Как пишется динозавр тирекс

Тиранноза́вр (лат. Tyrannosaurus — «ящер-тиран», от др.-греч. τύραννος [tyrannos] — «тиран» и σαῦρος [sauros] — «ящер, ящерица», также встречается неправильное написание тиранозавр) — монотипический род плотоядных динозавров из группы целурозавров подотряда тероподов, включающий единственный валидный вид — Tyrannosaurus rex (лат. rex — «царь»). Обитал в западной части Северной Америки, которая в те времена представляла собой остров Ларамидию, и был наиболее распространённым из тираннозавридов. Окаменелые останки тираннозавров находят в различных геологических формациях, датирующихся маастрихтским веком мелового периода, около 67—65,5 миллионов лет назад. Был одним из последних ящеротазовых динозавров, существовавших перед катаклизмом, положившим конец эре динозавров (мел-палеогеновым вымиранием).

Все значения слова «тираннозавр»

• ящер
• динозавр
• брахиозавр
• диплодок
• ихтиозавр
• (ещё синонимы…)

В разделе «Рептилии и земноводные» мы впервые решили рассказать о таком животном, которое раньше, несомненно, было царем зверей, если можно так назвать. Для начала мы узнаем, что означает тирранозавр в переводе с латинского, назовем ближайших родственников этого хищника. Затем подробнее поговорим о его внешности и размерах. Конечно же, статья про тирранозавра была бы не полной, если бы мы не рассказали, на кого он охотился, где и когда обитал на Земле.

Распространенный вопрос: зачем тирранозавру маленькие передние лапы? Из данного обзора вы узнаете мнение ученых на этот счет. А сейчас предлагаем поближе познакомиться с этим огромным динозавром на нашем сайте про животных, а здесь смотрите дополнительную информацию про динозавров.

Тираннозавр Рекс — один из самых известных хищных динозавров. Даже современный лев с ним не сравнится. Частично своей популярностью он обязан средствам массовой информации, особенно выходу в свет фильма «Парк Юрского периода». В Американском музее естественной истории в Нью-Йорке это самый любимый посетителями экспонат.

Значение имени тирранозавра и его ближайшие родственники

Тираннозавр — дословно в переводе с латинского «ящер-тиран». Это название произошло от древнегреческих слов [tyrannos] — «тиран» и [sauros] — «ящерица, ящер». Рекс (rex) означает «царь». Так назвал и впервые описал этого динозавра в 1905 году известный американский биолог и палеонтолог Генри Ферфилд Осборн, который в то время являлся президентом Американского музея естественной истории в Нью-Йорке.

Род Тираннозавр относится к семейству Тираннозавровые и состоит лишь из одного вида животных — Тираннозавра Рекса — крупного плотоядного динозавра. Помимо него Тирранозавриды включают еще одно подсемейство, в которое входят альбертозавр, алектрозавр, алиорам, хингканкоузавр (Chingkankousaurus), дасплетозавры, эотиранн, горгозавр, нанотираннус и тарбозавр.

Размеры, внешность и особенности строения Тирекса

Самый крупный и наиболее полный скелет Тирекса из всех когда-либо найденных был назван Сью — в честь его первооткрывателя — палеонтолога Сью Хендриксона. Тщательно измерив кости Сью, ученые пришли к выводу, что Тирекс являлся одним из самых крупных хищных динозавров. В высоту он был до 4 метров (13 футов), а в длину — 12,3 м (40 футов). Недавно проведенный анализ Сью, результаты которого были опубликованы в 2011 году в журнале PLoS ONE, показывает, что Тирранозавр весил целых 9 тонн (точнее — 8160 кг).

У Тирекса были мощные бедра и длинный сильный хвост, который служил не только смертельным оружием, но в первую очередь противовесом его большой голове (череп Сью — 1,5 м, или 5 футов в длину) и позволял динозавру быстро передвигаться. В 2011 году были проведены исследования, которые смогли смоделировать распределение мышечной ткани по скелету ящера. Согласно полученным результатам можно предположить, что этот хищный динозавр мог развивать скорость от 17 до 40 км/ч (10-25 миль в час).

Передние лапы с двумя пальцами были настольно ничтожны, что становится очень маловероятным, что Т. Рекс мог их использовать для охоты или с их помощью подносить еду ко рту. «Мы не знаем, зачем ему нужны были эти небольшие лапы», — честно сказал палеонтолог Канзасского университета Дэвид Бернхэм.

У Тирранозавра самые сильные укусы из всех животных

Исследования массивного черепа Тирекса, проводимые в 2011 году и опубликованные в журнале Biology Letter, показали, что укус этого динозавра можно было бы по праву считать самым сильным из укусов всех животных, когда-либо обитавших на Земле. Эти показатели достигли внушительной цифры — 12814 фунтов-силы (57000 Ньютонов).

У Тирекса были самые крепкие и острые зубы, крупнейшие из которых доходили до 12 дюймов в длину. Но согласно исследованиям, проведенным в 2012 году и опубликованным в журнале «Науки о Земле», не все зубы выполняли одну и ту же функцию. В частности, передними зубами динозавр захватывал еду, боковые разрывали ее на части, а задние уже перемалывали и отправляли кусочки пищи дальше по пищевому тракту. Нужно отметить, что передние зубы были плоскими и прилегали гораздо плотнее между собой, чем боковые. Это исключало возможность сломать зуб во время захвата жертвы, когда она еще пыталась сопротивляться и вырваться.

На кого охотился Тирранозавр?

Это огромный хищник, который в первую очередь охотился на травоядных динозавров, в том числе на эдмонтозавра и трицератопса. «Постоянно охотясь, этот хищник съедал сотни фунтов мяса в течение своей жизни», — сказал Бернем.

«Вполне возможно, что Тирекс делился своей добычей, но делал это неохотно», сказал Бернхэм. «У него была тяжелая жизнь, он был постоянно голоден и поэтому все время охотился». На заметку: стрекозы тоже все время должны охотиться, об этом вы можете прочитать в статье про стрекоз.

«На протяжении многих лет собирались доказательства того, что основным занятием Тирранозавра являлась охота ради пропитания. Все они были косвенными и базировались лишь на следах укусов, на выпавших зубах, найденных возле останков других динозавров, а также наличии следов и даже целых охотничьих троп Тираннозавра», — сказал Бернхэм. Но в 2013 году в официальном журнале «Труды Национальной академии наук» Бернхэмом и его коллегами были наконец-то представлены прямые доказательства хищной природы Тирекса. Они обнаружили зуб Тираннозавра, застрявший между хвостовыми позвонками утконосого динозавра. Причем жертве удалось уйти от Тирекса, и со временем эта рана с зубом так и зарубцевалась.

«Мы нашли дымящийся пистолет!» — говорит Бернхэм. «Благодаря этому открытию мы теперь точно знаем, что монстр из наших снов действительно существовал».

В журнале «PLoS ONE» в 2010 году были опубликованы результаты анализов глубоких укусов и порезов, полученных от зубов Тираннозавра. И все же неясно, были ли Тираннозавры подвержены каннибализму, сражаясь на смерть с другими сородичами, или просто же поедали их останки.

Ученые уверены, что Тираннозавры охотились как в одиночку, так и вместе с другими динозаврами. В 2014 году в Скалистых горах Британской Колумбии были обнаружены следы, которые принадлежали трем динозаврам из семейства Тираннозавровых. Предположительно это были альбертозавр, горгозавр и дасплетозавр. В исследовании, опубликованном в журнале «PLoS ONE», говорится, что по крайней мере родственники Т. Рекса охотились стаями.

В каких местах и в какое время обитал Тирекс?

Ископаемые останки динозавра можно обнаружить в различных горных породах, относящихся к маастрихтскому ярусу позднемелового периода, который был около 65-67 миллионов лет назад, в конце мезозойской эры. Тирранозавр был одним из последних динозавров, которые не эволюционировали в птиц, и жил вплоть до мел-палеогенового вымирания, во время которого и исчезли динозавры.

Тирранозавр Рекс, в отличие от других наземных динозавров, постоянно бродил по всей западной части Северной Америки, которая в то время являлась огромным островом — Ларамидией. Согласно данным National Geographic, были обнаружены более 50 скелетов Тирекса, некоторые из них очень хорошо сохранились. На них видны даже остатки кожи и мышц.

Охотник за ископаемыми Барнум Браун обнаружил первый частичный скелет Тирранозавра Рекса в Хелл-Крик (Монтана) в 1902 году и спустя некоторое время продал его в Музей естественной истории Карнеги в Питсбурге. Другие останки Тираннозавра находятся в Американском музее естественной истории в Нью-Йорке.

В 2007 году ученые обнаружили след Т. Рекса в Хелл-крик и опубликовали это открытие в журнале «Палаиос». Но если этот отпечаток действительно принадлежит Тираннозавру, то он будет уже вторым, который нашли палеонтологи. Первый след был обнаружен в 1993 году в Нью-Мексико.

Тиранноза́вр, или тиранозавр (лат. Tyrannosaurus) — род плотоядных динозавров конца мелового периода. Наиболее крупный и известный в популярной культуре представитель рода — Tyrannosaurus rex, один из крупнейших наземных хищников: длина тела — до 13 м, масса — около 7 тонн.^[1][2] Открыт в 1902 году в США. Описано более 30 находок T. rex, все они принадлежат формациям возрастом примерно 68—65 млн лет назад.

Во время своего обитания — в конце мелового периода, тираннозавр был крупнейшим наземным плотоядным животным. Если же сравнивать всех известных науке динозавров, то тираннозавр является четвёртым по длине среди плотоядных динозавров, уступая лишь хищным динозаврам середины мелового периода — спинозавру, гиганотозавру и кархародонтозавру.

Название происходит от двух греческих слов: др.-греч. τύραννος — тиран и σαῦρος — ящер, ящерица.

Общее описание

Передние двупалые конечности в сравнении с могучими ногами сравнительно невелики. Хвост длинный и тяжёлый. Позвоночник состоит из 10 шейных, 12 грудных, пяти крестцовых и около 40 хвостовых позвонков.^[3] Шея, как и у других теропод, имеет S-образную форму, но короткая и толстая для удержания массивной головы. Некоторые кости скелета имеют пустоты, уменьшая таким образом общую массу тела без значительной потери прочности.

Самый крупный из известных черепов тираннозавра имеет 1,5 м в длину.^[4] Форма черепа имеет значительные отличия в сравнении с тероподами из других семейств: чрезвычайно широкий сзади, череп спереди сильно сужается. По мнению специалистов^[5][6], с таким строением черепа тираннозавры обладали необыкновенно хорошим бинокулярным зрением. Особенности строения костей черепа в семействе тираннозаврид делает их укус несравнимо более мощным по сравнению с прочими тероподами. Верхушка верхних челюстей имеет U-образную форму (у большинства других хищных тероподов она V-образная), что увеличивает объем мяса и костей, которое тираннозавр может оторвать за один укус, хотя и за счёт добавочной нагрузки на передние зубы.^{[7] [8]}

Зубы тираннозавра различаются по форме. D-образные в поперечном сечении передние зубы плотно прилегают друг к другу. Они загнуты внутрь пасти и с задней стороны усилены гребнями. Расположение и форма передних зубов снижают риск их вырывания во время кусаний и рывков. Внутренние зубы имеют скорее форму банана, а не кинжала. Они более широко расставлены, но тоже имеют усиливающие прочность гребни с задней стороны. Полная (включая корень) длина крупнейшего из найденных зубов оценивается в 30 см. Это самый длинный зуб среди зубов всех хищных динозавров.^[9]

Тираннозавр передвигался на задних конечностях, как и другие представители семейства тираннозаврид.

Способ питания

Окончательно не установлено, были ли тираннозавры хищниками или же они питались падалью.

Версия: Тираннозавры — падальщики. Один из палеонтологов, американский эксперт Джек Хорнер (англ. Jack Horner) утверждает, что тираннозавры были исключительно падальщиками и вовсе не принимали участие в охоте^[10]. Его гипотеза строится на следующих утверждениях:

• тираннозавры имели большие (относительно к размеру мозга) обонятельные рецепторы, что предполагает хорошо развитое обоняние, которое предположительно служило для нахождения гниющих останков на огромных расстояниях;
• мощные зубы длиной в 18 см каждый позволяют дробить кости, что требуется не столько для убийства, сколько для извлечения как можно больше пищи из того, что осталось от туши, включая костный мозг;
• если принимать, что тираннозавры ходили, а не бегали (см. ниже), а их добыча двигалась гораздо быстрее их, то это может служить доказательством в пользу питания падалью.

Версия: Тираннозавры — хищники. Существуют доказательства и в пользу хищнического образа жизни тираннозавра:

• глазные впадины расположены таким образом, что глаза могли смотреть вперёд, обеспечивая тираннозавра бинокулярным зрением (позволяя точно оценивать расстояния), что требуется в первую очередь хищнику (хотя есть много исключений);
• следы укусов на других животных и даже других тираннозаврах;
• сравнительная редкость находок останков тираннозавров, в любой экосистеме количество крупных хищников значительно меньше их жертв.

