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恐爪龍屬:修订间差异

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一副恐爪龍的骨骼,包括原先標本的骨頭,都在美國自然歷史博物館中展覽,而其他的則在[[哈佛大學]]的[[比較動物學博物館]]中展覽。這兩個標本是在與耶魯大學標本不同的位點發現的。
一副恐爪龍的骨骼,包括原先標本的骨頭,都在美國自然歷史博物館中展覽,而其他的則在[[哈佛大學]]的[[比較動物學博物館]]中展覽。這兩個標本是在與耶魯大學標本不同的位點發現的。


===化石蛋===
===更多發現===
In 1974 Ostrom published another monograph on the shoulder of ''Deinonychus'' in which he realized that the pubis that he had described was actually a coracoid – a shoulder element.<ref name="Ostrom74">Ostrom, John H.,. (1974). " The Pectoral Girdle and Forelimb Function of Deinonychus (Reptilia: Saurischia) : A Correction". '' Postilla, Peabody Museum of Natural History Bulletin '', 165:1-11.</ref> In that same year, another specimen of ''Deinonychus'' was excavated in Montana by a Harvard University expedition headed by Farish Jenkins. This discovery added several new elements; well preserved femurs, pubes, a sacrum, and better ilia, as well as elements of the pes and metatarsus. Ostrom described this specimen and revised his skeletal restoration of ''Deinonychus''. This time it showed the very long pubes, and Ostrom began to suspect that they may have even been a little retroverted like those of birds.<ref name=”ostrom 1976”>{{cite journal|last=Ostrom|first=J.H. |year=1976 |title= On a new specimen of the Lower Cretaceous theropod dinosaur Deinonychus antirrhopus |journal=Breviora|volume=439 |pages=1-21}}</ref>
在[[約翰·奧斯特倫姆|奧斯特倫姆]]描述了[[巴納姆·布郎|布郎]]發現的原先標本後,仍有幾個細小陷在石灰的標本未被處理而擺放在[[美國自然歷史博物館]]。這些大部份都是獨立的骨頭及骨骼碎片,包括了標本埋葬時周圍的岩石。[[2000年]]時就這些標本的研究發現一些有趣及被忽略了的資料。當中的幾條幼長骨頭,原先被認為是用作令恐爪龍尾巴堅硬的骨頭,其實是[[胃骨片]]。而且亦在原先恐爪龍標本的周圍,發現了一些被忽略的蛋殼[[化石]]。<ref name="makovicky&grellet-tinner2000">{{cite journal|last=Makovicky|first=P. J.|coauthors=Grellet-Tinner, G.|year=2000|title=Association between a specimen of ''Deinonychus antirrhopus'' and theropod eggshell|journal= In Bravo, A.M. and T. Reyes (eds.) ''1st international symposium on dinosaur eggs and babies'',Isona i Conca Dellà Catalonia, Spain, 23–26 September 1999|pages=123–128}}</ref>在後期詳細的報告中,這些蛋差不多肯定是屬於恐爪龍的,故是首批發現[[馳龍科]]的蛋。<ref name="grellet-tinner&makovicky2006"/>再者,其中一個蛋殼的外表面緊接恐爪龍的胃骨片,可見恐爪龍可能正在孵蛋。由此推斷恐爪龍是用身體的[[熱量]]來孵蛋,且與現今[[鳥類]]相似是[[溫血動物]]。<ref>{{cite journal|last=Grellet-Tinner|first=Gerard|year=2006|title=Oology And The Evolution Of Thermophysiology In Saurischian Dinosaurs: Homeotherm And Endotherm Deinonychosaurians?|journal=Papeis Avulsos de Zoologia|volume=46|issue=1|pages=1-10}}</ref>

A skeleton of ''Deinonychus'' including bones from the original (and most complete) specimen can be seen on display at the American Museum of Natural History,<ref>{{cite web | author = American Museum of Natural History | title = Deinonychus | work = http://www.amnh.org | publisher = American Museum of Natural History | date = 2007 | url = http://www.amnh.org/exhibitions/expeditions/treasure_fossil/Fossils/Specimens/deinonychus.html
| accessdate = 2007-07-13}}</ref> with another specimen on display at the [[Museum of Comparative Zoology]] at [[Harvard University]]. The American Museum and Harvard specimens are from a different locality than the Yale specimens. Even these two skeletal mounts are lacking elements including the sterna, sternal ribs, furcula, and gastralia.

