動物定義
三(san)角龍(long)(屬(shu)名(ming):Triceratops) 是(shi)鳥臀目角龍(long)下(xia)目角龍(long)科的(de)(de)植食性恐(kong)龍(long)的(de)(de)一屬(shu),化石發現于北美洲(zhou)的(de)(de)晚白堊紀晚馬斯垂克階(jie)地層(ceng),約6800萬(wan)年(nian)前到6500萬(wan)年(nian)前。三(san)角龍(long)是(shi)最晚出現的(de)(de)恐(kong)龍(long)之一,經常被作為晚白堊紀的(de)(de)代表化石。
三角(jiao)龍是(shi)一種中等大小的(de)(de)四(si)足(zu)恐龍,全(quan)長6—8米(mi)、高2.4-2.8米(mi)、重5—10噸。他們有(you)(you)非(fei)常(chang)大的(de)(de)頭盾(dun),以及(ji)三根角(jiao)狀(zhuang)物,令人聯想起(qi)現代犀(xi)牛。雖(sui)然沒有(you)(you)發現過(guo)三角(jiao)龍的(de)(de)完整(zheng)骨(gu)骸(hai)(hai),他們仍因大量從1887年起(qi)發現的(de)(de)部份骨(gu)骸(hai)(hai)標本(ben)而著(zhu)名。長久以來,關(guan)于它們三根角(jiao)以及(ji)頭盾(dun)的(de)(de)功能(neng)(neng)處于爭(zheng)論中。傳統上這些結(jie)構(gou)被認為是(shi)用(yong)來抵抗掠食者的(de)(de)武器,但最近(jin)的(de)(de)理論認為這些結(jie)構(gou)可能(neng)(neng)用(yong)在求偶(ou),以及(ji)展示支配地位,如(ru)同現代馴鹿、山羊、獨角(jiao)仙的(de)(de)角(jiao)狀(zhuang)物。
三角(jiao)(jiao)龍最顯(xian)著的(de)特(te)征(zheng)是它們的(de)角(jiao)(jiao)和(he)頸(jing)盾。它們的(de)頭盾可長至超過(guo)1.5米。三角(jiao)(jiao)龍的(de)口鼻部鼻孔(kong)上(shang)方(fang)有(you)一根角(jiao)(jiao)狀(zhuang)物(wu);以(yi)及(ji)一對位在眼睛上(shang)方(fang)的(de)角(jiao)(jiao)狀(zhuang)物(wu),超過(guo)80公分。頭顱(lu)后方(fang)則是相對短(duan)的(de)骨質頭盾。
三(san)角龍(long)(long)四(si)肢(zhi)短,前腳掌有五個(ge)短腳趾、后(hou)腳掌則有四(si)個(ge)短腳趾。雖(sui)然三(san)角龍(long)(long)確定是(shi)四(si)足(zu)動物,它們的(de)(de)姿(zi)勢(shi)長久(jiu)以(yi)來處(chu)于爭(zheng)論(lun)(lun)中。三(san)角龍(long)(long)的(de)(de)前肢(zhi)起(qi)初(chu)被認為是(shi)從胸(xiong)部(bu)(bu)往兩側(ce)伸(shen)展(zhan),以(yi)助于承擔頭部(bu)(bu)的(de)(de)重量。這種站立(li)方式可(ke)見于查爾斯·耐特(Charles R. Knight)與(yu)魯(lu)道夫·札林格(Rudolph F. Zallinger)的(de)(de)繪畫中。然而,角龍(long)(long)類的(de)(de)足(zu)跡化石證據,以(yi)及如今的(de)(de)骨骸重建,顯示三(san)角龍(long)(long)在正常行(xing)走時保持者(zhe)直立(li)姿(zi)勢(shi),但肘部(bu)(bu)稍微彎曲,居于完全直立(li)與(yu)完全伸(shen)展(zhan)(現代(dai)犀牛)兩種說法的(de)(de)中間。但這種結(jie)論(lun)(lun)無法排(pai)除三(san)角龍(long)(long)抵抗或進食(shi)時會采伸(shen)展(zhan)姿(zi)態。
科屬分類
三(san)(san)角(jiao)龍(long)(long)(long)(long)(long)(long)(long)是角(jiao)龍(long)(long)(long)(long)(long)(long)(long)科(ke)(ke)(ke)中最著名屬(shu),角(jiao)龍(long)(long)(long)(long)(long)(long)(long)科(ke)(ke)(ke)是群中型北(bei)美洲(zhou)角(jiao)龍(long)(long)(long)(long)(long)(long)(long)類(lei)恐龍(long)(long)(long)(long)(long)(long)(long)。多年(nian)以來,三(san)(san)角(jiao)龍(long)(long)(long)(long)(long)(long)(long)于(yu)(yu)角(jiao)龍(long)(long)(long)(long)(long)(long)(long)下目中的(de)位置處于(yu)(yu)爭論(lun)中。混淆來自于(yu)(yu)三(san)(san)角(jiao)龍(long)(long)(long)(long)(long)(long)(long)的(de)短(duan)、堅硬頭(tou)盾類(lei)似尖角(jiao)龍(long)(long)(long)(long)(long)(long)(long)亞(ya)(ya)科(ke)(ke)(ke),而長(chang)的(de)額角(jiao)類(lei)似角(jiao)龍(long)(long)(long)(long)(long)(long)(long)科(ke)(ke)(ke)(或稱開角(jiao)龍(long)(long)(long)(long)(long)(long)(long)亞(ya)(ya)科(ke)(ke)(ke))。在(zai)第一(yi)個(ge)對于(yu)(yu)角(jiao)龍(long)(long)(long)(long)(long)(long)(long)類(lei)的(de)研(yan)(yan)究中,理察·史旺(wang)·魯爾(Richard Swann Lull)提出(chu)(chu)出(chu)(chu)兩個(ge)支系,一(yi)個(ge)往(wang)三(san)(san)角(jiao)龍(long)(long)(long)(long)(long)(long)(long)發展的(de)支系包(bao)括(kuo)(kuo)獨角(jiao)龍(long)(long)(long)(long)(long)(long)(long)、尖角(jiao)龍(long)(long)(long)(long)(long)(long)(long),另(ling)一(yi)個(ge)支系包(bao)括(kuo)(kuo)角(jiao)龍(long)(long)(long)(long)(long)(long)(long)、牛(niu)角(jiao)龍(long)(long)(long)(long)(long)(long)(long),這(zhe)個(ge)假設使三(san)(san)角(jiao)龍(long)(long)(long)(long)(long)(long)(long)屬(shu)于(yu)(yu)尖角(jiao)龍(long)(long)(long)(long)(long)(long)(long)亞(ya)(ya)科(ke)(ke)(ke)。