При изучении одного из тираннозавров палеонтолог Питер Ларсон (Peter Larson) обнаружил заживший перелом малоберцовой кости и одного позвонка, царапины на лицевых костях и зуб другого тираннозавра, внедрившийся в шейный позвонок. Если предположения верны, то это свидетельствует об агрессивном поведении тираннозавров по отношению друг к другу, хотя остаются неясными мотивы: было ли это конкуренцией за пищу/партнёра или же примером каннибализма. Однако более поздние исследования этих ран показали, что большинство их имеют не травматический, а инфекционный характер, или же могли быть нанесены уже после смерти.

Многие крупные растительноядные динозавры имели защиту на спине, что свидетельствует об опасности нападения высокого хищника с мощными челюстями.

Тираннозавры — хищники и падальщики. Многие учёные считают, что тираннозавры могли иметь смешанный рацион, как, например, современные львы — хищники, но могут поедать останки от убитых гиенами животных.

Способ передвижения

Спорным вопросом остаётся способ передвижения тираннозавра. Часть учёных склоняется к версии, что они могли бегать, достигая скорости 40—70 км/ч. Другие считают, что тираннозавры ходили, а не бегали.

«По всей видимости, — пишет Герберт Уэллс в знаменитых „Очерках истории цивилизации“, — тираннозавры передвигались, как кенгуру, опираясь на массивный хвост и задние ноги. Некоторые учёные даже предполагают, что тираннозавр двигался прыжками — в таком случае, он должен был обладать совершенно невероятными мускулами. Прыгающий слон куда меньше поражал бы воображение. Скорее всего, тираннозавр охотился на травоядных рептилий — обитателей болот. Наполовину погрузившись в жидкую болотную грязь, он преследовал свою жертву по протокам и озерцам заболоченных равнин, вроде нынешних Норфолкских болот или болот Эверглейдс во Флориде».

Мнение о двуногих динозаврах — подобиях кенгуру было широко распространено до середины XX века. Изучение следов, тем не менее, не показало наличия отпечатков хвоста. Все хищные динозавры при ходьбе держали тело горизонтально, хвост служил противовесом и балансиром. В целом тираннозавр близок по облику к огромной бегающей птице.

Недавние исследования белков, найденных при исследовании ископаемого бедра тираннозавра показали близость динозавров к птицам. Тираннозавр происходит от мелких хищных динозавров конца юрской эпохи, а не от карнозавров. Известные в настоящее время мелкие предки тираннозавра (например, дилонг из раннего мела Китая) были оперены тонкими волосовидными перьями. У самого тираннозавра перьев могло и не быть (известные отпечатки кожи бедра тираннозавра несут типичный для динозавров рисунок из полигональных чешуй).

Тираннозавр в массовой культуре

Благодаря внушительным размерам, огромным зубам и прочим внушительным габаритам, за весь XX век тираннозавр стал одним из самых узнаваемых динозавров в мире. Кроме того, именно по этому он часто становился главным динозавром убийцей в кинофильмах, таких как Затерянный мир, Кинг Конг. Самым главным и самым запоминающимся фильмом с участием тираннозавра считается фильм Стивена Спилберга Парк Юрского периода. Ни в одном из предыдущих фильмах Тираннозавр еще не внушал такого ужаса, как в этом. В фильме он подвергся тщательной проработке и по этому выглядел он очень внушительно. В фильме Парк юрского периода 2 присутствовала целая семья тираннозавров, что значительно снизило их отрицательную роль. В фильме Парк Юрского периода 3 разработчикам понадобился новый динозавр на роль главного злодея, и их выбор пал на египетского спинозавра. Сам Тираннозавр появился в фильме лишь на мгновение и был сразу убит спинозавром. До сих пор Тираннозавр считается одним из самых знаменитых динозавров и довольно часто встречается в рассказах, мультфильмах, почтовых марках и прочих атрибутах массовой культуры.

Интересные факты

• Учёные Гарвардского университета, изучив некоторые белки, полученные из костей тираннозавра, обитавшего 68 млн лет назад, и сравнив их структуру с современными птицами, пришли к выводу, что птицы имеют потенциально доказуемое филогенетическое родство с этим динозавром. Более того, птицы могут являться более близкими родственниками древнего ящера, нежели современные рептилии, такие как ящерицы и аллигаторы. Это подтверждает теорию о том, что птицы произошли от одной из групп динозавров. Чтобы подтвердить данное предположение, группа учёных продолжила данное исследование^[11][12].
• Примерно в одно время с тираннозавром на территории нынешней Азии обитал практически неотличимый от него вид — тарбозавр. Тарбозавры имели чуть более изящное строение и чуть меньшие размеры.

Примечания

1. ↑ Erickson, Gregory M.; Makovicky, Peter J.; Currie, Philip J.; Norell, Mark A.; Yerby, Scott A.; & Brochu, Christopher A. (2004). «Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs». Nature 430 (7001): 772–775. DOI:10.1038/nature02699.
2. ↑ Christopher A. Brochu Osteology of Tyrannosaurus Rex: Insights from a Nearly Complete Skeleton and High-resolution Computed Tomographic Analysis of the Skull. — Northbrook, Illinois: Society of Vertebrate Paleontology, 2003.
3. ↑ Tyrannosaurus, — Northern State University:: Aberdeen, SD
4. ↑ Montana State University (2006-04-07). Museum unveils world’s largest T-rex skull. Пресс-релиз. Проверено 2008-09-13.
5. ↑ Stevens, Kent A. (June 2006). «Binocular vision in theropod dinosaurs» (PDF). Journal of Vertebrate Paleontology 26 (2): 321–330. DOI:[321:BVITD2.0.CO;2 10.1671/0272-4634(2006)26[321:BVITD]2.0.CO;2].
6. ↑ Jaffe, Eric (2006-07-01). «Sight for ‘Saur Eyes: T. rex vision was among nature’s best». Science News 170 (1): 3. DOI:10.2307/4017288. Проверено 2008-10-06.
7. ↑ Holtz, Thomas R. (1994). «The Phylogenetic Position of the Tyrannosauridae: Implications for Theropod Systematics». Journal of Palaeontology 68 (5): 1100–1117. Проверено 2008-10-08.
8. ↑ Predatory dinosaurs of the world: a complete illustrated guide. — New York: Simon and Schuster, 1988. — ISBN 0-671-61946-2Шаблон:Pn
9. ↑ Sue’s vital statistics. Sue at the Field Museum. Field Museum of Natural History. Проверено 15 сентября 2007.
10. ↑ Horner, J.R. and Lessem, D. (1993). The Complete T. rex: How Stunning New Discoveries Are Changing Our Understanding of the World’s Most Famous Dinosaur. New York: Simon & Schuster.
11. ↑ Chris L. Organ, Mary H. Schweitzer, Wenxia Zheng, Lisa M. Freimark, Lewis C. Cantley, John M. Asara Molecular Phylogenetics of Mastodon and Tyrannosaurus rex. Science 25 April 2008. p. 499 онлайн
12. ↑ Newsru.com «Американские ученые выяснили, что куры произошли от тираннозавров» Прочитано 2008-04-25

Tyrannosaurus (pronounced or , meaning ‘tyrant lizard’) was a genus of theropod dinosaur. The species Tyrannosaurus rex (‘rex’ meaning ‘king’ in Latin), commonly abbreviated to T. rex, is a fixture in popular culture. It lived throughout what is now western North America, with a much wider range than other tyrannosaurids. Fossils are found in a variety of rock formations dating to the last three million years of the Cretaceous Period, approximately 68 to 65 million years ago. It was among the last non-avian dinosaurs to exist prior to the Cretaceous–Tertiary extinction event.

Like other tyrannosaurids, Tyrannosaurus was a bipedal carnivore with a massive skull balanced by a long, heavy tail. Relative to the large and powerful hindlimbs, Tyrannosaurus forelimbs were small, though unusually powerful for their size, and bore two clawed digits. Although other theropods rivaled or exceeded Tyrannosaurus rex in size, it was the largest known tyrannosaurid and one of the largest known land predators, measuring up to 13 metres (43 ft) in length, up to 4 metres (13 ft) tall at the hips, and up to 6.8 metric tons (7.5 short tons) in weight. By far the largest carnivore in its environment, Tyrannosaurus rex may have been an apex predator, preying upon hadrosaurs and ceratopsians, although some experts have suggested it was primarily a scavenger. The debate over Tyrannosaurus as apex predator or scavenger is among the longest running debates in paleontology.

More than 30 specimens of Tyrannosaurus rex have been identified, some of which are nearly complete skeletons. Soft tissue and proteins have been reported in at least one of these specimens. The abundance of fossil material has allowed significant research into many aspects of its biology, including life history and biomechanics. The feeding habits, physiology and potential speed of Tyrannosaurus rex are a few subjects of debate. Its taxonomy is also controversial, with some scientists considering Tarbosaurus bataar from Asia to represent a second species of Tyrannosaurus and others maintaining Tarbosaurus as a separate genus. Several other genera of North American tyrannosaurids have also been synonymized with Tyrannosaurus.

Tyrannosaurus
[1]
Scientific classification
Kingdom: Animalia Phylum: Chordata Class: Reptilia Superorder: Dinosauria Order: Saurischia Suborder: Theropoda (unranked): Coelurosauria Superfamily: Tyrannosauroidea Family: Tyrannosauridae Subfamily: Tyrannosaurinae Genus: TyrannosaurusOsborn, 1905
Species
T. rex
Synonyms
Manospondylus Dynamosaurus ?Nanotyrannus Stygivenator Dinotyrannus

Description

Tyrannosaurus rex was one of the largest land carnivores of all time; the largest complete specimen, FMNH PR2081 («Sue»), measured 12.8 metres (42 ft) long, and was 4.0 metres (13 ft) tall at the hips. Mass estimates have varied widely over the years, from more than 7.2 metric tons (7.9 short tons), to less than 4.5 metric tons (5.0 short tons), with most modern estimates ranging between 5.4 and 6.8 metric tons (6.0 and 7.5 short tons). Although Tyrannosaurus rex was larger than the well known Jurassic theropod Allosaurus, it was slightly smaller than Cretaceous carnivores Spinosaurus and Giganotosaurus.

The neck of Tyrannosaurus rex formed a natural S-shaped curve like that of other theropods, but was short and muscular to support the massive head. The forelimbs had only two clawed fingers, along with an additional small metacarpal representing the remnant of a third digit. In contrast the hind limbs were among the longest in proportion to body size of any theropod. The tail was heavy and long, sometimes containing over forty vertebrae, in order to balance the massive head and torso. To compensate for the immense bulk of the animal, many bones throughout the skeleton were hollow, reducing its weight without significant loss of strength.

The largest known Tyrannosaurus rex skulls measure up to 5 feet (1.5 m) in length. Large fenestrae (openings) in the skull reduced weight and provided areas for muscle attachment, as in all carnivorous theropods. But in other respects Tyrannosaurus’ skull was significantly different from those of large non-tyrannosauroid theropods. It was extremely wide at the rear but had a narrow snout, allowing unusually good binocular vision. The skull bones were massive and the nasals and some other bones were fused, preventing movement between them; but many were pneumatized (contained a «honeycomb» of tiny air spaces) which may have made the bones more flexible as well as lighter. These and other skull-strengthening features are part of the tyrannosaurid trend towards an increasingly powerful bite, which easily surpassed that of all non-tyrannosaurids. The tip of the upper jaw was U-shaped (most non-tyrannosauroid carnivores had V-shaped upper jaws), which increased the amount of tissue and bone a tyrannosaur could rip out with one bite, although it also increased the stresses on the front teeth.The teeth of Tyrannosaurus rex displayed marked heterodonty (differences in shape). The premaxillary teeth at the front of the upper jaw were closely packed, D-shaped in cross-section, had reinforcing ridges on the rear surface, were incisiform (their tips were chisel-like blades) and curved backwards. The D-shaped cross-section, reinforcing ridges and backwards curve reduced the risk that the teeth would snap when Tyrannosaurus bit and pulled. The remaining teeth were robust, like «lethal bananas» rather than daggers; more widely spaced and also had reinforcing ridges. Those in the upper jaw were larger than those in all but the rear of the lower jaw. The largest found so far is estimated to have been 30 centimetres (12 in) long including the root when the animal was alive, making it the largest tooth of any carnivorous dinosaur.

Classification

Tyrannosaurus is the type genus of the superfamily Tyrannosauroidea, the family Tyrannosauridae, and the subfamily Tyrannosaurinae; in other words it is the standard by which paleontologists decide whether to include other species in the same group. Other members of the tyrannosaurine subfamily include the North American Daspletosaurus and the Asian Tarbosaurus, both of which have occasionally been synonymized with Tyrannosaurus. Tyrannosaurids were once commonly thought to be descendants of earlier large theropods such as megalosaurs and carnosaurs, although more recently they were reclassified with the generally smaller coelurosaurs.In 1955, Soviet paleontologist Evgeny Maleev named a new species, Tyrannosaurus bataar, from Mongolia. By 1965, this species had been renamed Tarbosaurus bataar. Despite the renaming, many phylogenetic analyses have found Tarbosaurus bataar to be the sister taxon of Tyrannosaurus rex, and it has often been considered an Asian species of Tyrannosaurus. A recent redescription of the skull of Tarbosaurus bataar has shown that it was much narrower than that of Tyrannosaurus rex and that during a bite, the distribution of stress in the skull would have been very different, closer to that of Alioramus, another Asian tyrannosaur. A related cladistic analysis found that Alioramus, not Tyrannosaurus, was the sister taxon of Tarbosaurus, which, if true, would suggest that Tarbosaurus and Tyrannosaurus should remain separate.