Even after all of Ostrom’s work, several small blocks of lime-encased material remained unprepared in storage at the American Museum. These consisted mostly of isolated bones and bone fragments, including the original matrix, or surrounding rock in which the specimens were initially buried. An examination of these unprepared blocks by Gerald Grellet-Tinner and Peter Makovicky in 2000 revealed an interesting, overlooked feature. Several long, thin bones identified on the blocks as ossified tendons (structures which helped stiffen the tail of ''Deinonychus'') turned out to actually represent [[Gastralium|gastralia]] (abdominal ribs). More significantly, a large number of previously unnoticed fossilized eggshells were discovered in the rock matrix which had surrounded the original ''Deinonychus'' specimen.<ref name="makovicky&grellet-tinner2000">{{cite book|last=Makovicky|first=P. J. |coauthors= Grellet-Tinner, G. |year=2000 |chapter=Association between a specimen of ''Deinonychus antirrhopus'' and theropod eggshell |editor= Bravo, A.M. and T. Reyes|title= First international symposium on dinosaur eggs and babies,Isona i Conca Dellà Catalonia, Spain, 23–26 September 1999| pages=123–128}}</ref>

In a subsequent, more detailed report on the eggshells, Grellet-Tinner and Makovicky concluded that the [[Egg (biology)|egg]] almost certainly belonged to ''Deinonychus'', representing the first dromaeosaurid egg to be identified.<ref name="grellet-tinner&makovicky2006"/> Moreover, the external surface of one eggshell was found in close contact with the gastralia suggesting that ''Deinonychus'' might have [[Avian incubation|brooded]] its eggs. This implies that ''Deinonychus'' used body heat transfer as a mechanism for egg incubation, and indicates an [[endothermy]] similar to modern birds.<ref>{{cite journal|last=Grellet-Tinner|first=Gerard|year=2006|title=Oology And The Evolution Of Thermophysiology In Saurischian Dinosaurs: Homeotherm And Endotherm Deinonychosaurians? |journal=Papeis Avulsos de Zoologia |volume=46 |issue=1 |pages=1-10| url=http://www.scielo.br/scielo.php?pid=S0031-10492006000100001&script=sci_arttext| accessdate=2007-07-07}}</ref> Further study by Gregory Erickson and colleagues finds that this individual was 13 or 14 years old at death and its growth had plateaued. Unlike other theropods in their study of specimens found associated with eggs or nests, it had finished growing at the time of its death.<ref name=GMEetal07>{{cite journal |last=Erickson |first=Gregory M. |authorlink=Gregory M. Erickson |coauthors=Curry Rogers, Kristina; Varricchio, David J.; Norell, Mark A.; and Xu, Xing |year=2007 |title=Growth patterns in brooding dinosaurs reveals the timing of sexual maturity in non-avian dinosaurs and genesis of the avian condition |journal=Biology Letters |volume=published online |url=http://www.journals.royalsoc.ac.uk/content/6508252h00612424/fulltext.pdf |format=pdf |doi=10.1098/rsbl.2007.0254 |accessdate=2007-07-26}}</ref>


===影響===
===影響===
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恐爪龍蛋的微結構評估發現它們是屬於獸腳亞目中,因為它們有著其他獸腳亞目蛋的特徵,並與[[鳥臀目]]及[[蜥腳下目]]蛋的不同。與其他[[手盜龍類]]比較,恐爪龍蛋較像竊蛋龍科的蛋,而非傷齒龍科的蛋。雖然該蛋經已碎裂而很難評估其大小,但憑其經過的小骨盆腔可知[[直徑]]約為7厘米。大小與最大的[[葬火龍]]蛋的7.2厘米接近。再者,葬火龍蛋殼及恐爪龍蛋殼厚度差不多完全一致,由於蛋殼厚度影響著蛋的[[體積]],故此有指牠們蛋的大小是完全一樣的。<ref name="grellet-tinner&makovicky2006"/>
恐爪龍蛋的微結構評估發現它們是屬於獸腳亞目中,因為它們有著其他獸腳亞目蛋的特徵,並與[[鳥臀目]]及[[蜥腳下目]]蛋的不同。與其他[[手盜龍類]]比較,恐爪龍蛋較像竊蛋龍科的蛋,而非傷齒龍科的蛋。雖然該蛋經已碎裂而很難評估其大小,但憑其經過的小骨盆腔可知[[直徑]]約為7厘米。大小與最大的[[葬火龍]]蛋的7.2厘米接近。再者,葬火龍蛋殼及恐爪龍蛋殼厚度差不多完全一致,由於蛋殼厚度影響著蛋的[[體積]],故此有指牠們蛋的大小是完全一樣的。<ref name="grellet-tinner&makovicky2006"/>

