Monte San Giorgio is a Swiss mountain and UNESCO World Heritage Site near the border between Switzerland and Italy . It is part of the Lugano Prealps , overlooking Lake Lugano in the Swiss Canton of Ticino .
162-510: Tanystropheus ( Ancient Greek : τανυ ~ 'long' + στροφευς 'hinged') is an extinct genus of archosauromorph reptile which lived during the Triassic Period in Europe, Asia, and North America. It is recognisable by its extremely elongated neck, longer than the torso and tail combined. The neck was composed of 13 vertebrae strengthened by extensive cervical ribs . Tanystropheus is one of
324-594: A nomen dubium possibly synonymous with T. hydroides . Over 500 " Tanystropheus conspicuus " specimens have been recovered from a Lower Keuper bonebed near the Silesian village of Miedary . This is the largest known concentration of Tanystropheus fossils, more than double the number collected from Monte San Giorgio. Though the Miedary specimens are individually limited to isolated postcranial bones, they are preserved in three dimensions and show great potential for elucidating
486-436: A Tanystropheus individual of that length, the weight estimate varied between 32.9 kg (72.5 lbs) and 74.8 kg (164.9 lbs), depending on the volume estimation method. This was significantly lighter than crocodiles of the same length, and more similar to large lizards. The skull of Tanystropheus longobardicus is roughly triangular when seen from the side and top, narrowing towards the snout. Each premaxilla (the toothed bone at
648-566: A junior synonym of T. longobardicus . A 2019 revision of Tanystropheus found that T. longobardicus and T. antiquus were the only valid species in the genus . Tanystropheus specimens from Monte San Giorgio have long been segregated into two morphotypes based on their tooth structure. Smaller specimens bear tricuspid teeth at the back of the jaw while larger specimens have a set of single-pointed fangs. The two morphotypes were originally considered to represent juvenile and adult specimens of T. longobardicus , though many studies have supported
810-678: A nomen dubium . Several more von Huene species, including " Procerosaurus cruralis ", " Thecodontosaurus latespinatus ", and " Thecodontosaurus primus ", have been reconsidered as indeterminate material of Tanystropheus or other archosauromorphs . One of Von Huene's species appears to be valid: T. antiquus , from the Gogolin Formation of Poland, was based on cervical vertebrae which were proportionally shorter than those of other Tanystropheus species. Long considered destroyed in World War II, several T. antiquus fossils were rediscovered in
972-537: A pitch accent . In Modern Greek, all vowels and consonants are short. Many vowels and diphthongs once pronounced distinctly are pronounced as /i/ ( iotacism ). Some of the stops and glides in diphthongs have become fricatives , and the pitch accent has changed to a stress accent . Many of the changes took place in the Koine Greek period. The writing system of Modern Greek, however, does not reflect all pronunciation changes. The examples below represent Attic Greek in
1134-466: A European species from the latest part of the Early Triassic (late Olenekian stage). T. antiquus had a proportionally shorter neck than other Tanystropheus species, so some paleontologists consider that T. antiquus deserves a separate genus, Protanystropheus . The lifestyle of Tanystropheus has been the subject of much debate. Tanystropheus is unknown from drier environments and its neck
1296-424: A broad furrow on the underside. The parietals are strongly modified in T. hydroides . They are fused into a single X-shaped bone, somewhat resembling the parietals of erythrosuchids . This shape may have resulted from fusion between the parietals' anterolateral processes (front branches) and the postfrontals, which are separate bones in T. longobardicus but not apparent in T. hydroides . A prominent pineal foramen
1458-511: A cervical count of 13, Tanystropheus acquired four additional elongated cervicals in the front half of the neck, in addition to a stout vertebra which shifted from the dorsal series into the base of the neck, transforming into the 13th cervical. Tanystropheids are unusual among reptiles in that they acquire their long necks without prolonged somitogenesis (an increase in the overall number of presacral vertebrae during early development). Instead, their overall number of presacral vertebrae remains at
1620-426: A constant count of 25, the same as their shorter-necked ancestors. This would require a shift in regionalization, encouraging the development of new cervical vertebrae rather than dorsals. There are 12 dorsal (torso) vertebrae . This count is very low among early archosauromorphs: Protorosaurus has up to 19, Prolacerta has 18, and Macrocnemus has 17. Tanystropheus' s dorsals are smaller and less specialized than
1782-461: A deeper basinal environment. These basin sediments are preserved as the Moltrasio Limestone , a thick sheet of micrite (fine-grained limestone) with abundant cherty and marly beds created by turbidites (mudslides). Jurassic sediments are preserved to the east, south, and west of Monte San Giorgio; the position of the modern mountain would have been an island or shallow environment during
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#17328561805881944-465: A fatal hindrance to terrestrial locomotion. The hindlimbs and the base of the tail were large and muscular, capable of short bursts of active swimming in shallow water. Tanystropheus was most likely a piscivorous ambush predator : the narrow subtriangular skull of T. longobardicus is supplied with three-cusped teeth suited for holding onto slippery prey, while the broader skull of T. hydroides bears an interlocking set of large curved fangs similar to
2106-438: A few modern lizard species. Some individuals of T. longobardicus have tricuspid teeth along their entire maxilla, while in others up to seven maxillary teeth are single-cusped fangs similar to the premaxillary teeth. The front edge of each orbit (eye socket) is marked by two bones: the prefrontal and lacrimal . The prefrontal is tall and projects a low vertical ridge in front of the orbit. The small, sliver-shaped lacrimal
2268-555: A fortress and settlement which was continuously inhabited by artisans from the Neolithic up until the 14th century . Artifacts, architecture, and other evidence of Roman and medieval activity are abundant on and around the mountain. Productive limestone quarries were active during this period and beyond in Italy ( Viggiù and Saltrio ) and Switzerland ( Arzo ). The mountain's fossil fuel deposits were exploited more recently. Motivated by
2430-472: A lack of contemporaneous evidence. Several theories exist about what Hellenic dialect groups may have existed between the divergence of early Greek-like speech from the common Proto-Indo-European language and the Classical period. They have the same general outline but differ in some of the detail. The only attested dialect from this period is Mycenaean Greek , but its relationship to the historical dialects and
2592-448: A larger portion of the internal dentary than in T. longobardicus . In addition, the rear of the dentary overlaps a large portion of the surangular, rather than the surangular acting as the overlapping bone where they meet. The surangular internally bears a large fossa for the jaw's adductor (vertical biting) muscles, and a prominent surangular foramen is positioned in front of the jaw joint. The most recognisable feature of Tanystropheus
2754-419: A lesser degree. Pamphylian Greek , spoken in a small area on the southwestern coast of Anatolia and little preserved in inscriptions, may be either a fifth major dialect group, or it is Mycenaean Greek overlaid by Doric, with a non-Greek native influence. Regarding the speech of the ancient Macedonians diverse theories have been put forward, but the epigraphic activity and the archaeological discoveries in
2916-537: A long and slender posterior process (rear branch) that projects under the rounded orbit. There are 15 teeth in the maxilla, increasing in size up to the eighth tooth, which is about as large as the premaxillary fangs. T.hydroides is not known to possess a septomaxilla , a neomorphic bone at the rear tip of the naris in some reptiles. The nasals are broad and plate-like, with a depressed central portion. The lacrimal and prefrontal, though incompletely known, were likely similar to those of T. longobardicus . T. hydroides has
3078-475: A loose, strongly overlapping connection to the ectopterygoids (linking bones between the pterygoid and maxilla). The epipterygoids (vertical bones in front of the braincase) are tall and flattened from the side. T. hydroides is a rare example of an early archosauromorph with a three-dimensionally preserved braincase. The basioccipital (rear lower component of the braincase) was small, with inset basitubera (vertical plates connecting to neck muscles) linked by
3240-454: A more basal type of reptile unrelated to Archosauromorpha. The following cladogram is from Dilkes (1998), a study with a small sample of "prolacertiforms" but closer resemblance to most analyses of the 2000s and 2010s: Ancient Greek language Ancient Greek ( Ἑλληνῐκή , Hellēnikḗ ; [hellɛːnikɛ́ː] ) includes the forms of the Greek language used in ancient Greece and
