Hamipterus is an extinct genus of pteranodontoid pterosaur from the Early Cretaceous of northwestern China. It is known from a single type species, Hamipterus tianshanensis.

 Уникальную находку сделали китайские палеонтологи, изучающие ископаемую фауну хребта Тянь-Шань. Среди десятков скелетов нового вида птерозавров они обнаружили несколько сохранивших свою исходную форму яиц этих рептилий. Ничего подобного ученый мир до сих пор не видел.

 Пять продолговатых яиц небольшого размера стали настоящей сенсацией. До сих пор науке были известны лишь четыре птерозавровых яйца, причем все они в процессе окаменения были смяты в лепешку. Обнаруженные же на южных склонах Тянь-Шаня яйца исключительно хорошо сохранились и внешне напоминают яйца некоторых современных змей и ящериц, рассказали ученые.

 Вместе с ними были обнаружены и остатки по меньшей мере 40 особей нового вида птерозавров, названного Hamipterus tianshanensis. "Большинство окаменелостей оказались относительно нетронутыми, что благоприятствовало сборам полных скелетов этих птерозавров", – рассказал палеонтолог Ван Сяолинь, автор открытия. Как показали его исследования, хамиптерусы достигали четырех метров в размахе крыльев, имели заостренные зубы и удлиненный череп с гребнем на макушке. Кроме того, самцы и самки различались своим внешним видом, являясь довольно редким в каменной летописи примером полового диморфизма. Питались эти птерозавры, по всей вероятности, рыбой.

 "Данные окаменелости проливают свет на репродуктивную стратегию, онтогенез и поведение птерозавров, – констатируют авторы исследования. – Они демонстрируют половой диморфизм, благодаря которому самцы и самки отличались размерами гребня, его формой и прочностью... Мы полагаем, что эти новые птерозавры образовывали колонии и обладали стайным поведением, которое, возможно, являлось общим трендом, по крайней мере, для птеродактилоидных птерозавров".

 Директор Института палеонтологии позвоночных и палеоантропологии Китайской академии наук Чжунхэ Чжоу назвал место находки Hamipterus tianshanensis "самым важным местонахождением птерозавров изо всех когда-либо найденных".

 Исходя из характера отложений, исследователи полагают, что крупная колония птерозавров была уничтожена сильным штормом, разразившимся в этих местах в начале мелового периода, около 120 млн лет назад. До этой катастрофы летающие ящеры большой шумной колонией населяли окрестности живописного озера.

 "Это настоящий почетный трофей, который может считаться одним из лучших среди известных находок птерозавров. Он превращает эти места в основной район для исследований этих животных, – рассказал доктор Дэйв Хон. – Имеющиеся данные, естественно, на сегодняшний день далеко не полны, но сам факт находки вместе остатков многочисленных особей и яиц зафиксирован впервые в истории".

 По предварительным оценкам, в этих местах могут быть найдены остатки и других меловых позвоночных, в том числе и птиц.

 Репродукции (1, 2, 3):

 Ископаемые останки (1, 2, 3, 4, 5):

Tags: Вымершие рептилии, Мел, авеметатарзалии, архозавроморфы, архозавры, диапсиды, монофенестраты, орнитохейройды, птеранодонтойды, птеродактили, птерозавры

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 Moganopterus is an extinct genus of boreopterid pterosaur from the Early Cretaceous of western Liaoning Province, China.

 The fossil of Moganopterus was discovered at the village of Xiaosanjiazi near the town of Lamadong in Liaoning Province. In 2012 it was named and described by Lü Junchang, Pu Hanyong, Xu Li, Wu Yanhua and Wei Xuefang as the type species Moganopterus zhuiana. The generic name is derived from the legendary sword couple Gan Jiang and Mo Ye, in reference to the blade-like jaws, and a Latinised Greek πτερόν, pteron, "wing". The specific name honours Ms. Zhu Haifen, who made the specimen available to science.

 The holotype, 41HIII0419, was uncovered in a layer of the Yixian Formation, dating from the Aptian, about 125 million years old. It consists of an almost complete skull with lower jaws and the second to fourth neck vertebrae. The fossil is compressed on a slab and counterslab, the splitting of the two plates having damaged some bones. The specimen is part of the collection of the Geological Museum of Henan.

 Moganopterus is a large pterosaur. The skull has a preserved length of about ninety-five centimetres and the longest preserved neck vertebra, the fourth, a length of 14.5 centimetres. The skull is the largest known of any toothed pterosaur. The size of skull and neck indicates a wing span of at least five metres and probably of over 7 metres (23 ft), making Moganopterus one of the largest known pterosaurs.

 Apart from the size, the describers established some diagnostic traits. The jaws are very elongated and have straight edges. The total number of teeth in the skull is at least sixty-two. The large skull opening, the fenestra nasoantorbitalis, is rectangular and represents 22% of the snout length. The back of the skull bears a long and narrow parietal crest, sticking out at an angle of 15° to the longitudinal skull axis. Not taking into account the crest, the skull is 11.5 times longer than tall. The neck vertebrae are five times longer than high.

 Moganopterus shows an extreme elongation of the upper and lower jaws. The back of the skull is just six centimetres high and its top gradually descends towards the pointed snout tip. On the front of the snout a low triangular crest is present, five centimetres long and six millimetres tall. The profile of the skull is continued by a narrow crest sticking out at the back. It is unknown whether this crest was flat or rod-like; its length cannot be determined because it reaches the edge of the slabs. The lower jaws, lacking a keel, have a length of 68.5 centimetres. They are about as tall as the snout and have a pointed tip.

 The jaws are lined with long conical pointed teeth, up to thirty-one millimetres in length, slightly recurved and more or less oriented vertically. The describers estimated there were fifteen teeth in the upper jaw and seventeen in the lower jaw for a total of sixty-four, which closely matches the number of sixty-two actually found. The teeth rows stretch from the very front of the head until the back edge of the fenestra nasoantorbitalis. They are associated with oblique cellular structures visible in the bone of the upper and lower jaws, the nature of which has not been determined. Hollow structures, reinforced by struts, can also be seen in the parietal crest and the vertebrae. The preserved neck vertebrae have a length of eleven millimetres, eleven centimetres and 14.5 centimetres respectively. The fourth cervical is 7.25 times as long as it is tall.

 Moganopterus was in 2012 assigned to the Boreopteridae, forming a Moganopterinae with its sister taxon Feilongus.

 Репродукции (1, 2, 3):

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Tags: Вымершие рептилии, Мел, авеметатарзалии, архозавроморфы, архозавры, диапсиды, ктенохазматоидеи, монофенестраты, птеродактили, птерозавры

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 Volgadraco ("Volga River dragon") is a genus of azhdarchid pterodactyloid pterosaur from the Upper Cretaceous of European Russia. It is known from lower beak (holotype SGU, no. 46/104a) and postcranial fragments from the early Campanian-age Rybushka Formation of Saratov, Russia. The size of this animal, and the development of blood supply in the lower jaw, are intermediate between older Santonian or Turonian azhdarchids like Azhdarcho and Bakonydraco and later Maastrichtian azhdarchids like Quetzalcoatlus. Volgadraco was described in 2008 by Averianov, Arkhangelsky, and Pervushov. The type species is V. bogolubovi, the specific name honouring Russian paleontologist Nikolai Nikolaevich Bogolubov. The authors consider the earlier named genus Bogolubovia to be a nomen dubium that in fact might be identical to Volgadraco.

