triassic and would survive the extinction event, allowing them to radiate during the Jurassic. Amphibians were primarily represented by the temnospondyls , giant aquatic predators that had survived the end-Permian extinction and saw a new burst of diversification in the Triassic, before going extinct by the end; however, early crown-group lissamphibians (including stem-group frogs , salamanders and caecilians ) also became more common during

2774-449: A strandflat of Bømlo , southern Norway, have shown that landscape there became weathered in Late Triassic times ( c. 210 million years ago) with the landscape likely also being shaped during that time. Eustatic sea level in the Triassic was consistently low compared to the other geological periods. The beginning of the Triassic was around present sea level, rising to about 10–20 metres (33–66 ft) above present-day sea level during

2920-585: A supercontinent has less shoreline compared to a series of smaller continents, Triassic marine deposits are relatively uncommon on a global scale. A major exception is in Western Europe , where the Triassic was first studied. The northeastern margin of Gondwana was a stable passive margin along the Neo-Tethys Ocean, and marine sediments have been preserved in parts of northern India and Arabia . In North America , marine deposits are limited to

3066-449: A supraneural plate that, however, would not contact the notarium. The tails of pterosaurs were always rather slender. This means that the caudofemoralis retractor muscle which in most basal Archosauria provides the main propulsive force for the hindlimb, was relatively unimportant. The tail vertebrae were amphicoelous, the vertebral bodies on both ends being concave. Early species had long tails, containing up to fifty caudal vertebrae,

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3212-464: A thousand bristle-like teeth. Dsungaripteridae covered their teeth with jawbone tissue for a crushing function. If teeth were present, they were placed in separate tooth sockets. Replacement teeth were generated behind, not below, the older teeth. The public image of pterosaurs is defined by their elaborate head crests. This was influenced by the distinctive backward-pointing crest of the well-known Pteranodon . The main positions of such crests are

3358-416: A unique, complex circulatory system of looping blood vessels. The combination of actinofibrils and muscle layers may have allowed the animal to adjust the wing slackness and camber . As shown by cavities in the wing bones of larger species and soft tissue preserved in at least one specimen, some pterosaurs extended their system of respiratory air sacs into the wing membrane. The pterosaur wing membrane

3504-408: A weight of up to 250 kilograms (550 pounds) for the largest species. Compared to the other vertebrate flying groups, the birds and bats, pterosaur skulls were typically quite large. Most pterosaur skulls had elongated jaws. Their skull bones tend to be fused in adult individuals. Early pterosaurs often had heterodont teeth, varying in build, and some still had teeth in the palate. In later groups

3650-453: A wide range of adult sizes , from the very small anurognathids to the largest known flying creatures, including Quetzalcoatlus and Hatzegopteryx , which reached wingspans of at least nine metres. The combination of endothermy , a good oxygen supply and strong muscles made pterosaurs powerful and capable flyers. Pterosaurs are often referred to by popular media or the general public as "flying dinosaurs", but dinosaurs are defined as

3796-482: A wingspan no less than 25 centimetres (10 inches). The most sizeable forms represent the largest known animals ever to fly, with wingspans of up to 10–11 metres (33–36 feet). Standing, such giants could reach the height of a modern giraffe . Traditionally, it was assumed that pterosaurs were extremely light relative to their size. Later, it was understood that this would imply unrealistically low densities of their soft tissues. Some modern estimates therefore extrapolate

3942-659: Is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya ), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is the first and shortest period of the Mesozoic Era and the seventh period of the Phanerozoic Eon . Both the start and end of the period are marked by major extinction events . The Triassic Period

4088-468: Is a recent study of North American faunas. In the Petrified Forest of northeast Arizona there is a unique sequence of late Carnian-early Norian terrestrial sediments. An analysis in 2002 found no significant change in the paleoenvironment. Phytosaurs , the most common fossils there, experienced a change-over only at the genus level, and the number of species remained the same. Some aetosaurs ,

4234-555: Is common in warm-blooded animals who need insulation to prevent excessive heat-loss. Pycnofibers were flexible, short filaments, about five to seven millimetres long and rather simple in structure with a hollow central canal. Pterosaur pelts might have been comparable in density to many Mesozoic mammals. Pterosaur filaments could share a common origin with feathers, as speculated in 2002 by Czerkas and Ji. In 2009, Kellner concluded that pycnofibers were structured similarly to theropod proto-feathers . Others were unconvinced, considering

4380-422: Is curved to behind, resulting in a rounded wing tip, which reduces induced drag . The wingfinger is also bent somewhat downwards. When standing, pterosaurs probably rested on their metacarpals, with the outer wing folded to behind. In this position, the "anterior" sides of the metacarpals were rotated to the rear. This would point the smaller fingers obliquely to behind. According to Bennett, this would imply that

4526-421: Is divided into three basic units. The first, called the propatagium ("fore membrane"), was the forward-most part of the wing and attached between the wrist and shoulder, creating the "leading edge" during flight. The brachiopatagium ("arm membrane") was the primary component of the wing, stretching from the highly elongated fourth finger of the hand to the hindlimbs. Finally, at least some pterosaur groups had

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4672-508: Is likely a paraphyletic group rather than a true clade. Tanystropheids were a family of protorosaurs which elevated their neck size to extremes, with the largest genus Tanystropheus having a neck longer than its body. The protorosaur family Sharovipterygidae used their elongated hindlimbs for gliding. Other archosauromorphs, such as rhynchosaurs and allokotosaurs , were mostly stocky-bodied herbivores with specialized jaw structures. Rhynchosaurs, barrel-gutted herbivores, thrived for only

4818-469: Is no evidence of glaciation at or near either pole; in fact, the polar regions were apparently moist and temperate , providing a climate suitable for forests and vertebrates, including reptiles. Pangaea's large size limited the moderating effect of the global ocean; its continental climate was highly seasonal, with very hot summers and cold winters. The strong contrast between the Pangea supercontinent and

4964-448: Is one in a series of these failed rifts. During its rupture, tholeiitic basaltic lava erupted, producing a sequence of continental tholeiitic basalts. These tholeiitic basalts form a volcanic ridge known as North Mountain . 45°8′N 65°35′W  /  45.133°N 65.583°W  / 45.133; -65.583 Triassic The Triassic ( / t r aɪ ˈ æ s ɪ k / try- ASS -ik ; sometimes symbolized 🝈 )