Other tyrannosaurid fossils found in the same formations as Tyrannosaurus rex were originally classified as separate taxa, including Aublysodon and Albertosaurus megagracilis, the latter being named Dinotyrannus megagracilis in 1995. However, these fossils are now universally considered to belong to juvenile Tyrannosaurus rex. A small but nearly complete skull from Montana, 60 centimetres (2.0 ft) long, may be an exception. This skull was originally classified as a species of Gorgosaurus (G. lancensis) by Charles W. Gilmore in 1946, but was later referred to a new genus, Nanotyrannus. Opinions remain divided on the validity of N. lancensis. Many paleontologists consider the skull to belong to a juvenile Tyrannosaurus rex. There are minor differences between the two species, including the higher number of teeth in N. lancensis, which lead some scientists to recommend keeping the two genera separate until further research or discoveries clarify the situation.

Manospondylus

The first fossil specimen which can be attributed to Tyrannosaurus rex consists of two partial vertebrae (one of which has been lost) found by Edward Drinker Cope in 1892 and described as Manospondylus gigas. Osborn recognized the similarity between M. gigas and Tyrannosaurus rex as early as 1917 but, due to the fragmentary nature of the Manospondylus vertebrae, he could not synonymize them conclusively.

In June 2000, the Black Hills Institute located the type locality of M. gigas in South Dakota and unearthed more tyrannosaur bones there. These were judged to represent further remains of the same individual, and to be identical to those of Tyrannosaurus rex. According to the rules of the International Code of Zoological Nomenclature (ICZN), the system that governs the scientific naming of animals, Manospondylus gigas should therefore have priority over Tyrannosaurus rex, because it was named first. However, the Fourth Edition of the ICZN, which took effect on 1 January 2000, states that «the prevailing usage must be maintained» when «the senior synonym or homonym has not been used as a valid name after 1899» and «the junior synonym or homonym has been used for a particular taxon, as its presumed valid name, in at least 25 works, published by at least 10 authors in the immediately preceding 50 years …» Tyrannosaurus rex may qualify as the valid name under these conditions and would most likely be considered a nomen protectum («protected name») under the ICZN if it was ever challenged, which it has not yet been. Manospondylus gigas would then be deemed a nomen oblitum («forgotten name»).

Paleobiology

Life history

The identification of several specimens as juvenile Tyrannosaurus rex has allowed scientists to document ontogenetic changes in the species, estimate the lifespan, and determine how quickly the animals would have grown. The smallest known individual (LACM 28471, the «Jordan theropod») is estimated to have weighed only 29.9 kg (66 lb), while the largest, such as FMNH PR2081 («Sue») most likely weighed over 5400 kg (6 short tons). Histologic analysis of Tyrannosaurus rex bones showed LACM 28471 had aged only 2 years when it died, while «Sue» was 28 years old, an age which may have been close to the maximum for the species.

Histology has also allowed the age of other specimens to be determined. Growth curves can be developed when the ages of different specimens are plotted on a graph along with their mass. A Tyrannosaurus rex growth curve is S-shaped, with juveniles remaining under 1800 kg (2 short tons) until approximately 14 years of age, when body size began to increase dramatically. During this rapid growth phase, a young Tyrannosaurus rex would gain an average of 600 kg (1,300 lb) a year for the next four years. At 18 years of age, the curve plateaus again, indicating that growth slowed dramatically. For example, only 600 kg (1,300 lb) separated the 28-year-old «Sue» from a 22-year-old Canadian specimen (RTMP 81.12.1). Another recent histological study performed by different workers corroborates these results, finding that rapid growth began to slow at around 16 years of age. This sudden change in growth rate may indicate physical maturity, a hypothesis which is supported by the discovery of medullary tissue in the femur of a 16 to 20-year-old Tyrannosaurus rex from Montana (MOR 1125, also known as «B-rex»). Medullary tissue is found only in female birds during ovulation, indicating that «B-rex» was of reproductive age. Further study indicates an age of 18 for this specimen. Other tyrannosaurids exhibit extremely similar growth curves, although with lower growth rates corresponding to their lower adult sizes.

Over half of the known Tyrannosaurus rex specimens appear to have died within six years of reaching sexual maturity, a pattern which is also seen in other tyrannosaurs and in some large, long-lived birds and mammals today. These species are characterized by high infant mortality rates, followed by relatively low mortality among juveniles. Mortality increases again following sexual maturity, partly due to the stresses of reproduction. One study suggests that the rarity of juvenile Tyrannosaurus rex fossils is due in part to low juvenile mortality rates; the animals were not dying in large numbers at these ages, and so were not often fossilized. However, this rarity may also be due to the incompleteness of the fossil record or to the bias of fossil collectors towards larger, more spectacular specimens.

Sexual dimorphism

As the number of specimens increased, scientists began to analyze the variation between individuals and discovered what appeared to be two distinct body types, or morphs, similar to some other theropod species. As one of these morphs was more solidly built, it was termed the ‘robust’ morph while the other was termed ‘gracile.’ Several morphological differences associated with the two morphs were used to analyze sexual dimorphism in Tyrannosaurus rex, with the ‘robust’ morph usually suggested to be female. For example, the pelvis of several ‘robust’ specimens seemed to be wider, perhaps to allow the passage of eggs. It was also thought that the ‘robust’ morphology correlated with a reduced chevron on the first tail vertebra, also ostensibly to allow eggs to pass out of the reproductive tract, as had been erroneously reported for crocodiles.

In recent years, evidence for sexual dimorphism has been weakened. A 2005 study reported that previous claims of sexual dimorphism in crocodile chevron anatomy were in error, casting doubt on the existence of similar dimorphism between Tyrannosaurus rex genders. A full-sized chevron was discovered on the first tail vertebra of «Sue,» an extremely robust individual, indicating that this feature could not be used to differentiate the two morphs anyway. As Tyrannosaurus rex specimens have been found from Saskatchewan to New Mexico, differences between individuals may be indicative of geographic variation rather than sexual dimorphism. The differences could also be age-related, with ‘robust’ individuals being older animals.

Only a single Tyrannosaurus rex specimen has been conclusively shown to belong to a specific gender. Examination of «B-rex» demonstrated the preservation of soft tissue within several bones. Some of this tissue has been identified as medullary tissue, a specialized tissue grown only in modern birds as a source of calcium for the production of eggshell during ovulation. As only female birds lay eggs, medullary tissue is only found naturally in females, although males are capable of producing it when injected with female reproductive hormones like estrogen. This strongly suggests that «B-rex» was female, and that she died during ovulation. Recent research has shown that medullary tissue is never found in crocodiles, which are thought to be the closest living relatives of dinosaurs, aside from birds. The shared presence of medullary tissue in birds and theropod dinosaurs is further evidence of the close evolutionary relationship between the two.

Posture

Like many bipedal dinosaurs, Tyrannosaurus rex was historically depicted as a ‘living tripod’, with the body at 45 degrees or less from the vertical and the tail dragging along the ground, similar to a kangaroo. This concept dates from Joseph Leidy’s 1865 reconstruction of Hadrosaurus, the first to depict a dinosaur in a bipedal posture. Henry Fairfield Osborn, former president of the American Museum of Natural History (AMNH) in New York City, who believed the creature stood upright, further reinforced the notion after unveiling the first complete Tyrannosaurus rex skeleton in 1915. It stood in this upright pose for nearly a century, until it was dismantled in 1992. By 1970, scientists realized this pose was incorrect and could not have been maintained by a living animal, as it would have resulted in the dislocation or weakening of several joints, including the hips and the articulation between the head and the spinal column. The inaccurate AMNH mount inspired similar depictions in many films and paintings (such as Rudolph Zallinger’s famous mural The Age Of Reptiles in Yale University’s Peabody Museum of Natural History) until the 1990s, when films such as Jurassic Park introduced a more accurate posture to the general public. Modern representations in museums, art, and film show Tyrannosaurus rex with its body approximately parallel to the ground and tail extended behind the body to balance the head.

Arms

When Tyrannosaurus rex was first discovered, the humerus was the only element of the forelimb known. For the initial mounted skeleton as seen by the public in 1915, Osborn substituted longer, three-fingered forelimbs like those of Allosaurus. However, a year earlier, Lawrence Lambe described the short, two-fingered forelimbs of the closely related Gorgosaurus. This strongly suggested that Tyrannosaurus rex had similar forelimbs, but this hypothesis was not confirmed until the first complete Tyrannosaurus rex forelimbs were identified in 1989, belonging to MOR 555 (the «Wankel rex»). The remains of «Sue» also include complete forelimbs. Tyrannosaurus rex arms are very small relative to overall body size, measuring only 1 metre (3.3 ft) long. However, they are not vestigial but instead show large areas for muscle attachment, indicating considerable strength. This was recognized as early as 1906 by Osborn, who speculated that the forelimbs may have been used to grasp a mate during copulation. It has also been suggested that the forelimbs were used to assist the animal in rising from a prone position. Another possibility is that the forelimbs held struggling prey while it was dispatched by the tyrannosaur’s enormous jaws. This hypothesis may be supported by biomechanical analysis.

Tyrannosaurus rex forelimb bones exhibit extremely thick cortical bone, indicating that they were developed to withstand heavy loads. The biceps brachii muscle of a full-grown Tyrannosaurus rex was capable of lifting 199 kilograms (439 lb) by itself; this number would only increase with other muscles (like the brachialis) acting in concert with the biceps. A Tyrannosaurus rex forearm also had a reduced range of motion, with the shoulder and elbow joints allowing only 40 and 45 degrees of motion, respectively. In contrast, the same two joints in Deinonychus allow up to 88 and 130 degrees of motion, respectively, while a human arm can rotate 360 degrees at the shoulder and move through 165 degrees at the elbow. The heavy build of the arm bones, extreme strength of the muscles, and limited range of motion may indicate a system designed to hold fast despite the stresses of a struggling prey animal.

Soft tissue

In the March 2005 issue of Science, Mary Higby Schweitzer of North Carolina State University and colleagues announced the recovery of soft tissue from the marrow cavity of a fossilized leg bone, from a 68-million-year-old Tyrannosaurus. The bone had been intentionally, though reluctantly, broken for shipping and then not preserved in the normal manner, specifically because Schweitzer was hoping to test it for soft tissue. Designated as the Museum of the Rockies specimen 1125, or MOR 1125, the dinosaur was previously excavated from the Hell Creek Formation. Flexible, bifurcating blood vessels and fibrous but elastic bone matrix tissue were recognized. In addition, microstructures resembling blood cells were found inside the matrix and vessels. The structures bear resemblance to ostrich blood cells and vessels. Whether an unknown process, distinct from normal fossilization, preserved the material, or the material is original, the researchers do not know, and they are careful not to make any claims about preservation. If it is found to be original material, any surviving proteins may be used as a means of indirectly guessing some of the DNA content of the dinosaurs involved, because each protein is typically created by a specific gene. The absence of previous finds may merely be the result of people assuming preserved tissue was impossible, therefore simply not looking. Since the first, two more tyrannosaurs and a hadrosaur have also been found to have such tissue-like structures. Research on some of the tissues involved has suggested that birds are closer relatives to tyrannosaurs than other modern animals.

In studies reported in the journal Science in April 2007, Asara and colleagues concluded that seven traces of collagen proteins detected in purified Tyrannosaurus rex bone most closely match those reported in chickens, followed by frogs and newts. The discovery of proteins from a creature tens of millions of years old, along with similar traces the team found in a mastodon bone at least 160,000 years old, upends the conventional view of fossils and may shift paleontologists’ focus from bone hunting to biochemistry. Until these finds, most scientists presumed that fossilization replaced all living tissue with inert minerals. Paleontologist Hans Larsson of McGill University in Montreal, who was not part of the studies, called the finds «a milestone», and suggested that dinosaurs could «enter the field of molecular biology and really slingshot paleontology into the modern world».

Subsequent studies in April 2008 confirmed the close connection of Tyrannosaurus rex to modern birds. Postdoctoral biology researcher Chris Organ at Harvard University announced, «With more data, they would probably be able to place T. rex on the evolutionary tree between alligators and chickens and ostriches.» Co-author John M. Asara added, «We also show that it groups better with birds than modern reptiles, such as alligators and green anole lizards.»

The presumed soft tissue was called into question by Thomas Kaye of the University of Washington and his co-authors in 2008. They contend that what was really inside the tyrannosaur bone was slimy biofilm created by bacteria that coated the voids once occupied by blood vessels and cells. The researchers found that what previously had been identified as remnants of blood cells, because of the presence of iron, were actually framboids, microscopic mineral spheres bearing iron. They found similar spheres in a variety of other fossils from various periods, including an ammonite. In the ammonite they found the spheres in a place where the iron they contain could not have had any relationship to the presence of blood.