較晚(wan)的(de)研(yan)(yan)究支持(chi)這(zhe)個(ge)觀點,并將這(zhe)個(ge)短(duan)頭(tou)盾支系正式命名為尖角(jiao)龍(long)(long)(long)(long)(long)(long)(long)亞(ya)(ya)科(ke)(ke)(ke)(包(bao)含三(san)(san)角(jiao)龍(long)(long)(long)(long)(long)(long)(long)),另(ling)一(yi)長(chang)頭(tou)盾支系為開角(jiao)龍(long)(long)(long)(long)(long)(long)(long)亞(ya)(ya)科(ke)(ke)(ke)。
在1949年,查爾(er)斯(si)(si)·斯(si)(si)騰伯格(ge)(Charles Mortram Sternberg)首次對這(zhe)假(jia)設提出質(zhi)疑(yi),并基于頭顱(lu)與角(jiao)的特征,而(er)認為(wei)(wei)三(san)角(jiao)龍與無鼻角(jiao)龍、開角(jiao)龍關系較近,使得三(san)角(jiao)龍屬為(wei)(wei)三(san)角(jiao)龍亞(ya)科(ke)(尖(jian)角(jiao)龍亞(ya)科(ke))。然而(er),斯(si)(si)騰伯格(ge)的分類與約翰·奧斯(si)(si)特倫姆(John Ostrom)、大(da)衛·諾曼(David Norman)將三(san)角(jiao)龍置于尖(jian)角(jiao)龍亞(ya)科(ke)不同。
后來的發(fa)現與(yu)研(yan)究(jiu)支持了(le)斯騰伯(bo)格的觀(guan)點(dian),Lehman在1990年(nian)為(wei)兩個亞科(ke)(ke)定義,并基于數個形態(tai)上的特征,而將三角龍歸于角龍亞科(ke)(ke)。事實上,除了(le)短頭盾以外,三角龍相當符(fu)合角龍亞科(ke)(ke)的特征。彼(bi)得·達德森(Peter Dodson)在1990年(nian)的親緣(yuan)分支分類法研(yan)究(jiu)與(yu)1993年(nian)的形態(tai)學研(yan)究(jiu),加強三角龍歸類為(wei)角龍亞科(ke)(ke)的論點(dian)。
種系分類
在種系發生學的分類中,三角龍通常作為恐龍定義中的一個參考點;恐龍(long)被定義為三(san)角龍(long)與(yu)新鳥亞綱(gang)(現代鳥類)與(yu)共(gong)同祖(zu)先(xian),的(de)(de)所(suo)有后代。而(er)(er)鳥臀目的(de)(de)定義為:所(suo)有親緣關(guan)系與(yu)三(san)角龍(long)接近(jin),而(er)(er)離現代鳥類較遠的(de)(de)擁有共(gong)同祖(zu)先(xian)之物種(zhong)。
物種起源
很多年(nian)(nian)來三角(jiao)(jiao)(jiao)龍(long)(long)的起源(yuan)非常(chang)不明(ming)確。在(zai)1922年(nian)(nian),新發(fa)(fa)現的原角(jiao)(jiao)(jiao)龍(long)(long)被亨利·費(fei)爾費(fei)爾德·奧斯本(Henry Fairfield Osborn)認為(wei)是(shi)三角(jiao)(jiao)(jiao)龍(long)(long)的祖(zu)先(xian)(xian)。然(ran)而(er),如今發(fa)(fa)現數種與三角(jiao)(jiao)(jiao)龍(long)(long)祖(zu)先(xian)(xian)有關系的物種。發(fa)(fa)現于90年(nian)(nian)代晚期的祖(zu)先(xian)(xian)角(jiao)(jiao)(jiao)龍(long)(long),是(shi)角(jiao)(jiao)(jiao)龍(long)(long)下(xia)目中已知(zhi)最早有額角(jiao)(jiao)(jiao)的恐(kong)龍(long)(long)。而(er)2005年(nian)(nian)發(fa)(fa)現的隱龍(long)(long),是(shi)已知(zhi)唯一(yi)的侏羅紀角(jiao)(jiao)(jiao)龍(long)(long)下(xia)目恐(kong)龍(long)(long)。
這(zhe)些新(xin)發現非(fei)常重要,并描繪出角(jiao)龍(long)類(lei)恐(kong)龍(long)的起源,它們起源于(yu)侏(zhu)羅紀的亞洲(zhou),而(er)真正有角(jiao)的角(jiao)龍(long)類(lei)出現在(zai)晚白堊紀之初。三(san)(san)角(jiao)龍(long)越來越常被認為是(shi)角(jiao)龍(long)亞科的一個成(cheng)員(yuan),三(san)(san)角(jiao)龍(long)的祖(zu)先(xian)可能外(wai)表(biao)類(lei)似開角(jiao)龍(long),開角(jiao)龍(long)生存(cun)時間早(zao)于(yu)三(san)(san)角(jiao)龍(long)約500萬(wan)年。
發現與種
命名