5110-418: Is short but powerfully built. It sports a large deltopectoral crest, to which the major flight muscles are attached. Despite the considerable forces exerted on it, the humerus is hollow or pneumatised inside, reinforced by bone struts. The long bones of the lower arm, the ulna and radius , are much longer than the humerus. They were probably incapable of pronation . A bone unique to pterosaurs, known as

5256-605: Is subdivided into three epochs: Early Triassic , Middle Triassic and Late Triassic . The Triassic began in the wake of the Permian–Triassic extinction event , which left the Earth's biosphere impoverished; it was well into the middle of the Triassic before life recovered its former diversity. Three categories of organisms can be distinguished in the Triassic record: survivors from the extinction event, new groups that flourished briefly, and other new groups that went on to dominate

5402-460: Is superimposed by 22 sea level drop events widespread in the geologic record, mostly of minor (less than 25-metre (82 ft)) and medium (25–75-metre (82–246 ft)) magnitudes. A lack of evidence for Triassic continental ice sheets suggest that glacial eustasy is unlikely to be the cause of these changes. The Triassic continental interior climate was generally hot and dry, so that typical deposits are red bed sandstones and evaporites . There

5548-411: Is usually divided into Early , Middle , and Late Triassic Epochs , and the corresponding rocks are referred to as Lower, Middle, or Upper Triassic. The faunal stages from the youngest to oldest are: During the Triassic, almost all the Earth's land mass was concentrated into a single supercontinent , Pangaea ( lit.   ' entire land ' ). This supercontinent was more-or-less centered on

5694-578: The Carnian (early part of the Late Triassic), some advanced cynodonts gave rise to the first mammals . During the Triassic, archosaurs displaced therapsids as the largest and most ecologically prolific terrestrial amniotes. This "Triassic Takeover" may have contributed to the evolution of mammals by forcing the surviving therapsids and their mammaliaform successors to live as small, mainly nocturnal insectivores . Nocturnal life may have forced

5840-614: The Jurassic , when the temnospondyls had become very rare. Most of the Reptiliomorpha , stem-amniotes that gave rise to the amniotes, disappeared in the Triassic, but two water-dwelling groups survived: Embolomeri that only survived into the early part of the period, and the Chroniosuchia , which survived until the end of the Triassic. The Permian–Triassic extinction devastated terrestrial life. Biodiversity rebounded as

5986-568: The Lake Lugano region of northern Italy and southern Switzerland , was in Middle Triassic times a lagoon behind reefs with an anoxic bottom layer, so there were no scavengers and little turbulence to disturb fossilization, a situation that can be compared to the better-known Jurassic Solnhofen Limestone lagerstätte . The remains of fish and various marine reptiles (including the common pachypleurosaur Neusticosaurus , and

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6132-509: The Mesozoic Era. Reptiles , especially archosaurs , were the chief terrestrial vertebrates during this time. A specialized group of archosaurs, called dinosaurs , first appeared in the Late Triassic but did not become dominant until the succeeding Jurassic Period. Archosaurs that became dominant in this period were primarily pseudosuchians , relatives and ancestors of modern crocodilians , while some archosaurs specialized in flight,

6278-658: The Newark Supergroup . Rift basins are also common in South America, Europe, and Africa. Terrestrial environments are particularly well-represented in the South Africa, Russia, central Europe, and the southwest United States. Terrestrial Triassic biostratigraphy is mostly based on terrestrial and freshwater tetrapods, as well as conchostracans ("clam shrimps"), a type of fast-breeding crustacean which lived in lakes and hypersaline environments. Because

6424-474: The Olenekian and Anisian of Gondwana . Both kannemeyeriiform dicynodonts and gomphodont cynodonts remained important herbivores during much of the period. Therocephalians included both large predators ( Moschorhinus ) and herbivorous forms ( bauriids ) until their extinction midway through the period. Ecteniniid cynodonts played a role as large-sized, cursorial predators in the Late Triassic. During

6570-477: The patagium , and the presence of both aktinofibrils and filaments on Jeholopterus ningchengensis and Sordes pilosus . The various forms of filament structure present on the anurognathids in the 2018 study would also require a form of decomposition that would cause the different 'filament' forms seen. They therefore conclude that the most parsimonious interpretation of the structures is that they are filamentous protofeathers. But Liliana D'Alba points out that

6716-735: The surviving species repopulated empty terrain, but these were short-lived. Diverse communities with complex food-web structures took 30 million years to reestablish. Archosauromorph reptiles, which had already appeared and diversified to an extent in the Permian Period, exploded in diversity as an adaptive radiation in response to the Permian-Triassic mass extinction. By the Early Triassic, several major archosauromorph groups had appeared. Long-necked, lizard-like early archosauromorphs were known as protorosaurs , which

6862-468: The thecodonts ) disappeared, as did most of the large labyrinthodont amphibians, groups of small reptiles, and most synapsids. Some of the early, primitive dinosaurs also became extinct, but more adaptive ones survived to evolve into the Jurassic. Surviving plants that went on to dominate the Mesozoic world included modern conifers and cycadeoids. The cause of the Late Triassic extinction is uncertain. It

7008-427: The thorax . It was probably covered by thick muscle layers. The upper bone, the shoulder blade , was a straight bar. It was connected to a lower bone, the coracoid that is relatively long in pterosaurs. In advanced species, their combined whole, the scapulocoracoid, was almost vertically oriented. The shoulder blade in that case fitted into a recess in the side of the notarium, while the coracoid likewise connected to

7154-437: The traversodont cynodonts—were much reduced in the northern half of Pangaea ( Laurasia ). These extinctions within the Triassic and at its end allowed the dinosaurs to expand into many niches that had become unoccupied. Dinosaurs became increasingly dominant, abundant and diverse, and remained that way for the next 150 million years. The true "Age of Dinosaurs" is during the following Jurassic and Cretaceous periods, rather than