===化石蛋===
在[[約翰·奧斯特倫姆|奧斯特倫姆]]描述了[[巴納姆·布郎|布郎]]發現的原先標本後,仍有幾個細小陷在石灰的標本未被處理而擺放在[[美國自然歷史博物館]]。這些大部份都是獨立的骨頭及骨骼碎片,包括了標本埋葬時周圍的岩石。[[2000年]]時就這些標本的研究發現一些有趣及被忽略了的資料。當中的幾條幼長骨頭,原先被認為是用作令恐爪龍尾巴堅硬的骨頭,其實是[[胃骨片]]。而且亦在原先恐爪龍標本的周圍,發現了一些被忽略的蛋殼[[化石]]。<ref name="makovicky&grellet-tinner2000">{{cite journal|last=Makovicky|first=P. J.|coauthors=Grellet-Tinner, G.|year=2000|title=Association between a specimen of ''Deinonychus antirrhopus'' and theropod eggshell|journal= In Bravo, A.M. and T. Reyes (eds.) ''1st international symposium on dinosaur eggs and babies'',Isona i Conca Dellà Catalonia, Spain, 23–26 September 1999|pages=123–128}}</ref>在後期詳細的報告中,這些蛋差不多肯定是屬於恐爪龍的,故是首批發現[[馳龍科]]的蛋。<ref name="grellet-tinner&makovicky2006"/>再者,其中一個蛋殼的外表面緊接恐爪龍的胃骨片,可見恐爪龍可能正在孵蛋。由此推斷恐爪龍是用身體的[[熱量]]來孵蛋,且與現今[[鳥類]]相似是[[溫血動物]]。<ref>{{cite journal|last=Grellet-Tinner|first=Gerard|year=2006|title=Oology And The Evolution Of Thermophysiology In Saurischian Dinosaurs: Homeotherm And Endotherm Deinonychosaurians?|journal=Papeis Avulsos de Zoologia|volume=46|issue=1|pages=1-10}}</ref>


==參考==
==參考==

2008年1月23日 (三) 06:31的版本

恐爪龍
化石時期: 下白堊紀
平衡恐爪龍的骨骼
平衡恐爪龍的骨骼
保护状况
化石
科學分類
界: 動物界 Animalia
門: 脊索動物門 Chordata
綱: 蜥形綱 Sauropsida
超目: 恐龍總目 Dinosauria
目: 蜥臀目 Saurischia
亞目: 獸腳亞目 Theropoda
科: 馳龍科 Dromaeosauridae
亞科: 伶盜龍亞科 Velociraptorinae
屬: 恐爪龍屬 Deinonychus
種: 平衡恐爪龍 D. antirrhopus
二名法
Deinonychus antirrhopus
Ostrom, 1969

恐爪龍學名Deinonychus),是一生活於下白堊紀阿普第階阿爾布階(即距今121-98.9百萬年前)的恐龍。牠的學名是來自古希臘文的「δεινος」(意即「恐怖」)及「ονυξ/ονυχος」(意即「爪」)而來,因牠的後肢二趾上有非常大,呈鐮刀狀的爪,在行走時這趾可能會縮起,來讓三趾及四趾行走。一般認為恐爪龍會用其鐮刀爪來割傷獵物,但近年就迅猛龍重組的測試顯示這爪作刺戳之用。[1]就像其他的馳龍科,恐爪龍的堅硬尾巴是由一連串的長骨頭組成。這可提供恐爪龍更好的平衡及轉彎能力,故其模式種稱為平衡恐爪龍(D. antirrhopus)。