Skin and feathers

In 2004, the scientific journal Nature published a report describing an early tyrannosauroid, Dilong paradoxus, from the famous Yixian Formation of China. As with many other theropods discovered in the Yixian, the fossil skeleton was preserved with a coat of filamentous structures which are commonly recognized as the precursors of feathers. It has also been proposed that Tyrannosaurus and other closely related tyrannosaurids had such protofeathers. However, skin impressions from large tyrannosaurid specimens show mosaic scales. While it is possible that protofeathers existed on parts of the body which have not been preserved, a lack of insulatory body covering is consistent with modern multi-ton mammals such as elephants, hippopotamus, and most species of rhinoceros. As an object increases in size, its ability to retain heat increases due to its decreasing surface area-to-volume ratio. Therefore, as large animals evolve in or disperse into warm climates, a coat of fur or feathers loses its selective advantage for thermal insulation and can instead become a disadvantage, as the insulation traps excess heat inside the body, possibly overheating the animal. Protofeathers may also have been secondarily lost during the evolution of large tyrannosaurids like Tyrannosaurus, especially in warm Cretaceous climates.

Thermoregulation

Tyrannosaurus, like most dinosaurs, was long thought to have an ectothermic («cold-blooded») reptilian metabolism. The idea of dinosaur ectothermy was challenged by scientists like Robert T. Bakker and John Ostrom in the early years of the «Dinosaur Renaissance», beginning in the late 1960s. Tyrannosaurus rex itself was claimed to have been endothermic («warm-blooded»), implying a very active lifestyle. Since then, several paleontologists have sought to determine the ability of Tyrannosaurus to regulate its body temperature. Histological evidence of high growth rates in young Tyrannosaurus rex, comparable to those of mammals and birds, may support the hypothesis of a high metabolism. Growth curves indicate that, as in mammals and birds, Tyrannosaurus rex growth was limited mostly to immature animals, rather than the indeterminate growth seen in most other vertebrates.

Oxygen isotope ratios in fossilized bone are sometimes used to determine the temperature at which the bone was deposited, as the ratio between certain isotopes correlates with temperature. In one specimen, the isotope ratios in bones from different parts of the body indicated a temperature difference of no more than 4 to 5°C (7 to 9°F) between the vertebrae of the torso and the tibia of the lower leg. This small temperature range between the body core and the extremities was claimed by paleontologist Reese Barrick and geochemist William Showers to indicate that Tyrannosaurus rex maintained a constant internal body temperature (homeothermy) and that it enjoyed a metabolism somewhere between ectothermic reptiles and endothermic mammals. Other scientists have pointed out that the ratio of oxygen isotopes in the fossils today does not necessarily represent the same ratio in the distant past, and may have been altered during or after fossilization (diagenesis). Barrick and Showers have defended their conclusions in subsequent papers, finding similar results in another theropod dinosaur from a different continent and tens of millions of years earlier in time (Giganotosaurus). Ornithischian dinosaurs also showed evidence of homeothermy, while varanid lizards from the same formation did not. Even if Tyrannosaurus rex does exhibit evidence of homeothermy, it does not necessarily mean that it was endothermic. Such thermoregulation may also be explained by gigantothermy, as in some living sea turtles.

Footprints

Two isolated fossilized footprints have been tentatively assigned to Tyrannosaurus rex. The first was discovered at Philmont Scout Ranch, New Mexico, in 1983 by American geologist Charles Pillmore. Originally thought to belong to a hadrosaurid, examination of the footprint revealed a large ‘heel’ unknown in ornithopod dinosaur tracks, and traces of what may have been a hallux, the dewclaw-like fourth digit of the tyrannosaur foot. The footprint was published as the ichnogenus Tyrannosauripus pillmorei in 1994, by Martin Lockley and Adrian Hunt. Lockley and Hunt suggested that it was very likely the track was made by a Tyrannosaurus rex, which would make it the first known footprint from this species. The track was made in what was once a vegetated wetland mud flat. It measures 83 centimetres (33 in) long by 71 centimetres (28 in) wide.

A second footprint that may have been made by a Tyrannosaurus was first reported in 2007 by British paleontologist Phil Manning, from the Hell Creek Formation of Montana. This second track measures 76 centimetres (30 in) long, shorter than the track described by Lockley and Hunt. Whether or not the track was made by Tyrannosaurus is unclear, though Tyrannosaurus and Nanotyrannus are the only large theropods known to have existed in the Hell Creek Formation. Further study of the track (a full description has not yet been published) will compare the Montana track with the one found in New Mexico.

Locomotion

There are two main issues concerning the locomotory abilities of Tyrannosaurus: how well it could turn; and what its maximum straight-line speed was likely to have been. Both are relevant to the debate about whether it was a hunter or a scavenger (see below).

Tyrannosaurus may have been slow to turn, possibly taking one to two seconds to turn only 45°—an amount that humans, being vertically oriented and tail-less, can spin in a fraction of a second. The cause of the difficulty is rotational inertia, since much of Tyrannosaurus’ mass was some distance from its center of gravity, like a human carrying a heavy timber—although it might have reduced the average distance by arching its back and tail and pulling its head and forelimbs close to its body, rather like the way ice skaters pull their arms closer in order to spin faster.

Scientists have produced a wide range of maximum speed estimates, mostly around 11 metres per second (25 mph), but a few as low as 5–11 metres per second (11–25 mph), and a few as high as 20 metres per second (45 mph). Researchers have to rely on various estimating techniques because, while there are many tracks of very large theropods walking, so far none have been found of very large theropods running—and this absence may indicate that they did not run. Scientists who think that Tyrannosaurus was able to run point out that hollow bones and other features that would have lightened its body may have kept adult weight to a mere 5 tons or so, or that other animals like ostriches and horses with long, flexible legs are able to achieve high speeds through slower but longer strides. Additionally, some have argued that Tyrannosaurus had relatively larger leg muscles than any animal alive today, which could have enabled fast running 40–70 kilometres per hour (25–43 mph).

Jack Horner and Don Lessem argued in 1993 that Tyrannosaurus was slow and probably could not run (no airborne phase in mid-stride), because its ratio of femur (thigh bone) to tibia (shin bone) length was greater than 1, as in most large theropods and like a modern elephant. However, Holtz (1998) noted that tyrannosaurids and some closely related groups had significantly longer distal hindlimb components (shin plus foot plus toes) relative to the femur length than most other theropods), and that tyrannosaurids and their close relatives had a tightly interlocked metatarsus that more effectively transmitted locomotory forces from the foot to the lower leg than in earlier theropods («metatarsus» means the foot bones, which function as part of the leg in digitigrade animals). He therefore concluded that tyrannosaurids and their close relatives were the fastest large theropods.Christiansen (1998) estimated that the leg bones of Tyrannosaurus were not significantly stronger than those of elephants, which are relatively limited in their top speed and never actually run (there is no airborne phase), and hence proposed that the dinosaur’s maximum speed would have been about 11 metres per second (25 mph), which is about the speed of a human sprinter. But he also noted that such estimates depend on many dubious assumptions.

Farlow and colleagues (1995) have argued that a Tyrannosaurus weighing 6 short tons (5.4 t) to 8 short tons (7.3 t) would have been critically or even fatally injured if it had fallen while moving quickly, since its torso would have slammed into the ground at a deceleration of 6 g (six times the acceleration due to gravity, or about 60 meters/s²) and its tiny arms could not have reduced the impact. However, giraffes have been known to gallop at 50 kilometres per hour (31 mph), despite the risk that they might break a leg or worse, which can be fatal even in a «safe» environment such as a zoo. Thus it is quite possible that Tyrannosaurus also moved fast when necessary and had to accept such risks.

Most recent research on Tyrannosaurus locomotion does not support speeds faster than 40 kilometres per hour (25 mph), i.e. moderate-speed running. For example, a 2002 paper in the journal Nature used a mathematical model (validated by applying it to three living animals, alligators, chickens, and humans; additionally later eight more species including emus and ostriches) to gauge the leg muscle mass needed for fast running (over 40 kilometres per hour (25 mph)). They found that proposed top speeds in excess of 40 kilometres per hour (25 mph) were unfeasible, because they would require very large leg muscles (more than approximately 40–86% of total body mass). Even moderately fast speeds would have required large leg muscles. This discussion is difficult to resolve, as it is unknown how large the leg muscles actually were in Tyrannosaurus. If they were smaller, only 18 kilometres per hour (11 mph) walking/jogging might have been possible.

A study in 2007 used computer models to estimate running speeds, based on data taken directly from fossils, and claimed that Tyrannosaurus rex had a top running speed of 8 metres per second (18 mph). An average professional football (soccer) player would be slightly slower, while a human sprinter can reach 12 metres per second (27 mph). Note that these computer models predict a top speed of 17.8 metres per second (40 mph) for a 3 kilograms (6.6 lb) Compsognathus (probably a juvenile individual).

Those who argue that Tyrannosaurus was incapable of running estimate the top speed of Tyrannosaurus at about 17 kilometres per hour (11 mph). This is still faster than its most likely prey species, hadrosaurids and ceratopsians. In addition, some advocates of the idea that Tyrannosaurus was a predator claim that tyrannosaur running speed is not important, since it may have been slow but still faster than its probable prey. However, Paul and Christiansen (2000) argued that at least the later ceratopsians had upright forelimbs and the larger species may have been as fast as rhinos. Healed Tyrannosaurus bite wounds on ceratopsian fossils are interpreted as evidence of attacks on living ceratopsians (see below). If the ceratopsians that lived alongside Tyrannosaurus were fast, that casts doubt on the argument that Tyrannosaurus did not have to be fast to catch its prey.

Feeding strategies

The debate about whether Tyrannosaurus was a predator or a pure scavenger is as old as the debate about its locomotion. Lambe (1917) described a good skeleton of Tyrannosaurus’ close relative Gorgosaurus and concluded that it and therefore also Tyrannosaurus was a pure scavenger, because the Gorgosaurus’ teeth showed hardly any wear. This argument is no longer taken seriously, because theropods replaced their teeth quite rapidly. Ever since the first discovery of Tyrannosaurus most scientists have agreed that it was a predator, although like modern large predators it would have been happy to scavenge or steal another predator’s kill if it had the opportunity.

Noted hadrosaur expert Jack Horner is currently the major advocate of the idea that Tyrannosaurus was exclusively a scavenger and did not engage in active hunting at all. Horner has presented several arguments to support the pure scavenger hypothesis:

• Tyrannosaur arms are short when compared to other known predators. Horner argues that the arms were too short to make the necessary gripping force to hold on to prey.
• Tyrannosaurs had large olfactory bulbs and olfactory nerves (relative to their brain size). These suggest a highly developed sense of smell which could sniff out carcasses over great distances, as modern vultures do. Research on the olfactory bulbs of dinosaurs has shown that Tyrannosaurus had the most highly developed sense of smell of 21 sampled dinosaurs. Opponents of the pure scavenger hypothesis have used the example of vultures in the opposite way, arguing that the scavenger hypothesis is implausible because the only modern pure scavengers are large gliding birds, which use their keen senses and energy-efficient gliding to cover vast areas economically. However, researchers from Glasgow concluded that an ecosystem as productive as the current Serengeti would provide sufficient carrion for a large theropod scavenger, although the theropod might have had to be cold-blooded in order to get more calories from carrion than it spent on foraging (see Warm-bloodedness of dinosaurs). They also suggested that modern ecosystems like Serengeti have no large terrestrial scavengers because gliding birds now do the job much more efficiently, while large theropods did not face competition for the scavenger ecological niche from gliding birds.
• Tyrannosaur teeth could crush bone, and therefore could extract as much food (bone marrow) as possible from carcass remnants, usually the least nutritious parts. Karen Chin and colleagues have found bone fragments in coprolites (fossilized dung) that they attribute to tyrannosaurs, but point out that a tyrannosaur’s teeth were not well adapted to systematically chewing bone like hyenas do to extract marrow.
• Since at least some of Tyrannosaurus’s potential prey could move quickly, evidence that it walked instead of ran could indicate that it was a scavenger. On the other hand, recent analyses suggest that Tyrannosaurus, while slower than large modern terrestrial predators, may well have been fast enough to prey on large hadrosaurs and ceratopsians.Other evidence suggests hunting behavior in Tyrannosaurus. The eye-sockets of tyrannosaurs are positioned so that the eyes would point forward, giving them binocular vision slightly better than that of modern hawks. He also pointed out that the tyrannosaur lineage had a history of steadily improving binocular vision. It is hard to see how natural selection would have favored this long-term trend if tyrannosaurs had been pure scavengers, which would not have needed the advanced depth perception that stereoscopic vision provides. In modern animals, binocular vision is found mainly in predators.

A skeleton of the hadrosaurid Edmontosaurus annectens has been described from Montana with healed tyrannosaur-inflicted damage on its tail vertebrae. The fact that the damage seems to have healed suggests that the Edmontosaurus survived a tyrannosaur’s attack on a living target, i.e. the tyrannosaur had attempted active predation. There is also evidence for an aggressive interaction between a Triceratops and a Tyrannosaurus in the form of partially healed tyrannosaur tooth marks on a Triceratops brow horn and squamosal (a bone of the neck frill); the bitten horn is also broken, with new bone growth after the break. It is not known what the exact nature of the interaction was, though: either animal could have been the aggressor. When examining Sue, paleontologist Pete Larson found a broken and healed fibula and tail vertebrae, scarred facial bones and a tooth from another Tyrannosaurus embedded in a neck vertebra. If correct, these might be strong evidence for aggressive behavior between tyrannosaurs but whether it would have been competition for food and mates or active cannibalism is unclear. However, further recent investigation of these purported wounds has shown that most are infections rather than injuries (or simply damage to the fossil after death) and the few injuries are too general to be indicative of intraspecific conflict.