3402-435: A more comprehensive view of the species' anatomy. A small but well-preserved skull and neck, specimen PIMUZ T 3901, was found in the slightly younger Meride Limestone at Monte San Giorgio. Wild (1980) gave it a new species, T. meridensis , based on a set of skull and vertebral traits proposed to differ from T. longobardicus . Later reinvestigations failed to confirm the validity of these differences, rendering T. meridensis
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#17328561805883564-447: A pair of atlantal neural arches (prong-like upper components). There does not appear to be a proatlas , which slots between the atlas and skull in some other reptiles. The intercentrum and pleurocentrum are not fused to each other, unlike the single-part atlas of allokotosaurs . The tiny crescent-shaped intercentrum is overlain by a semicircular pleurocentrum, which acts as a base to the backswept neural arches. The axis (second cervical)
3726-483: A pair of segmented rods which intermingle at the midline. The two sacral (hip) vertebrae are low but robust, bridging over to the hip with expanded sacral ribs. The latter sacral rib is a single unit without a bifurcated structure. The tail is long, with at least 30 and possible up to 50 caudal vertebrae . The first few caudals are large, with closely interlinked zygapophyses and widely projecting pleurapophyses (a term for transverse processes lacking ribs). The length of
3888-415: A particularly large nasolacrimal duct , a tubular channel opening out of the rear of the lacrimal. The frontals are quite wide and form much of the upper edge of the orbit, a condition akin to T. longobardicus . However, the paired frontals meet along a straight suture with a low ridge on the lower (internal) surface, in contrast to T. longobardicus , where the frontals meet at an interdigitating suture with
4050-689: A period of rifting which would eventually break up Pangea . By the time of the Rhaetian stage (~208 to 201 Ma), the Dolomia Principale was buried by a shorter but more stable sequence of shallow-water marl and carbonate, the Tremona Series . Rifting continued into the Early Jurassic , alongside marine sedimentation. From the Hettangian to Pliensbachian stages (201 to 183 Ma), the area reacquired
4212-543: A prefix /e-/, called the augment . This was probably originally a separate word, meaning something like "then", added because tenses in PIE had primarily aspectual meaning. The augment is added to the indicative of the aorist, imperfect, and pluperfect, but not to any of the other forms of the aorist (no other forms of the imperfect and pluperfect exist). The two kinds of augment in Greek are syllabic and quantitative. The syllabic augment
4374-442: A prominent neural spine and robust zygapophyses, also unlike its predecessors. The 13th vertebra has long been assumed to be the first dorsal (torso) vertebra. This was justified by its general stout shape and supposedly dichocephalous (two-headed) rib facets, unlike the cervicals. However, specimen GMPKU-P-1527 has shown that the 13th vertebra's rib simply has a single wide articulation and an unconnected forward branch, more similar to
4536-429: A quantitative manner, by collecting a set of characteristics in sampled species and then using computational models to find the simplest ( most parsimonious ) path evolution could take to produce that character distribution. Cladistics stabilized and defined a fundamental split in the family tree of reptiles: one side of the family tree, Lepidosauromorpha , leads to lepidosaurs such as squamates (lizards and snakes) and
4698-561: A sandy coastline dotted with deltas and floodplains . As the Anisian stage continued, the coastal sandstone of the Bellano Formation was replaced with calcareous marine deposits. These were the first of many massive carbonate platforms building up on a branch of the Tethys Sea which was expanding westwards. The shallow carbonate platform of Monte San Giorgio and surrounding areas
4860-524: A search for furnace and lamp oil for Milan , mining projects attempted to establish themselves in the late 18th and early 19th centuries, focusing on the bituminous shale of the Grenzbitumenzone (Besano Formation). Though these early efforts did not last very long, exploitation of the Grenzbitumenzone ramped up in the early 20th century once its pharmaceutical properties were discovered. In 1908,
5022-671: A separate historical stage, though its earliest form closely resembles Attic Greek , and its latest form approaches Medieval Greek , and Koine may be classified as Ancient Greek in a wider sense. There were several regional dialects of Ancient Greek; Attic Greek developed into Koine. Ancient Greek was a pluricentric language , divided into many dialects. The main dialect groups are Attic and Ionic , Aeolic , Arcadocypriot , and Doric , many of them with several subdivisions. Some dialects are found in standardized literary forms in literature , while others are attested only in inscriptions. There are also several historical forms. Homeric Greek
Tanystropheus - Misplaced Pages Continue
5184-437: A series of large and systematic excavations in 1924, greatly expanding both the number of known fossil sites and the number of geological layers known to preserve fossils. Peyer's excavations continued until 1938, discovering many new species of fossil animals in the process. World War II paused both Saurol production and fossil collection. The Museo Civico di Storia Naturale di Milano (Milan Civic Museum of Natural History, MSNM)
5346-407: A shallowly concave outer edge. Like the frontals, the paired parietals are seemingly separate bones, unfused to each other in every member of the species. A large hole, the pineal foramen (sometimes called the parietal foramen), is present at the midline of the skull between the front part of each parietal. When seen from below, a pair of curved crests along the frontals and parietals mark the edge of
5508-423: A single central narial opening. Unlike T. longobardicus , T. hydroides has a nearly vertical rear edge of the premaxilla, without a postnarial process. The maxilla is low, with a large and rectangular front portion. There is a perforation near the front of the bone, which would have been penetrated by the tenth dentary tooth when the mouth was closed. Towards the rear, the maxilla develops a concave edge overlooking
5670-507: A single row of 15 relatively large curved teeth along the outer edge of the bone, adjacent to the elongated choanae (internal openings of the nasal cavity ). Most other archosauromorphs, T. longobardicus included, have restricted vomers with rows of minuscule teeth. The rest of the palate is completely toothless in T. hydroides , even the palatines and pterygoids, which bear tooth rows in most early archosauromorphs. The pterygoids are also unusual for their broad palatal ramus (front plate) and
5832-433: A small fourth distal tarsal and a minuscule third distal tarsal. There are five closely appressed metatarsals (foot bones), with the fourth and third being the longest. Though the first four metatarsals are slender and similar in length, the fifth (outermost) is very stout and subtly hooked, slotting into the ankle along a smooth joint. The estimated phalangeal formula (joints per toe) is 2-3-4-5-4. The first phalange of
5994-615: A small number of museums, primarily the PIMUZ, MSNM, and MCSN. Local museums in Besano, Meride, and Induno Olona also play a role in promotion of the site and its fossils. The Museo dei fossili del Monte San Giorgio (Museum of fossils from Monte San Giorgio) in Meride was first opened 1973, receiving a 2012 redesign and expansion courtesy of Ticinese architect Mario Botta . The geological layers of Monte San Giorgio span more than 100 million years, from
6156-428: A smaller and less specialized reptile found in the same geological strata. Beyond this conclusion, Peyer initially suggested that Tanystropheus was related to other long-necked Triassic reptiles. Sauropterygians such as plesiosaurs and nothosaurs were one possibility, and another was the fragmentary German reptile Trachelosaurus . Later, Peyer classified Tanystropheus and Macrocnemus closer to "protorosaurs",
6318-528: A specific species. The type species of Tanystropheus is T. conspicuus , a dubious name applied to particularly large fossils from Germany and Poland . Complete skeletons are common in the Besano Formation at Monte San Giorgio , on the border of Italy and Switzerland . Monte San Giorgio fossils belong to two species: the smaller T. longobardicus and the larger T. hydroides . These two species were formally differentiated in 2020 primarily on
6480-609: A standard subject of study in educational institutions of the Western world since the Renaissance . This article primarily contains information about the Epic and Classical periods of the language, which are the best-attested periods and considered most typical of Ancient Greek. From the Hellenistic period ( c. 300 BC ), Ancient Greek was followed by Koine Greek , which is regarded as
6642-480: A steep north edge sloping towards Lake Lugano and a more shallow South Slope extending towards the Po Plain . The eastern (Swiss) side of the mountain, between the municipalities of Brusino Arsizio , Riva San Vitale , and Meride , was listed as a World Heritage Site in 2003. This was in recognition of its cultural, biological, and especially paleontological significance. The site is renowned for its fossil content, one of