古生物學家約翰·奧斯特倫姆(John Ostrom)於1960年代末就恐爪龍的研究有突破的性的發現,引發究竟牠是否恆溫動物的爭論。在這之前,一般認為恐爪龍是巨大爬行動物。奧斯特倫姆指從牠那輕的骨頭及硬的鍵,可見牠是活躍及輕巧的獵食者。

恐爪龍的遺骸是從美國蒙大拿州懷俄明州Cloverly組[2][3]奧克拉荷馬州Antlers組[4]中被發現。而在馬利蘭州大西洋區海岸平原波多馬克組發現的牙齒亦可能屬於此屬。[5]在Cloverly組及Antlers組發現的恐爪龍遺骸與鳥腳下目腱龍有密切的關聯。從與腱龍一同被發現的牙齒標本,可見牠正被恐爪龍所獵殺。[6][4]

描述

人類與恐爪龍的比例。

就恐爪龍最大的標本而言,牠的體型可達3.4公尺,頭顱骨最大可達41公分長,臀部高度為0.87公尺,而體重最高可達73公斤。[7]牠的頭顱骨有強壯的顎骨,及約60根彎曲劍形的牙齒。奧斯特倫姆最初將恐爪龍的顱骨重建成三角形、寬廣、平坦的頭部,類似異特龍。在發現更多、更良好的恐爪龍與其近親的化石後,[8]顯示恐爪龍的上顎較呈拱形,口鼻部較狹窄,顴骨寬廣,使頭部看起來較為立體,與奧斯特倫姆的重建不同。恐爪龍顱骨與伶盜龍不同,頭頂較堅固,類似馳龍;伶盜龍的鼻部上側往下凹,這點與恐爪龍不同。[9]頭顱骨及下顎骨都有孔洞來減輕頭部重量,而恐爪龍的眶前孔是特別的大。[8]按頭顱骨來推算,眼睛主要是向兩側的。

就像其他的馳龍科,恐爪龍的大手有三指。拇指最短,而第二指最長。每隻後肢的第二趾都有鐮刀般的趾爪,長度約13公分,有可能是作為捕獵用的。牠可以先向前刺戳,並向下割來撕破獵物。相對於恐爪龍的體型,這些趾爪來的相當地大。恐爪龍的身體是靠尾巴脊骨人字骨,在高速轉向時來維持穩定及平衡。

分類

恐爪龍是最著名的馳龍科恐龍之一,[10]且是迅猛龍的近親,迅猛龍的體型較小,生存於上白堊紀蒙古[11][12]牠們的都屬於伶盜龍亞科。這個亞科最初是由巴思缽(Rinchen Barsbold)於1983年所提出,[13]而當時只有單一的迅猛龍屬。後來菲力·柯爾(Phil Currie)將大部份的馳龍科都放在這亞科中。[14]生存於上白堊紀蒙古白魔龍[11]以及同時代在北美洲蜥鳥盜龍都可能是恐爪龍的近親,[15]但後者因缺乏資料而難以分類。[11]迅猛龍及其親屬都是多以牠們的爪來作為獵殺工具,而不像馳龍是以厚實的頭顱骨來作為武器。[15]馳龍科傷齒龍科一起組成了恐爪龍下目,是鳥類的姊妹分類。在親緣上,恐爪龍下目代表了一類非鳥類,而卻與鳥類是近親的恐龍。[16]

發現與命名

恐爪龍的化石出土於美國蒙大拿州懷俄明州的Cloverly組,[3]以及奧克拉荷馬州鹿角組[4]此外,馬里蘭州大西洋沿岸平原地帶的波多馬克組,發現了一些可能屬於恐爪龍的牙齒,年代為阿普第階[17]

恐爪龍的第一副化石是由巴納姆·布郎(Barnum Brown)所帶領的隊伍於1931年蒙大拿州南部的Cloverly組發現。布郎當時主要是想發掘並處理腱龍的遺骸,但在他交予美國自然歷史博物館的1931年報告中,指出發現了一細小的肉食性恐龍,發現位置接近腱龍化石,但因陷在石灰中而難以作清潔處理。[18]他非正式地將之命名為Daptosaurus,並準備描述及展示其骨骼,但他卻沒有完成。[19]另外,布朗將一個恐爪龍的牙齒,與另一種較小型獸腳類恐龍的骨骸混淆在一起,並非正式命名為Megadontosaurus