第(di)一個(ge)(ge)被命名為(wei)三(san)角(jiao)(jiao)龍(long)的(de)(de)(de)標本,是在(zai)1887年發(fa)(fa)現(xian)(xian)(xian)于(yu)科羅(luo)拉(la)多州(zhou)丹佛市(shi)附近,由(you)一個(ge)(ge)頭(tou)顱(lu)(lu)骨頂部,與(yu)附著在(zai)上面的(de)(de)(de)一對額角(jiao)(jiao)所構成(cheng)。這(zhe)個(ge)(ge)標本被交給奧塞(sai)內爾·查利斯(si)·馬(ma)什(Othniel Charles Marsh),他認為(wei)該化石(shi)的(de)(de)(de)所處地層年代為(wei)上新世,而(er)該化石(shi)屬(shu)于(yu)一種(zhong)特(te)別大(da)的(de)(de)(de)北(bei)(bei)(bei)美(mei)野(ye)牛(niu),因此將它們命名為(wei)長角(jiao)(jiao)北(bei)(bei)(bei)美(mei)野(ye)牛(niu)(Bison alticornis)。第(di)二年,馬(ma)什根(gen)據一些破碎的(de)(de)(de)化石(shi),發(fa)(fa)現(xian)(xian)(xian)了(le)(le)有角(jiao)(jiao)恐龍(long)的(de)(de)(de)存在(zai),因此建立了(le)(le)角(jiao)(jiao)龍(long)屬(shu);但他仍認為(wei)長角(jiao)(jiao)北(bei)(bei)(bei)美(mei)野(ye)牛(niu)是種(zhong)上新世的(de)(de)(de)哺乳類。直到(dao)第(di)三(san)個(ge)(ge)更完(wan)整(zheng)的(de)(de)(de)角(jiao)(jiao)龍(long)類頭(tou)顱(lu)(lu)骨,才改變他的(de)(de)(de)想法。這(zhe)個(ge)(ge)由(you)約(yue)翰·貝爾·海徹(che)爾(John Bell Hatcher)在(zai)1888年于(yu)懷俄明(ming)(ming)州(zhou)蘭斯(si)組發(fa)(fa)現(xian)(xian)(xian)的(de)(de)(de)標本,起初被敘述成(cheng)角(jiao)(jiao)龍(long)屬(shu)的(de)(de)(de)另外一個(ge)(ge)種(zhong),但馬(ma)什經過熟慮(lv)之后,他將這(zhe)個(ge)(ge)標本命名為(wei)三(san)角(jiao)(jiao)龍(long)(Triceratops),并將原(yuan)本的(de)(de)(de)長角(jiao)(jiao)北(bei)(bei)(bei)美(mei)野(ye)牛(niu)改歸類于(yu)角(jiao)(jiao)龍(long)的(de)(de)(de)一個(ge)(ge)種(zhong)(后來(lai)也(ye)成(cheng)為(wei)三(san)角(jiao)(jiao)龍(long)的(de)(de)(de)一種(zhong))。三(san)角(jiao)(jiao)龍(long)的(de)(de)(de)結實頭(tou)顱(lu)(lu)骨使得許(xu)多頭(tou)顱(lu)(lu)骨被保存下來(lai),允許(xu)科學(xue)家們研究不同種(zhong)與(yu)個(ge)(ge)體間的(de)(de)(de)變化。除了(le)(le)科羅(luo)拉(la)多州(zhou)與(yu)懷俄明(ming)(ming)州(zhou)之外,隨后在(zai)美(mei)國(guo)的(de)(de)(de)蒙大(da)拿州(zhou)與(yu)南達科他州(zhou)、加拿大(da)的(de)(de)(de)亞伯達省(sheng)與(yu)薩克其萬省(sheng)也(ye)發(fa)(fa)現(xian)(xian)(xian)了(le)(le)三(san)角(jiao)(jiao)龍(long)的(de)(de)(de)化石(shi)。
種
在三(san)角(jiao)(jiao)龍(long)第一次被(bei)命(ming)(ming)名的(de)前十(shi)年內,發現了不(bu)(bu)同(tong)(tong)(tong)的(de)頭(tou)顱(lu)(lu)骨(gu),這(zhe)些(xie)(xie)骨(gu)骸與(yu)馬什最初命(ming)(ming)名的(de)恐怖(bu)(bu)三(san)角(jiao)(jiao)龍(long)(T. horridus)頭(tou)顱(lu)(lu)骨(gu)有或多或少的(de)不(bu)(bu)同(tong)(tong)(tong);恐怖(bu)(bu)三(san)角(jiao)(jiao)龍(long)的(de)種名horridus在拉丁(ding)語中其實(shi)意為「凹凸不(bu)(bu)平的(de)」,意指原型標本的(de)凹凸不(bu)(bu)平表面,該標本后來(lai)被(bei)確認(ren)為幼年個(ge)體。這(zhe)些(xie)(xie)頭(tou)顱(lu)(lu)骨(gu)的(de)差(cha)異可歸類出三(san)種不(bu)(bu)同(tong)(tong)(tong)尺寸,這(zhe)些(xie)(xie)差(cha)異來(lai)自于不(bu)(bu)同(tong)(tong)(tong)年齡與(yu)性別的(de)差(cha)異,以及化石化過程中的(de)不(bu)(bu)同(tong)(tong)(tong)程度或壓力方向。這(zhe)些(xie)(xie)不(bu)(bu)同(tong)(tong)(tong)頭(tou)顱(lu)(lu)骨(gu)被(bei)命(ming)(ming)名為個(ge)別的(de)種(可見于以下列表),并(bing)形成數個(ge)系統發生學研究。
魯(lu)爾發現這些種可分為(wei)(wei)兩(liang)個(ge)(ge)生(sheng)物群(qun),但他(ta)并(bing)沒(mei)有(you)說明如何(he)分辨它們;其中一群(qun)由(you)恐(kong)怖三(san)角(jiao)(jiao)龍(long)(long)(long)(long)、T. prorsus、短角(jiao)(jiao)三(san)角(jiao)(jiao)龍(long)(long)(long)(long)(T. brevicornus)所(suo)構(gou)成,另一群(qun)由(you)T . elatus、T. calicornis所(suo)構(gou)成。鋸齒三(san)角(jiao)(jiao)龍(long)(long)(long)(long)(T. serratus)與(yu)(yu)(yu)(yu)扇形(xing)三(san)角(jiao)(jiao)龍(long)(long)(long)(long)(T. flabellatus)則不屬于這兩(liang)個(ge)(ge)生(sheng)物群(qun)。