Tanystropheus - Misplaced Pages Continue
6804-530: A term initially used for Permian reptiles such as Protorosaurus and Araeoscelis . In the early and mid-20th century, it was commonplace for Permian and Triassic reptiles of uncertain affinity to intermingle together in classification schemes. Names such as " Eosuchia ", " Euryapsida ", " Younginiformes ", " Protorosauria ", and others were all applied by different authors with little consistency. The Early Triassic reptile Prolacerta , from South Africa , also became involved upon its discovery. Prolacerta
6966-441: A thin-shelled bivalve adapted to low oxygen. Fossils of free-swimming animals are more diverse, with marine reptiles , fish, and shelled cephalopods being the most prominent. Terrestrial and shallow-water organisms such as shrimps , conifer branches ( Voltzia ), and land reptiles ( Ticinosuchus ) were occasionally washed into the basin as well. The basin responsible for the Grenzbitumenzone continued to persist through
7128-416: A three-lobed contact with the nasals. The sutures between the frontals and their neighboring bones are coarse and interdigitating (interlocking). A small triangular bone, the postfrontal , wedges behind the rear outer corner of each frontal. A pair of larger plate-like bones, the parietals , sit directly behind the frontals on the skull roof. In T. longobardicus , the parietals are fairly broad and flat, with
7290-406: A transverse ridge, similar to allokotosaurs and archosauriforms . The parabasisphenoid (front lower component) is less specialized; it lies flat and tapers forwards to a blade-like cultriform process. The rear part of the bone has a deep triangular excavation (known as a median pharyngeal recess) on its underside, flanked by low crests and a pair of small basipterygoid processes (knobs connecting to
7452-510: A vowel or /n s r/ ; final stops were lost, as in γάλα "milk", compared with γάλακτος "of milk" (genitive). Ancient Greek of the classical period also differed in both the inventory and distribution of original PIE phonemes due to numerous sound changes, notably the following: The pronunciation of Ancient Greek was very different from that of Modern Greek . Ancient Greek had long and short vowels ; many diphthongs ; double and single consonants; voiced, voiceless, and aspirated stops ; and
7614-408: A whole. The lower jaw is slender, and most of its length is devoted to the toothed dentary bone. The dentary is downturned at its tip and its outer surface is dotted with a row of prominent foramina (blood vessel pits). There are up to 19 teeth in the dentary. Most commonly, the first six teeth are prominent conical fangs, akin to the premaxilla, while the remainder are small and tricuspid, akin to
7776-556: Is a literary form of Archaic Greek (derived primarily from Ionic and Aeolic) used in the epic poems , the Iliad and the Odyssey , and in later poems by other authors. Homeric Greek had significant differences in grammar and pronunciation from Classical Attic and other Classical-era dialects. The origins, early form and development of the Hellenic language family are not well understood because of
7938-835: Is a vertebra from the lower Carnian Fusea site in Friuli, Italy . In 2015, a large Tanystropheus cervical vertebra was described from the Economy Member of the Wolfville Formation , in the Bay of Fundy of Nova Scotia, Canada . The Wolfville Formation spans the Anisian to Carnian stages, and the Economy Member is likely Middle Triassic (Anisian-Ladinian) in age. It is a rare example of predominantly freshwater strata preserving Tanystropheus fossils. Tanystropheus -like tanystropheid fossils are known from another freshwater formation in North America:
8100-418: Is added to stems beginning with consonants, and simply prefixes e (stems beginning with r , however, add er ). The quantitative augment is added to stems beginning with vowels, and involves lengthening the vowel: Some verbs augment irregularly; the most common variation is e → ei . The irregularity can be explained diachronically by the loss of s between vowels, or that of the letter w , which affected
8262-471: Is broader and flatter than that of T. longobardicus . The first five of six teeth in the premaxilla are very large and fang-like, forming an interlocking "fish trap" similar to Dinocephalosaurus and many sauropterygians such as plesiosaurs and nothosaurs . All teeth in the skull have a single cusp which is sharp, curved, and unserrated. They have an oval-shaped cross section and shallow subthecodont implantation. Like T. longobardicus , T. hydroides has
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#17328561805888424-644: Is called 'East Greek'. Arcadocypriot apparently descended more closely from the Mycenaean Greek of the Bronze Age. Boeotian Greek had come under a strong Northwest Greek influence, and can in some respects be considered a transitional dialect, as exemplified in the poems of the Boeotian poet Pindar who wrote in Doric with a small Aeolic admixture. Thessalian likewise had come under Northwest Greek influence, though to
8586-448: Is considered by some linguists to have been closely related to Greek . Among Indo-European branches with living descendants, Greek is often argued to have the closest genetic ties with Armenian (see also Graeco-Armenian ) and Indo-Iranian languages (see Graeco-Aryan ). Ancient Greek differs from Proto-Indo-European (PIE) and other Indo-European languages in certain ways. In phonotactics , ancient Greek words could end only in
8748-403: Is floored and reinforced by a similar bone: the prearticular . In Tanystropheus species with known skull material, both the articular and prearticular contribute equally to a segment of the jaw extending back beyond the level of the jaw joint. This projection, known as a retroarticular process, is enlarged to a similar degree to that of early rhynchosaurs. The skull of Tanystropheus hydroides
8910-413: Is formed by the supraoccipitals , which were presumably fused together as a continuous surface sloping smoothly down to the foramen magnum. In the lower jaw, the dentaries meet each other at a robust symphysis with an interdigitating suture. The front end of the dentary hosts a prominent keel on its lower edge, a unique trait of the species. There are at least 18 dentary teeth; the first three are by far
9072-447: Is its hyperelongate neck, equivalent to the combined length of the body and tail. Tanystropheus has 13 cervical (neck) vertebrae , most of which are massive, though the two closest to the head are smaller and less strongly developed. The atlas (first cervical), which connects to the skull, is a small, four-part bone complex. It consists of an atlantal intercentrum (small lower component) and pleurocentrum (large lower component), and
9234-635: Is known as the Salvatore platform, which is now preserved as the San Salvatore Dolomite . It reconstructs a warm, tropical environment, with the most common fossils belonging to algae and shelled invertebrates . Only the lower portion of the San Salvatore Dolomite is preserved on Monte San Giorgio, corresponding to a particularly shallow and saline period in the history of the platform. Stromatolites and other algal laminations are generally
9396-671: Is larger, with a small axial intercentrum followed by a much larger axial pleurocentrum. The axial pleurocentrum is longer than tall, has a low neural spine set forwards, and small prezygapophyses (front articular plates). The large postzygophyses (rear articular plates) are separated by a broad trough and support pointed epipophyses (additional projections). The third to eleventh cervicals are hyperelongate in T. longobardicus and T. hydroides , ranging from three to 15 times longer than tall. They are somewhat less elongated in T. antiquus , less than 6 times longer than tall. The cervicals gradually increase in size and proportional length, with
9558-508: Is low and extends to a tapered point at the rear. The pubis (lower front hip blade) is vertically oriented, with a small but distinct obturator foramen and a concave rear edge. The lower front tip of the large, fan-shaped ischium (lower rear hip blade) converges towards the pubis, but does not contact it. The large oval-shaped gap between the pubis and ischium is known as the thyroid foramen . Two pairs of large, curved bones, known as heterotopic ossifications or postcloacal bones, sit behind
9720-462: Is nestled further down along the maxilla. The lower edge of the orbit is formed by the jugal , a bone with a slender anterior process (front branch) and a somewhat broader dorsal process (upper branch). There is also a very short pointed posterior process (rear branch) which ends freely and fails to contact any other bone. The shape of the jugal in Tanystropheus is typical for early archosauromorphs;
9882-412: Is positioned near the straight contact with the frontals, one of the few similarities with T. longobardicus . Strong supratemporal fossae excavate into the outer edge of the parietal and define a low sagittal crest along the midline of the skull. This trend is shared with other large archosauromorphs, like Dinocephalosaurus and Azendohsaurus . The supratemporal fenestrae (upper skull holes behind