7300-428: The 1990s, pterosaur finds and histological and ultraviolet examination of pterosaur specimens have provided incontrovertible proof: pterosaurs had pycnofiber coats. Sordes pilosus (which translates as "hairy demon") and Jeholopterus ninchengensis show pycnofibers on the head and body. The presence of pycnofibers strongly indicates that pterosaurs were endothermic (warm-blooded). They aided thermoregulation, as

7446-832: The Anisian to Ladinian of the Tethysian domain, and from the Carnian and Rhaetian of a larger area that includes also the Boreal domain (e.g., Svalbard Islands), the North American continent, the South China block and Argentina . The best-studied of such episodes of humid climate, and probably the most intense and widespread, was the Carnian Pluvial Event . The Early Triassic was the hottest portion of

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7592-405: The Carnian and include early sauropodomorphs and theropods. Most Triassic dinosaurs were small predators and only a few were common, such as Coelophysis , which was 1 to 2 metres (3.3 to 6.6 ft) long. Triassic sauropodomorphs primarily inhabited cooler regions of the world. The large predator Smok was most likely also an archosaur, but it is uncertain if it was a primitive dinosaur or

7738-563: The Early Triassic, forming small patches of reefs of modest extent compared to the great reef systems of Devonian or modern times. At the end of the Carnian, a reef crisis occurred in South China. Serpulids appeared in the Middle Triassic. Microconchids were abundant. The shelled cephalopods called ammonites recovered, diversifying from a single line that survived the Permian extinction. Bivalves began to rapidly diversify during

7884-402: The Early Triassic, while others (e.g. capitosaurs ) remained successful throughout the whole period, or only came to prominence in the Late Triassic (e.g. Plagiosaurus , metoposaurs ). The first Lissamphibians (modern amphibians) appear in the Triassic, with the progenitors of the first frogs already present by the Early Triassic. However, the group as a whole did not become common until

8030-562: The Early and Middle Triassic. Sea level rise accelerated in the Ladinian, culminating with a sea level up to 50 metres (164 ft) above present-day levels during the Carnian. Sea level began to decline in the Norian, reaching a low of 50 metres (164 ft) below present sea level during the mid-Rhaetian. Low global sea levels persisted into the earliest Jurassic. The long-term sea level trend

8176-403: The Jurassic. The Triassic was named in 1834 by Friedrich August von Alberti , after a succession of three distinct rock layers (Greek triás meaning 'triad') that are widespread in southern Germany : the lower Buntsandstein (colourful sandstone ) , the middle Muschelkalk (shell-bearing limestone ) and the upper Keuper (coloured clay ). On the geologic time scale , the Triassic

8322-613: The Jurassic. There were many types of marine reptiles. These included the Sauropterygia , which featured pachypleurosaurus and nothosaurs (both common during the Middle Triassic, especially in the Tethys region), placodonts , the earliest known herbivorous marine reptile Atopodentatus , and the first plesiosaurs . The first of the lizardlike Thalattosauria ( askeptosaurs ) and the highly successful ichthyopterygians , which appeared in Early Triassic seas, soon diversified. By

8468-600: The Latest Olenekian Cooling (LOC), from 248 to 247 Ma, temperatures cooled by about 6 °C. The Middle Triassic was cooler than the Early Triassic, with temperatures falling over most of the Anisian, with the exception of a warming spike in the latter portion of the stage. From 242 to 233 Ma, the Ladinian-Carnian Cooling (LCC) ensued. At the beginning of the Carnian, global temperatures continued to be relatively cool. The eruption of

8614-486: The Middle Triassic, becoming highly abundant in the oceans. Aquatic insects rapidly diversified during the Middle Triassic, with this time interval representing a crucial diversification for Holometabola , the clade containing the majority of modern insect species. In the wake of the Permian-Triassic mass extinction event , the fish fauna was remarkably uniform, with many families and genera exhibiting

8760-618: The Middle Triassic, some ichthyopterygians were achieving very large body masses. Among other reptiles, the earliest turtles , like Proganochelys and Proterochersis , appeared during the Norian Age (Stage) of the Late Triassic Period. The Lepidosauromorpha , specifically the Sphenodontia , are first found in the fossil record of the earlier Carnian Age, though the earliest lepidosauromorphs likely occurred in

8906-450: The Permian extinction, Archaeplastida (red and green algae) had been the major marine phytoplanktons since about 659–645 million years ago, when they replaced marine planktonic cyanobacteria , which first appeared about 800 million years ago, as the dominant phytoplankton in the oceans. In the Triassic, secondary endosymbiotic algae became the most important plankton. In marine environments , new modern types of corals appeared in

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9052-475: The Permian. The Procolophonidae , the last surviving parareptiles , were an important group of small lizard-like herbivores. The drepanosaurs were a clade of unusual, chameleon-like arboreal reptiles with birdlike heads and specialised claws. Three therapsid groups survived into the Triassic: dicynodonts , therocephalians , and cynodonts . The cynodont Cynognathus was a characteristic top predator in

9198-408: The Triassic (teleosts are by far the most diverse group of fish today). Predatory actinopterygians such as saurichthyids and birgeriids , some of which grew over 1.2 m (3.9 ft) in length, appeared in the Early Triassic and became widespread and successful during the period as a whole. Lakes and rivers were populated by lungfish (Dipnoi), such as Ceratodus , which are mainly known from

9344-497: The Triassic and survived the extinction event. The earliest known neopterygian fish, including early holosteans and teleosts , appeared near the beginning of the Triassic, and quickly diversified to become among the dominant groups of fish in both freshwater and marine habitats. The vast supercontinent of Pangaea dominated the globe during the Triassic, but in the latest Triassic ( Rhaetian ) and Early Jurassic it began to gradually rift into two separate landmasses: Laurasia to

9490-609: The Triassic, enlarging the Neo-Tethys Ocean which formed in their wake. At the same time, they forced the Paleo-Tethys Ocean to shrink as it was being subducted under Asia. By the end of the Triassic, the Paleo-Tethys Ocean occupied a small area and the Cimmerian terranes began to collide with southern Asia. This collision, known as the Cimmerian Orogeny , continued into the Jurassic and Cretaceous to produce

9636-584: The Triassic. Pterosaurs Ornithosauria Seeley , 1870 Pterosaurs are an extinct clade of flying reptiles in the order Pterosauria . They existed during most of the Mesozoic : from the Late Triassic to the end of the Cretaceous (228 to 66 million years ago). Pterosaurs are the earliest vertebrates known to have evolved powered flight . Their wings were formed by