Some researchers argue that if Tyrannosaurus were a scavenger, another dinosaur had to be the top predator in the Amerasian Upper Cretaceous. Top prey was the larger marginocephalians and ornithopods. The other tyrannosaurids share so many characteristics that only small dromaeosaurs remain as feasible top predators. In this light, scavenger hypothesis adherents have suggested that the size and power of tyrannosaurs allowed them to steal kills from smaller predators. Most paleontologists accept that Tyrannosaurus was both an active predator and a scavenger like all large carnivores.

History

Henry Fairfield Osborn, president of the American Museum of Natural History, named Tyrannosaurus rex in 1905. The generic name is derived from the Greek words τυραννος (tyrannos, meaning «tyrant») and σαυρος (sauros, meaning «lizard»). Osborn used the Latin word rex, meaning «king», for the specific name. The full binomial therefore translates to «tyrant lizard king,» emphasizing the animal’s size and perceived dominance over other species of the time.

Earliest finds

Teeth from what is now documented as a Tyrannosaurus rex were found in 1874 by A. Lakes near Golden, Colorado. In the early 1890s, J. B. Hatcher collected postcranial elements in eastern Wyoming. The fossils were believed to be from a large species of Ornithomimus (O. grandis) but are now considered Tyrannosaurus rex. Vertebral fragments found by E. D. Cope in western South Dakota in 1892 and named as Manospondylus gigas have also been reclassified as Tyrannosaurus rex. Barnum Brown, assistant curator of the American Museum of Natural History, found the first partial skeleton of Tyrannosaurus rex in eastern Wyoming in 1900. H. F. Osborn originally named this skeleton Dynamosaurus imperiosus in a paper in 1905. Brown found another partial skeleton in the Hell Creek Formation in Montana in 1902. Osborn used this holotype to describe Tyrannosaurus rex in the same paper in which D. imperiosus was described. Had it not been for page order, Dynamosaurus would have become the official name. The original Dynamosaurus material resides in the collections of the Natural History Museum, London.

In total, Brown found five Tyrannosaurus partial skeletons. In 1941, Brown’s 1902 find was sold to the Carnegie Museum of Natural History in Pittsburgh, Pennsylvania. Brown’s fourth and largest find, also from Hell Creek, is on display in the American Museum of Natural History in New York.

Although there are numerous skeletons in the world, only one track has been documented — at Philmont Scout Ranch in northeast New Mexico. It was discovered in 1983 and identified and documented in 1994.

Notable specimens

Sue Hendrickson, amateur paleontologist, discovered the most complete (approximately 85%) and, until 2001, the largest, Tyrannosaurus fossil skeleton known in the Hell Creek Formation near Faith, South Dakota, on 12 August 1990. This Tyrannosaurus, nicknamed «Sue» in her honor, was the object of a legal battle over its ownership. In 1997 this was settled in favor of Maurice Williams, the original land owner. The fossil collection was purchased by the Field Museum of Natural History at auction for USD 7.6 million, making it the most expensive dinosaur skeleton to date. From 1998 to 1999 Field Museum of Natural History preparators spent 30,000 hours taking the rock off each of the bones. The bones were then shipped off to New Jersey where the mount was made. The finished mount was then taken apart, and along with the bones, shipped back to Chicago for the final assembly. The mounted skeleton opened to the public on May 17, 2000 in the great hall (Stanley Field Hall) at the Field Museum of Natural History. A study of this specimen’s fossilized bones showed that «Sue» reached full size at age 19 and died at age 28, the longest any tyrannosaur is known to have lived. Early speculation that Sue may have died from a bite to the back of the head was not confirmed. Though subsequent study showed many pathologies in the skeleton, no bite marks were found. Damage to the back of the skull may have been caused by post-mortem trampling.

Another Tyrannosaurus, nicknamed «Stan», in honor of amateur paleontologist Stan Sacrison, was found in the Hell Creek Formation near Buffalo, South Dakota, in the spring of 1987. After 30,000 hours of digging and preparing, a 65% complete skeleton emerged. Stan is currently on display in the Black Hills Museum of Natural History Exhibit in Hill City, South Dakota, after an extensive world tour. This tyrannosaur, too, was found to have many bone pathologies, including broken and healed ribs, a broken (and healed) neck and a spectacular hole in the back of its head, about the size of a Tyrannosaurus tooth. Both Stan and Sue were examined by Peter Larson.

In the summer of 2000, Jack Horner discovered five Tyrannosaurus skeletons near the Fort Peck Reservoir in Montana. One of the specimens, dubbed «C. rex,» was reported to be perhaps the largest Tyrannosaurus ever found.In 2001, a 50% complete skeleton of a juvenile Tyrannosaurus was discovered in the Hell Creek Formation in Montana, by a crew from the Burpee Museum of Natural History of Rockford, Illinois. Dubbed «Jane the Rockford T-Rex,» the find was initially considered the first known skeleton of the pygmy tyrannosaurid Nanotyrannus but subsequent research has revealed that it is more likely a juvenile Tyrannosaurus. It is the most complete and best preserved juvenile example known to date. Jane has been examined by Jack Horner, Pete Larson, Robert Bakker, action=edit redlink=1 Greg Erickson and several other renowned paleontologists, because of the uniqueness of her age. Jane is currently on exhibit at the Burpee Museum of Natural History in Rockford, Illinois.

In a press release on 7 April 2006, Montana State University revealed that it possessed the largest Tyrannosaurus skull yet discovered. Discovered in the 1960s and only recently reconstructed, the skull measures 59 inches (150 cm) long compared to the 55.4 inches (141 cm) of “Sue’s” skull, a difference of 6.5%.

Appearances in popular culture

Since it was first described in 1905, Tyrannosaurus rex has become the most widely recognized dinosaur species in popular culture. It is the only dinosaur that is commonly known to the general public by its full scientific name (binomial name) (Tyrannosaurus rex), and the scientific abbreviation T. rex has also come into wide usage (commonly abbreviated «T-Rex»). Robert T. Bakker notes this in The Dinosaur Heresies and explains that a name like «Tyrannosaurus rex is just irresistible to the tongue.»

Gallery

Tyrannosaurus (pronounced or , meaning ‘tyrant lizard’) was a genus of theropod dinosaur. The species Tyrannosaurus rex (‘rex’ meaning ‘king’ in Latin), commonly abbreviated to T. rex, is a fixture in popular culture. It lived throughout what is now western North America, with a much wider range than other tyrannosaurids. Fossils are found in a variety of rock formations dating to the last three million years of the Cretaceous Period, approximately 68 to 65 million years ago. It was among the last non-avian dinosaurs to exist prior to the Cretaceous–Tertiary extinction event.

Like other tyrannosaurids, Tyrannosaurus was a bipedal carnivore with a massive skull balanced by a long, heavy tail. Relative to the large and powerful hindlimbs, Tyrannosaurus forelimbs were small, though unusually powerful for their size, and bore two clawed digits. Although other theropods rivaled or exceeded Tyrannosaurus rex in size, it was the largest known tyrannosaurid and one of the largest known land predators, measuring up to 13 metres (43 ft) in length, up to 4 metres (13 ft) tall at the hips, and up to 6.8 metric tons (7.5 short tons) in weight. By far the largest carnivore in its environment, Tyrannosaurus rex may have been an apex predator, preying upon hadrosaurs and ceratopsians, although some experts have suggested it was primarily a scavenger. The debate over Tyrannosaurus as apex predator or scavenger is among the longest running debates in paleontology.

More than 30 specimens of Tyrannosaurus rex have been identified, some of which are nearly complete skeletons. Soft tissue and proteins have been reported in at least one of these specimens. The abundance of fossil material has allowed significant research into many aspects of its biology, including life history and biomechanics. The feeding habits, physiology and potential speed of Tyrannosaurus rex are a few subjects of debate. Its taxonomy is also controversial, with some scientists considering Tarbosaurus bataar from Asia to represent a second species of Tyrannosaurus and others maintaining Tarbosaurus as a separate genus. Several other genera of North American tyrannosaurids have also been synonymized with Tyrannosaurus.

Tyrannosaurus
[1]
Scientific classification
Kingdom: Animalia Phylum: Chordata Class: Reptilia Superorder: Dinosauria Order: Saurischia Suborder: Theropoda (unranked): Coelurosauria Superfamily: Tyrannosauroidea Family: Tyrannosauridae Subfamily: Tyrannosaurinae Genus: TyrannosaurusOsborn, 1905
Species
T. rex
Synonyms
Manospondylus Dynamosaurus ?Nanotyrannus Stygivenator Dinotyrannus

Description

Tyrannosaurus rex was one of the largest land carnivores of all time; the largest complete specimen, FMNH PR2081 («Sue»), measured 12.8 metres (42 ft) long, and was 4.0 metres (13 ft) tall at the hips. Mass estimates have varied widely over the years, from more than 7.2 metric tons (7.9 short tons), to less than 4.5 metric tons (5.0 short tons), with most modern estimates ranging between 5.4 and 6.8 metric tons (6.0 and 7.5 short tons). Although Tyrannosaurus rex was larger than the well known Jurassic theropod Allosaurus, it was slightly smaller than Cretaceous carnivores Spinosaurus and Giganotosaurus.

The neck of Tyrannosaurus rex formed a natural S-shaped curve like that of other theropods, but was short and muscular to support the massive head. The forelimbs had only two clawed fingers, along with an additional small metacarpal representing the remnant of a third digit. In contrast the hind limbs were among the longest in proportion to body size of any theropod. The tail was heavy and long, sometimes containing over forty vertebrae, in order to balance the massive head and torso. To compensate for the immense bulk of the animal, many bones throughout the skeleton were hollow, reducing its weight without significant loss of strength.

The largest known Tyrannosaurus rex skulls measure up to 5 feet (1.5 m) in length. Large fenestrae (openings) in the skull reduced weight and provided areas for muscle attachment, as in all carnivorous theropods. But in other respects Tyrannosaurus’ skull was significantly different from those of large non-tyrannosauroid theropods. It was extremely wide at the rear but had a narrow snout, allowing unusually good binocular vision. The skull bones were massive and the nasals and some other bones were fused, preventing movement between them; but many were pneumatized (contained a «honeycomb» of tiny air spaces) which may have made the bones more flexible as well as lighter. These and other skull-strengthening features are part of the tyrannosaurid trend towards an increasingly powerful bite, which easily surpassed that of all non-tyrannosaurids. The tip of the upper jaw was U-shaped (most non-tyrannosauroid carnivores had V-shaped upper jaws), which increased the amount of tissue and bone a tyrannosaur could rip out with one bite, although it also increased the stresses on the front teeth.The teeth of Tyrannosaurus rex displayed marked heterodonty (differences in shape). The premaxillary teeth at the front of the upper jaw were closely packed, D-shaped in cross-section, had reinforcing ridges on the rear surface, were incisiform (their tips were chisel-like blades) and curved backwards. The D-shaped cross-section, reinforcing ridges and backwards curve reduced the risk that the teeth would snap when Tyrannosaurus bit and pulled. The remaining teeth were robust, like «lethal bananas» rather than daggers; more widely spaced and also had reinforcing ridges. Those in the upper jaw were larger than those in all but the rear of the lower jaw. The largest found so far is estimated to have been 30 centimetres (12 in) long including the root when the animal was alive, making it the largest tooth of any carnivorous dinosaur.

Classification

Tyrannosaurus is the type genus of the superfamily Tyrannosauroidea, the family Tyrannosauridae, and the subfamily Tyrannosaurinae; in other words it is the standard by which paleontologists decide whether to include other species in the same group. Other members of the tyrannosaurine subfamily include the North American Daspletosaurus and the Asian Tarbosaurus, both of which have occasionally been synonymized with Tyrannosaurus. Tyrannosaurids were once commonly thought to be descendants of earlier large theropods such as megalosaurs and carnosaurs, although more recently they were reclassified with the generally smaller coelurosaurs.In 1955, Soviet paleontologist Evgeny Maleev named a new species, Tyrannosaurus bataar, from Mongolia. By 1965, this species had been renamed Tarbosaurus bataar. Despite the renaming, many phylogenetic analyses have found Tarbosaurus bataar to be the sister taxon of Tyrannosaurus rex, and it has often been considered an Asian species of Tyrannosaurus. A recent redescription of the skull of Tarbosaurus bataar has shown that it was much narrower than that of Tyrannosaurus rex and that during a bite, the distribution of stress in the skull would have been very different, closer to that of Alioramus, another Asian tyrannosaur. A related cladistic analysis found that Alioramus, not Tyrannosaurus, was the sister taxon of Tarbosaurus, which, if true, would suggest that Tarbosaurus and Tyrannosaurus should remain separate.

Other tyrannosaurid fossils found in the same formations as Tyrannosaurus rex were originally classified as separate taxa, including Aublysodon and Albertosaurus megagracilis, the latter being named Dinotyrannus megagracilis in 1995. However, these fossils are now universally considered to belong to juvenile Tyrannosaurus rex. A small but nearly complete skull from Montana, 60 centimetres (2.0 ft) long, may be an exception. This skull was originally classified as a species of Gorgosaurus (G. lancensis) by Charles W. Gilmore in 1946, but was later referred to a new genus, Nanotyrannus. Opinions remain divided on the validity of N. lancensis. Many paleontologists consider the skull to belong to a juvenile Tyrannosaurus rex. There are minor differences between the two species, including the higher number of teeth in N. lancensis, which lead some scientists to recommend keeping the two genera separate until further research or discoveries clarify the situation.