在1933年(nian),魯(lu)爾將之(zhi)前他(ta)與(yu)(yu)(yu)(yu)海徹(che)爾、馬什(shen)先后完成的(de)角(jiao)(jiao)龍(long)(long)(long)(long)下目專題論文重(zhong)新出版(ban),他(ta)維持原(yuan)本的(de)兩(liang)個(ge)(ge)生(sheng)物群(qun)與(yu)(yu)(yu)(yu)兩(liang)個(ge)(ge)未定(ding)種的(de)分類(lei)法,并(bing)增加(jia)了第(di)(di)(di)三(san)個(ge)(ge)支系,由(you)鈍頭(tou)三(san)角(jiao)(jiao)龍(long)(long)(long)(long)(T. obtusus)與(yu)(yu)(yu)(yu)海氏三(san)角(jiao)(jiao)龍(long)(long)(long)(long)(T. hatcheri)所(suo)構(gou)成,特征是非常小(xiao)的(de)鼻(bi)(bi)(bi)角(jiao)(jiao)。恐(kong)怖三(san)角(jiao)(jiao)龍(long)(long)(long)(long)、T. prorsus、以及短角(jiao)(jiao)三(san)角(jiao)(jiao)龍(long)(long)(long)(long)所(suo)構(gou)成的(de)生(sheng)物群(qun),此時被(bei)認為(wei)(wei)是最傳(chuan)統的(de)支系,頭(tou)顱骨(gu)較大,鼻(bi)(bi)(bi)角(jiao)(jiao)較小(xiao)。而T. elatus、T. calicornis所(suo)構(gou)成的(de)第(di)(di)(di)二(er)個(ge)(ge)生(sheng)物群(qun),有(you)大型額角(jiao)(jiao)與(yu)(yu)(yu)(yu)小(xiao)型鼻(bi)(bi)(bi)角(jiao)(jiao)。查爾斯·斯騰伯格作了些調整(zheng),他(ta)將寬頭(tou)三(san)角(jiao)(jiao)龍(long)(long)(long)(long)(T. eurycephalus)視為(wei)(wei)第(di)(di)(di)二(er)與(yu)(yu)(yu)(yu)第(di)(di)(di)三(san)個(ge)(ge)生(sheng)物群(qun)之(zhi)間的(de)連(lian)結,而非恐(kong)怖三(san)角(jiao)(jiao)龍(long)(long)(long)(long)所(suo)屬的(de)第(di)(di)(di)一生(sheng)物群(qun)。這個(ge)(ge)分類(lei)法持續用(yong)到80年(nian)代與(yu)(yu)(yu)(yu)90年(nian)代。
不同角(jiao)龍類的(de)(de)頭顱骨代(dai)表者單(dan)一(yi)種(zhong)(zhong)(或兩(liang)(liang)個(ge)(ge)(ge)種(zhong)(zhong))之內的(de)(de)個(ge)(ge)(ge)體(ti)(ti)變(bian)化,這個(ge)(ge)(ge)論點逐漸普(pu)及。在1986年(nian),奧斯(si)特倫姆與(yu)(yu)彼(bi)得·沃爾赫(he)費爾(Peter Wellnhofer)公布一(yi)份研究,他們宣稱三(san)(san)(san)角(jiao)龍屬只有一(yi)個(ge)(ge)(ge)種(zhong)(zhong),恐(kong)(kong)怖三(san)(san)(san)角(jiao)龍。其中一(yi)個(ge)(ge)(ge)理由(you)是在一(yi)個(ge)(ge)(ge)地區中,通常(chang)只存在者單(dan)一(yi)或兩(liang)(liang)個(ge)(ge)(ge)大型(xing)動物(wu)群(qun);例如現(xian)代(dai)非洲(zhou)的(de)(de)非洲(zhou)象與(yu)(yu)長頸鹿。 Lehman在魯(lu)爾與(yu)(yu)斯(si)騰伯格的(de)(de)支系研究中,加進兩(liang)(liang)性異(yi)形與(yu)(yu)年(nian)齡變(bian)化,他認(ren)為由(you)恐(kong)(kong)怖三(san)(san)(san)角(jiao)龍、T. prorsus、短(duan)角(jiao)三(san)(san)(san)角(jiao)龍所(suo)構(gou)(gou)成(cheng)的(de)(de)第一(yi)個(ge)(ge)(ge)生物(wu)群(qun)是雌性個(ge)(ge)(ge)體(ti)(ti),而T. elatus、T. calicornis所(suo)構(gou)(gou)成(cheng)的(de)(de)第二個(ge)(ge)(ge)生物(wu)群(qun)是雄(xiong)性個(ge)(ge)(ge)體(ti)(ti),而鈍頭三(san)(san)(san)角(jiao)龍與(yu)(yu)海氏三(san)(san)(san)角(jiao)龍所(suo)構(gou)(gou)成(cheng)的(de)(de)第三(san)(san)(san)個(ge)(ge)(ge)生物(wu)群(qun)是年(nian)老的(de)(de)雄(xiong)性個(ge)(ge)(ge)體(ti)(ti)。他的(de)(de)理由(you)是雄(xiong)性個(ge)(ge)(ge)體(ti)(ti)的(de)(de)體(ti)(ti)型(xing)較(jiao)高、頭角(jiao)較(jiao)直、頭顱骨較(jiao)大,而雌性個(ge)(ge)(ge)體(ti)(ti)的(de)(de)頭顱骨較(jiao)小(xiao)、頭角(jiao)較(jiao)短(duan)。
數(shu)年后,凱薩琳·佛斯特(te)(Catherine Forster)對奧(ao)斯特(te)倫姆與(yu)沃爾赫費爾的研(yan)究(jiu)提出(chu)質疑,佛斯特(te)對三(san)角(jiao)龍(long)(long)的化石(shi)材料做了更廣(guang)泛(fan)的研(yan)究(jiu),并認為三(san)角(jiao)龍(long)(long)只(zhi)有(you)兩個(ge)種(zhong):恐(kong)怖三(san)角(jiao)龍(long)(long)、T. prorsus;而(er)海氏(shi)三(san)角(jiao)龍(long)(long)因(yin)擁有(you)獨特(te)頭顱骨(gu),足(zu)以成立新的屬,已改(gai)為海氏(shi)雙(shuang)角(jiao)龍(long)(long)(Diceratops hatcheri)。她發現數(shu)個(ge)種(zhong)其實屬于(yu)(yu)恐(kong)怖三(san)角(jiao)龍(long)(long),而(er)T. prorsus與(yu)短角(jiao)三(san)角(jiao)龍(long)(long)是同一個(ge)種(zhong),因(yin)為有(you)許(xu)多種(zhong)被分(fen)類于(yu)(yu)第一個(ge)生物群,佛斯特(te)提出(chu)前兩個(ge)生物群其實分(fen)別(bie)代表恐(kong)怖三(san)角(jiao)龍(long)(long)與(yu)T. prorsus。但(dan)在這(zhe)個(ge)新分(fen)類法之下,這(zhe)兩個(ge)種(zhong)仍有(you)可能是兩性(xing)異形的結果。