9782-839: The Wrangellia Large Igneous Province around 234 Ma caused abrupt global warming, terminating the cooling trend of the LCC. This warming was responsible for the Carnian Pluvial Event and resulted in an episode of widespread global humidity. The CPE ushered in the Mid-Carnian Warm Interval (MCWI), which lasted from 234 to 227 Ma. At the Carnian-Norian boundary occurred a positive δ C excursion believed to signify an increase in organic carbon burial. From 227 to 217 Ma, there

9928-428: The ankle, sometimes reducing total length to a third. Typically, it was fused to the shinbone. The ankle was a simple, "mesotarsal", hinge. The, rather long and slender, metatarsus was always splayed to some degree. The foot was plantigrade, meaning that during the walking cycle the sole of the metatarsus was pressed onto the soil. There was a clear difference between early pterosaurs and advanced species regarding

10074-484: The ankles. The exact curvature of the trailing edge, however, is still equivocal. While historically thought of as simple leathery structures composed of skin, research has since shown that the wing membranes of pterosaurs were highly complex dynamic structures suited to an active style of flight. The outer wings (from the tip to the elbow) were strengthened by closely spaced fibers called actinofibrils . The actinofibrils themselves consisted of three distinct layers in

10220-427: The anterior surface of the distal syncarpal. The medial carpal bears a deep concave fovea that opens anteriorly, ventrally and somewhat medially, within which the pteroid articulates, according to Wilkinson. In derived pterodactyloids like pteranodontians and azhdarchoids , metacarpals I-III are small and do not connect to the carpus, instead hanging in contact with the fourth metacarpal. With these derived species,

10366-455: The bizarre long-necked archosauromorph Tanystropheus ), along with some terrestrial forms like Ticinosuchus and Macrocnemus , have been recovered from this locality. All these fossils date from the Anisian and Ladinian ages (about 242 Ma ago). The Triassic Period ended with a mass extinction, which was particularly severe in the oceans; the conodonts disappeared, as did all

10512-413: The breastbone. This way, both sides together made for a rigid closed loop, able to withstand considerable forces. A peculiarity was that the breastbone connections of the coracoids often were asymmetrical, with one coracoid attached in front of the other. In advanced species the shoulder joint had moved from the shoulder blade to the coracoid. The joint was saddle-shaped and allowed considerable movement to

10658-447: The broad ischium into an ischiopubic blade. Sometimes, the blades of both sides were also fused, closing the pelvis from below and forming the pelvic canal. The hip joint was not perforated and allowed considerable mobility to the leg. It was directed obliquely upwards, preventing a perfectly vertical position of the leg. The front of the pubic bones articulated with a unique structure, the paired prepubic bones. Together these formed

10804-474: The clade Anurognathidae ( Anurognathus , Jeholopterus , Vesperopterylus ) is debated. Anurognathids were highly specialized. Small flyers with shortened jaws and a wide gape, some had large eyes suggesting nocturnal or crepuscular habits, mouth bristles, and feet adapted for clinging. Parallel adaptations are seen in birds and bats that prey on insects in flight. Pterosaurs had a wide range of sizes, though they were generally large. The smallest species had

10950-445: The clades Ornithocheiroidea ( Istiodactylus , Ornithocheirus , Pteranodon ), Ctenochasmatoidea ( Ctenochasma , Pterodactylus ), Dsungaripteroidea ( Germanodactylus , Dsungaripterus ), and Azhdarchoidea ( Tapejara , Tupuxuara , Quetzalcoatlus ). The two groups overlapped in time, but the earliest pterosaurs in the fossil record are basal pterosaurs, and the latest pterosaurs are pterodactyloids. The position of

11096-506: The dental plates, abundant in the fossils record. Hybodonts , a group of shark-like cartilaginous fish , were dominant in both freshwater and marine environments throughout the Triassic. Last survivors of the mainly Palaeozoic Eugeneodontida are known from the Early Triassic. Temnospondyl amphibians were among those groups that survived the Permian–Triassic extinction. Once abundant in both terrestrial and aquatic environments,

11242-502: The descendants of the last common ancestor of the Saurischia and Ornithischia , which excludes the pterosaurs. Pterosaurs are nonetheless more closely related to birds and other dinosaurs than to crocodiles or any other living reptile, though they are not bird ancestors. Pterosaurs are also colloquially referred to as pterodactyls , particularly in fiction and journalism. However, technically, pterodactyl may refer to members of

11388-431: The description of the preserved integumentary structures on the two anurognathid specimens is still based upon gross morphology. She also points out that Pterorhynchus was described to have feathers to support the claim that feathers had a common origin with Ornithodirans but was argued against by several authors. The only method to assure if it was homologous to feathers is to use a scanning electron microscope. In 2022,

11534-544: The difference with the "quills" found on many of the bird-like maniraptoran specimens too fundamental. A 2018 study of the remains of two small Jurassic -age pterosaurs from Inner Mongolia , China , found that pterosaurs had a wide array of pycnofiber shapes and structures, as opposed to the homogeneous structures that had generally been assumed to cover them. Some of these had frayed ends, very similar in structure to four different feather types known from birds or other dinosaurs but almost never known from pterosaurs prior to

11680-428: The dominant carnivores in the early Triassic. Phytosaurs were a particularly common group which prospered during the Late Triassic. These long-snouted and semiaquatic predators resemble living crocodiles and probably had a similar lifestyle, hunting for fish and small reptiles around the water's edge. However, this resemblance is only superficial and is a prime-case of convergent evolution. True archosaurs appeared in

11826-410: The down feathers found on both avian and some non-avian dinosaurs , suggesting that early feathers evolved in the common ancestor of pterosaurs and dinosaurs, possibly as insulation. They were warm-blooded (endothermic), active animals. The respiratory system had efficient unidirectional "flow-through" breathing using air sacs , which hollowed out their bones to an extreme extent. Pterosaurs spanned