約30年後,從1964年8月開始,古生物學家約翰·奧斯特倫姆(John Ostrom)率領一個耶魯大學皮博迪自然史博物館的挖掘團隊,在其後的兩年內發現了超過1000個骨頭,來自於至少三個個體。因為很難判斷這些化石的正確位置,所以恐爪龍的正模標本(編號YPM 5205)只限於完整的左足部,與部分的右足部,但都確定屬於同一個體。[2]其他標本則在耶魯大學皮博迪自然史博物館區分、標示成50個不同的項目。

在接下來幾年,奧斯特倫姆及格蘭特·邁耶(Grant E. Meyer)研究這些新發現化石,以及布朗所命名的Daptosaurus化石,發現牠們是同種生物,並於1969年以平衡恐爪龍(Deinonychus antirrhopus)發表了他們的發現。[2]種名antirrhopus意為「平衡」,指的是牠們堅挺尾巴的可能用途。

奧斯特倫姆同時重新檢驗布朗所命名的Megadontosauru,他發現那隻牙齒是屬於恐爪龍,而骨骼則是來自另一類動物。他在1970年,將骨骼部分命名為小獵龍[20]

雖然在1969年發現大量的恐爪龍化石,但許多重要部位的骨頭並沒有發現,剩下的難以確認。這些化石包含少數眼眶後顱骨、沒有股骨、沒有薦骨、沒有叉骨胸骨、也沒有脊椎,以及一個被奧斯特倫姆認為是喙骨的小型碎片。奧斯特倫姆將恐爪龍重建成具有獨特的骨盆,恥骨呈梯形且平坦,與坐骨長度一樣。

一副恐爪龍的骨骼,包括原先標本的骨頭,都在美國自然歷史博物館中展覽,而其他的則在哈佛大學比較動物學博物館中展覽。這兩個標本是在與耶魯大學標本不同的位點發現的。

更多發現

In 1974 Ostrom published another monograph on the shoulder of Deinonychus in which he realized that the pubis that he had described was actually a coracoid – a shoulder element.[21] In that same year, another specimen of Deinonychus was excavated in Montana by a Harvard University expedition headed by Farish Jenkins. This discovery added several new elements; well preserved femurs, pubes, a sacrum, and better ilia, as well as elements of the pes and metatarsus. Ostrom described this specimen and revised his skeletal restoration of Deinonychus. This time it showed the very long pubes, and Ostrom began to suspect that they may have even been a little retroverted like those of birds.[22]

A skeleton of Deinonychus including bones from the original (and most complete) specimen can be seen on display at the American Museum of Natural History,[23] with another specimen on display at the Museum of Comparative Zoology at Harvard University. The American Museum and Harvard specimens are from a different locality than the Yale specimens. Even these two skeletal mounts are lacking elements including the sterna, sternal ribs, furcula, and gastralia.

Even after all of Ostrom’s work, several small blocks of lime-encased material remained unprepared in storage at the American Museum. These consisted mostly of isolated bones and bone fragments, including the original matrix, or surrounding rock in which the specimens were initially buried. An examination of these unprepared blocks by Gerald Grellet-Tinner and Peter Makovicky in 2000 revealed an interesting, overlooked feature. Several long, thin bones identified on the blocks as ossified tendons (structures which helped stiffen the tail of Deinonychus) turned out to actually represent gastralia (abdominal ribs). More significantly, a large number of previously unnoticed fossilized eggshells were discovered in the rock matrix which had surrounded the original Deinonychus specimen.[24]

In a subsequent, more detailed report on the eggshells, Grellet-Tinner and Makovicky concluded that the egg almost certainly belonged to Deinonychus, representing the first dromaeosaurid egg to be identified.[18] Moreover, the external surface of one eggshell was found in close contact with the gastralia suggesting that Deinonychus might have brooded its eggs. This implies that Deinonychus used body heat transfer as a mechanism for egg incubation, and indicates an endothermy similar to modern birds.[25] Further study by Gregory Erickson and colleagues finds that this individual was 13 or 14 years old at death and its growth had plateaued. Unlike other theropods in their study of specimens found associated with eggs or nests, it had finished growing at the time of its death.[26]