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#173285618058810044-428: Is rather stiff and ungainly, suggesting a reliance on water. Conversely, the limbs and tail lack most adaptations for swimming and closely resemble their equivalents in terrestrial reptiles. Recent studies have supported an intermediate position, reconstructing Tanystropheus as an animal equally capable on land and in the water. Despite its length, the neck was lightweight and stabilized by tendons , so it would not been
10206-448: Is seen in few other archosauromorphs , namely rhynchosaurs , most allokotosaurs , modern crocodilians , and Teyujagua . The maxilla is triangular, reaching its maximum height at mid-length and tapering to the front and rear. There are up to 14 or 15 teeth in the maxilla, though some individuals have fewer. T. longobardicus is a reptile with heterodont dentition, meaning that it had more than one type of tooth shape. In contrast to
10368-442: Is similar in shape to the humerus and lacks a distinct olecranon (elbow projection). There are four carpals (wrist bones): the ulnare , radiale , and two distal carpals. The ulnare and radiale are large and cuboid, enclosing a small foramen (gap) between them. The larger outer distal carpal connects to metacarpals III and IV, while the much smaller inner distal carpal connects to metacarpals II and III. Metacarpals III and IV are
10530-792: Is the only Swiss stronghold for Microtus savii (Savi's pine vole), and hosts breeding sites for amphibians such as Bufo bufo (common toad), Rana temporaria (common frog), Rana dalmatina (agile frog), Hyla intermedia (Italian tree frog), and other species. Invertebrates are even more diverse, including some species which are very rare in Switzerland, such as Pyrgus armoricanus (Oberthur's grizzled skipper), Euchorthippus declivus (Jersey grasshopper), and Pholidoptera littoralis insubrica (littoral dark bush-cricket). The dry meadows are especially diverse, hosting several species of previously undiscovered or undocumented spiders . Isolated populations of crustaceans and millipedes inhabit
10692-698: The Greek region of Macedonia during the last decades has brought to light documents, among which the first texts written in Macedonian , such as the Pella curse tablet , as Hatzopoulos and other scholars note. Based on the conclusions drawn by several studies and findings such as Pella curse tablet , Emilio Crespo and other scholars suggest that ancient Macedonian was a Northwest Doric dialect , which shares isoglosses with its neighboring Thessalian dialects spoken in northeastern Thessaly . Some have also suggested an Aeolic Greek classification. The Lesbian dialect
10854-665: The Middle Triassic epoch ( Anisian and Ladinian stages), with some exceptions. For example, a vertebra from Nova Scotia was recovered from primarily freshwater sediments. The youngest fossils in the genus are a pair of well-preserved skeletons from the Zhuganpo Formation , a geological unit in China which dates to the earliest part of the Late Triassic (early Carnian stage). The oldest putative fossils belong to "T. antiquus" ,
11016-515: The Middle Triassic , from the latest Anisian to middle Ladinian stages. Though the fossils were initially given the name Macroscelosaurus by Count Georg Zu Münster , the publication containing this name is lost and its genus is considered a nomen oblitum . In 1855, Hermann von Meyer supplied the name Tanystropheus conspicuus , the type species of Tanystropheus , to the fossils. They were later regarded as Tanystropheus fossils undiagnostic relative to other species, rendering T. conspicuus
11178-454: The Permian to Jurassic periods. The rocks forming the mountain dip southwards, with older rocks exposed as one travels north and younger rocks exposed as one travels south. The oldest rocks are Permian volcanic basement material on the mountain's steep north slope. These are followed by Triassic sediments and carbonates at higher elevations on the mountain. Middle Triassic layers are
11340-522: The Villány Mountains of Hungary . The most well-preserved Tanystropheus fossils outside of Monte San Giorgio come from the Guizhou province of China , as described by Li (2007) and Rieppel (2010). They are also among the youngest and easternmost fossils in the genus, hailing from the upper Ladinian or lower Carnian Zhuganpo Formation . Although the postcrania is complete and indistinguishable from
11502-600: The ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek ( c. 1400–1200 BC ), Dark Ages ( c. 1200–800 BC ), the Archaic or Epic period ( c. 800–500 BC ), and the Classical period ( c. 500–300 BC ). Ancient Greek was the language of Homer and of fifth-century Athenian historians, playwrights, and philosophers . It has contributed many words to English vocabulary and has been
11664-402: The coracoid (lower shoulder blade), which is a large oval-shaped plate with a broad glenoid facet (shoulder socket). The humerus (upper arm bone) is straight and slightly constricted at the middle. Near the elbow it is expanded and twisted, with an ectepicondylar groove on its outer edge. The radius (outer forearm bone) is slender and somewhat curved, while the ulna (inner forearm bone)
11826-414: The forebrain , as defined by a bulbous central hollow. The eye was supported by more than 10 rectangular ossicles (tiny plate-like bones) connecting into a scleral ring , though a full reconstruction of the ring, with 18 ossicles, is conjectural. Few details of the braincase and palate (bony roof of the mouth) are known for T. longobardicus . The scant available evidence suggests that these regions of
11988-501: The present , future , and imperfect are imperfective in aspect; the aorist , present perfect , pluperfect and future perfect are perfective in aspect. Most tenses display all four moods and three voices, although there is no future subjunctive or imperative. Also, there is no imperfect subjunctive, optative or imperative. The infinitives and participles correspond to the finite combinations of tense, aspect, and voice. The indicative of past tenses adds (conceptually, at least)
12150-829: The sister taxon to Tanytrachelos , a much smaller tanystropheid from Virginia . Another small tanystropheid, Cosesaurus from Spain , is allied with the Tanystropheus + Tanytrachelos clade in many analyses of the 1980s and 1990s. Within Archosauromorpha, "prolacertiforms" are joined by several other groups. The clade Archosauriformes is a diverse archosauromorph subset including crown group archosaurs and their predatory close relatives such as Euparkeria and Proterosuchus . Stocky Triassic herbivores like rhynchosaurs , Trilophosaurus , and azendohsaurids additionally qualify as archosauromorphs. The bizarre chameleon -like drepanosaurs were also included by many analyses, though more recently they have been reinterpreted as
12312-585: The tuatara . The other side, Archosauromorpha , leads to archosaurs. Cladistics was one of many lines of evidence that helped to demonstrate the dinosaurian origin of birds. This left crocodilians and birds as the two surviving archosaur groups. A series of phylogenetic analyses in the late 1980s and 1990s strongly supported the proposal of Gow (1975). Tanystropheus , Macrocnemus , Protorosaurus , and Prolacerta were always placed as members of Archosauromorpha, closer to archosaurs than to squamates. "Protorosauria" and "Prolacertiformes" were used interchangeably for
12474-412: The 1970s, not long after the field of paleontology was reinvigorated by the " dinosaur renaissance " in the 1960s and beyond. In the 1980s, the advent of cladistics saw a paradigm shift in the field of taxonomy , emphasizing monophyletic clades (all-encompassing groups defined by shared ancestry) over other categorization styles. Phylogenetic analyses were invented to evaluate reptile evolution in
12636-1031: The 5th century BC. Ancient pronunciation cannot be reconstructed with certainty, but Greek from the period is well documented, and there is little disagreement among linguists as to the general nature of the sounds that the letters represent. /oː/ raised to [uː] , probably by the 4th century BC. Greek, like all of the older Indo-European languages , is highly inflected. It is highly archaic in its preservation of Proto-Indo-European forms. In ancient Greek, nouns (including proper nouns) have five cases ( nominative , genitive , dative , accusative , and vocative ), three genders ( masculine , feminine , and neuter ), and three numbers (singular, dual , and plural ). Verbs have four moods ( indicative , imperative , subjunctive , and optative ) and three voices (active, middle, and passive ), as well as three persons (first, second, and third) and various other forms. Verbs are conjugated through seven combinations of tenses and aspect (generally simply called "tenses"):
12798-655: The Anisian-age Moenkopi Formation of Arizona and New Mexico. Several new tanystropheid genera have been named from former Tanystropheus fossils. Fossils from the Anisian Röt Formation in Germany, previously referred to Tanystropheus antiquus, were named as a new genus and species in 2006: Amotosaurus rotfeldensis . In 2011, fossils from the Lipovskaya Formation of Russia were given