11972-405: The early Triassic, splitting into two branches: Avemetatarsalia (the ancestors to birds) and Pseudosuchia (the ancestors to crocodilians). Avemetatarsalians were a minor component of their ecosystems, but eventually produced the earliest pterosaurs and dinosaurs in the Late Triassic. Early long-tailed pterosaurs appeared in the Norian and quickly spread worldwide. Triassic dinosaurs evolved in

12118-640: The entire Phanerozoic, seeing as it occurred during and immediately after the discharge of titanic volumes of greenhouse gases from the Siberian Traps. The Early Triassic began with the Permian-Triassic Thermal Maximum (PTTM) and was followed by the brief Dienerian Cooling (DC) from 251 to 249 Ma, which was in turn followed by the Latest Smithian Thermal Maximum (LSTT) around 249 to 248 Ma. During

12264-467: The equator and extended between the poles, though it did drift northwards as the period progressed. Southern Pangea, also known as Gondwana , was made up by closely-appressed cratons corresponding to modern South America , Africa , Madagascar , India , Antarctica , and Australia . North Pangea, also known as Laurussia or Laurasia , corresponds to modern-day North America and the fragmented predecessors of Eurasia . The western edge of Pangea lay at

12410-404: The extent of their wing membranes and it is possible that, like these groups, different species of pterosaur had different wing designs. Indeed, analysis of pterosaur limb proportions shows that there was considerable variation, possibly reflecting a variety of wing-plans. The bony elements of the arm formed a mechanism to support and extend the wing. Near the body, the humerus or upper arm bone

12556-480: The extinct family Cheirolepidiaceae , which first appeared in the Late Triassic, and would be prominent throughout most of the rest of the Mesozoic. No known coal deposits date from the start of the Triassic Period. This is known as the Early Triassic "coal gap" and can be seen as part of the Permian–Triassic extinction event . Possible explanations for the coal gap include sharp drops in sea level at

12702-676: The feather-specific melanosome signaling found in extant birds are possibly homologous with those found in pterosaurs. Pterosaur fossils are very rare, due to their light bone construction. Complete skeletons can generally only be found in geological layers with exceptional preservation conditions, the so-called Lagerstätten . The pieces from one such Lagerstätte , the Late Jurassic Solnhofen Limestone in Bavaria , became much sought after by rich collectors. In 1784, Italian naturalist Cosimo Alessandro Collini

12848-432: The fifth toes as hooks. Another hypothesis held that they stretched the brachiopatagia, but in articulated fossils the fifth digits are always flexed towards the tail. Later it became popular to assume that these toes extended an uropatagium or cruropatagium between them. As the fifth toes were on the outside of the feet, such a configuration would only have been possible if these rotated their fronts outwards in flight. Such

12994-406: The first time among vertebrates, becoming the pterosaurs . Therapsids , the dominant vertebrates of the preceding Permian period, saw a brief surge in diversification in the Triassic, with dicynodonts and cynodonts quickly becoming dominant, but they declined throughout the period with the majority becoming extinct by the end. However, the first stem-group mammals ( mammaliamorphs ), themselves

13140-450: The forces caused by flapping the wings. The notarium included three to seven vertebrae, depending on the species involved but also on individual age. These vertebrae could be connected by tendons or a fusion of their neural spines into a "supraneural plate". Their ribs also would be tightly fused into the notarium. In general, the ribs are double headed. The sacrum consisted of three to ten sacral vertebrae. They too, could be connected via

13286-400: The forelimb digits besides the wingfinger have been lost altogether. The wingfinger accounts for about half or more of the total wing length. It normally consists of four phalanges. Their relative lengths tend to vary among species, which has often been used to distinguish related forms. The fourth phalanx is usually the shortest. It lacks a claw and has been lost completely by nyctosaurids. It

13432-417: The form of the fifth digit. Originally, the fifth metatarsal was robust and not very shortened. It was connected to the ankle in a higher position than the other metatarsals. It bore a long, and often curved, mobile clawless fifth toe consisting of two phalanges. The function of this element has been enigmatic. It used to be thought that the animals slept upside-down like bats, hanging from branches and using

13578-415: The fourth metacarpal has been enormously elongated, typically equalling or exceeding the length of the long bones of the lower arm. The fifth metacarpal had been lost. In all species, the first to third fingers are much smaller than the fourth, the "wingfinger", and contain two, three and four phalanges respectively. The smaller fingers are clawed, with the ungual size varying among species. In nyctosaurids

13724-473: The front of the snout, as an outgrowth of the premaxillae, or the rear of the skull as an extension of the parietal bones in which case it is called a "supraoccipital crest". Front and rear crests can be present simultaneously and might be fused into a single larger structure, the most expansive of which is shown by the Tapejaridae . Nyctosaurus sported a bizarre antler-like crest. The crests were only

13870-429: The genus Pterodactylus , and more broadly to members of the suborder Pterodactyloidea of the pterosaurs. Pterosaurs had a variety of lifestyles. Traditionally seen as fish-eaters, the group is now understood to have also included hunters of land animals, insectivores, fruit eaters and even predators of other pterosaurs. They reproduced by eggs , some fossils of which have been discovered. The anatomy of pterosaurs

14016-521: The global ocean triggered intense cross-equatorial monsoons , sometimes referred to as the Pangean megamonsoons . The Triassic may have mostly been a dry period, but evidence exists that it was punctuated by several episodes of increased rainfall in tropical and subtropical latitudes of the Tethys Sea and its surrounding land. Sediments and fossils suggestive of a more humid climate are known from

14162-683: The ground, they walked well on all four limbs with an upright posture, standing plantigrade on the hind feet and folding the wing finger upward to walk on the three-fingered "hand". They could take off from the ground, and fossil trackways show that at least some species were able to run, wade, and/or swim. Their jaws had horny beaks, and some groups lacked teeth. Some groups developed elaborate head crests with sexual dimorphism . Pterosaurs sported coats of hair-like filaments known as pycnofibers , which covered their bodies and parts of their wings. Pycnofibers grew in several forms, from simple filaments to branching down feathers . These may be homologous to

14308-460: The ground, they would have had an awkward sprawling posture, but the anatomy of their joints and strong claws would have made them effective climbers, and some may have even lived in trees. Basal pterosaurs were insectivores or predators of small vertebrates. Later pterosaurs ( pterodactyloids ) evolved many sizes, shapes, and lifestyles. Pterodactyloids had narrower wings with free hind limbs, highly reduced tails, and long necks with large heads. On