影響

恐爪龍的手(左)與始祖鳥的手(右)比較。

奧斯特倫姆1969年就恐爪龍的描述近認為是20世紀中期最重要的恐龍發現。[27]因為這個新發現的活躍、敏捷恐龍,改變了很多對恐龍的科學(或民間)認知,且開啟了對恐龍可能是溫血動物的辯論。這個認知的變化稱為恐龍文藝復興。幾年後,奧斯特倫姆發現了恐爪龍與鳥類的手的相似性,使他認為鳥類是從恐龍所演化而來。[28]三十年後,這個觀念幾乎是被普遍地接受。事實上,恐爪龍及其他馳龍科都很像鳥類,故亦有指牠們本身就是鳥類。

中國發現的恐爪龍近親,中國鳥龍小盜龍的遺骸在發現時都是有像羽毛的結構,故恐爪龍亦有可能有羽毛。[29]這兩類恐龍的特徵顯示牠們較恐爪龍更為原始,加上測定其年份要較恐爪龍早約1千萬年,加強了恐爪龍有羽毛的說法。[10]在其他獸腳亞目中,如尾羽龍鳥面龍北票龍帝龍,亦有發現羽毛的輪廓,在親緣分支分類法上令人更相信所有馳龍科都是有羽毛的。[30]

在2007年,恐爪龍的近親迅猛龍被發現前肢具有羽莖瘤。現代的鳥類也具有羽莖瘤,是羽毛的附著處。這個發現支持了所有馳龍科都具有羽毛的假設。[31]

古生物學

棲息地

平衡恐爪龍的重組,羽毛是根據相關物種而來的。

地質學證據顯示恐爪龍可在氾濫平原沼澤中發現。[10]Cloverly組及Antlers組的古生態環境包含有森林三角洲礁湖,並不像今天的路易斯安那州[32]其他同時期的恐龍包括有草食性楯甲龍鳥腳下目西風龍腱龍。恐爪龍的生態系統亦包括了巨大獸腳亞目高棘龙蜥腳下目波塞東龍、以及稜角鱗鱷雀鱔屬[32]

獵食行為

基於在同一位點發現大量恐爪龍的骨骼,且在腱龍的骨骼中發現恐爪龍的牙齒,估計恐爪龍是獵食腱龍的。奧斯特倫姆更以此推斷恐爪龍是成群生活及獵食的。[6]但是,最近的研究基於現今肉食性動物的獵食方式及腱龍位點的埋葬學,質疑了恐爪龍在群獵時的合作性行為。現今的初龍鳥類鱷魚)及科莫多龍有小數的合作性捕獵,不過牠們主要是單獨獵食,或是被吸引至已死的屍骸。例如,當一群科莫多龍一同攝食時,最大的會先吃,且會攻擊其他嘗試吃食的科莫多龍;如果細小的科莫多龍被殺死,牠們會吃食同類。當這些資料套用在腱龍位點時,這似乎與恐爪龍的攝食模式很吻合。在這些位點發現的恐爪龍骨骼都是接近成年的,而失去了的部位可能是被其他恐爪龍吃了。[33]

平衡恐爪龍的圖繪。

恐爪龍似鐮刀的爪在不同的標本都有所不同。模式種的爪是非常彎曲的,而於1976年描述的標本則較直,且與其他的爪很相似。[34]奧斯特倫姆認為在不同的個體、性別或年紀,這隻鐮刀爪的大小及形狀都有所不同。他猜測恐爪龍是以牠的手來捉住獵物,並以鐮刀爪來割破肚子。[2]但是後來發現這些鐮刀爪並非用作割開獵物的,而是刺戳獵物的。[35][1]生物力學確認了恐爪龍的前肢是捉住獵物的,前肢的長度較其他獸腳亞目更易於捕捉。大及長的喙突顯示牠有強壯的前肢肌肉,更加強了這個解釋。[36]不過,恐爪龍是不能掘曲牠的手臂,這與過往的說法有所出入。[37][15]其後的研究指恐爪龍的前肢未必只用來捕捉獵物,亦可以將物體移向胸部[38]

幼年及接近成年的恐爪龍標本與成年有著一些形態上的不同。例如,幼龍的手臂在比例上較成年的為長,顯示可能幼龍與成年的行為亦有所不同。[39][40]