 Ученые из Саратовского государственного технического университета во время полевых работ в верхнемеловых отложениях южной части саратовского правобережья обнаружили кости летающих ящеров-птерозавров. 

"По мнению ведущего российского специалиста по птерозаврам профессора Александра Аверьянова (Зоологический институт РАН, Санкт-Петербург), кости относятся к представителям семейства аждархид. Это самые крупные летающие создания на нашей планете, обитавшие в конце мелового периода. 

 Как выяснилось, ранее останки Volgadraco bogolubovi «Волжский дракон Боголюбова» уже находили. Они являлись самыми крупными летающими созданиями на нашей планете, обитавшими в конце мелового периода. Их размах крыльев был более 10 метров и пропитание они добывали, пикируя на поверхностью водоемов, высматривая рыбу плавающую на небольшой глубине.

 Новые находки были сделаны доцентом кафедры "Геоэкология и инженерная геология" Саратовского технического университета Максимом Архангельским и участником исследовательской группы "Искатели" Сергеем Меркуловым. 

 "Вместе с останками птерозавров обнаружены и кости морских рептилий - плезиозавров, мозазавров и черепах. В ближайшее время экспедиционные работы по сбору ископаемых останков мезозойских рептилий будут продолжены", - цитирует сообщение РИА "Новости".

 Репродукция:

 Сравнение размеров с другими аждархидами и человеком, обозначен буквой I:

Tags: Вымершие рептилии, Мел, авеметатарзалии, аждархиды, аждархойды, архозавроморфы, архозавры, диапсиды, монофенестраты, орнитохейройды, птеродактили, птерозавры

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 Zhejiangopterus is a genus of azhdarchid pterosaur known from one species, which lived in China during the late Cretaceous Period.

 The genus was named in 1994 by Chinese paleontologists Cai Zhengquan and Wei Feng. The type species is Zhejiangopterus linhaiensis. The genus name refers to Zhejiang Province and a Latinised Greek pteron, "wing". The specific name refers to the city of Linhai.

 In 1986 a young chalkstone quarry worker named Xu Chengfa, found a large fossil near the village of Aolicun. Xu by letter informed the director of the Zhejiang Museum of Natural History at Hangzhou, Ming Hua, who from it understood the remains were those of an unknown pterosaur. Therefore he sent a team consisting of the describers and Wu Weitang to investigate. They secured the fossil, instructing the local population to be alert for possible further finds. Xu himself managed to find three more specimens before being killed in an accident in 1988; another worker found a complete skull.

 In the early nineties in total six larger fossils had been recovered from the Tangshang Formation, an 81.5 million year old layer from the Campanian. Among those was the holotype, ZMNH M1330, the impression of the skull of a juvenile individual. Several paratypes were referred: ZMNH M1325, a skeleton lacking the skull; ZMNH M1328, an almost complete skeleton and ZMNH M1329, a fragmentary skeleton.

 Zhejiangopterus was a moderately large pterosaur. Its wingspan was first estimated at 5 metres (16.4 feet). Later estimates reduced this to about 3.5 metres (11.5 ft). Its skull was long, low, perfectly arched, and lacked a "keel" or any other crest sometimes seen in related species. The nasal opening and the large opening typically present between the nose and eye openings of archosaurs (the "antorbital fenestra") had joined together in species such as this to create a single oval opening that occupied nearly one half the length of the skull. The beak was long, thin, sharply pointed, and lacked teeth. The cervical vertebrae were elongated. The first six dorsal vertebrae had fused into a notarium. Several pairs of belly ribs were preserved. Its upper leg bone was half the size of its upper arm bone, and strong and thin. The wings were short but robust.

 Zhejiangopterus was by the original describers classified as a member of the Nyctosauridae, because of the two edentulous pterosaurs they possessed good descriptions of, Pteranodon and Nyctosaurus, it showed the most resemblance to the latter. They deplored lacking good data on Quetzalcoatlus. Indeed in 1997 David Unwin determined that Zhejiangopterus was more closely related to this giant American form and thus belonged to the Azhdarchidae. No other azhdarchid is known from such complete skeletal material.

 Репродукции (1, 2, 3, 4, 5, 6):

 Размеры тела в сравнении с человеком:

 Ископаемые останки:

Tags: Вымершие рептилии, Мел, авеметатарзалии, аждархиды, аждархойды, архозавроморфы, архозавры, диапсиды, монофенестраты, орнитохейройды, птеродактили, птерозавры

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 Tupuxuara is a genus of large, crested, toothless pterodactyloid pterosaur.

 The genus was named and described by Alexander Kellner and Diógenes de Almeida Campos in 1988. The type species is Tupuxuara longicristatus. The genus name refers to a familiar spirit from the mythology of the Tupi. The specific name means "long-crested" in Latin.

 The holotype, MN 6591-V, was found in the Cretaceous Santana Formation of Brazil. It consists of a snout and some partial wing bones. Mature individuals of T. longicristatus had a back-swept crest arising from the snout. Much more fossil material has later been found, showing considerable variation in morphology. Some researchers explain this as intra-specific variability, being caused by a difference in age or sex. Others, however, assume there are different species present.

 In 1994 a second species was named by Kellner: Tupuxuara leonardii. The specific name honours Giuseppe Leonardi. The holotype is MN 6592-V, a fragmentary skull with a more rounded crest. Other such material has been referred to T. leonardii. The largest skulls have a length of 130 centimetres indicating a wingspan of 5.5 metres (18 ft).

 In 2009 a third species was named, by Mark Paul Witton: Tupuxuara deliridamus. The holotype is SMNK PAL 6410, a skull. Another skull is the paratype: KPMNH DL 84. The specific name is derived from Latin delirus, "insane" or "crazy", and adamas, "invincible" but also the word from which "diamond" is derived. The species has a distinctive diamond-shaped skull opening and low eye sockets. The name is a tribute to the song "Shine On You Crazy Diamond" by Pink Floyd, one of Witton's favourite bands.

 Tupuxuara is a member of the group Azhdarchoidea. Kellner assigned it to the Tapejaridae within Azhdarchoidea. According to some analyses however, Tupuxuara is closer to the Azhdarchidae (the group that includes the giant Texan form Quetzalcoatlus) than to Tapejara and its relatives.

 It has been suggested that Tupuxuara was a fish eater at the coasts of South America. Other hypotheses include the possibility it was a fruit eater.

 A subadult described by David Martill and Darren Naish from the University of Portsmouth in 2006 had not yet fully developed its crest, which supports the suggestion that the crest was a marker for sexual maturity.

 Comparisons between the scleral rings of Tupuxuara and modern birds and reptiles suggest that it may have been diurnal.

 Для чего доисторическим летающим рептилиям были нужны гребни на головах, выяснили британские ученые - команда палеонтологов во главе с доктором Дэрреном Нэйшем из университета Портсмута.

 Редкий экземпляр черепа, найденный в Бразилии, "рассказал" исследователям, что гребень использовался птерозаврами для привлечения внимания противоположного пола - существа щеголяли своим "головным убором", демонстрируя половую зрелость. Современные павлины с той же целью используют хвост.

 "Эта поразительная структура яркого цвета походила на гребень гигантского петуха. Мы не знаем этого наверняка, но полагаем, что гребень применялся для привлечения других птерозавров", - сообщил Нэйш.