12960-490: The Archaic period of ancient Greek (see Homeric Greek for more details): Μῆνιν ἄειδε, θεά, Πηληϊάδεω Ἀχιλῆος οὐλομένην, ἣ μυρί' Ἀχαιοῖς ἄλγε' ἔθηκε, πολλὰς δ' ἰφθίμους ψυχὰς Ἄϊδι προΐαψεν ἡρώων, αὐτοὺς δὲ ἑλώρια τεῦχε κύνεσσιν οἰωνοῖσί τε πᾶσι· Διὸς δ' ἐτελείετο βουλή· ἐξ οὗ δὴ τὰ πρῶτα διαστήτην ἐρίσαντε Ἀτρεΐδης τε ἄναξ ἀνδρῶν καὶ δῖος Ἀχιλλεύς. The beginning of Apology by Plato exemplifies Attic Greek from
13122-557: The Classical period of ancient Greek. (The second line is the IPA , the third is transliterated into the Latin alphabet using a modern version of the Erasmian scheme .) Ὅτι [hóti Hóti μὲν men mèn ὑμεῖς, hyːmêːs hūmeîs, Monte San Giorgio Monte San Giorgio is a wooded mountain, rising to 1,097 m (3,600 feet) above sea level. It has a roughly pyramidal shape, with
13284-512: The Grenzbitumenzone (in Switzerland). It represents the first of several sections on the mountain enriched with well-preserved fossils. The Grenzbitumenzone, especially its shale layers, is enriched with organic material derived from cyanobacteria . This accumulation of organic material presumably made the bottom of the basin anoxic or dysoxic, with low oxygen levels in the seawater. The only fossils of seabed-living organisms belong to Daonella ,
13446-633: The Jurassic. Its Jurassic sediments are now eroded away to reveal older Triassic and Permian rocks. Conversely, Monte Generoso , immediately to the east of Monte San Giorgio, is composed mostly of Jurassic basinal sediments. Outcrops of Jurassic sediments are also seen close to the Po Plain, at the south edge of Monte San Giorgio (in a broad sense). The productive " marble " quarries found south of Monte San Giorgio actually mined non- metamorphosed limestone, rather than true marble. These limestone units were formed at
13608-548: The Ladinian, though the Grenzbitumenzone itself transitioned into a less fossiliferous formation known as the San Giorgio Dolomite . This formation has lower organic content, no shale, and only a few fragmentary fossils. Higher organic content and finer laminations return a short while later, forming the lower part of the fossil-rich Meride Limestone . The Meride Limestone probably represents a period of increased instability on
13770-484: The Museo Cantonale di Scienze Naturali di Lugano (MCSN). Rupert Wild reviewed and redescribed all specimens known at the time via several large monographs in 1973/4 and 1980. In 2005, Silvio Renesto described a T. longobardicus specimen from Switzerland which preserved the impressions of skin and other soft tissue. Five new MSNM specimens of T. longobardicus were described by Stefania Nosotti in 2007, allowing for
13932-468: The Spinirolo plant was built for the purpose of processing the shale into saurol, an ichthyol -like skin ointment. Saurol production and mining continued until the 1950s, and operations went bankrupt in 1960. Minerals such as barite , fluorite , and galena were also prospected on the mountain during the 20th century. Italian paleontologist Giulio Curioni first mentioned that fossils were present on
14094-616: The Ticino dry meadows, a unique biome with over 100 plant and species, 38 of which are rare or endangered within Switzerland. Carex humilis (dwarf sedge) and Molinia caerulea arundinacea (tall moor grass) are the most common grasses , while Monte San Giorgio supports the few Swiss populations of wildflowers such as Adenophora liliifolia , Gladiolus imbricatus , Iris graminea , Lotus herbaceus , and Danthonia alpina . 102 species of vertebrates are found on Monte San Giorgio, 37 of which are endangered in Switzerland. The mountain
14256-526: The World Heritage Site was expanded further, adding 240.34 ha of land from the Italian communes of Besano , Porto Ceresio , and Viggiù. These communes, alongside Clivio and Saltrio, were also included within an 1818.45 ha Italian buffer zone. This additional land brings the total area of UNESCO protected property to 1089.34 ha and the total buffer zone area to 3207.45 ha. Inclusion of the Italian territory
14418-496: The aftermath of the Variscan orogeny . These volcanic rocks are mainly reddish rhyolite and andesite with a porphyritic texture, produce large crystals of quartz, barite, and fluorite. The Permian basement rocks are terminated by an unconformity , an erosional surface succeeded by Triassic sediments. These following Triassic sediments are siliciclastic and terrestrial in origin, mainly sandstone and conglomerate eroded from
14580-550: The aorist. Following Homer 's practice, the augment is sometimes not made in poetry , especially epic poetry. The augment sometimes substitutes for reduplication; see below. Almost all forms of the perfect, pluperfect, and future perfect reduplicate the initial syllable of the verb stem. (A few irregular forms of perfect do not reduplicate, whereas a handful of irregular aorists reduplicate.) The three types of reduplication are: Irregular duplication can be understood diachronically. For example, lambanō (root lab ) has
14742-414: The archosauromorph subgroup encompassing these superficially lizard-like reptiles. Some authors preferred "Protorosauria" for its priority . Most others used "Prolacertiformes" arguing that "Protorosauria" was a name that carried too much historical baggage, since it had previously encompassed non-archosauromorph "euryapsids" like Araeoscelis . As a "prolacertiform", Tanystropheus is typically considered
14904-419: The augment when it was word-initial. In verbs with a preposition as a prefix, the augment is placed not at the start of the word, but between the preposition and the original verb. For example, προσ(-)βάλλω (I attack) goes to προσ έ βαλoν in the aorist. However compound verbs consisting of a prefix that is not a preposition retain the augment at the start of the word: αὐτο(-)μολῶ goes to ηὐ τομόλησα in
15066-458: The basis of their strongly divergent skull anatomy. When T. longobardicus was first described in 1886, it was initially mistaken for a pterosaur and given the name " Tribelesodon ". Starting in the 1920s, systematic excavations at Monte San Giorgio unearthed many more Tanystropheus fossils, revealing that the putative wing bones of "Tribelesodon" were actually neck vertebrae. Most Tanystropheus fossils hail from marine or coastal deposits of
15228-527: The best known records of marine life in the Middle Triassic period. The Italian region west of Poncione d'Arzo ( Porto Ceresio ) was added as an extension to the World Heritage Site in 2010. Humans have inhabited Monte San Giorgio at least since the area's equivalent of the Neolithic Period , around 6,000 years ago. The south side of the mountain is home to Tremona-Castello Archaeological Park ,
15390-438: The center of Greek scholarship, this division of people and language is quite similar to the results of modern archaeological-linguistic investigation. One standard formulation for the dialects is: West vs. non-West Greek is the strongest-marked and earliest division, with non-West in subsets of Ionic-Attic (or Attic-Ionic) and Aeolic vs. Arcadocypriot, or Aeolic and Arcado-Cypriot vs. Ionic-Attic. Often non-West
15552-532: The cervical ribs than the dorsal ribs. The elongation of Tanystropheus 's neck is mostly a consequence of particular vertebrae lengthening. This is a contrast to trachelosaurids such as Dinocephalosaurus , which achieve a long neck by the addition of numerous cervicals, for a total cervical count exceeding 30. Nevertheless, Tanystropheus does have more vertebrae in its neck than typical archosauromorphs. Protorosaurus , for example, has only seven cervicals, while Macrocnemus and Prolacerta have eight. To achieve
15714-410: The cervicals. Though their neural spines are taller than those of the cervicals, they are still rather short. The dorsal ribs are double-headed close to the shoulder and single-headed in the rest of the torso, sitting on stout transverse processes projecting outwards from the front half of each vertebra. More than 20 angled rows of gastralia extend along the belly, each gastral element represented by
15876-611: The dialect of Sparta ), and Northern Peloponnesus Doric (including Corinthian ). All the groups were represented by colonies beyond Greece proper as well, and these colonies generally developed local characteristics, often under the influence of settlers or neighbors speaking different Greek dialects. After the conquests of Alexander the Great in the late 4th century BC, a new international dialect known as Koine or Common Greek developed, largely based on Attic Greek , but with influence from other dialects. This dialect slowly replaced most of
16038-440: The edge of the cranium. In T. longobardicus , the paroccipital processes are shorter and narrower at their base. The stapes , a bone which transmits vibrations from the ear to the braincase, is slender and splits into two small prongs where it contacts the opisthotic. The opisthotic merges forwards into the prootic , which extensively contacts the parabasisphenoid and hosts a range of larger nerve foramina. The prootic forms much of
16200-428: The eye) are wide and semi-triangular, exposed almost entirely from above. The postorbital has large and blocky ventral and medial processes (lower and inward branches), which meet at a sharper angle than in any other early archosauromorph. The jugal, conversely, is basically indistinguishable from that of T. longobardicus . The squamosal is deep and rectangular when viewed from the side, with little differentiation between