Manospondylus

The first fossil specimen which can be attributed to Tyrannosaurus rex consists of two partial vertebrae (one of which has been lost) found by Edward Drinker Cope in 1892 and described as Manospondylus gigas. Osborn recognized the similarity between M. gigas and Tyrannosaurus rex as early as 1917 but, due to the fragmentary nature of the Manospondylus vertebrae, he could not synonymize them conclusively.

In June 2000, the Black Hills Institute located the type locality of M. gigas in South Dakota and unearthed more tyrannosaur bones there. These were judged to represent further remains of the same individual, and to be identical to those of Tyrannosaurus rex. According to the rules of the International Code of Zoological Nomenclature (ICZN), the system that governs the scientific naming of animals, Manospondylus gigas should therefore have priority over Tyrannosaurus rex, because it was named first. However, the Fourth Edition of the ICZN, which took effect on 1 January 2000, states that «the prevailing usage must be maintained» when «the senior synonym or homonym has not been used as a valid name after 1899» and «the junior synonym or homonym has been used for a particular taxon, as its presumed valid name, in at least 25 works, published by at least 10 authors in the immediately preceding 50 years …» Tyrannosaurus rex may qualify as the valid name under these conditions and would most likely be considered a nomen protectum («protected name») under the ICZN if it was ever challenged, which it has not yet been. Manospondylus gigas would then be deemed a nomen oblitum («forgotten name»).

Paleobiology

Life history

The identification of several specimens as juvenile Tyrannosaurus rex has allowed scientists to document ontogenetic changes in the species, estimate the lifespan, and determine how quickly the animals would have grown. The smallest known individual (LACM 28471, the «Jordan theropod») is estimated to have weighed only 29.9 kg (66 lb), while the largest, such as FMNH PR2081 («Sue») most likely weighed over 5400 kg (6 short tons). Histologic analysis of Tyrannosaurus rex bones showed LACM 28471 had aged only 2 years when it died, while «Sue» was 28 years old, an age which may have been close to the maximum for the species.

Histology has also allowed the age of other specimens to be determined. Growth curves can be developed when the ages of different specimens are plotted on a graph along with their mass. A Tyrannosaurus rex growth curve is S-shaped, with juveniles remaining under 1800 kg (2 short tons) until approximately 14 years of age, when body size began to increase dramatically. During this rapid growth phase, a young Tyrannosaurus rex would gain an average of 600 kg (1,300 lb) a year for the next four years. At 18 years of age, the curve plateaus again, indicating that growth slowed dramatically. For example, only 600 kg (1,300 lb) separated the 28-year-old «Sue» from a 22-year-old Canadian specimen (RTMP 81.12.1). Another recent histological study performed by different workers corroborates these results, finding that rapid growth began to slow at around 16 years of age. This sudden change in growth rate may indicate physical maturity, a hypothesis which is supported by the discovery of medullary tissue in the femur of a 16 to 20-year-old Tyrannosaurus rex from Montana (MOR 1125, also known as «B-rex»). Medullary tissue is found only in female birds during ovulation, indicating that «B-rex» was of reproductive age. Further study indicates an age of 18 for this specimen. Other tyrannosaurids exhibit extremely similar growth curves, although with lower growth rates corresponding to their lower adult sizes.

Over half of the known Tyrannosaurus rex specimens appear to have died within six years of reaching sexual maturity, a pattern which is also seen in other tyrannosaurs and in some large, long-lived birds and mammals today. These species are characterized by high infant mortality rates, followed by relatively low mortality among juveniles. Mortality increases again following sexual maturity, partly due to the stresses of reproduction. One study suggests that the rarity of juvenile Tyrannosaurus rex fossils is due in part to low juvenile mortality rates; the animals were not dying in large numbers at these ages, and so were not often fossilized. However, this rarity may also be due to the incompleteness of the fossil record or to the bias of fossil collectors towards larger, more spectacular specimens.

Sexual dimorphism

As the number of specimens increased, scientists began to analyze the variation between individuals and discovered what appeared to be two distinct body types, or morphs, similar to some other theropod species. As one of these morphs was more solidly built, it was termed the ‘robust’ morph while the other was termed ‘gracile.’ Several morphological differences associated with the two morphs were used to analyze sexual dimorphism in Tyrannosaurus rex, with the ‘robust’ morph usually suggested to be female. For example, the pelvis of several ‘robust’ specimens seemed to be wider, perhaps to allow the passage of eggs. It was also thought that the ‘robust’ morphology correlated with a reduced chevron on the first tail vertebra, also ostensibly to allow eggs to pass out of the reproductive tract, as had been erroneously reported for crocodiles.

In recent years, evidence for sexual dimorphism has been weakened. A 2005 study reported that previous claims of sexual dimorphism in crocodile chevron anatomy were in error, casting doubt on the existence of similar dimorphism between Tyrannosaurus rex genders. A full-sized chevron was discovered on the first tail vertebra of «Sue,» an extremely robust individual, indicating that this feature could not be used to differentiate the two morphs anyway. As Tyrannosaurus rex specimens have been found from Saskatchewan to New Mexico, differences between individuals may be indicative of geographic variation rather than sexual dimorphism. The differences could also be age-related, with ‘robust’ individuals being older animals.

Only a single Tyrannosaurus rex specimen has been conclusively shown to belong to a specific gender. Examination of «B-rex» demonstrated the preservation of soft tissue within several bones. Some of this tissue has been identified as medullary tissue, a specialized tissue grown only in modern birds as a source of calcium for the production of eggshell during ovulation. As only female birds lay eggs, medullary tissue is only found naturally in females, although males are capable of producing it when injected with female reproductive hormones like estrogen. This strongly suggests that «B-rex» was female, and that she died during ovulation. Recent research has shown that medullary tissue is never found in crocodiles, which are thought to be the closest living relatives of dinosaurs, aside from birds. The shared presence of medullary tissue in birds and theropod dinosaurs is further evidence of the close evolutionary relationship between the two.

Posture

Like many bipedal dinosaurs, Tyrannosaurus rex was historically depicted as a ‘living tripod’, with the body at 45 degrees or less from the vertical and the tail dragging along the ground, similar to a kangaroo. This concept dates from Joseph Leidy’s 1865 reconstruction of Hadrosaurus, the first to depict a dinosaur in a bipedal posture. Henry Fairfield Osborn, former president of the American Museum of Natural History (AMNH) in New York City, who believed the creature stood upright, further reinforced the notion after unveiling the first complete Tyrannosaurus rex skeleton in 1915. It stood in this upright pose for nearly a century, until it was dismantled in 1992. By 1970, scientists realized this pose was incorrect and could not have been maintained by a living animal, as it would have resulted in the dislocation or weakening of several joints, including the hips and the articulation between the head and the spinal column. The inaccurate AMNH mount inspired similar depictions in many films and paintings (such as Rudolph Zallinger’s famous mural The Age Of Reptiles in Yale University’s Peabody Museum of Natural History) until the 1990s, when films such as Jurassic Park introduced a more accurate posture to the general public. Modern representations in museums, art, and film show Tyrannosaurus rex with its body approximately parallel to the ground and tail extended behind the body to balance the head.

Arms

When Tyrannosaurus rex was first discovered, the humerus was the only element of the forelimb known. For the initial mounted skeleton as seen by the public in 1915, Osborn substituted longer, three-fingered forelimbs like those of Allosaurus. However, a year earlier, Lawrence Lambe described the short, two-fingered forelimbs of the closely related Gorgosaurus. This strongly suggested that Tyrannosaurus rex had similar forelimbs, but this hypothesis was not confirmed until the first complete Tyrannosaurus rex forelimbs were identified in 1989, belonging to MOR 555 (the «Wankel rex»). The remains of «Sue» also include complete forelimbs. Tyrannosaurus rex arms are very small relative to overall body size, measuring only 1 metre (3.3 ft) long. However, they are not vestigial but instead show large areas for muscle attachment, indicating considerable strength. This was recognized as early as 1906 by Osborn, who speculated that the forelimbs may have been used to grasp a mate during copulation. It has also been suggested that the forelimbs were used to assist the animal in rising from a prone position. Another possibility is that the forelimbs held struggling prey while it was dispatched by the tyrannosaur’s enormous jaws. This hypothesis may be supported by biomechanical analysis.

Tyrannosaurus rex forelimb bones exhibit extremely thick cortical bone, indicating that they were developed to withstand heavy loads. The biceps brachii muscle of a full-grown Tyrannosaurus rex was capable of lifting 199 kilograms (439 lb) by itself; this number would only increase with other muscles (like the brachialis) acting in concert with the biceps. A Tyrannosaurus rex forearm also had a reduced range of motion, with the shoulder and elbow joints allowing only 40 and 45 degrees of motion, respectively. In contrast, the same two joints in Deinonychus allow up to 88 and 130 degrees of motion, respectively, while a human arm can rotate 360 degrees at the shoulder and move through 165 degrees at the elbow. The heavy build of the arm bones, extreme strength of the muscles, and limited range of motion may indicate a system designed to hold fast despite the stresses of a struggling prey animal.

Soft tissue

In the March 2005 issue of Science, Mary Higby Schweitzer of North Carolina State University and colleagues announced the recovery of soft tissue from the marrow cavity of a fossilized leg bone, from a 68-million-year-old Tyrannosaurus. The bone had been intentionally, though reluctantly, broken for shipping and then not preserved in the normal manner, specifically because Schweitzer was hoping to test it for soft tissue. Designated as the Museum of the Rockies specimen 1125, or MOR 1125, the dinosaur was previously excavated from the Hell Creek Formation. Flexible, bifurcating blood vessels and fibrous but elastic bone matrix tissue were recognized. In addition, microstructures resembling blood cells were found inside the matrix and vessels. The structures bear resemblance to ostrich blood cells and vessels. Whether an unknown process, distinct from normal fossilization, preserved the material, or the material is original, the researchers do not know, and they are careful not to make any claims about preservation. If it is found to be original material, any surviving proteins may be used as a means of indirectly guessing some of the DNA content of the dinosaurs involved, because each protein is typically created by a specific gene. The absence of previous finds may merely be the result of people assuming preserved tissue was impossible, therefore simply not looking. Since the first, two more tyrannosaurs and a hadrosaur have also been found to have such tissue-like structures. Research on some of the tissues involved has suggested that birds are closer relatives to tyrannosaurs than other modern animals.

In studies reported in the journal Science in April 2007, Asara and colleagues concluded that seven traces of collagen proteins detected in purified Tyrannosaurus rex bone most closely match those reported in chickens, followed by frogs and newts. The discovery of proteins from a creature tens of millions of years old, along with similar traces the team found in a mastodon bone at least 160,000 years old, upends the conventional view of fossils and may shift paleontologists’ focus from bone hunting to biochemistry. Until these finds, most scientists presumed that fossilization replaced all living tissue with inert minerals. Paleontologist Hans Larsson of McGill University in Montreal, who was not part of the studies, called the finds «a milestone», and suggested that dinosaurs could «enter the field of molecular biology and really slingshot paleontology into the modern world».

Subsequent studies in April 2008 confirmed the close connection of Tyrannosaurus rex to modern birds. Postdoctoral biology researcher Chris Organ at Harvard University announced, «With more data, they would probably be able to place T. rex on the evolutionary tree between alligators and chickens and ostriches.» Co-author John M. Asara added, «We also show that it groups better with birds than modern reptiles, such as alligators and green anole lizards.»

The presumed soft tissue was called into question by Thomas Kaye of the University of Washington and his co-authors in 2008. They contend that what was really inside the tyrannosaur bone was slimy biofilm created by bacteria that coated the voids once occupied by blood vessels and cells. The researchers found that what previously had been identified as remnants of blood cells, because of the presence of iron, were actually framboids, microscopic mineral spheres bearing iron. They found similar spheres in a variety of other fossils from various periods, including an ammonite. In the ammonite they found the spheres in a place where the iron they contain could not have had any relationship to the presence of blood.

Skin and feathers

In 2004, the scientific journal Nature published a report describing an early tyrannosauroid, Dilong paradoxus, from the famous Yixian Formation of China. As with many other theropods discovered in the Yixian, the fossil skeleton was preserved with a coat of filamentous structures which are commonly recognized as the precursors of feathers. It has also been proposed that Tyrannosaurus and other closely related tyrannosaurids had such protofeathers. However, skin impressions from large tyrannosaurid specimens show mosaic scales. While it is possible that protofeathers existed on parts of the body which have not been preserved, a lack of insulatory body covering is consistent with modern multi-ton mammals such as elephants, hippopotamus, and most species of rhinoceros. As an object increases in size, its ability to retain heat increases due to its decreasing surface area-to-volume ratio. Therefore, as large animals evolve in or disperse into warm climates, a coat of fur or feathers loses its selective advantage for thermal insulation and can instead become a disadvantage, as the insulation traps excess heat inside the body, possibly overheating the animal. Protofeathers may also have been secondarily lost during the evolution of large tyrannosaurids like Tyrannosaurus, especially in warm Cretaceous climates.