 Свою теорию ученые основывают на исследовании черепа представителя вида, известного как тупуксуара (Tupuxuara). Как оказалось, этот череп принадлежал птерозавру-подростку.

 Вместо единого большого треугольного гребня от морды до затылка у "юнца" гребень состоял из двух частей, растущих навстречу друг другу. Палеонтологи считают, что когда эти части соединялись, птерозавр достигал половой зрелости. И это становилось заметным для потенциальных партнеров.

 Репродукции (1, 2, 3, 4, 5):

 Размеры тела в сравнении с человеком:

 Ископаемые останки и реплики (1, 2, 3, 4):

Tags: Вымершие рептилии, Мел, авеметатарзалии, аждархойды, архозавроморфы, архозавры, диапсиды, монофенестраты, орнитохейройды, птеродактили, птерозавры, талассодромиды, тапежариды

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 Tupandactylus (meaning "Tupan finger", in reference to the Tupi thunder god) is a genus of tapejarid pterodactyloid pterosaur from the Early Cretaceous Crato Formation of Brazil. It is notable for its large cranial crest, composed partly of bone and partly of soft tissue. The Tupandactylus genus possibly contains two species, both bearing differently sized/shaped crests that may have been used to signal and display for other Tupandactylus, much as toucans use their bright bills to signal to one another. Tupandactylus crests consisted of a semicircular crest over the snout, and in the case of the type species T. imperator, a bony prong which extended back behind the head. A second species, T. navigans, lacked this prong, and had a much more vertical crest. Soft tissue impressions also show that the small bony crests were extended by a much larger structure made of a keratinous material. The complete crest of T. navigans rose in a sharp, sail-like "dome" high above the rest of the skull.

 Tupandactylus imperator is known from four nearly complete skulls. The holotype specimen is MCT 1622-R, a skull and partial lower jaw, found in the Crato Formation, dating to the boundary of the Aptian-Albian stages of the early Cretaceous period, about 112 Ma ago. It was initially described as a species of Tapejara, but later research has indicated it warrants its own genus. The skull was toothless and had a prominent sagittal crest, only the base of which was bony: the front of the crest featured a tall bony rod extending up and back, and the rear of the crest had a long prong of bone projecting behind it. The bulk of the crest was made up of soft tissue similar to keratin, supported by the two bony struts. An additional skull described in 2011, specimen CPCA 3590, preserved more of the lower jaw, showing that like Tapejara, T. imperator had a large, asymmetrical "keel"-like crest on the underside of the lower jaw tip.

 Some Tupandactylus specimens preserve evidence of a keratinous beak at the jaw tips. However, this was restricted to the crested portion of the lower jaw, as one specimen also preserves pycnofibres (simple feather-like filaments) covering the jaws further back.

 Beginning in 2006, several researchers, including Kellner and Campos (who named Tupandactylus), had found that the three species traditionally assigned to the genus Tapejara (T. wellnhofferi, T. imperator, and T. navigans) are in fact distinct both in anatomy and in their relationships to other tapejarid pterosaurs, and thus needed to be given new generic names. However, just how the species should be split proved controversial. Kellner and Campos considered only T. imperator to warrant a new name, creating Tupandactylus. However, another study published in 2007 by Unwin and Martill found that T. navigans, previously assigned to Tapejara, was actually most closely related to T. imperator and belonged with it in a new genus separate from Tapejara. In 2007, at a symposium held in honor of renowned pterosaur researcher Peter Wellnhofer, Unwin and Martill announced the new genus name Ingridia, in honor of Wellnhofer's late wife Ingrid. However, when they published this name in a 2007 volume, they assigned imperator as the type species of their new genus, rather than navigans, which they also included as a species of Ingridia. Furthermore, Unwin and Martill's paper was not published until several months after the similar paper by Kellner and Campos. Therefore, because both sets of authors used imperator as the type, Ingridia is considered a junior objective synonym of Tupandactylus. It was not until 2011 that T. navigans was formally reclassified in the genus Tupandactylus, in a subsequent study supporting the conclusions of Unwin and Martill in 2007.

 Репродукции (1, 2, 3, 4, 5, 6):

 Размеры тела в сравнении с человеком:

 Ископаемые останки (1, 2, 3):

Tags: Вымершие рептилии, Мел, авеметатарзалии, аждархойды, архозавроморфы, архозавры, диапсиды, монофенестраты, орнитохейройды, птеродактили, птерозавры, тапежариды

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 Tropeognathus is a genus of large pterosaurs from the late Cretaceous Period of South America. It was member of the Ornithocheiridae (alternately Anhangueridae), a group of pterosaurs known for their keel-tipped snouts, and was closely related to species of the genus Anhanguera. It is known primarily for the species Tropeognathus mesembrinus, though a second species, Tropeognathus robustus, has been named in the genus.

 In the 1980s the German Bayerische Staatssammlung für Paläontologie und historische Geologie at Munich acquired a pterosaur skull from Brazilian fossil dealers, that had probably been found in Ceará, in the Chapade do Araripe. In 1987 it was named and described as the type species Tropeognathus mesembrinus by Peter Wellnhofer. The generic name is derived from Greek τρόπις, tropis, "keel", and γνάθος, gnathos, "jaw". The specific name is derived from Koine mesembrinos, "of the noontide", "southern", in reference to the provenance from the Southern hemisphere.

 The holotype, BSP 1987 I 46, was discovered in a layer of the Romualdo Member of the Santana Formation, dating from the Aptian-Albian. It consists of a skull with lower jaws. A second specimen was referred by André Jacques Veldmeijer in 2002: SMNS 56994, consisting of partial lower jaws. In 2013, Alexander Wilhelm Armin Kellner referred a third, larger, specimen: MN 6594-1, a skeleton with skull, with extensive elements of all body parts, except the tail and the lower hindlimbs.

 After Tropeognathus mesembrinus was named by Peter Wellnhofer in 1987 other researchers tended to consider it part of several other genera, leading to an enormous taxonomic confusion. It was considered an Anhanguera mesembrinus by Alexander Kellner in 1989, a Coloborhynchus mesembrinus by Veldmeijer in 1998 and a Criorhynchus mesembrinus by Michael Fastnacht in 2001. In 2001, David Unwin referred the Tropeognathus material to Ornithocheirus simus, making Tropeognathus mesembrinus a junior synonym, though he again reinstated a Ornithocheirus mesembrinus in 2003. Veldmeijer in 2003 accepted that Tropeognathus and Ornithocheirus were cogeneric but rejecting O. simus as the type species of Ornithocheirus in favor of O. compressirostris (named Lonchodectes by Unwin), used the names Criorhynchus simus and Criorhynchus mesembrinus. In 2000, Kellner again began to use the original name Tropeognathus mesembrinus. In 2013, Taissa Rodrigues and Kellner concluded Tropeognathus to be valid, and containing only T. mesembrinus.

 In 1987 Wellnhofer named a second species, Tropeognathus robustus, based on specimen BSP 1987 I 47, a more robust lower jaw. Today, this is no longer considered cogeneric with Tropeognathus mesembrinus.

 Tropeognathus mesembrinus is known to have reached wingspans of up to 8.2 m (27 ft), as can be inferred from the size of specimen MN 6594-1. T. mesembrinus bore distinctive convex "keeled" crests on its snout and underside of the lower jaws. The upper crests arose from the snout tip and extended back to the fenestra nasoantorbitalis, the large opening in the skull side. An additional, smaller crest projected down from the lower jaws at their symphysis ("chin" area). While many ornithocheirids had a small, rounded bony crest projecting from the back of the skull, this was particularly large and well-developed in Tropeognathus. The first five dorsal vertebrae are fused in to a notarium. Five sacral vertebrae are fused into a synsacrum. The third and fourth sacral vertebrae are keeled. The front blade of the ilium is strongly directed upwards.