已確定種
*恐怖三角龍(T. horridus):奧思尼爾·馬什1889 (模式種)
* T. prorsus:馬什1890
可疑種
*亞伯(bo)達三(san)角龍(long)(T. albertensis):斯騰(teng)伯(bo)格1949
*長角三(san)角龍(long)(T. alticornis):馬(ma)什1887
*寬頭三角(jiao)龍(T. eurycephalus):施賴克(ke)耶爾1935
* T. galeus:馬什1889
*碩大(da)三角(jiao)龍(T. ingens):魯(lu)爾1915
*大(da)三角龍(T. maximus):布朗1933
* T. sulcatus:馬什1890
錯誤歸類
*短(duan)角三角龍(T. brevicornis):海徹爾1905 ( = T. prorsus)
* T. calicornus:馬什1898 ( =恐怖三(san)角龍(long))
* T. elatus:馬什1891 ( =恐怖三角龍)
*扇(shan)形三角(jiao)龍(T. flabellatus):馬什1889 ( =恐怖三角(jiao)龍)
*海氏(shi)三角龍(long)(T. hatcheri):魯爾1907 ( =海氏(shi)雙角龍(long))
* T. mortuarius:愛德華·科普1874 ( =大師(shi)龍(long),疑名)
*鈍(dun)頭三角龍(T. obtusus):馬什1898 ( =恐怖三角龍)
*鋸齒三角龍(long)(T. serratus):馬什1890 ( =恐怖三角龍(long))
*森(sen)林(lin)三(san)角龍(long)(T. sylvestris):科普1872 ( =森(sen)林(lin)奇(qi)跡龍(long),疑名)
古生物學
雖然三(san)角(jiao)龍(long)(long)常(chang)被(bei)描述成(cheng)群居動物(wu),但沒有直接(jie)證據(ju)顯示(shi)它們(men)為群居動物(wu)。但有些角(jiao)龍(long)(long)類恐(kong)龍(long)(long)的(de)(de)(de)發(fa)現(xian)地點常(chang)有數十或數百個個體。明尼蘇達科(ke)學博物(wu)館的(de)(de)(de)古(gu)生物(wu)學家(jia)Bruce Erickson宣(xuan)稱(cheng)在(zai)蒙大拿州海爾(er)河(he)組(zu)發(fa)現(xian)了(le)200個T. prorsus的(de)(de)(de)標本。巴納姆·布郎(lang)(Barnum Brown)則宣(xuan)稱(cheng)在(zai)該處發(fa)現(xian)了(le)超(chao)過500個頭(tou)顱(lu)骨。因為在(zai)北美洲西(xi)部的(de)(de)(de)蘭斯組(zu)(晚馬斯垂克階,6800萬(wan)年前(qian)到(dao)6500萬(wan)年前(qian))發(fa)現(xian)了(le)豐富的(de)(de)(de)三(san)角(jiao)龍(long)(long)牙齒、角(jiao)狀物(wu)碎片、頭(tou)盾碎片、以及三(san)角(jiao)龍(long)(long)其(qi)他(ta)的(de)(de)(de)破(po)碎頭(tou)顱(lu)骨。在(zai)1986年,羅伯特·巴克(Robert Bakker)估計(ji)在(zai)白(bai)堊紀,三(san)角(jiao)龍(long)(long)的(de)(de)(de)數量占了(le)蘭斯組(zu)動物(wu)群的(de)(de)(de)5/6。三(san)角(jiao)龍(long)(long)的(de)(de)(de)頭(tou)骨較常(chang)被(bei)發(fa)現(xian),而(er)非身(shen)體部分,與大部分動物(wu)不(bu)同。
三角(jiao)(jiao)龍(long)是白(bai)堊紀-第三紀滅絕事件之前最后出現(xian)的角(jiao)(jiao)龍(long)類之一(yi)。三角(jiao)(jiao)龍(long)的近親(qin)雙(shuang)角(jiao)(jiao)龍(long)與牛(niu)角(jiao)(jiao)龍(long)、以及(ji)遠親(qin)纖角(jiao)(jiao)龍(long)也生存在同一(yi)時期,但它們的化石(shi)較少(shao)被發現(xian)。
齒列
三(san)角(jiao)(jiao)龍是植(zhi)食性動物,但它(ta)們(men)也可能使用(yong)頭角(jiao)(jiao)、喙(hui)狀嘴、以身(shen)體來撞倒(dao)較高的(de)植(zhi)被(bei)來食用(yong)。三(san)角(jiao)(jiao)龍的(de)顎(e)部前(qian)端具(ju)有(you)長(chang)、狹窄(zhai)的(de)喙(hui)狀嘴,被(bei)認(ren)為較適合抓取、拉(la)扯(che),而非咬(yao)合。
三(san)(san)角(jiao)龍的牙齒(chi)(chi)排列(lie)(lie)成齒(chi)(chi)系(Tooth batteries),每列(lie)(lie)由36到40個牙齒(chi)(chi)群所構成,上下顎兩側(ce)各有(you)3到5列(lie)(lie)牙齒(chi)(chi)群,牙齒(chi)(chi)群的牙齒(chi)(chi)數量(liang)依照動物體(ti)型(xing)而改(gai)變。三(san)(san)角(jiao)龍總共擁有(you)432到 800顆牙齒(chi)(chi),其(qi)(qi)中大部分都是(shi)正在(zai)使用,而三(san)(san)角(jiao)龍的牙齒(chi)(chi)是(shi)不(bu)斷地生長(chang)并取代(dai)。這些牙齒(chi)(chi)以(yi)垂直或接(jie)近垂直的方向來(lai)切割(ge)食物。三(san)(san)角(jiao)龍的眾多牙齒(chi)(chi),顯示它(ta)們以(yi)體(ti)積(ji)大的棕櫚植物為食。三(san)(san)角(jiao)龍的食物中可能(neng)包(bao)含(han)棕櫚科(ke)與(yu)蘇鐵,而其(qi)(qi)他人(ren)員則認為包(bao)含(han)草原上的蕨類。
角與頭盾