Thermoregulation

Tyrannosaurus, like most dinosaurs, was long thought to have an ectothermic («cold-blooded») reptilian metabolism. The idea of dinosaur ectothermy was challenged by scientists like Robert T. Bakker and John Ostrom in the early years of the «Dinosaur Renaissance», beginning in the late 1960s. Tyrannosaurus rex itself was claimed to have been endothermic («warm-blooded»), implying a very active lifestyle. Since then, several paleontologists have sought to determine the ability of Tyrannosaurus to regulate its body temperature. Histological evidence of high growth rates in young Tyrannosaurus rex, comparable to those of mammals and birds, may support the hypothesis of a high metabolism. Growth curves indicate that, as in mammals and birds, Tyrannosaurus rex growth was limited mostly to immature animals, rather than the indeterminate growth seen in most other vertebrates.

Oxygen isotope ratios in fossilized bone are sometimes used to determine the temperature at which the bone was deposited, as the ratio between certain isotopes correlates with temperature. In one specimen, the isotope ratios in bones from different parts of the body indicated a temperature difference of no more than 4 to 5°C (7 to 9°F) between the vertebrae of the torso and the tibia of the lower leg. This small temperature range between the body core and the extremities was claimed by paleontologist Reese Barrick and geochemist William Showers to indicate that Tyrannosaurus rex maintained a constant internal body temperature (homeothermy) and that it enjoyed a metabolism somewhere between ectothermic reptiles and endothermic mammals. Other scientists have pointed out that the ratio of oxygen isotopes in the fossils today does not necessarily represent the same ratio in the distant past, and may have been altered during or after fossilization (diagenesis). Barrick and Showers have defended their conclusions in subsequent papers, finding similar results in another theropod dinosaur from a different continent and tens of millions of years earlier in time (Giganotosaurus). Ornithischian dinosaurs also showed evidence of homeothermy, while varanid lizards from the same formation did not. Even if Tyrannosaurus rex does exhibit evidence of homeothermy, it does not necessarily mean that it was endothermic. Such thermoregulation may also be explained by gigantothermy, as in some living sea turtles.

Footprints

Two isolated fossilized footprints have been tentatively assigned to Tyrannosaurus rex. The first was discovered at Philmont Scout Ranch, New Mexico, in 1983 by American geologist Charles Pillmore. Originally thought to belong to a hadrosaurid, examination of the footprint revealed a large ‘heel’ unknown in ornithopod dinosaur tracks, and traces of what may have been a hallux, the dewclaw-like fourth digit of the tyrannosaur foot. The footprint was published as the ichnogenus Tyrannosauripus pillmorei in 1994, by Martin Lockley and Adrian Hunt. Lockley and Hunt suggested that it was very likely the track was made by a Tyrannosaurus rex, which would make it the first known footprint from this species. The track was made in what was once a vegetated wetland mud flat. It measures 83 centimetres (33 in) long by 71 centimetres (28 in) wide.

A second footprint that may have been made by a Tyrannosaurus was first reported in 2007 by British paleontologist Phil Manning, from the Hell Creek Formation of Montana. This second track measures 76 centimetres (30 in) long, shorter than the track described by Lockley and Hunt. Whether or not the track was made by Tyrannosaurus is unclear, though Tyrannosaurus and Nanotyrannus are the only large theropods known to have existed in the Hell Creek Formation. Further study of the track (a full description has not yet been published) will compare the Montana track with the one found in New Mexico.

Locomotion

There are two main issues concerning the locomotory abilities of Tyrannosaurus: how well it could turn; and what its maximum straight-line speed was likely to have been. Both are relevant to the debate about whether it was a hunter or a scavenger (see below).

Tyrannosaurus may have been slow to turn, possibly taking one to two seconds to turn only 45°—an amount that humans, being vertically oriented and tail-less, can spin in a fraction of a second. The cause of the difficulty is rotational inertia, since much of Tyrannosaurus’ mass was some distance from its center of gravity, like a human carrying a heavy timber—although it might have reduced the average distance by arching its back and tail and pulling its head and forelimbs close to its body, rather like the way ice skaters pull their arms closer in order to spin faster.

Scientists have produced a wide range of maximum speed estimates, mostly around 11 metres per second (25 mph), but a few as low as 5–11 metres per second (11–25 mph), and a few as high as 20 metres per second (45 mph). Researchers have to rely on various estimating techniques because, while there are many tracks of very large theropods walking, so far none have been found of very large theropods running—and this absence may indicate that they did not run. Scientists who think that Tyrannosaurus was able to run point out that hollow bones and other features that would have lightened its body may have kept adult weight to a mere 5 tons or so, or that other animals like ostriches and horses with long, flexible legs are able to achieve high speeds through slower but longer strides. Additionally, some have argued that Tyrannosaurus had relatively larger leg muscles than any animal alive today, which could have enabled fast running 40–70 kilometres per hour (25–43 mph).

Jack Horner and Don Lessem argued in 1993 that Tyrannosaurus was slow and probably could not run (no airborne phase in mid-stride), because its ratio of femur (thigh bone) to tibia (shin bone) length was greater than 1, as in most large theropods and like a modern elephant. However, Holtz (1998) noted that tyrannosaurids and some closely related groups had significantly longer distal hindlimb components (shin plus foot plus toes) relative to the femur length than most other theropods), and that tyrannosaurids and their close relatives had a tightly interlocked metatarsus that more effectively transmitted locomotory forces from the foot to the lower leg than in earlier theropods («metatarsus» means the foot bones, which function as part of the leg in digitigrade animals). He therefore concluded that tyrannosaurids and their close relatives were the fastest large theropods.Christiansen (1998) estimated that the leg bones of Tyrannosaurus were not significantly stronger than those of elephants, which are relatively limited in their top speed and never actually run (there is no airborne phase), and hence proposed that the dinosaur’s maximum speed would have been about 11 metres per second (25 mph), which is about the speed of a human sprinter. But he also noted that such estimates depend on many dubious assumptions.

Farlow and colleagues (1995) have argued that a Tyrannosaurus weighing 6 short tons (5.4 t) to 8 short tons (7.3 t) would have been critically or even fatally injured if it had fallen while moving quickly, since its torso would have slammed into the ground at a deceleration of 6 g (six times the acceleration due to gravity, or about 60 meters/s²) and its tiny arms could not have reduced the impact. However, giraffes have been known to gallop at 50 kilometres per hour (31 mph), despite the risk that they might break a leg or worse, which can be fatal even in a «safe» environment such as a zoo. Thus it is quite possible that Tyrannosaurus also moved fast when necessary and had to accept such risks.

Most recent research on Tyrannosaurus locomotion does not support speeds faster than 40 kilometres per hour (25 mph), i.e. moderate-speed running. For example, a 2002 paper in the journal Nature used a mathematical model (validated by applying it to three living animals, alligators, chickens, and humans; additionally later eight more species including emus and ostriches) to gauge the leg muscle mass needed for fast running (over 40 kilometres per hour (25 mph)). They found that proposed top speeds in excess of 40 kilometres per hour (25 mph) were unfeasible, because they would require very large leg muscles (more than approximately 40–86% of total body mass). Even moderately fast speeds would have required large leg muscles. This discussion is difficult to resolve, as it is unknown how large the leg muscles actually were in Tyrannosaurus. If they were smaller, only 18 kilometres per hour (11 mph) walking/jogging might have been possible.

A study in 2007 used computer models to estimate running speeds, based on data taken directly from fossils, and claimed that Tyrannosaurus rex had a top running speed of 8 metres per second (18 mph). An average professional football (soccer) player would be slightly slower, while a human sprinter can reach 12 metres per second (27 mph). Note that these computer models predict a top speed of 17.8 metres per second (40 mph) for a 3 kilograms (6.6 lb) Compsognathus (probably a juvenile individual).

Those who argue that Tyrannosaurus was incapable of running estimate the top speed of Tyrannosaurus at about 17 kilometres per hour (11 mph). This is still faster than its most likely prey species, hadrosaurids and ceratopsians. In addition, some advocates of the idea that Tyrannosaurus was a predator claim that tyrannosaur running speed is not important, since it may have been slow but still faster than its probable prey. However, Paul and Christiansen (2000) argued that at least the later ceratopsians had upright forelimbs and the larger species may have been as fast as rhinos. Healed Tyrannosaurus bite wounds on ceratopsian fossils are interpreted as evidence of attacks on living ceratopsians (see below). If the ceratopsians that lived alongside Tyrannosaurus were fast, that casts doubt on the argument that Tyrannosaurus did not have to be fast to catch its prey.

Feeding strategies

The debate about whether Tyrannosaurus was a predator or a pure scavenger is as old as the debate about its locomotion. Lambe (1917) described a good skeleton of Tyrannosaurus’ close relative Gorgosaurus and concluded that it and therefore also Tyrannosaurus was a pure scavenger, because the Gorgosaurus’ teeth showed hardly any wear. This argument is no longer taken seriously, because theropods replaced their teeth quite rapidly. Ever since the first discovery of Tyrannosaurus most scientists have agreed that it was a predator, although like modern large predators it would have been happy to scavenge or steal another predator’s kill if it had the opportunity.

Noted hadrosaur expert Jack Horner is currently the major advocate of the idea that Tyrannosaurus was exclusively a scavenger and did not engage in active hunting at all. Horner has presented several arguments to support the pure scavenger hypothesis:

• Tyrannosaur arms are short when compared to other known predators. Horner argues that the arms were too short to make the necessary gripping force to hold on to prey.
• Tyrannosaurs had large olfactory bulbs and olfactory nerves (relative to their brain size). These suggest a highly developed sense of smell which could sniff out carcasses over great distances, as modern vultures do. Research on the olfactory bulbs of dinosaurs has shown that Tyrannosaurus had the most highly developed sense of smell of 21 sampled dinosaurs. Opponents of the pure scavenger hypothesis have used the example of vultures in the opposite way, arguing that the scavenger hypothesis is implausible because the only modern pure scavengers are large gliding birds, which use their keen senses and energy-efficient gliding to cover vast areas economically. However, researchers from Glasgow concluded that an ecosystem as productive as the current Serengeti would provide sufficient carrion for a large theropod scavenger, although the theropod might have had to be cold-blooded in order to get more calories from carrion than it spent on foraging (see Warm-bloodedness of dinosaurs). They also suggested that modern ecosystems like Serengeti have no large terrestrial scavengers because gliding birds now do the job much more efficiently, while large theropods did not face competition for the scavenger ecological niche from gliding birds.
• Tyrannosaur teeth could crush bone, and therefore could extract as much food (bone marrow) as possible from carcass remnants, usually the least nutritious parts. Karen Chin and colleagues have found bone fragments in coprolites (fossilized dung) that they attribute to tyrannosaurs, but point out that a tyrannosaur’s teeth were not well adapted to systematically chewing bone like hyenas do to extract marrow.
• Since at least some of Tyrannosaurus’s potential prey could move quickly, evidence that it walked instead of ran could indicate that it was a scavenger. On the other hand, recent analyses suggest that Tyrannosaurus, while slower than large modern terrestrial predators, may well have been fast enough to prey on large hadrosaurs and ceratopsians.Other evidence suggests hunting behavior in Tyrannosaurus. The eye-sockets of tyrannosaurs are positioned so that the eyes would point forward, giving them binocular vision slightly better than that of modern hawks. He also pointed out that the tyrannosaur lineage had a history of steadily improving binocular vision. It is hard to see how natural selection would have favored this long-term trend if tyrannosaurs had been pure scavengers, which would not have needed the advanced depth perception that stereoscopic vision provides. In modern animals, binocular vision is found mainly in predators.

A skeleton of the hadrosaurid Edmontosaurus annectens has been described from Montana with healed tyrannosaur-inflicted damage on its tail vertebrae. The fact that the damage seems to have healed suggests that the Edmontosaurus survived a tyrannosaur’s attack on a living target, i.e. the tyrannosaur had attempted active predation. There is also evidence for an aggressive interaction between a Triceratops and a Tyrannosaurus in the form of partially healed tyrannosaur tooth marks on a Triceratops brow horn and squamosal (a bone of the neck frill); the bitten horn is also broken, with new bone growth after the break. It is not known what the exact nature of the interaction was, though: either animal could have been the aggressor. When examining Sue, paleontologist Pete Larson found a broken and healed fibula and tail vertebrae, scarred facial bones and a tooth from another Tyrannosaurus embedded in a neck vertebra. If correct, these might be strong evidence for aggressive behavior between tyrannosaurs but whether it would have been competition for food and mates or active cannibalism is unclear. However, further recent investigation of these purported wounds has shown that most are infections rather than injuries (or simply damage to the fossil after death) and the few injuries are too general to be indicative of intraspecific conflict.

Some researchers argue that if Tyrannosaurus were a scavenger, another dinosaur had to be the top predator in the Amerasian Upper Cretaceous. Top prey was the larger marginocephalians and ornithopods. The other tyrannosaurids share so many characteristics that only small dromaeosaurs remain as feasible top predators. In this light, scavenger hypothesis adherents have suggested that the size and power of tyrannosaurs allowed them to steal kills from smaller predators. Most paleontologists accept that Tyrannosaurus was both an active predator and a scavenger like all large carnivores.

History

Henry Fairfield Osborn, president of the American Museum of Natural History, named Tyrannosaurus rex in 1905. The generic name is derived from the Greek words τυραννος (tyrannos, meaning «tyrant») and σαυρος (sauros, meaning «lizard»). Osborn used the Latin word rex, meaning «king», for the specific name. The full binomial therefore translates to «tyrant lizard king,» emphasizing the animal’s size and perceived dominance over other species of the time.