 In 1987 Wellnhofer assigned Tropeognathus to a Tropeognathidae. This concept was not adopted by other workers; Brazilian researchers place Tropeognathus mesembrinus in the Anhangueridae, their European colleagues tend to prefer the Ornithocheiridae.

 Тропеогнат был одним из крупнейших птерозавров мелового периода и ярким представителем группы Ornithocheiridae, отличавшихся своим крупными килеобразными выростами на конце челюстей от других птеродактилей.

 Впервые тропеогнат был описан в 1980 г., когда немецкий музей Bayerische Staatssammlung fur Palaontologie und Historische Geologie выкупил у бразильских продавцов окаменелостей огромного летающего ящера, который был обнаружен в местонахождении Сеара на севере Бразилии. Вид в 1987 г. получил типовое наименование Tropeognathus mesembrinus.

 Тропеогнаты достигали достаточно внушительных размеров: размах крыльев этих ящеров достигал 8,2 метров. Как отмечалось ранее, отличительная черта орнитохейрид: килевые гребни на конце морды, верхний из которых шел от конца челюсти и до начала анторбитального отверстия (antorbitalis fenestra). Нижний гребень был немного меньше и представлял собой расширение подбородочной кости.

 Грудные позвонки у тропеогната были мощными и были слиты в единый костный выступ, поддерживающий объемные мышечные маховые связки. Крестцовые позвонки также слиты в одно костное образование.

 В сериале ВВС «Прогулки с динозаврами» одна из серий посвящена жизни летающего гиганта Ornithocheirus, который, на самом деле, является представителем Tropeognathus mesembrinus. В сериале использовались данные по окаменелостям летающего ящера, обнаруженного в 1998 г. на севере Бразилии. Расчетные показатели свидетельствовали о том, что этот бразильский гигант мог достигать размаха крыльев в 12 метров и был весом до 100 кг, что делало его одним из самых крупных обнаруженных птерозавров.

 Однако типовые экземпляры Ornithocheirus не превышают 6 метров в размахе крыльев. Окаменелости из Бразилии еще находились на стадии изучения и только в 2012 г. Дэйв Мартилл и Хайнц Петер Бредов опубликовали окончательную оценку размеров найденной особи, согласно которой этот экземпляр тропеогната не мог превышать 8,2 м. в размахе крыльев. Х.П. Бредов пояснил: «Вероятно, при работе над сериалом были использованы самые максимальные из возможных показателей, чтобы сериал казался более захватывающим и зрелищным».

 У этих летающих ящеров был достаточно развит половой диморфизм: самки были мельче в размерах и имели менее выраженные выросты на челюстях.

 Анализ строения нижних конечностей тропеогната свидетельствует о том, что этот птерозавр вполне был способен перемещаться по плоской поверхности, а не только по скальным выступам и утесам, как большинство других птерозавров.

 Длинные, острые и изогнутые зубы, разного размера говорят о рыбоядной специализации тропеогната. Предполагается, что ящер парил над поверхностью воды, вылавливая рыбу из верхних слоев океана, причем в данном случае, выступающие гребни на челюстях служили своеобразным стабилизатором при полете.

 В меловом периоде птерозавры, в целом, характеризуются увеличением своих размеров, размах крыльев некоторых видов достигает поистине неимоверных величин, в то же самое время видовое разнообразие летающих ящеров становится очень разнообразным. Поэтому тропеогнат, наряду с гигантскими аждархидами, является ярким представителем гигантизма и разнообразия летающих ящеров в меловом периоде.

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 Талассодромеус (Thalassodromeus sethi, от др.-греч. θαλασσο- +δρομεύς «морской бегун», sethi — от имени Сета, из-за формы гребня) — птеродактиль из раннего мела Бразилии (формация Сантана, возраст около 122 − 109 млн.л.н.). Описан А. Келльнером и Д. Кампосом в 2002 году. Длина черепа до 1420 мм, череп высокий, имеется огромный затылочный гребень. Гребень очень тонкий, со следами кровеносных сосудов. Возможно, он служил для терморегуляции и для брачной демонстрации. На нижней челюсти в подбородочной области имеется небольшой вырост, который мог служить своеобразным килем-рассекателем, когда птерозавр опускал нижнюю челюсть в воду во время охоты. Способ питания талассодромеуса мог быть сходен со способом питания чаек-водорезов. Ящер летел низко над водой и опущенной нижней челюстью «резал» воду, выхватывая мелкую живность. В отличие от водорезов, талассодромеус мог достигать 4,5 метров в размахе крыльев. Это самое крупное известное животное, питавшееся по типу водореза. Тем не менее, недавние опыты на натурных моделях показали, что подобный способ питания для столь крупного животного был бы невозможен — челюсть создавала бы такое сопротивление, что ящер не мог бы удержать себя в воздухе. Родственные связи талассодромеуса не вполне ясны, он может быть родственником птеродактилей-тупуксар. В свою очередь, тупуксары близки к аждархидам.

 Thalassodromeus was a large pterodactyloid pterosaur found in northeastern Brazil.

 The genus was named in 2002 by Alexander Kellner and Diogenes de Almeida Campos. The type species is Thalassodromeus sethi. The genus name is derived from Greek thalasse, "sea" and dromaios, "runner", in reference to its presumed life style as a skimmer. The specific name refers to the Egyptian god Seth because of the similarity in head form. In 2006 André Jacques Veldmeijer suggested Kellner had confused Seth with the god Amun whose crown shows a remarkable resemblance with the Thalassodromeus head crest.

 The genus is based on holotype DGM 1476-R, a damaged partial skull, found in the Santana Formation. Thalassodromeus lived in the Early Cretaceous, roughly 108 million years ago. It shared the skies with its smaller cousin Tapejara. It is particularly notable for its immense head crest, beginning at the tip of the snout and ending far behind the braincase, which accounts for seventy-five percent of the surface of its 1.42 metre (4.6 ft) long skull. The jaws were pointed and toothless. It had a wing span of roughly 4.5 metres (14.7 ft). The function of the crest is unknown, but it may have been used for sexual display, species recognition, or thermoregulation. A lower jaw fragment referred to Thalassodromeus, DGM 1476-M, indicates a larger example with a wingspan of 5.3 metres (17.4 ft).

 Another jaw fragment, SAO 251093, was unofficially suggested to be a new species, "Thalassodromeus oberli" (referring to the Urs Oberli collection), by Veldmeijer in 2006, after having referred the specimen to T. sethi in 2005. In 2014 this was made a separate genus Banguela.

 Thalassodromeus was believed by Kellner to have fed in a similar way to modern skimmers; trailing its lower jaw in the water while it flew. However, later research on its jaw and neck anatomy suggested that for this and other larger pterosaurs it would not be feasible to skim because of the drag: the energy expenditure would be too high. Rather, Thalassodromeus appears to have had specialisation for terrestrial foraging like Azhdarchidae, even converging on leg proportions, and it's powerful jaws might suggest raptorial tendencies akin to those of phorusrhacids.