16362-419: The fifth toe was very long, filling a metatarsal-like role as seen in other tanystropheids. Knowledge on the anatomy of Tanystropheus was transformed by Bernhard Peyer 's discoveries in the 1920s and 1930s, but its relationship to other reptiles remained enigmatic for much of the 20th century. Most paleontologists (including modern authorities) agree that Tanystropheus was closely related to Macrocnemus ,
16524-421: The forelimbs, though similar in overall structure and proportions. The femur (thigh bone) is long, slender, and sigmoid (curved at both ends). It has a longitudinal muscle crest for muscle attachment (the internal trochanter ) on its underside, and it contacts the acetabulum (hip socket) at a broad smooth joint. The tibia and fibula (shin bones) are straight, with the former much thicker and more expanded at
16686-655: The fossil-rich basin and carbonate system of the Middle Triassic. During the Carnian (around 237 to 227 Ma), the first stage of the Late Triassic, carbonate platforms were replaced by shallow-water and coastal sediments. This formation, the Pizzella Marls , is diagnosed by a higher amount of siliciclastics (sediments eroded down from terrestrial rocks) and evaporites (mineral deposits from dried water), such as gypsum . In
16848-627: The fossils of Monte San Giorgio, no skull material is preserved, and their younger age precludes unambiguous placement into any Tanystropheus species. The Chinese material includes a large morphotype ( T. hydroides? ) specimen, GMPKU-P-1527, and an indeterminate juvenile skeleton, IVPP V 14472. Indeterminate Tanystropheus remains are also known from the Jilh Formation of Saudi Arabia and various Anisian-Ladinian sites in Spain , France , Italy, and Switzerland. The youngest Tanystropheus fossil in Europe
17010-425: The front edge of the interclavicle , a plate-like bone at the center of the chest with a rhombic (broad, diamond-shaped) front region followed by a long stalk at the rear. The interclavicle is rarely preserved and its connections to the rest of the pectoral girdle are mostly inferred from Macrocnemus . The scapula (upper shoulder blade) has the form of a large semicircular plate on a short, broad stalk. It lies above
17172-404: The front edge of the paroccipital process, akin to the condition in archosauriforms. Another archosauriform-like feature is the presence of a laterosphenoid , an additional braincase component in front of the prootic and above the exit hole for the trigeminal nerve (also known as cranial nerve V). The laterosphenoid is small, similar to that of Azendohsaurus . The upper rear part of the braincase
17334-575: The fully aquatic plesiosaurs . 19th century excavations at Monte San Giorgio , a UNESCO world heritage site on the Italy - Switzerland border, revealed a fragmentary fossil of an animal with three-cusped (tricuspid) teeth and elongated bones. Monte San Giorgio preserves the Besano Formation (also known as the Grenzbitumenzone), a late Anisian -early Ladinian lagerstätte recognised for its spectacular fossils. In 1886, Francesco Bassani interpreted
17496-529: The growing carbonate platforms, sending surges of carbonate grains into the basin. Skeletons tend to be even better preserved than in the Grenzbitumenzone, suggesting that the basin deepened further or acquired extensive microbial mats. A section of dolomite, the "Dolomitband", forms the top of the Lower Meride Limestone. It also marks the start of the Upper Meride Limestone, which is similar to
17658-470: The hips in about half of known specimens preserving the area. They occupy the base of the tail, a region which lacks chevrons. These bones are possibly sexually dimorphic , and have also been reported in the small American tanystropheid Tanytrachelos . Heterotopic ossifications may be linked to reproductive biology, supporting reproductive organs (if they belong to males) or an egg pouch (if they belong to females). The hindlimbs are significantly larger than
17820-556: The historical Dorians . The invasion is known to have displaced population to the later Attic-Ionic regions, who regarded themselves as descendants of the population displaced by or contending with the Dorians. The Greeks of this period believed there were three major divisions of all Greek people – Dorians, Aeolians, and Ionians (including Athenians), each with their own defining and distinctive dialects. Allowing for their oversight of Arcadian, an obscure mountain dialect, and Cypriot, far from
17982-472: The historical circumstances of the times imply that the overall groups already existed in some form. Scholars assume that major Ancient Greek period dialect groups developed not later than 1120 BC, at the time of the Dorian invasions —and that their first appearances as precise alphabetic writing began in the 8th century BC. The invasion would not be "Dorian" unless the invaders had some cultural relationship to
18144-405: The hypothesis that they represent separate species. A 2020 study found numerous differences between the skulls of large and small specimens, formalizing the proposal to divide the two into separate species. Moreover, a histological investigation revealed that one small specimen, PIMUZ T 1277, was a skeletally mature adult at a length of only 1.5 meters (4.9 ft). The larger one-cusped morphotype
18306-423: The jaw. Although it is plausible that a small coronoid bone could be present in front of the surangular, evidence is ambiguous at best for all Tanystropheus species. A sheathe-like bone, the angular , is well-exposed under the dentary and surangular, though sutures between these bones are difficult to interpret with certainty. The joint at the back of the jaw lies on the articular , a lumpy rectangular bone which
18468-426: The knee. The large proximal tarsals (ankle or heel bones contacting the shin) consist of a rounded calcaneum and a blocky astragalus , which meet each other along a straight or shallowly indented contact in most specimens. Like most non-aquatic reptiles, a set of small pebble-shaped distal tarsals are present between the proximal tarsals and the foot bones. Tanystropheus has a reduced number of distal tarsals: only
18630-403: The largest bones in the hand, followed closely by metacarpal II. Metacarpals I and V are both short. The hand's phalangeal formula (joints per finger) is 2-3-4-4-3. The terminal phalanges (fingertips) may have formed thick, blunt claws. The components of the pelvis (hip) are proportionally small, though their shape is unremarkable relative to other tanystropheids. The ilium (upper hip blade)
18792-402: The largest teeth in the skull, forming the lower half of the interlocking "fish trap" with the premaxilla. Most other teeth in the dentary are small, with the exception of the tenth tooth, which juts up to pierce the maxilla. The remainder of the jaw contains the same set of bones as in T. longobardicus , but some details differ in T. hydroides . For example, the splenial is plate-like and covers
18954-479: The late 2010s. The proportions of T. antiquus fossils are easily distinguishable, and it is currently considered a valid species of archosauromorph, though its referral to the genus Tanystropheus has been questioned . The Gogolin Formation ranges from the upper Olenekian (latest part of the Early Triassic ) to the lower Anisian in age. Assuming they belong within Tanystropheus , the fossils of T. antiquus may be
19116-431: The lower part of the formation but has only a few fossiliferous sections. The Upper Meride Limestone eventually becomes dominated by very finely-laminated marls and shales with increased clay content. This clay-rich interval, indicative of increased terrestrial runoff within the shrinking basin, is known as the "Kalkschieferzone". By the beginning of the Late Triassic, a major marine regression (sea level fall) threatened
19278-413: The maxilla. There is some variation in the number of each tooth shape, and some individuals may have up to 11 conical teeth. The inner surface of the dentary is joined by a splint-shaped bone, the splenial , at its lower edge. The splenial was most likely not visible in lateral view. At its rear, the dentary seems to be partially overlapped by the surangular , a bone which comprises much of the rear part of
19440-456: The middle Ladinian Erfurt Formation (Lettenkeuper) of Germany was described in 1846 as one of several fossils gathered under the name " Zanclodon laevis ". Though likely the first Tanystropheus fossil to be discovered, the vertebra is now lost, and surviving jaw fragments and other fossil scraps of " Zanclodon laevis" represent indeterminate archosauriforms with no relation to Tanystropheus . Tanystropheus vertebrae have also been found in
19602-500: The morphology of the genus. The Miedary locality represents an isolated brackish body of water close to the coast, and the abundance of Tanystropheus fossils suggests that it was an animal well-suited for this kind of habitat. In the late 1900s, Friedrich von Huene named several dubious Tanystropheus species from Germany and Poland. T. posthumus , from the Norian of Germany, was later reevaluated as an indeterminate theropod vertebra and
19764-469: The most fossiliferous and extraordinary from a global perspective, and are encompassed by the protected area north of Meride. South of Meride, they are replaced by Late Triassic coastal sediments which give way to Early Jurassic limestone overlooking the Po Plain. The stratigraphically lowest rocks exposed on Monte San Giorgio are Lower Permian in age, around 290-280 Ma (million years old). They are remnants of early rifting and volcanic activity in