Earliest finds

Teeth from what is now documented as a Tyrannosaurus rex were found in 1874 by A. Lakes near Golden, Colorado. In the early 1890s, J. B. Hatcher collected postcranial elements in eastern Wyoming. The fossils were believed to be from a large species of Ornithomimus (O. grandis) but are now considered Tyrannosaurus rex. Vertebral fragments found by E. D. Cope in western South Dakota in 1892 and named as Manospondylus gigas have also been reclassified as Tyrannosaurus rex. Barnum Brown, assistant curator of the American Museum of Natural History, found the first partial skeleton of Tyrannosaurus rex in eastern Wyoming in 1900. H. F. Osborn originally named this skeleton Dynamosaurus imperiosus in a paper in 1905. Brown found another partial skeleton in the Hell Creek Formation in Montana in 1902. Osborn used this holotype to describe Tyrannosaurus rex in the same paper in which D. imperiosus was described. Had it not been for page order, Dynamosaurus would have become the official name. The original Dynamosaurus material resides in the collections of the Natural History Museum, London.

In total, Brown found five Tyrannosaurus partial skeletons. In 1941, Brown’s 1902 find was sold to the Carnegie Museum of Natural History in Pittsburgh, Pennsylvania. Brown’s fourth and largest find, also from Hell Creek, is on display in the American Museum of Natural History in New York.

Although there are numerous skeletons in the world, only one track has been documented — at Philmont Scout Ranch in northeast New Mexico. It was discovered in 1983 and identified and documented in 1994.

Notable specimens

Sue Hendrickson, amateur paleontologist, discovered the most complete (approximately 85%) and, until 2001, the largest, Tyrannosaurus fossil skeleton known in the Hell Creek Formation near Faith, South Dakota, on 12 August 1990. This Tyrannosaurus, nicknamed «Sue» in her honor, was the object of a legal battle over its ownership. In 1997 this was settled in favor of Maurice Williams, the original land owner. The fossil collection was purchased by the Field Museum of Natural History at auction for USD 7.6 million, making it the most expensive dinosaur skeleton to date. From 1998 to 1999 Field Museum of Natural History preparators spent 30,000 hours taking the rock off each of the bones. The bones were then shipped off to New Jersey where the mount was made. The finished mount was then taken apart, and along with the bones, shipped back to Chicago for the final assembly. The mounted skeleton opened to the public on May 17, 2000 in the great hall (Stanley Field Hall) at the Field Museum of Natural History. A study of this specimen’s fossilized bones showed that «Sue» reached full size at age 19 and died at age 28, the longest any tyrannosaur is known to have lived. Early speculation that Sue may have died from a bite to the back of the head was not confirmed. Though subsequent study showed many pathologies in the skeleton, no bite marks were found. Damage to the back of the skull may have been caused by post-mortem trampling.

Another Tyrannosaurus, nicknamed «Stan», in honor of amateur paleontologist Stan Sacrison, was found in the Hell Creek Formation near Buffalo, South Dakota, in the spring of 1987. After 30,000 hours of digging and preparing, a 65% complete skeleton emerged. Stan is currently on display in the Black Hills Museum of Natural History Exhibit in Hill City, South Dakota, after an extensive world tour. This tyrannosaur, too, was found to have many bone pathologies, including broken and healed ribs, a broken (and healed) neck and a spectacular hole in the back of its head, about the size of a Tyrannosaurus tooth. Both Stan and Sue were examined by Peter Larson.

In the summer of 2000, Jack Horner discovered five Tyrannosaurus skeletons near the Fort Peck Reservoir in Montana. One of the specimens, dubbed «C. rex,» was reported to be perhaps the largest Tyrannosaurus ever found.In 2001, a 50% complete skeleton of a juvenile Tyrannosaurus was discovered in the Hell Creek Formation in Montana, by a crew from the Burpee Museum of Natural History of Rockford, Illinois. Dubbed «Jane the Rockford T-Rex,» the find was initially considered the first known skeleton of the pygmy tyrannosaurid Nanotyrannus but subsequent research has revealed that it is more likely a juvenile Tyrannosaurus. It is the most complete and best preserved juvenile example known to date. Jane has been examined by Jack Horner, Pete Larson, Robert Bakker, action=edit redlink=1 Greg Erickson and several other renowned paleontologists, because of the uniqueness of her age. Jane is currently on exhibit at the Burpee Museum of Natural History in Rockford, Illinois.

In a press release on 7 April 2006, Montana State University revealed that it possessed the largest Tyrannosaurus skull yet discovered. Discovered in the 1960s and only recently reconstructed, the skull measures 59 inches (150 cm) long compared to the 55.4 inches (141 cm) of “Sue’s” skull, a difference of 6.5%.

Appearances in popular culture

Since it was first described in 1905, Tyrannosaurus rex has become the most widely recognized dinosaur species in popular culture. It is the only dinosaur that is commonly known to the general public by its full scientific name (binomial name) (Tyrannosaurus rex), and the scientific abbreviation T. rex has also come into wide usage (commonly abbreviated «T-Rex»). Robert T. Bakker notes this in The Dinosaur Heresies and explains that a name like «Tyrannosaurus rex is just irresistible to the tongue.»

Gallery

(Tyrannosaurus rex) – настоящий король динозавров. Тирекс неспроста стал иконой палеонтологии. Огромный семитонный хищник вобрал в себя огромное множество прогрессивных черт. Давайте рассмотрим их более детально.

Размеры

Размеры тираннозавра впечатляют. 12 метров в длину, 4 метра в высоту бедра, и около 7 тонн веса. Огромнейший череп, усаженный десятками длинных зубов, мог насчитывать 1,5-2 метра в длину и полностью уместить в себя человека. Зубы тираннозавра являются абсолютными рекордсменами среди зубов всех сухопутных хищников. Длина с корнем достигала 30 см. Даже гипертрофированные верхние клыки саблезубых смилодонов были хоть и немного, но короче.

Сила укуса тираннозавра в разы больше всех современных хищников. Конечно же, её реконструкция не идеальна и ещё будут перерасчёты, но на сегодня мы имеем впечатляющие цифры: 250-300 кН. Это делает челюсти тираннозавра одними из самых мощных в истории сухопутных животных.

Навыки охотника

Вопрос, был ли тирекс охотником или падальщиком, до сих пор не решён. Но набор прогрессивных черт тираннозавра всё больше склоняет палеонтологов к охотничьему образу жизни. Рекс – обладатель хорошо развитого бинокулярного зрения. Бинокулярное зрение – способность фокусироваться на объекте обоими глазами, что даёт более точную оценку расстояния. Такое зрение более свойственно активным охотникам, нежели падальщикам. Отдел мозга тираннозавра, отвечающий за обоняние, также хорошо развит. Можно сослаться, что это свойственно падальщикам, но хорошее обоняние есть у всех плотоядных: как любителей падали, так и превосходных охотников. Также, тираннозавр — обладатель укреплённой челюсти и глубоко посаженных зубов. Что примечательно, челюсть имеет дополнительную подвижность, относительно других тиранозаврид, что обеспечивает повышенную устойчивость при боковых нагрузках. Такая адаптация тоже больше подходит охотнику на крупных животных, нежели падальщику, чья еда не сопротивляется и не может повредить челюсть. Челюсти и зубы тираннозавра будто специально созданы для удержания сопротивляющейся многотонной жертвы. Тираннозавр со своим весом, конечно же, не мог бегать за жертвой, что подтверждают свежие исследования, но выслеживать был вполне в состоянии. Правда тираннозавру и незачем было бегать: его основной рацион – медленные трицератопсы и гадрозавры с углом разворота как у поезда. Об этом свидетельствуют прижизненные следы укусов гигантских челюстей на воротниках и хвостах жертв.

Поведение

Тираннозавры были крайне агрессивными динозаврами. Эта агрессия проявлялась и внутри вида, что ставит под вопрос их социальный образ жизни. Мы можем наблюдать за борьбой гигантских тероподов, живших 66 млн лет назад, по почти полным скелетам (60-70% найденных костей относительно полного скелета). Такие взрослые особи тираннозавров как Стен и Сью имели большое количество прижизненных травм, похожих на следы укусов. Возможно, такие укусы могли быть следами борьбы в период брачных игр. Самки тираннозавров вступали в половую зрелость к 14-16 годам, а средняя продолжительность жизни тирекса составляла 30 лет.

Лапки

Мало кто из изучающих динозавров не любит обсуждать назначение лапок тираннозавра. На самом деле, многим тероподам свойственны рудиментарные передние конечности. Этим славятся практически все тираннозавриды, гиганотозавры, карнотавры. Короткие лапки тираннозавру могли быть полезными разве что в совсем юном возрасте. Если в твоём арсенале чуть ли не самые мощные челюсти в истории Земли, нужда в когтях и мощных лапах отпадает сама собой. Из тысячелетия в тысячелетие передние конечности тираннозавра уменьшались, пока не достигли столь крохотных размеров. Проживи король чуть больше, возможно и вовсе лишился бы их.

Тираннозавр как икона палеонтологии

Ни для кого не секрет, что первое животное, приходящее на ум при упоминании палеонтологии в целом и динозавров в частности, это тираннозавр. Ему посвящают книги и фильмы. Профессионалов он манит своими адаптациями, которых нет у сородичей, любителей – ужасающими размерами. И казалось бы, что в XXI веке, когда уже найдены десятки тираннозавров, в том числе и хорошей сохранности, острые вопросы должны разрешиться. Но это только поверхностный взгляд на животное. Вопросы охоты, поведения, кожных покровов всё ещё актуальны. Тираннозавр стал главным антагонистом в книге Майкла Крайтона «Парк юрского периода», а после и в фильмах. Тирекс настолько впечатлил зрителей, что на Западе сегодня специалистов по динозаврам нет счёту. Огромный двенадцатиметровый хищник – настоящая икона палеонтологии.

Итог

Как сегодня королями хищников считаются львы и другие представители рода Panthera за их размеры и прогрессивность, так и в верхнем мелу королём был тираннозавр. История мезозоя знала много гигантских хищников: спинозавры, гиганотозавры, кархородонтозавры, но тираннозавр, кроме размеров, мог похвастаться массивным телосложением, увеличенной прочностью челюстей и зубов, хорошо развитыми органами чувств и огромным множеством других деталей, делающих его самым узнаваемым образом в палеонтологии.

Тиранноза́вр (лат. Tyrannosaurus — «ящер-тиран»)  — монотипический род плотоядных динозавров из группы целурозавров подотряда тероподов, включающий единственный валидный вид — Tyrannosaurus rex. Обитал в основном в западной части Северной Америки, и был наиболее распространённым из тираннозавридов.

Окаменелые останки тираннозавров находят в различных геологических формациях, датирующихся маастрихтским веком мелового периода, около 67—65,5 миллионов лет назад. Был одним из последних ящеротазовых динозавров, существовавших перед катаклизмом, положившим конец эре динозавров.

Описание[]

Размеры[]

Тираннозавр был одним из самых крупных сухопутных существ того времени. Длина голотипа оценивается в 10,6 м, а высота до бёдер — 3,4 м. Самый большой из относительно полных скелетов, FMNH PR2081 по прозвищу «Сью», в длину достигает 12,3 метра и имеет высоту на уровне бедра в 4 метра. Оценка массы тела этого образца в течение многих лет сильно изменялись: обычно от 5,4 тонн до 6,8 тонны, реже — более чем 7,2 тонны или менее чем 4,5 тонны, самая современная же оценка равняется примерно 9,5 тонны.

В 2009 году палеонтолог Пакард и его коллеги протестировали метод измерения массы динозавров на слонах и пришли к выводу, что прежние техники измерения были весьма несовершенны и масса динозавров зачастую переоценивалась, соответственно, масса тираннозавра могла оказаться гораздо меньше, чем было принято считать. Однако, более современные измерения (выполненные при помощи метода GDI, в котором объём животного вычисляется с помощью построения множества сечений и умножения их средней площади на длину тела) позволили заключить, что масса тела самого большого из найденных типовых экземпляров приближается либо даже превышает 9,5 тонны.

Существуют фрагментарные окаменелости, принадлежавшие, возможно, ещё более крупным тираннозаврам. Так, Грегори С. Пол оценивает длину образца UCMP 118742 (81 см верхнечелюстная кость) приблизительно в 13,6 м, высоту на уровне бёдер — в 4,4 метра, а массу — в 12 тонн.

Телосложение[]

Шея тираннозавра, как и у других теропод, имела S-образную форму, но была короткой и мускулистой, что позволяло ей удерживать массивную голову. Передние конечности имели только два когтистых пальца вместе с маленькой пястной костью, рудиментом третьего пальца. Задние конечности, напротив, были самыми длинными (по отношению к телу) среди всех теропод. Позвоночник образован 10 шейными, 12 грудными, пятью крестцовыми и около 40 хвостовыми позвонками. Хвост был тяжёлым и длинным, поскольку выполнял функцию балансира, уравновешивая массивную голову и грузное туловище. Чтобы компенсировать невероятную громоздкость животного, многие кости скелета были полыми, что снижало их вес, но не сильно влияло на прочность.

Галерея[]

Тираннозавр с риском напал на Анкилозавра

Тираннозавр убивает Трицератопса