 Kellner assigned Thalassodromeus to the Tapejaridae. Other analyses however, showed that it was, joined with Tupuxuara in a Thalassodrominae, more closely related to the Azhdarchidae.

 Thalassodromidae (meaning "sea runners") is a family of pterodactyloid pterosaurs from the early Cretaceous period of Brazil. It contains two genera, Thalassodromeus and Tupuxuara.

 The classification of thalassodromids is controversial. Some studies, including one by Lü and colleagues in 2008, have found that the thalassodromids are more closely related to the azhdarchids than to the tapejarids, and have placed them in their own family (which has sometimes been referred to as Tupuxuaridae, though Thalassodrominae was named first). Alternately, they have been considered a subfamily (Thalassodrominae) within the Tapejaridae.

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 Тапежара, или тапейяра (Tapejara — от тупийского «древнее существо») — род птерозавров, обитавших в раннем меловом периоде, 121—112 млн лет назад. Отличался причудливым гребнем на голове. Размах крыльев — до 5 м. Хищник. Ископаемые находки были найдены в Южной Америке (Бразилия).

 Тапежара — древнейший из известных ученым птерозавр с беззубым клювом. Его ископаемые останки были обнаружены в бразильских горах Сантана, содержащих окаменелости и многих других причудливых видов птерозавров. Родственные связи тапежары с многочисленными видами беззубых птерозавров позднего мелового периода изучены недостаточно.

 Крупный птерозавр тапежара охотился на рыбу у побережья Южной Америки в меловой период. Своей известностью тапежара обязана огромному, высотой до 1 м, гребню на голове. Гребень, которым обладали только самцы, доставлял животным большие неудобства: он делал их голову очень тяжёлой, а полёт медленным и неровным. Очевидно, ярко окрашенный гребень использовался самцами тапежары во время зрелищных брачных демонстраций и служил для привлечения самок. Также существует предположение Санкара Чхаттердже, что гребень при плавании в воде использовался в качестве своеобразного паруса.

 Тапежары гнездились крупными колониями недалеко от моря, где родители добывали корм для своих детёнышей.

 Tapejara (from a Tupi word meaning "the old being") is a genus of Brazilian pterosaur from the Cretaceous Period (Santana Formation, dating to about 108 Ma ago). Tapejara crests consisted of a semicircular crest over the snout, and a bony prong which extended back behind the head.

 The type species and only one currently recognized as valid by most researchers, T. wellnhoferi, is the smallest species to have been assigned to Tapejara and does not preserve evidence of soft-tissue crest extensions. The specific name honours German paleontologist Peter Wellnhofer. A second species, originally named Tapejara imperator ("emperor"), is much larger and possessed a crest made up of distinctively long prongs, projecting from the rounded snout crest and the back of the skull, which supported a large, possibly rounded sail-like crest of keratin. A third species, Tapejara navigans ("sailing"), was mid-sized and sported a similar crest to T. imperator, though narrower and more dome-shaped, that lacked the backwards-pointing bony support prong.

 Several studies in 2007 showed that T. imperator and T. navigans are too different from T. wellnhoferi and therefore require their own genus names. The species T. imperator was given its own genus, Tupandactylus, by Kellner and Campos. Unwin and Martill found that T. imperator and T. navigans belong in the same genus, and named them Ingridia imperator and I. navigans, respectively. The genus name honoured Wellnhofer's late wife Ingrid. Because Tupandactylus was named first, it retains priority over the name Ingridia. To complicate matters, both the name Tupandactylus and Ingridia used the former Tapejara imperator as their type species. The scientists who described Tupandactylus did not name a Tupandactylus navigans (but instead suggested it was synonymous to Tupandactylus imperator), and Tapejara navigans was not formally reclassified as a distinct species of Tupandactylus until 2011.

 Comparisons between the scleral rings of Tapejara and modern birds and reptiles suggest that it may have been cathemeral, active throughout the day at short intervals.

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 Sinopterus (meaning "Chinese wing") is a genus of tapejarid pterodactyloid pterosaur from the Aptian-age Lower Cretaceous Jiufotang Formation of Chaoyang, Liaoning, China. Three species have been classified in this genus, though only two are generally considered to be valid. Sinopterus is known for its proportionally large skull, which has a birdlike pointed beak, a long bony crest that starts with a tall premaxilla and goes back along the middle of the skull to form a point overhanging the rear of the skull, and its lack of teeth.

 The type species, S. dongi, is based on IVPP V13363, an articulated, nearly complete skeleton. The skull of this individual was 17 centimeters (6.7 inches) long, and the wingspan was estimated to be 1.2 meters (3.9 feet). The authors suggested that it was an omnivore, and noted that it was the first record of a tapejarid outside of Brazil, and the earliest and most complete tapejarid.

 A second species, S. gui, was named by Li, Lü, and Zhang in 2003 based on BPV-077, another nearly complete skeleton from the Jiufotang Formation. It was said to differ from S. dongi mainly in its smaller size (only about half the size of S. dongi) and the presence of a notarium, though this was later disproved. Most later studies have found S. gui to simply represent a younger specimen of S. dongi.

 A third species was referred to Sinopterus in 2007, S. jii. This species was first named by Lü & Yuan in 2005 as the type species of a new genus, which they named Huaxiapterus. However, two later studies in 2007 and 2011 both showed that H. jii was in fact more closely related to Sinopterus than to two other species also assigned to Huaxiapterus, "H." corollatus and "H." benxiensis. Both groups of researchers concluded that Huaxiapterus jii should therefore be reclassified as Sinopterus jii, and that the other two species of "Huaxiapterus" require a new genus name.

 Тапежариды (Tapejaridae Kellner, 1989, Tupuxuaridae Martill, Bechly & Heads, 2007) — семейство птерозавров раннего мелового периода. Известны находки из Китая и Бразилии. Роды из Китая более примитивны, что указывает на азиатское происхождение семейства.

 Филогения тапежарид остаётся спорной, имеется несколько конкурирующих кладограмм. В частности, нет общепринятой точки зрения на то, принадлежит ли талассодромеус к данному семейству.

 Tapejaridae (meaning "the old beings") are a family of pterodactyloid pterosaurs from the early Cretaceous period. Members are currently known from Brazil, Morocco, Spain and China, where the most primitive genera are found, indicating that the family has an Asian origin.

 Tapejarids were small to medium-sized pterosaurs with several unique, shared characteristics, mainly relating to the skull. Most tapejarids possessed a bony crest arising from the snout (formed mostly by the premaxillary bones of the upper jaw tip). In some species, this bony crest is known to have supported an even larger crest of softer, fibrous tissue that extends back along the skull. Tapejarids are also characterized by their large nasoantorbital fenestra, the main opening in the skull in front of the eyes, which spans at least half the length of the entire skull in this family. Their eye sockets were small and pear-shaped. Studies of tapejarid brain cases show that they had extremely good vision, more so than in other pterosaur groups, and probably relied nearly exclusively on vision when hunting or interacting with other members of their species. Tapejarids had unusually reduced shoulder girdles that would have been slung low on the torso, resulting in wings that protruded from near the belly rather than near the back, a "bottom decker" arrangement reminiscent of some planes.