在(zai)2005年,BBC的(de)(de)(de)(de)(de)(de)(de)(de)電視節(jie)(jie)目(mu)(mu)(mu)《恐龍(long)(long)(long)(long)(long)(long)(long)兇(xiong)面目(mu)(mu)(mu)》(The Truth About Killer Dinosaurs)之(zhi)中,節(jie)(jie)目(mu)(mu)(mu)單位測試三角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)龍(long)(long)(long)(long)(long)(long)(long)將如何抵抗大(da)(da)型(xing)掠食者如暴(bao)龍(long)(long)(long)(long)(long)(long)(long)的(de)(de)(de)(de)(de)(de)(de)(de)攻擊(ji)。為(wei)(wei)了(le)(le)(le)了(le)(le)(le)解(jie)三角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)龍(long)(long)(long)(long)(long)(long)(long)是(shi)(shi)(shi)否如同(tong)現代(dai)(dai)(dai)犀牛(niu)般沖撞敵人(ren)(ren),節(jie)(jie)目(mu)(mu)(mu)單位制作(zuo)了(le)(le)(le)一個(ge)(ge)(ge)人(ren)(ren)工(gong)的(de)(de)(de)(de)(de)(de)(de)(de)三角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)龍(long)(long)(long)(long)(long)(long)(long)頭顱(lu),并(bing)以(yi)每(mei)小(xiao)時(shi)24公里的(de)(de)(de)(de)(de)(de)(de)(de)時(shi)速撞向模擬(ni)的(de)(de)(de)(de)(de)(de)(de)(de)暴(bao)龍(long)(long)(long)(long)(long)(long)(long)皮膚。三角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)龍(long)(long)(long)(long)(long)(long)(long)的(de)(de)(de)(de)(de)(de)(de)(de)額角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)刺穿的(de)(de)(de)(de)(de)(de)(de)(de)模擬(ni)皮膚,但是(shi)(shi)(shi)額角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)與(yu)(yu)喙狀嘴則無法(fa)刺穿,而(er)(er)且(qie)頭顱(lu)的(de)(de)(de)(de)(de)(de)(de)(de)前段(duan)斷裂。結論(lun)是(shi)(shi)(shi)三角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)龍(long)(long)(long)(long)(long)(long)(long)無法(fa)利用(yong)沖撞敵人(ren)(ren)來(lai)自我(wo)防衛,如果它們(men)遭到攻擊(ji)時(shi),應該(gai)會采取堅守策(ce)略,當敵人(ren)(ren)接(jie)近(jin)時(shi),使(shi)(shi)用(yong)它們(men)的(de)(de)(de)(de)(de)(de)(de)(de)角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)來(lai)牴(di)刺敵人(ren)(ren)。除(chu)了(le)(le)(le)將頭角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)用(yong)于抵抗掠食者以(yi)外,三角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)龍(long)(long)(long)(long)(long)(long)(long)可能會使(shi)(shi)用(yong)頭角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)互相碰撞。研(yan)究顯(xian)示(shi)(shi)(shi)這(zhe)(zhe)種互相碰撞的(de)(de)(de)(de)(de)(de)(de)(de)行(xing)為(wei)(wei)是(shi)(shi)(shi)合理的(de)(de)(de)(de)(de)(de)(de)(de)、可行(xing)的(de)(de)(de)(de)(de)(de)(de)(de),但沒(mei)有(you)(you)(you)證(zheng)(zheng)據顯(xian)示(shi)(shi)(shi)三角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)龍(long)(long)(long)(long)(long)(long)(long)擁(yong)(yong)有(you)(you)(you)這(zhe)(zhe)種行(xing)為(wei)(wei)。三角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)龍(long)(long)(long)(long)(long)(long)(long)與(yu)(yu)其(qi)他角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)龍(long)(long)(long)(long)(long)(long)(long)科的(de)(de)(de)(de)(de)(de)(de)(de)頭顱(lu)骨(gu)上(shang)的(de)(de)(de)(de)(de)(de)(de)(de)瘡孔(kong)、洞孔(kong)、損害、以(yi)及(ji)其(qi)他傷(shang)口,常被認為(wei)(wei)是(shi)(shi)(shi)以(yi)頭角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)互相戰斗造成的(de)(de)(de)(de)(de)(de)(de)(de)傷(shang)痕(hen)。