14454-464: The head and torso. The term "pycnofiber", meaning "dense filament", was coined by palaeontologist Alexander Kellner and colleagues in 2009. Pycnofibers were unique structures similar to, but not homologous (sharing a common origin) with, mammalian hair, an example of convergent evolution . A fuzzy integument was first reported from a specimen of Scaphognathus crassirostris in 1831 by Georg August Goldfuss , but had been widely doubted. Since

14600-412: The head making only a small angle with the shaft. This implies that the legs were not held vertically below the body but were somewhat sprawling. The shinbone was often fused with the upper ankle bones into a tibiotarsus that was longer than the thighbone. It could attain a vertical position when walking. The calf bone tended to be slender, especially at its lower end that in advanced forms did not reach

14746-420: The impact. So, the evidence suggests the Manicouagan impact preceded the end of the Triassic by approximately 10±2 Ma. It could not therefore be the immediate cause of the observed mass extinction. The number of Late Triassic extinctions is disputed. Some studies suggest that there are at least two periods of extinction towards the end of the Triassic, separated by 12 to 17 million years. But arguing against this

14892-401: The jaw joint was in a more forward position. The front lower jaw bones, the dentaries or ossa dentalia , were at the tip tightly fused into a central symphysis. This made the lower jaws function as a single connected whole, the mandible . The symphysis was often very thin transversely and long, accounting for a considerable part of the jaw length, up to 60%. If a crest was present on the snout,

15038-401: The keystone predators of most Triassic terrestrial ecosystems. Over 25 species have been found, including giant quadrupedal hunters, sleek bipedal omnivores, and lumbering beasts with deep sails on their backs. They probably occupied the large-predator niche later filled by theropods. "Rauisuchians" were ancestral to small, lightly-built crocodylomorphs, the only pseudosuchians which survived into

15184-544: The landscape, covering much of southern Nova Scotia. Sections of the flood basalts have been eroded away, but still form a basaltic mountain range known as North Mountain . The rift valley eventually failed ( see aulacogen ) as the Mid-Atlantic Ridge continued to separate North America, Europe, and Africa. As in all rifting environments, grabens formed. Many of these grabens were created, but for some of them, extension stopped before full rifting occurred. Where only partial rifting occurred, basins formed. By definition,

15330-547: The legs. There has been considerable argument among paleontologists about whether the main wing membranes (brachiopatagia) attached to the hindlimbs, and if so, where. Fossils of the rhamphorhynchoid Sordes , the anurognathid Jeholopterus , and a pterodactyloid from the Santana Formation seem to demonstrate that the wing membrane did attach to the hindlimbs, at least in some species. However, modern bats and flying squirrels show considerable variation in

15476-545: The mammaliaforms to develop fur and a higher metabolic rate . Two Early Triassic lagerstätten (high-quality fossil beds), the Dienerian aged Guiyang biota and the earliest Spathian aged Paris biota stand out due to their exceptional preservation and diversity . They represent the earliest lagerstätten of the Mesozoic era and provide insight into the biotic recovery from the Permian-Triassic mass extinction event. The Monte San Giorgio lagerstätte, now in

15622-573: The margin of an enormous ocean, Panthalassa ( lit.   ' entire sea ' ), which roughly corresponds to the modern Pacific Ocean . Practically all deep-ocean crust present during the Triassic has been recycled through the subduction of oceanic plates, so very little is known about the open ocean from this time period. Most information on Panthalassan geology and marine life is derived from island arcs and rare seafloor sediments accreted onto surrounding land masses, such as present-day Japan and western North America. The eastern edge of Pangea

15768-453: The marine reptiles except ichthyosaurs and plesiosaurs . Invertebrates like brachiopods and molluscs (such as gastropods ) were severely affected. In the oceans, 22% of marine families and possibly about half of marine genera went missing. Though the end-Triassic extinction event was not equally devastating in all terrestrial ecosystems, several important clades of crurotarsans (large archosaurian reptiles previously grouped together as

15914-402: The membrane from the ground. In Pterodactyloidea, the fifth metatarsal was much reduced and the fifth toe, if present, little more than a stub. This suggests that their membranes were split, increasing flight maneuverability. The first to fourth toes were long. They had two, three, four and five phalanges respectively. Often the third toe was longest; sometimes the fourth. Flat joints indicate

16060-421: The middle ones stiffened by elongated articulation processes, the zygapophyses , and chevrons . Such tails acted as rudders, sometimes ending at the rear in a vertical diamond-shaped or oval vane. In pterodactyloids, the tails were much reduced and never stiffened, with some species counting as few as ten vertebrae. The shoulder girdle was a strong structure that transferred the forces of flapping flight to

16206-575: The neck is typically longer than the torso. This length is not caused by an increase of the number of vertebrae, which is invariably seven. Some researchers include two transitional "cervicodorsals" which brings the number to nine. Instead, the vertebrae themselves became more elongated, up to eight times longer than wide. Nevertheless, the cervicals were wider than high, implying a better vertical than horizontal neck mobility. Pterodactyloids have lost all neck ribs. Pterosaur necks were probably rather thick and well-muscled, especially vertically. The torso

16352-430: The next most common tetrapods, and early dinosaurs, passed through unchanged. However, both phytosaurs and aetosaurs were among the groups of archosaur reptiles completely wiped out by the end-Triassic extinction event. It seems likely then that there was some sort of end-Carnian extinction, when several herbivorous archosauromorph groups died out, while the large herbivorous therapsids —the kannemeyeriid dicynodonts and

16498-512: The north and Gondwana to the south. The global climate during the Triassic was mostly hot and dry, with deserts spanning much of Pangaea's interior. However, the climate shifted and became more humid as Pangaea began to drift apart. The end of the period was marked by yet another major mass extinction, the Triassic–Jurassic extinction event , that wiped out many groups, including most pseudosuchians, and allowed dinosaurs to assume dominance in

16644-402: The order Isoetales (which contains living quillworts ), rose to prominence due to the environmental instability following the Permian-Triassic extinction, with one particularly notable example being the genus Pleuromeia , which grew in columnar like fashion, sometimes reaching a height of 2 metres (6.6 ft). The relevance of lycophytes declined from the Middle Triassic onwards, following