 Tapejaridae may be composed of two subfamilies: a Tapejarinae of "Huaxiapterus" corollatus, Sinopterus, Tapejara, Tupandactylus, Europejara, Caiuajara, and possibly Bakonydraco, and a Thalassodrominae of Thalassodromeus and Tupuxuara. Some studies, such as one by Lü and colleagues in 2008, have found that the thalassodromines are more closely related to the azhdarchids proper than to the tapejarids, and have placed them in their own family (which has sometimes been referred to as Tupuxuaridae, though Thalassodrominae was named first). At least one study has also found that the Chaoyangopteridae, often found to be closer to azhdarchids, represent a lineage within the Tapejaridae, more closely related to the tapejarines than to the thalassodromines. Felipe Pinheiro and colleagues (2011) reclassified the group as a subfamily of Tapejaridae, Chaoyangopterinae, for this reason.

 The exact relationships of tapejarids to one another and to other azhdarchoid pterosaurs has historically been unclear, with different studies producing significantly different cladograms (family trees). It is also unclear exactly which pterosaurs belong to the Tapejaridae; some researchers have found the thalassodromines and chaoyangopterines to be members of this family, while other studies have found them to be more closely related to the azhdarchids (in the clade Neoazhdarchia). Some studies have even allowed the possibility that the "tapejarids" as traditionally thought of are paraphyletic, that is, they may not form a natural group but instead represent sequential branches of the tree leading to the more advanced neoazhdarchians.

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 Santanadactylus (meaning "Santana Formation finger") was a genus of pterodactyloid pterosaur from the Aptian-age Romualdo Member of the Lower Cretaceous Santana Formation, of Barra do Jardim, Araripe Plateau, Ceará Province, Brazil. Four species have been named, but today it is doubted they are part of the same genus. It was a rather large pterosaur.

 The genus was named in 1980 by the Dutch paleontologist Paul de Buisonjé. The type species is S. brasilensis, the specific name referring to Brazil. It is based on holotype UvA M 4894 (Geological Institute of the University of Amsterdam), an upper part of the right humerus and a right scapulacoracoid; other remains were also included in the genus: this paratype consisted of two cervical vertebrae from a different individual, referred because they were found in the same lot of 25 chalk nodules bought from collectors. Additional remains, including a notarium (fused vertebrae supporting the shoulder) have been referred to it since then. The structure of the humerus suggests an ornithocheirid, but the long neck vertebrae argue against this.

 In 1985, Peter Wellnhofer, a German paleontologist who has written numerous scientific publications on pterosaurs, named three additional species: S. araripensis, S. pricei, and S. spixi. S. araripensis, named after the Araripe Plateau, was a large species based on BSP 1982 I 89, remains including a partial skull (missing the end of the jaws) and arms; the preserved skull section had no crest. S. pricei, named after Llewellyn Ivor Price, was the smallest of the three species; it was based on BSP 1980 I 122, a left wing from the elbow down, and additional arm material has been referred to it over the years. S. spixi, intermediate in size, was based on BSP 1980 I 121, another left wing, the name honouring Johann Baptist von Spix.

 Over the years, the species of this taxon have been reassessed. Chris Bennett suggested that S. brasilensis was a chimera of a pteranodontid and something else (in that the holotype and paratype belonged to different forms), S. araripensis and S. pricei were pteranodontids, and S. spixi was a dsungaripterid. Wellnhofer removed S. spixi from the genus as well, in 1991. In 1992, Alexander Kellner and Diogenes de Almeida Campos suggested that S. spixi was a tapejarid; David Unwin in 2003 thought that "Santanadactylus" spixi was a species of Tupuxuara. Kellner in 1990 renamed S. araripensis to Anhanguera araripensis, followed by Wang and colleagues in 2008, though Veldmeijer (2003) included it in Coloborhynchus. Part of the problem is the complicated taxonomy of Santana Formation pterosaurs and their English contemporaries, involving numerous genera, such as Amblydectes, Anhanguera, Araripesaurus, Criorhynchus, Coloborhynchus, Lonchodectes, Ornithocheirus, and Tropeognathus.

 Also the phylogeny is contentious. De Buisonjé first placed the genus into the Criorhynchidae, a concept that is no longer used. Wellnhofer considered it a member of the Ornithocheiridae. According to Bennett S. brasilensis belonged to Pteranodontidae sensu Bennett. Kellner, concluding that he could find but a single autapomorphy for Santanadactylus brasilensis, the straight ventral margin of the proximal part of the deltopectoral crest, at first thought no greater precision was possible than a Pterodactyloidae incertae sedis, but in 2000 narrowed it down to a Pteranodontoidea sensu Kellner. S. pricei according to Kellner belonged to a clade descending from the last common ancestor of Istiodactylus and the Anhangueridae. The same was in his analysis true for Araripesaurus, a genus of which he had previously thought S. pricei was a junior synonym.

 Santanadactylus is regarded as a large pterosaur, Wellnhofer for the various species indicating a wingspan of 2.9–5.7 m (9.5–18.7 ft). De Buisonjé thought Santanadactylus brasilensis had a wingspan of six metres. It may have been adapted for gliding over flapping flight.

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 Кетцалькоа́тль (Quetzalcoatlus) — крупнейший известный на сегодня представитель отряда птерозавров (Pterosauria). Типовой вид — Quetzalcoatlus northropi Lawson, 1975. Размах крыльев точно не известен из-за неполной сохранности останков, но по пропорциям птерозавров других видов оценивается приблизительно в 11 метров (по мнению некоторых палеонтологов — до 15 м). Обнаружен в позднемеловых отложениях Северной Америки. Время обитания — поздний меловой период, 68—65,5 миллионов лет назад.

 Весил, по разным оценкам, от 85 до 250 кг. Окаменелости кетцалькоатля обнаружены в Северной Америке. Название дано в честь ацтекского бога. В настоящее время кетцалькоатль вместе с другими гигантскими птерозаврами (Hatzegopteryx) являеся самым крупным известным летающим существом за всю историю жизни на планете. Кетцалькоатль и хатцегоптерикс были примерно одинакового размера, только первый был немного более массивным. Кетцалькоатль летал над сушей и питался падалью и мелкими позвоночными. Возможно, он мог поймать и небольшого динозавра весом до 30 килограммов.

 Изначально размах крыльев оценили в 15,9 метров, усреднив оценку через пропорции других птерозавров. Однако в ходе исследования 1981 года оценочный размер уменьшили до 11—12 метров. Более поздние исследования ещё уменьшили размер Q. до 10—11 метров.

 Оценить массу гигантских аждархидов очень сложно, поскольку нет существующих видов схожего размера или строения, поэтому в разных публикациях результаты разнятся. В то время как некоторые исследования традиционно указывают крайне низкую оценочную массу, как например 70 кг для 10-и метровой особи, большинство оценок, опубликованных с начала 2000-х, указывают на массу в районе 200 кг.

 О жизненном укладе Q. существует несколько предположений. Поскольку кости были найдены в сотнях километров от береговой линии, и не было найдено следов больших рек или глубоких озёр, Дуглас Лоусон в 1975 году отверг рыболовную натуру Q., предположив, что животное питалось падалью подобно африканскому марабу. Лоусон нашел останки гигантского птерозавра во время поисков костей аламозавра, бывшего важной частью экосистемы.

 В 1996 году Томас Леман и Лангстон указали, что строение нижней челюсти отличается от таковой у типичных птиц-падальщиков. Они предположили, что длинная шея и длинные беззубые челюсти позволяли Q. питаться примерно как современные водорезы, ловя рыбу во время полета над водой, прочесывая волны клювом.

 Исследование 2007 года показало, что для такого большого птерозавра подобный полет был бы слишком энергозатратным из-за сильного лобового сопротивления.