速度

馳龍科,尤其是恐爪龍,一直都被認為是奔走快速的恐龍,而奧斯特倫姆亦在原先的描述中有此推論。[2]但是在首次描述後,一隻完整的恐爪龍腳被發現,而奧斯特倫姆所估計的大腿骨長度被證實是過長。奧斯特倫姆在其後的研究指腳掌與脛骨的相對長度較大腿骨與脛骨的比例,在評估速度上更為重要。現今奔走快速的鳥類,如鴕鳥,腳掌與脛骨比例為0.95;而在恐龍中,似鴕龍是0.68,恐爪龍則是出奇的低,只有0.48。[34]他於是結論出恐爪龍相對於其他恐龍並不怎麼快速,且更不是現今不能飛的鳥類的對手。

恐爪龍的這個低比例部份是因牠非常短的中骨。細小個體的中骨,縱然是短,仍比較大個體的為長。奧斯特倫姆認為短的中骨是與鐮刀爪的功能有關,當年紀漸長時中骨會更為短小。他解釋所有這些特徵(短的二趾上有大的爪、短的中骨等)都是為使後肢成為攻擊性的武器,鐮刀爪可以向上及向下攻擊,而腳則同時向後及向下拉,把獵物割破及撕開。奧斯特倫姆指短的中骨減少了腳骨在攻擊時的整體壓力,而恐爪龍的特殊肌肉是用作獵食而非奔走。所以他斷定恐爪龍的腳是平衡奔跑,及減低作為武器時的壓力。[34]

繁殖

2000年發現與原先標本有關的恐爪龍蛋,可以與其他的獸腳亞目在蛋結構、巢穴及繁殖作出比較。在評估這些蛋殼標本的時候,一些可能性,如馳龍科是吃蛋的或這些蛋碎片的出現是巧合,都一一被否定。因為這些蛋殼是在恐爪龍的胃骨片與前肢之間,並非是在胃部之內,故這些蛋並非牠們的食物。再者,蛋殼裂開的方式顯示它在埋葬時並沒有改變,而是因化石化的過程中裂開的。另外,由於蛋周圍的骨骼並非分散或分開的(反而是與在生時差不多的姿勢),可見這個位點在保存期間並沒有受到影響,故巧合之說亦不成立。事實上,胃骨片一般很難發現是連接的,可見這些蛋是在埋葬時就在恐爪龍的腹下。這可能是恐爪龍的孵蛋或築巢行為(像傷齒龍科竊蛋龍科),或是仍在輸卵管之中。[18]

恐爪龍蛋的微結構評估發現它們是屬於獸腳亞目中,因為它們有著其他獸腳亞目蛋的特徵,並與鳥臀目蜥腳下目蛋的不同。與其他手盜龍類比較,恐爪龍蛋較像竊蛋龍科的蛋,而非傷齒龍科的蛋。雖然該蛋經已碎裂而很難評估其大小,但憑其經過的小骨盆腔可知直徑約為7厘米。大小與最大的葬火龍蛋的7.2厘米接近。再者,葬火龍蛋殼及恐爪龍蛋殼厚度差不多完全一致,由於蛋殼厚度影響著蛋的體積,故此有指牠們蛋的大小是完全一樣的。[18]

化石蛋

奧斯特倫姆描述了布郎發現的原先標本後,仍有幾個細小陷在石灰的標本未被處理而擺放在美國自然歷史博物館。這些大部份都是獨立的骨頭及骨骼碎片,包括了標本埋葬時周圍的岩石。2000年時就這些標本的研究發現一些有趣及被忽略了的資料。當中的幾條幼長骨頭,原先被認為是用作令恐爪龍尾巴堅硬的骨頭,其實是胃骨片。而且亦在原先恐爪龍標本的周圍,發現了一些被忽略的蛋殼化石[24]在後期詳細的報告中,這些蛋差不多肯定是屬於恐爪龍的,故是首批發現馳龍科的蛋。[18]再者,其中一個蛋殼的外表面緊接恐爪龍的胃骨片,可見恐爪龍可能正在孵蛋。由此推斷恐爪龍是用身體的熱量來孵蛋,且與現今鳥類相似是溫血動物[41]

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外部連結

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