16790-458: The original material. They may include horn crests, beaks or claw sheaths as well as the various flight membranes. Exceptionally, muscles were preserved. Skin patches show small round non-overlapping scales on the soles of the feet, the ankles and the ends of the metatarsals . They covered pads cushioning the impact of walking. Scales are unknown from other parts of the body. Most or all pterosaurs had hair -like filaments known as pycnofibers on

16936-405: The pteroid bone, which may itself be a modified distal carpal. The proximal carpals are fused together into a "syncarpal" in mature specimens, while three of the distal carpals fuse to form a distal syncarpal. The remaining distal carpal, referred to here as the medial carpal, but which has also been termed the distal lateral, or pre-axial carpal, articulates on a vertically elongate biconvex facet on

17082-402: The pteroid in articulation with the proximal syncarpal, suggesting that the pteroid articulated with the 'saddle' of the radiale (proximal syncarpal) and that both the pteroid and preaxial carpal were migrated centralia. The pterosaur wrist consists of two inner (proximal, at the side of the long bones of the arm) and four outer (distal, at the side of the hand) carpals (wrist bones), excluding

17228-441: The pteroid pointed forward, extending the forward membrane and allowing it to function as an adjustable flap . This view was contradicted in a 2007 paper by Chris Bennett, who showed that the pteroid did not articulate as previously thought and could not have pointed forward, but rather was directed inward toward the body as traditionally interpreted. Specimens of Changchengopterus pani and Darwinopterus linglongtaensis show

17374-488: The pteroid, connected to the wrist and helped to support the forward membrane (the propatagium) between the wrist and shoulder. Evidence of webbing between the three free fingers of the pterosaur forelimb suggests that this forward membrane may have been more extensive than the simple pteroid-to-shoulder connection traditionally depicted in life restorations. The position of the pteroid bone itself has been controversial. Some scientists, notably Matthew Wilkinson, have argued that

17520-487: The return of more stable environmental conditions. While having first appeared during the Permian, the extinct seed plant group Bennettitales first became a prominent element in global floras during the Late Triassic, a position they would hold for much of the Mesozoic. In the Southern Hemisphere landmasses of Gondwana, the tree Dicroidium , an extinct " seed fern " belong to the order Corystospermales

17666-586: The skull, the sutures between elements disappeared. In some later pterosaurs, the backbone over the shoulders fused into a structure known as a notarium , which served to stiffen the torso during flight, and provide a stable support for the shoulder blade . Likewise, the sacral vertebrae could form a single synsacrum while the pelvic bones fused also. Basal pterosaurs include the clades Dimorphodontidae ( Dimorphodon ), Campylognathididae ( Eudimorphodon , Campyognathoides ), and Rhamphorhynchidae ( Rhamphorhynchus , Scaphognathus ). Pterodactyloids include

17812-400: The study, suggesting homology. A response to this study was published in 2020, where it was suggested that the structures seen on the anurognathids were actually a result of the decomposition of aktinofibrils: a type of fibre used to strengthen and stiffen the wing. However, in a response to this, the authors of the 2018 paper point to the fact that the presence of the structures extend past

17958-405: The symphysis could feature a matching mandible crest, jutting out to below. Toothed species also bore teeth in their dentaries. The mandible opened and closed in a simple vertical or "orthal" up-and-down movement. The vertebral column of pterosaurs numbered between thirty-four and seventy vertebrae . The vertebrae in front of the tail were "procoelous": the cotyle (front of the vertebral body )

18104-405: The teeth mostly became conical. Front teeth were often longer, forming a "prey grab" in transversely expanded jaw tips, but size and position were very variable among species. With the derived Pterodactyloidea , the skulls became even more elongated, sometimes surpassing the combined neck and torso in length. This was caused by a stretching and fusion of the front snout bone, the premaxilla , with

18250-480: The term to the genus Pterodactylus or members of the Pterodactyloidea . In 1812 and 1817, Samuel Thomas von Soemmerring redescribed the original specimen and an additional one. He saw them as affiliated to birds and bats. Although he was mistaken in this, his "bat model" would be influential during the 19th century. In 1843, Edward Newman thought pterosaurs were flying marsupials . Ironically, as

18396-676: The terminus of the Triassic, there was an extreme warming event referred to as the End-Triassic Thermal Event (ETTE), which was responsible for the Triassic-Jurassic mass extinction. Bubbles of carbon dioxide in basaltic rocks dating back to the end of the Triassic indicate that volcanic activity from the Central Atlantic Magmatic Province helped trigger climate change in the ETTE. During the Early Triassic, lycophytes , particularly those of

18542-595: The terrestrial species had mostly died out during the extinction event. The Triassic survivors were aquatic or semi-aquatic, and were represented by Tupilakosaurus , Thabanchuia , Branchiosauridae and Micropholis , all of which died out in Early Triassic, and the successful Stereospondyli , with survivors into the Cretaceous Period. The largest Triassic stereospondyls, such as Mastodonsaurus , were up to 4 to 6 metres (13 to 20 ft) in length. Some lineages (e.g. trematosaurs ) flourished briefly in

18688-577: The time of the Permo-Triassic boundary; acid rain from the Siberian Traps eruptions or from an impact event that overwhelmed acidic swamps; climate shift to a greenhouse climate that was too hot and dry for peat accumulation; evolution of fungi or herbivores that were more destructive of wetlands; the extinction of all plants adapted to peat swamps, with a hiatus of several million years before new plant species evolved that were adapted to peat swamps; or soil anoxia as oxygen levels plummeted. Before

18834-453: The time, distinct from previously known contemporary integumentary structures and more similar to those reported from mammalian hair and avian feathers. The feather fossils obtained from this specimen also suggest the presence of Stage IIIa feathers, a new discovery that indicates more complex feather structures were present in pterosaurs. The study describing this specimen further clarifies the timeline of avian feather evolution and suggests that

18980-422: The true extent of these crests has only been uncovered using ultraviolet photography. While fossil crests used to be restricted to the more advanced Pterodactyloidea, Pterorhynchus and Austriadactylus show that even some early pterosaurs possessed them. Like the upper jaws, the paired lower jaws of pterosaurs were very elongated. In advanced forms, they tended to be shorter than the upper cranium because