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 Pterodaustro is a genus of Cretaceous pterodactyloid pterosaur from South America, which lived 105 million years ago.

 The first fossils, among them the holotype PLV 2571, a thigh bone, were in the late sixties discovered by Bonaparte in the Lagarcito Formation, situated in the San Luis Province of Patagonia, Argentina, and dating from the Albian. The genus has later also been found in Chile in the Santa Ana Formation. At the Argentine site, the just 50 m² large "Loma del Pterodaustro", since then during several expeditions over 750 Pterodaustro specimens have been collected, 288 of them having been catalogued until 2008. This makes the species one of the best known pterosaurs, with examples from all growth stages, from egg to adult.

 The genus was named in 1969 by José Bonaparte as an as yet undescribed nomen nudum. The first description followed in 1970, making the name valid, the type species being Pterodaustro guiñazui. The genus name is derived from Greek pteron, "wing" and Latin auster, "south (wind)". The elements are combined as a condensed pteron de austro, "wing from the south". The specific name honours paleontologist Román Guiñazú. It was emended in 1978 by Peter Wellnhofer into guinazui, because diacritical signs such as the tilde are not allowed in species names.

 Pterodaustro has a very elongated skull, up to 29 centimetres long. The portion in front of the eye sockets comprises 85% of skull length. The long snout and lower jaws curve strongly upwards; the tangent at the point of the snout is perpendicular to that of the jaw joint. Pterodaustro has about a thousand bristle-like modified teeth in its lower jaws that might have been used to strain crustaceans, plankton, algae, and other small creatures from the water. These teeth stand for the most part not in separate alveoli but in two long grooves parallel to the edges of the jaw. They have a length of three centimetres and are oval in cross-section, with a width of just 0.2 - 0.3 millimetres. At first it was suspected these structures were not true teeth at all, but later research established they were built like normal teeth, including enamel, dentine and a pulpa. Despite being made of very hard material, they might still have been flexible to some extent due to their extreme length-width ratio, a bend of up to 45° being possible. The upper jaws also carried teeth, but these were very small with a flat conical base and a spatula-formed crown. These teeth also do not have separate tooth sockets but were apparently held by ligaments in a special tooth pad, that was also covered with small ossicles, or bone plates.

 The back of the skull was also rather elongated and in a low position; there are some indications for a low parietal crest.

 Pterodaustro had an adult wingspan of approximately 250 centimetres (8.20 ft). Its hindlimbs are rather robust and its feet large. Its tail is uniquely elongated for a pterodactyloid, containing 22 caudal vertebrae, whereas other members of this group have at most sixteen.

 Pterodaustro probably waded in shallow water like flamingos, straining food with its tooth comb, a method called "filter feeding". Once it caught its food, Pterodaustro probably mashed it with the small, globular teeth present in its upper jaw.

 Robert Bakker incorrectly suggested that, like flamingos, this pterosaur's diet may have resulted in a pink hue. However, recent studies show that only Neoaves can sequester carotenoids for use as a pigment in the feathers rather than just the skin or beak, and even then they require to enhance it with structural colors, so a pink hue for any pterosaur seems extremely unlikely.

 At least two specimens of Pterodaustro have been found, MIC V263 and MIC V243, with gizzard stones in the stomach cavity, the first ever reported for any pterosaur. These clusters of small stones with angled edges support the idea that Pterodaustro ate mainly small, hard-shelled aquatic crustaceans using filter-feeding. Such invertebrates are abundant in the sediment of the fossil site.

 A study of the growth stages of Pterodaustro concluded that juveniles grew relatively fast in their first two years, attaining about half of the adult size. Then they reached sexual maturity, growing at a slower rate for four to five years until there was a determinate growth stop.

 In 2004 a Pterodaustro embryo in an egg was reported, specimen MHIN-UNSL-GEO-V246. The egg was elongated, six centimetres long and 22 millimetres across and its mainly flexible shell was covered with a thin layer, 0.3 mm thick, of calcite. Three-dimensionally preserved eggs were reported in 2014.

 Comparisons between the scleral rings of Pterodaustro and modern birds and reptiles suggest that it may have been nocturnal, and may have had similar activity patterns to modern anseriform birds that feed at night.

 Bonaparte in 1970 assigned Pterodaustro to the Pterodactylidae; in 1971 to a Pterodaustriidae. However, from 1996 cladistic studies by Alexander Kellner and David Unwin have shown a position in the Ctenochasmatidae, together with other filter feeders.

 The genus Puntanipterus might be a subjective junior synonym of Pterodaustro.

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 Muzquizopteryx is a genus of pterodactyloid pterosaur (flying reptile) from the Late Cretaceous of what is now Coahuila,Mexico.

 In the nineties José Martínez Vásquez, a worker at the chalk quarry of El Rosario, uncovered a skeleton of a pterosaur. This he handed to a quarry official, who had it bricked in on the face of an office wall as a decorative piece. After its unique scientific value had been recognised in 2002, the specimen was acquired by the Universidad Nacional Autónoma de México. Subsequently it was studied by a combined team of the University of Karlsruhe and the University of Heidelberg, in this financially supported by the Volkswagen company. The specimen was scientifically reported in 2004.

 In 2006 the type species Muzquizopteryx coahuilensis was named and described by Eberhard Frey, Marie-Céline Buchy,Wolfgang Stinnesbeck, Arturo González-González and Alfredo di Stefano. The generic name is derived from the Múzquizdistrict and a Greek πτέρυξ, pteryx, "wing". The specific name is derived from the state of Coahuila.

 Muzquizopteryx is based on holotype UNAM IGM 8621, found in the El Rosario layers, early Coniacian-aged rocks. It consists of a nearly complete, articulated skeleton that includes soft tissue remains, among them long fossilised tendons along both sides of both lower arms. The specimen represents an adult individual.

 In 2012 a second specimen was reported, MUDE CPC-494, again uncovered by a quarry worker, perhaps at the same site, and sold to a private collector. It was later acquired by the Museo del Desierto Saltillo. It consists of the right upper wing of a subadult individual, with about 81% of the length of the holotype. As its provenance probably consists of slightly older layers from the late Turonian and the remains are limited, it was referred to a Muzquizopteryx sp.

 Muzquizopteryx was relatively small for a pterodactyloid pterosaur, with a wingspan of around two metres (6.6 ft). It had an elongated head with a convex upper profile, ending at the back of the head in a backward pointing short rounded crest. The jaws were toothless. The arms were very robust with the humerus featuring a large hatchet-shaped deltopectoral crest, indicating a strong wing musculature. The pteroid bone was long and pointed towards the neck, supporting a flight membrane.

 Muzquizopteryx was by its describers assigned to the Nyctosauridae. It would then be the oldest known member of the group and the smallest known; indeed the smallest adult Late Cretaceous pterosaur discovered until 2006. As Nyctosaurus is sometimes included with the Pteranodontidae, Muzquizopteryx too might be considered a member of that group under some classifications.

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 Nyctosaurus is a genus of pterodactyloid pterosaur, the remains of which have been found in the Niobrara Formation of the mid-western United States, which, during the late Cretaceous Period, was covered in an extensive shallow sea. The genus Nyctosaurus has had numerous species referred to it, though how many of these may actually be valid requires further study. At least one species possessed an extraordinarily large antler-like cranial crest.