一個(ge)(ge)(ge)最近(jin)的(de)(de)(de)(de)(de)(de)(de)(de)研(yan)究則認為(wei)(wei)沒(mei)有(you)(you)(you)證(zheng)(zheng)據顯(xian)示(shi)(shi)(shi)這(zhe)(zhe)些傷(shang)痕(hen)是(shi)(shi)(shi)因為(wei)(wei)打斗而(er)(er)留下的(de)(de)(de)(de)(de)(de)(de)(de),也沒(mei)有(you)(you)(you)感染或(huo)痊愈的(de)(de)(de)(de)(de)(de)(de)(de)證(zheng)(zheng)據。而(er)(er)骨(gu)質流(liu)失(shi)、或(huo)不(bu)(bu)明的(de)(de)(de)(de)(de)(de)(de)(de)骨(gu)頭疾病,是(shi)(shi)(shi)這(zhe)(zhe)些傷(shang)痕(hen)的(de)(de)(de)(de)(de)(de)(de)(de)來(lai)源。三角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)龍(long)(long)(long)(long)(long)(long)(long)的(de)(de)(de)(de)(de)(de)(de)(de)大(da)(da)型(xing)頭盾(dun)(dun)(dun)可能用(yong)來(lai)增(zeng)加身體(ti)(ti)的(de)(de)(de)(de)(de)(de)(de)(de)表面積(ji),以(yi)協(xie)助(zhu)調節(jie)(jie)體(ti)(ti)溫。劍龍(long)(long)(long)(long)(long)(long)(long)的(de)(de)(de)(de)(de)(de)(de)(de)骨(gu)板也被推測擁(yong)(yong)有(you)(you)(you)類(lei)似的(de)(de)(de)(de)(de)(de)(de)(de)功能,但這(zhe)(zhe)個(ge)(ge)(ge)理論(lun)無法(fa)解(jie)釋角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)龍(long)(long)(long)(long)(long)(long)(long)科不(bu)(bu)同(tong)成員的(de)(de)(de)(de)(de)(de)(de)(de)頭盾(dun)(dun)(dun)形狀變化(hua)。頭盾(dun)(dun)(dun)與(yu)(yu)角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)是(shi)(shi)(shi)兩性異形特徵(zhi)的(de)(de)(de)(de)(de)(de)(de)(de)理論(lun),是(shi)(shi)(shi)由Davitashvili在(zai)1961年首(shou)次提出,并(bing)且(qie)逐漸獲(huo)得更多人(ren)(ren)的(de)(de)(de)(de)(de)(de)(de)(de)支(zhi)持。頭盾(dun)(dun)(dun)與(yu)(yu)角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)在(zai)求偶以(yi)及(ji)其(qi)他社會行(xing)為(wei)(wei)上(shang),被視為(wei)(wei)重(zhong)要(yao)的(de)(de)(de)(de)(de)(de)(de)(de)視覺辨認物(wu);這(zhe)(zhe)個(ge)(ge)(ge)理論(lun)可從不(bu)(bu)同(tong)角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)龍(long)(long)(long)(long)(long)(long)(long)類(lei)恐龍(long)(long)(long)(long)(long)(long)(long)擁(yong)(yong)有(you)(you)(you)不(bu)(bu)同(tong)的(de)(de)(de)(de)(de)(de)(de)(de)裝飾物(wu)而(er)(er)得到證(zheng)(zheng)實。而(er)(er)現代(dai)(dai)(dai)現代(dai)(dai)(dai)擁(yong)(yong)有(you)(you)(you)角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)狀物(wu)或(huo)裝飾物(wu)的(de)(de)(de)(de)(de)(de)(de)(de)動物(wu),也將它們(men)作(zuo)為(wei)(wei)視覺辨識物(wu)使(shi)(shi)用(yong)。最近(jin),一對于最小(xiao)型(xing)三角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)龍(long)(long)(long)(long)(long)(long)(long)頭顱(lu)骨(gu)的(de)(de)(de)(de)(de)(de)(de)(de)研(yan)究,確(que)定該(gai)頭顱(lu)骨(gu)屬于一個(ge)(ge)(ge)幼年體(ti)(ti),并(bing)顯(xian)示(shi)(shi)(shi)頭盾(dun)(dun)(dun)與(yu)(yu)角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)是(shi)(shi)(shi)在(zai)年紀非常小(xiao)的(de)(de)(de)(de)(de)(de)(de)(de)時(shi)期開始(shi)發展,早于性發育;因此三角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)龍(long)(long)(long)(long)(long)(long)(long)的(de)(de)(de)(de)(de)(de)(de)(de)頭盾(dun)(dun)(dun)與(yu)(yu)角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)可能作(zuo)為(wei)(wei)視覺辨認物(wu)使(shi)(shi)用(yong)。而(er)(er)該(gai)幼體(ti)(ti)化(hua)石的(de)(de)(de)(de)(de)(de)(de)(de)大(da)(da)型(xing)眼睛與(yu)(yu)較(jiao)短的(de)(de)(de)(de)(de)(de)(de)(de)頭盾(dun)(dun)(dun)與(yu)(yu)角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao),也顯(xian)示(shi)(shi)(shi)三角(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)(jiao)龍(long)(long)(long)(long)(long)(long)(long)的(de)(de)(de)(de)(de)(de)(de)(de)親代(dai)(dai)(dai)具有(you)(you)(you)親代(dai)(dai)(dai)養育的(de)(de)(de)(de)(de)(de)(de)(de)行(xing)為(wei)(wei)。