19926-590: The most well-described non- archosauriform archosauromorphs, known from numerous fossils, including nearly complete skeletons. Some species within the genus may have reached a total length of 6 meters (20 ft), making Tanystropheus the longest non-archosauriform archosauromorph as well. Tanystropheus is the namesake of the family Tanystropheidae , a clade collecting many long-necked Triassic archosauromorphs previously described as " protorosaurs " or " prolacertiforms ". Tanystropheus contains at least two valid species as well as fossils which cannot be referred to
20088-439: The mountain in 1847. The first paper focusing on Monte San Giorgio fossils in particular was published by Emilio Cornalia in 1854. Small excavations by Milanese paleontological societies in 1863 and 1878 provided more specific context on the paleontology of Monte San Giorgio. Shale extraction brought the fossil deposits to the attention of University of Zurich paleontologist Bernhard Peyer in 1919. Peyer and his associates began
20250-582: The new genus Coelophysis . Authentic Tanystropheus specimens from the Makhtesh Ramon in Israel were described as a new species, T. haasi , in 2001. However, this species may be dubious due to the difficulty of distinguishing its vertebrae from T. conspicuus or T. longobardicus . Another new species, T. biharicus , was described from Romania in 1975. It has also been considered possibly synonymous with T. longobardicus . A Tanystropheus -like vertebra from
20412-628: The new genus and species Augustaburiania vatagini by A.G. Sennikov. He also named the new genus Protanystropheus for T. antiquus, though a few studies continued to retain that species within Tanystropheus . Tanystropheus fossai, from the Norian -age Argillite di Riva di Solto in Italy, was given its own genus Sclerostropheus in 2019. Tanystropheus was one of the longest known non-archosauriform archosauromorphs. Vertebrae referred to " T. conspicuus " may correspond to an animal up to five or six meters (16.4 to 20 feet ) in length . T. hydroides
20574-549: The ninth cervical typically being the largest vertebra in the skeleton. In general structure, the elongated cervicals resemble the axial pleurocentrum. However, the axis also has a keel on its underside and an incomplete neural canal, unlike its immediate successors. In the rest of the cervicals, all but the front of each neural spine is so low that it is barely noticeable as a thin ridge. The zygapophyses are closely set and tightly connected between vertebrae. The epipophyses develop into hooked spurs. The cervicals are also compressed from
20736-499: The older dialects, although the Doric dialect has survived in the Tsakonian language , which is spoken in the region of modern Sparta. Doric has also passed down its aorist terminations into most verbs of Demotic Greek . By about the 6th century AD, the Koine had slowly metamorphosed into Medieval Greek . Phrygian is an extinct Indo-European language of West and Central Anatolia , which
20898-657: The oldest in the genus. Specimens likely referable to T. antiquus are also known from throughout Germany and the fossiliferous Winterswijk site in the Netherlands . In the 1880s, E.D. Cope named three supposed new Tanystropheus species ( T. bauri , T. willistoni , and T. longicollis ) from the Late Triassic Chinle Formation in New Mexico . However, these fossils were later determined to be tail vertebrae belonging to theropod dinosaurs, which were named under
21060-605: The only fossils found in the Lower Salvatore Dolomite. Near the end of the Anisian, the southern edge of the Salvatore platform deepens abruptly, giving way to a more sterile basin developed between carbonate platforms. The basin is now preserved as a relatively narrow band of dark dolomite and shale, running east to west along the edge of Monte San Giorgio. This formation has been called the Besano Formation (in Italy) or
21222-487: The perfect stem eilēpha (not * lelēpha ) because it was originally slambanō , with perfect seslēpha , becoming eilēpha through compensatory lengthening. Reduplication is also visible in the present tense stems of certain verbs. These stems add a syllable consisting of the root's initial consonant followed by i . A nasal stop appears after the reduplication in some verbs. The earliest extant examples of ancient Greek writing ( c. 1450 BC ) are in
21384-461: The pleurapophyses decreases until they disappear between the eighth and thirteenth caudal. The height of the neural spines also decreases gradually down the tail. A row of long chevrons is present under a short portion of the tail, though not immediately behind the hips. The pectoral girdle (shoulder girdle) has a fairly standard form shared with other tanystropheids . The clavicles (collarbones) were curved and slightly twisted rods. They lie along
21546-419: The premaxilla, which extends below and behind the nares (nostril holes). The nasals (bones at the top edge of the snout) are poorly known, but were likely narrow and flat. A 2020 reinvestigation revealed that the front part of the nasals and the inner spur of the premaxillae are too short to keep the nares divided. This leaves a single central narial opening for the nostrils, opening upwards. An undivided naris
21708-668: The present day, managed by the MSNM, University of Milan (UNIMI), and the Museo Cantonale di Storia Naturale di Lugano (Cantonal Museum of Natural History, MCSN). Over 21,000 fossil specimens have been collected in total by 2010. In 2003, the Monte San Giorgio was listed as a UNESCO World Heritage Site, with 849 ha ( hectares ) of protected land from the Swiss communes of Meride , Brusino Arsizio , and Riva San Vitale . This protected area
21870-444: The pterygoid). The remainder of the braincase is fully fused together into a strongly ossified composite bone, and its constituents must be estimated by comparison to other reptiles. The exoccipitals , which mostly encompass the foramen magnum (spinal cord hole), are perforated with nerve foramina. Each exoccipital merges outwards into the opisthotic , which sends out a straight, elongated paroccipital process (thick outer branch) to
22032-416: The quadrate at the rear lower corner of the skull. Nevertheless, a quadratojugal was likely present in the species, since it occurs in T. hydroides and nearly every other early archosauromorph. The paired frontals (skull roof bones above the orbits) have been described as "axe-shaped flanges", projecting broad curved plates above each orbit. Together, the frontals are narrowest at the front, terminating at
22194-411: The reptile family tree near the ancestry of archosaurs , a diverse group of reptiles with lightweight skulls and serrated teeth set in deep sockets. Dinosaurs are among the most famous subset of archosaurs, as are modern crocodilians and their prehistoric ancestors. Several newly discovered "prolacertiforms", including Tanystropheus -, Protorosaurus -, and Prolacerta -like species, were described in
22356-488: The same general time as the Moltrasio Limestone. The fauna and flora of Monte San Giorgio are diverse, with some species found nowhere else in Switzerland. The prevailing ecosystems are mixed broadleaf forests and meadows influenced by the mountain's sub-Mediterranean climate . Monte San Giorgio is one of the southernmost areas of Switzerland, with mild winters, high humidity, and many hours of sunshine. Due to
22518-421: The side, so they are taller than wide. Many specimens have a longitudinal lamina (ridge) on the side of each cervical. Ventral keels return to vertebrae in the rear half of the neck. All cervicals, except potentially the atlas, connected to holocephalous (single-headed) cervical ribs via facets at their front lower corner. Each cervical rib bears a short stalk connecting to two spurs running under and parallel to
22680-436: The simple fang-like premaxillary teeth, most or all of the maxillary teeth have a distinctive tricuspid shape, with the crown split into three stout triangular cusps (points). The cusps are arranged in a line from front-to-back, with the central cusp larger than the other two cusps. Among Triassic reptiles, early pterosaurs such as Eudimorphodon developed an equivalent tooth shape, and tricuspid teeth can also be found in
22842-429: The skull are rather unspecialized in this species. The vomers (front components of the palate) are narrow and dotted with at least nine tiny teeth. The succeeding palatine and pterygoid bones are also supplied with rows of teeth: up to six relatively large teeth in the former and at least 12 small teeth in the latter. Teeth on the vomers, palatines, and pterygoids are the norm for early archosauromorphs and reptiles as
23004-481: The skull). One idea was that Tanystropheus and kin (particularly Macrocnemus and Prolacerta ) were ancestral to " lacertilians ", an antequated term for lizards . This hypothesis was supported up until the 1980s by German and Swiss paleontologists, including Rupert Wild, and Peyer's successor at Zürich, Emil Kuhn-Schnyder . The other idea maintained that Tanystropheus was a "protorosaur", closer to Protorosaurus and Araeoscelis and unrelated to Prolacerta . This