 Nyctosaurus was similar in anatomy to its close relative and contemporary, Pteranodon. It had relatively long wings, similar in shape to modern seabirds. However, it was smaller overall than Pteranodon, with an adult wingspan of 2 meters (6.6 ft) and a maximum weight of about 1.86 kg. The overall body length was 37 cm. Some specimens preserve a distinctive crest, at least 55 cm tall in old adults, relatively gigantic compared to the rest of the body and over three times the length of the head. The crest is composed of two long, grooved spars, one pointed upward and the other backward, arising from a common base projecting up and back from the back of the skull. The two spars were nearly equal in length, and both were nearly as long or longer than the total length of the body. The upward-pointing crest spar was at least 42 cm long (1.3 ft) and the backward-pointing spar was at least 32 cm long (1 ft).

 The jaws of Nyctosaurus were long and extremely pointed. The jaw tips were thin and needle sharp, and are often broken off in fossil specimens, giving the appearance that one jaw is longer than the other, though in life they were probably equal in length.

 Nyctosaurus is the only pterosaur to have lost its clawed "fingers", with the exception of the wing finger (of which however the fourth phalanx was lost), which is likely to have impaired its movement on the ground, leading scientists to conjecture that it spent almost all of its time on the wing and rarely landed. In particular, the lack of claws with which to grip surfaces would have made climbing or clinging to cliffs and tree trunks impossible for Nyctosaurus.

 Nyctosaurus, like its relative Pteranodon, appears to have grown very rapidly after hatching. Fully adult specimens are no larger than some immature specimens such as P 25026, indicating that Nyctosaurus went from hatching to adult size (with wingspans of 2 m or more) in under a year. Some sub-adult specimens have been preserved with their skulls in nearly pristine condition, and lack any trace of a head crest, indicating that the distinctively large crest only began to develop after the first year of life. The crest may have continued to grow more elaborate as the animal aged, though no studies have examined the age of the fully adult, large-crested specimens. These individuals may have been 5 or even 10 years old at the time of their deaths.

 Only five relatively complete Nyctosaurus skulls have been found. Of those, one is juvenile and does not possess a crest (specimen FMNH P 25026), and two are more mature and may show signs of having had a crest but are too badly crushed to say for sure (FHSM 2148 and CM 11422). Two specimens (KJ1 and KJ2) described in 2003, however, preserved an enormous double-pronged crest.

 A few scientists had initially hypothesized that this crest, which resembles an enormous antler, may have supported a skin "headsail" used for stability in flight. While there is no fossil evidence for such a sail, studies have shown that a membranous attachment to the bony crest would have imparted aerodynamic advantages. However, in the actual description of the fossils, paleontologist Christopher Bennett argued against the possibility of a membrane or soft tissue extension to the crest. Bennett noted that the edges of each prong were smooth and rounded, and showed no evidence for any soft-tissue attachment points. He also compared Nyctosaurus with large-crested tapejarids, which do preserve soft tissue extensions supported by prongs, and showed that in those species, the attachment points were obvious, with jagged edges where the transition from bone to soft tissue occurred. Bennett concluded that the crest was most likely used solely for display, citing similar structures in modern animals. The 2009 study by Xing and colleagues testing the aerodynamics of the giant crest with a "headsail" also tested the aerodynamics of the same crest with no sail, and found that it added no significant negative factors, so a crest with no headsail would not have hindered normal flight. It is more likely that the crest acted mainly for display, and that any aerodynamic effects it may have had were secondary. Bennett also argued that the crest was probably not a sexually dimorphic character, as in most crested pterosaurs, including the related Pteranodon, both sexes are crested and it is only the size and shape of the crest that differs. The apparently non-crested Nyctosaurus specimens therefore probably came from sub-adults.

 Sankar Chatterjee and R.J. Templin used estimates based on complete Nyctosaurus specimens to determine weight and total wing area, and to calculate its total wing loading. They also estimated its total available flight power based on estimated musculature. Using these calculations, they estimated the cruising speed of Nyctosaurus gracilis as 9.6 meters/second (34.5 kilometers/hour or 21.4 miles/hour).

 All known Nyctosaurus fossils come from the Smokey Hills Chalk of Kansas, part of the Niobrara Formation. Specifically, they are found only within a narrow zone characterised by the abundance of ammonite fossils belonging to the species Spinaptychus sternbergi. These limestone deposits were laid down during a marine regression of the Western Interior Seaway that lasted between 85 and 84.5 million years ago. Therefore, Nyctosaurus was a relatively short-lived species, unlike its relative Pteranodon, which is found throughout almost all of the Niobrara layers into the overlying Pierre Shale Formation, and existed between 88 and 80.5 million years ago.

 The ecosystem preserved in this zone was unique in its abundance of vertebrate life. Nyctosaurus shared the sky with the bird Ichthyornis and with Pteranodon longiceps, though the second Niobrara Pteranodon species, P. sternbergi, had disappeared from the fossil record by this point. In the waters of the Western Interior Seaway below swam mosasaurs (Clidastes, Ectenosaurus and Tylosaurus), the flightless diving bird Parahesperornis, and a wide variety of fish, including swordfish-like Protosphyraena, the predatory Xiphactinus and the shark Cretolamna.

 The first Nyctosaurus fossils were described in 1876 by Othniel Charles Marsh, based on fragmentary material, holotype YPM 1178, from the Smoky Hill River site in Kansas. Marsh referred the specimen to a species of his new genus Pteranodon, as Pteranodon gracilis. Later that year, Marsh reclassified the species in its own genus, which he named Nyctosaurus, meaning "night lizard" or "bat lizard", in reference to the wing structure somewhat paralleling those of bats. In 1881, Marsh incorrectly assumed the name was preoccupied and changed it into Nyctodactylus, which thus is now a junior synonym. In 1902,Samuel Wendell Williston described the most complete skeleton then known (P 25026) discovered in 1901 by H. T. Martin. In 1903, Williston named a second species, N. leptodactylus, but this is today considered identical to N. gracilis.

  In 1953, Llewellyn Ivor Price named a partial humerus, DGM 238-R found in Brazil, N. lamegoi; the specific name honours the geologist Alberto Ribeiro Lamego. This species has an estimated wingspan of four metres; it is today generally considered to be a form different from Nyctosaurus, but has not yet been assigned its own genus name.

 In 1972 a new skeleton, FHSM VP-2148, in 1962 discovered by George Fryer Sternberg, was named N. bonneri; it is today generally seen as identical to N. gracilis.

 In 1978 Gregory Brown prepared the most complete Nyctosaurus skeleton currently known, UNSM 93000.

 In 1984 Robert Milton Schoch renamed Pteranodon nanus (Marsh 1881), "the dwarf", Nyctosaurus nanus. The question of this species validity is currently pending further study.

 In the early 2000s, Kenneth Jenkins of Ellis, Kansas collected two specimens of Nyctosaurus which were the first to demonstrate conclusively that not only was this species crested, the crest in mature specimens was very large and elaborate. The specimens were purchased by a private collector in Austin, Texas. Despite being in private hands rather than a museum collection, paleontologist Chris Bennett was able to study the specimens and gave them the manuscript reference numbers KJ1 and KJ2 (for Kenneth Jenkins). Bennett published a description of the specimens in 2003. Despite the unusual crests, the specimens were otherwise indistinguishable from other specimens of Nyctosaurus. However, the then-currently named species were extremely similar and Bennett declined to refer them to a specific one pending further study of the differences, or lack therof, between species of Nyctosaurus.

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