19126-416: The upper jawbone, the maxilla . Unlike most archosaurs , the nasal and antorbital openings of pterodactyloid pterosaurs merged into a single large opening, called the nasoantorbital fenestra . This feature likely evolved to lighten the skull for flight. In contrast, the bones behind the eye socket contracted and rotated, strongly inclining the rear skull and bringing the jaw joint forward. The braincase

19272-416: The wing, forming a crisscross pattern when superimposed on one another. The function of the actinofibrils is unknown, as is the exact material from which they were made. Depending on their exact composition (keratin, muscle, elastic structures, etc.), they may have been stiffening or strengthening agents in the outer part of the wing. The wing membranes also contained a thin layer of muscle, fibrous tissue, and

19418-402: The wing. It faced sideways and somewhat upwards. The breastbone, formed by fused paired sterna , was wide. It had only a shallow keel. Via sternal ribs, it was at its sides attached to the dorsal ribs. At its rear, a row of belly ribs or gastralia was present, covering the entire belly. To the front, a long point, the cristospina , jutted obliquely upwards. The rear edge of the breastbone

19564-475: The wingfinger, able to describe the largest arc of any wing element, up to 175°, was not folded by flexion but by an extreme extension. The wing was automatically folded when the elbow was bowed. A laser-simulated fluorescence scan on Pterodactylus also identified a membranous "fairing" (area conjunctioning the wing with the body at the neck), as opposed to the feathered or fur-composed "fairing" seen in birds and bats respectively. The pelvis of pterosaurs

19710-615: Was a dominant element in forest habitats across the region during the Middle-Late Triassic. During the Late Triassic, the Ginkgoales (which today are represented by only a single species, Ginkgo biloba ) underwent considerable diversification. Conifers were abundant during the Triassic, and included the Voltziales (which contains various lineages, probably including those ancestral to modern conifers), as well as

19856-673: Was a relatively cool period known as the Early Norian Cool Interval (ENCI), after which occurred the Mid-Norian Warm Interval (MNWI) from 217 to 209 Ma. The MNWI was briefly interrupted around 214 Ma by a cooling possibly related to the Manicouagan impact . Around 212 Ma, a 10 Myr eccentricity maximum caused a paludification of Pangaea and a reduction in the size of arid climatic zones. The Rhaetian Cool Interval (RCI) lasted from 209 to 201 Ma. At

20002-505: Was accompanied by huge volcanic eruptions that occurred as the supercontinent Pangaea began to break apart about 202 to 191 million years ago (40Ar/39Ar dates), forming the Central Atlantic Magmatic Province (CAMP), one of the largest known inland volcanic events since the planet had first cooled and stabilized. Other possible but less likely causes for the extinction events include global cooling or even

20148-401: Was concave and into it fitted a convex extension at the rear of the preceding vertebra, the condyle . Advanced pterosaurs are unique in possessing special processes projecting adjacent to their condyle and cotyle, the exapophyses , and the cotyle also may possess a small prong on its midline called a hypapophysis. The necks of pterosaurs were relatively long and straight. In pterodactyloids,

20294-484: Was encroached upon by a pair of extensive oceanic basins: The Neo-Tethys (or simply Tethys) and Paleo-Tethys Oceans . These extended from China to Iberia, hosting abundant marine life along their shallow tropical peripheries. They were divided from each other by a long string of microcontinents known as the Cimmerian terranes . Cimmerian crust had detached from Gondwana in the early Permian and drifted northwards during

20440-451: Was highly modified from their reptilian ancestors by the adaptation to flight. Pterosaur bones were hollow and air-filled, like those of birds . This provided a higher muscle attachment surface for a given skeletal weight. The bone walls were often paper-thin. They had a large and keeled breastbone for flight muscles and an enlarged brain able to coordinate complex flying behaviour. Pterosaur skeletons often show considerable fusion. In

20586-407: Was of moderate size compared to the body as a whole. Often the three pelvic bones were fused. The ilium was long and low, its front and rear blades projecting horizontally beyond the edges of the lower pelvic bones. Despite this length, the rod-like form of these processes indicates that the hindlimb muscles attached to them were limited in strength. The, in side view narrow, pubic bone fused with

20732-643: Was relatively large for reptiles. In some cases, fossilized keratinous beak tissue has been preserved, though in toothed forms, the beak is small and restricted to the jaw tips and does not involve the teeth. Some advanced beaked forms were toothless, such as the Pteranodontidae and Azhdarchidae , and had larger, more extensive, and more bird-like beaks. Some groups had specialised tooth forms. The Istiodactylidae had recurved teeth for eating meat. Ctenochasmatidae used combs of numerous needle-like teeth for filter feeding; Pterodaustro could have over

20878-403: Was relatively short and egg-shaped. The vertebrae in the back of pterosaurs originally might have numbered eighteen. With advanced species a growing number of these tended to be incorporated into the sacrum . Such species also often show a fusion of the front dorsal vertebrae into a rigid whole which is called the notarium after a comparable structure in birds. This was an adaptation to withstand

21024-482: Was that if such creatures were still alive, only the sea was a credible habitat; Collini suggested it might be a swimming animal that used its long front limbs as paddles. A few scientists continued to support the aquatic interpretation even until 1830, when German zoologist Johann Georg Wagler suggested that Pterodactylus used its wings as flippers and was affiliated with Ichthyosauria and Plesiosauria . In 1800, Johann Hermann first suggested that it represented

21170-456: Was the deepest point of the thorax. Clavicles or interclavicles were completely absent. Pterosaur wings were formed by bones and membranes of skin and other tissues. The primary membranes attached to the extremely long fourth finger of each arm and extended along the sides of the body. Where they ended has been very controversial but since the 1990s a dozen specimens with preserved soft tissue have been found that seem to show they attached to

21316-400: Was the first scientist to describe a pterosaur fossil. At that time the concepts of evolution and extinction were imperfectly developed. The bizarre build of the pterosaur was shocking, as it could not clearly be assigned to any existing animal group. The discovery of pterosaurs would thus play an important role in the progress of modern paleontology and geology. Scientific opinion at the time

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