23166-453: The species from Monte San Giorgio. Peyer's discoveries allowed Tribelesodon longobardicus to be recognised as a non-flying reptile, more than 40 years after its original description. Its supposed elongated finger bones were recognized as neck vertebrae, which compared favorably with those previously described as Tanystropheus from Germany and Poland. Thus, Tribelesodon longobardicus was renamed to Tanystropheus longobardicus and its anatomy
23328-417: The species have been reinterpreted as displaced postorbitals. The quadrate bone , which forms the rear edge of the skull and upper half of the jaw joint, is wide and tall. It has a strong lateral crest and a low pterygoid ramus (a vertical internal plate, articulating with the pterygoid bone in the roof of the mouth). No fossils of T. longobardicus preserve a quadratojugal , a bone which normally lies along
23490-477: The succeeding Norian stage (around 227 to ~208 Ma), carbonate platforms and rising sea levels were renewed with vigor, depositing a massive expanse of carbonate known as the Dolomia Principale or Hauptdolomit. The Dolomia Principale is a brittle, crystalline rock mass which was fractured by normal faults not long after it was first formed. This is an early pulse of an overall extensional tectonic regime,
23652-517: The syllabic script Linear B . Beginning in the 8th century BC, however, the Greek alphabet became standard, albeit with some variation among dialects. Early texts are written in boustrophedon style, but left-to-right became standard during the classic period. Modern editions of ancient Greek texts are usually written with accents and breathing marks , interword spacing , modern punctuation , and sometimes mixed case , but these were all introduced later. The beginning of Homer 's Iliad exemplifies
23814-538: The tall suture with the postorbital and the small suture with the quadratojugal. As a result, most of the posterior skull is clustered together, and the infratemporal fenestra is reduced to a small diagonal hole. The quadratojugal is a curved sliver of bone which twists back alongside the quadrate. Relative to T. longobardicus , the quadrate has a larger pterygoid ramus and a strongly hooked projection at its upper extent. The palate of T. hydroides has several unique traits. The vomers are wide and tongue-shaped, each hosting
23976-413: The tip of the snout) has a long tooth row, with six teeth. The premaxillary teeth are conical, fluted by longitudinal ridges, and have subthecodont implantation, meaning that the inner wall of each tooth socket is lower than the outer wall. The premaxilla meets the maxilla (the succeeding toothed bone) along a long, slanted contact. This shape is produced by an elongated postnarial process (rear prong) of
24138-465: The underdeveloped posterior process indicates that the margin of the infratemporal fenestra (lower skull hole behind the eye) was incomplete and open from below. The postorbital bone , which links the jugal to the top of the skull, was tall and roughly boomerang-shaped, though poor preservation obscures some details. The squamosal bone , which extends behind the postorbital, is also poorly known in T. longobardicus , and many supposed squamosal fossils in
24300-491: The underlying volcanic material. " Servino " is the name given to older sediments from the Early Triassic (about 252-247 Ma). Slightly younger sediments from the late Anisian (the first stage of the Middle Triassic, 247-242 Ma) are called the Bellano Formation . The Servino and Bellano Formation can be difficult to differentiate, but together they reconstruct a period of transgression (rising sea levels) encroaching onto
24462-559: The unusual tricuspid fossil as a pterosaur , which he named Tribelesodon longobardicus . The holotype specimen of Tribelesodon longobardicus was stored in the Museo Civico di Storia Naturale di Milano (Natural History Museum of Milan ), and was destroyed by allied bombing of Milan in World War II . Excavations by University of Zürich paleontologist Bernhard Peyer in the late 1920s and 1930s revealed many more complete fossils of
24624-700: The variation in underlying geology, both acidic and alkaline soils are developed, supporting different vegetation communities. The rhyolite -based northern slope is mostly covered by Castanea sativa (sweet chestnut), Quercus petraea (sessile oak), and Fraxinus excelsior (European ash). The dolomite -based southern slope is more diverse in its plant life and soil quality, with common plants including Carpinus betulus (common hornbeam), Ostrya carpinifolia (European hop-hornbeam), Tilia (linden), Asperula taurina , Quercus pubescens (pubescent oak), and Fraxinus ornus (manna ash). The driest and most alkaline soils of Monte San Giorgio are home to
24786-450: The vertebrae. The forward-projecting spurs were short and stubby, while the rear-projecting spurs were extremely narrow and elongated, up to three times longer than their respective vertebrae. This bundle of rod-like bones running along the neck afforded a large degree of rigidity. The 12th cervical and its corresponding ribs, though still longer than tall, are notably shorter (from front-to-back) than their predecessors. The 12th cervical has
24948-475: Was Aeolic. For example, fragments of the works of the poet Sappho from the island of Lesbos are in Aeolian. Most of the dialect sub-groups listed above had further subdivisions, generally equivalent to a city-state and its surrounding territory, or to an island. Doric notably had several intermediate divisions as well, into Island Doric (including Cretan Doric ), Southern Peloponnesus Doric (including Laconian ,
25110-422: Was around the same size, with the largest specimens at an estimated length of 5.25 meters (17.2 feet). T. longobardicus was significantly smaller, with an absolute maximum size of two meters (6.6 feet). Despite the large size of some Tanystropheus species, the animal was lightly built. One mass estimate used crocodiles as a density guideline for a 3.6 meter (11.8 feet)-long model of a Tanystropheus skeleton. For
25272-554: Was bombed in 1943, destroying its collection of Monte San Giorgio specimens. Fossil excavations resumed in 1950 under the helm of Emil Kuhn-Schnyder , Peyer's successor and former student. Kuhn-Schnyder established the Palaeontological Institute and Museum of the University of Zurich (PIMUZ) in 1956, which now hosts over 15,000 specimens of Monte San Giorgio fossils. Collection campaigns have continued intermittently up until
25434-494: Was motivated for its paleontological heritage. Each side of the site is managed by separate Swiss and Italian organizations, as well as a transnational board which moderates between the management organizations. The site is not in any particular danger from overutilization or degradation, so management is mainly related to closely-regulated fossil excavations, promotion, and maintenance of low-impact tourism facilities. Monte San Giorgio fossils are collected, curated, and displayed by
25596-605: Was named as a new species, Tanystropheus hydroides (referencing the Hydra of Greek mythology ), while the smaller tricuspid morphotype retains the name T. longobardicus . The first Tanystropheus specimens to be described were found in the mid-19th century. They included eight large vertebrae from the Upper Muschelkalk of Germany , and a partial skeleton from the Lower Keuper of Poland . These geological units occupy part of
25758-579: Was popular among American paleontologists like Alfred Romer . Some publications from the mid-20th century argued that "protorosaurs" were "euryapsids" (reptiles with only an upper temporal fenestra ) related to sauropterygians, though later accounts admitted that Euryapsida was likely polyphyletic , with its members lacking a common ancestor. In 1975, a paper by South African paleontologist C.E. Gow argued that none of these hypotheses were entirely correct. He proposed that Prolacerta , and by extension Macrocnemus and Tanystropheus , occupied an extinct spur on
25920-503: Was revised into a long-necked, non-pterosaur reptile. Specimen PIMUZ T 2791, which was discovered in 1929, has been designated as the neotype of the species. Well-preserved T. longobardicus fossils continue to be recovered from Monte San Giorgio up to the present day. Fossils from the mountain are primarily stored at the rebuilt Museo Civico di Storia Naturale di Milano (MSNM), the Paleontological Museum of Zürich (PIMUZ), and
26082-435: Was surrounded by a 1389 ha buffer zone overlapping six additional communes. The nomination of Monte San Giorgio was inspired by its exceptional paleontological value, with multiple fossiliferous levels preserving among the best records of Middle Triassic life in the world. Monte San Giorgio also presents a link between local geology and culture, as well as unique ecological heritage relative to the rest of Switzerland. In 2010,
26244-420: Was the namesake of yet another term introduced into the convoluted space of reptile taxonomy: " Prolacertiformes ". As the century progressed, two competing hypotheses for the affinities of Tanystropheus developed from the groundwork set by Peyer. Both hypotheses were justified by patterns of skull fenestration (the shape of holes in the skull behind the eye) and cranial kinesis (the flexibility of joints within
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