三角龍祖先
專門收集古動(dong)物遺骸的(de)(de)(de)(de)加拿(na)大(da)(da)化石公司在加拿(na)大(da)(da)艾伯塔省與(yu)美國蒙大(da)(da)拿(na)州交界地(di)帶以南的(de)(de)(de)(de)米爾克(ke)河附(fu)近發(fa)現了一副較為完(wan)整的(de)(de)(de)(de)恐(kong)(kong)龍(long)(long)(long)骨骼化石。加拿(na)大(da)(da)卡爾頓(dun)大(da)(da)學古生(sheng)物學教(jiao)授邁(mai)克(ke)爾·瑞(rui)安(an)等人經過多年的(de)(de)(de)(de)研究,才(cai)把(ba)這(zhe)(zhe)種恐(kong)(kong)龍(long)(long)(long)與(yu)外(wai)表兇猛的(de)(de)(de)(de)食(shi)草恐(kong)(kong)龍(long)(long)(long)家族(zu)的(de)(de)(de)(de)其(qi)他(ta)成員(yuan)區別開來。新(xin)發(fa)現的(de)(de)(de)(de)這(zhe)(zhe)種恐(kong)(kong)龍(long)(long)(long)長7米,重(zhong)約2噸。它生(sheng)活在白堊紀時代(dai),被認(ren)為是三角(jiao)龍(long)(long)(long)的(de)(de)(de)(de)祖先(xian),是角(jiao)龍(long)(long)(long)亞(ya)科恐(kong)(kong)龍(long)(long)(long)在北(bei)美最早的(de)(de)(de)(de)成員(yuan)。引人注目的(de)(de)(de)(de)是,其(qi)頭骨后面有(you)一個巨大(da)(da)的(de)(de)(de)(de)頭盾,邊緣(yuan)有(you)數(shu)個鉤(gou)角(jiao)。瑞(rui)安(an)認(ren)為,這(zhe)(zhe)個器官可能(neng)不(bu)是用來防御掠(lve)食(shi)者(zhe),而更(geng)有(you)可能(neng)是吸引配偶(ou)的(de)(de)(de)(de)炫耀物。
大眾文化
三角(jiao)龍(long)(long)獨特的(de)外(wai)形,使(shi)得它們經常出現(xian)(xian)(xian)在電影(ying)、電腦游戲、以及(ji)(ji)電視節目中(zhong)。在1993年的(de)電影(ying)《侏羅(luo)紀公園》中(zhong),出現(xian)(xian)(xian)了一(yi)只(zhi)因為(wei)不適應現(xian)(xian)(xian)代(dai)植(zhi)被(bei)而生病的(de)三角(jiao)龍(long)(long)。而在1933年的(de)電影(ying)《金剛(gang)之子》(Son of Kong)中(zhong),一(yi)只(zhi)類似三角(jiao)龍(long)(long)的(de)恐龍(long)(long)追(zhui)趕者(zhe)進入(ru)叢林的(de)人類,該只(zhi)三角(jiao)龍(long)(long)模型是由(you)早(zao)期(qi)的(de)電影(ying)特效專家Willis O'Brien所操作。三角(jiao)龍(long)(long)也(ye)出現(xian)(xian)(xian)在三個(ge)以恐龍(long)(long)為(wei)主題的(de)電視節目:《與恐龍(long)(long)同(tong)行》(Walking with the Dinosaurs)、《恐龍(long)(long)兇(xiong)面目》(The Truth About Killer Dinosaurs)以及(ji)(ji)《史前公園》(Prehistoric Park)。
三角龍(long)因為它(ta)們頭部的(de)三根大型(xing)角而著名(ming)。它(ta)們常在(zai)英文中被(bei)昵(ni)稱為「Three-horns」,例如動畫(hua)電(dian)影(ying)《歷(li)險(xian)小恐龍(long)》(The Land Before Time)與(yu)其續集。它(ta)們也常在(zai)兒童讀物、動畫(hua)節目中被(bei)簡稱為「Trike」,例如《哈利與(yu)小恐龍(long)》。
在兒童讀物中,經常出現三(san)角(jiao)龍(long)(long)與暴(bao)龍(long)(long)打斗的場景,因此這兩種恐(kong)龍(long)(long)普遍(bian)被認為是天(tian)敵或是敵手。但在1966年的電影《公元前(qian)一(yi)百萬年》(One Million Years BC)中,三(san)角(jiao)龍(long)(long)的打斗對象,從(cong)暴(bao)龍(long)(long)替換成(cheng)角(jiao)鼻龍(long)(long),但角(jiao)鼻龍(long)(long)與三(san)角(jiao)龍(long)(long)其實生存(cun)于不同的時期。
三(san)(san)角龍也(ye)出現(xian)在《侏羅紀(ji)公(gong)園(yuan)》周邊的電視游(you)戲,或(huo)是(shi)其(qi)他(ta)類似主題(ti)的游(you)戲,例(li)如1997年的電腦(nao)游(you)戲《Jurassic Park: Chaos Island》與(yu)《恐(kong)龍獵(lie)人(ren)》(Turok: Dinosaur Hunter)、以及2000年的電腦(nao)與(yu)Playstation游(you)戲《恐(kong)龍危機》(Dino Crisis 2)。三(san)(san)角龍也(ye)出現(xian)在任(ren)天堂的游(you)戲中(zhong),例(li)如《大金剛賽車》(Diddy Kong Racing)、《星戰(zhan)火狐大冒(mao)險》(Starfox Adventures)。三(san)(san)角龍也(ye)是(shi)南達科他(ta)州的官方州化石,以及懷俄(e)明州的官方州恐(kong)龍。