辽西义县组义县薄氏龙（Bolong yixianesis Wu et al.2010）及原始禽龙类的系统演化
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摘要
义县薄氏龙Bolong yixianensis Wu,Godefroit and Hu,2010是本文作者等最近在辽宁义县头台乡白台沟村义县组发现的原始禽龙类恐龙。其模式标本近完整保存,它是迄今在中国发现的最完整的原始禽龙类化石。
本文首次对义县薄氏龙模式标本的头骨和头后骨骼进行了详细的描述,新发现了义县薄氏龙的4个自近裔性状,并重新修订了义县薄氏龙的鉴定特征,其特征如下:上颌骨与泪骨之间,在相对于Iguanodon、Mantellisaurus和Ouranosaurus眶前孔的位置发育有一凹坑;前额骨后部在眼眶之上发育有一前后延长的凹陷;齿骨背侧的前齿骨关节面高度不足齿骨高度的2/3,齿骨前尖位于齿骨自下向上1/3高处;上颌骨齿齿冠主脊向后弯曲;尺骨和桡骨相对短粗,尺骨与桡骨的近端宽度与骨干长度的比值分别为0.28和0.33;桡骨近端和远端前后几乎对称膨大,外、内侧视呈三角形;肠骨后髋臼突腹背向窄(长/高=2);跖骨非常短,第III跖骨与胫骨长度比为0.18。
运用分支系统学研究方法,本文对禽龙类的32个类群的126个特征进行了系统发育分析。分析结果表明义县薄氏龙为最原始的鸭嘴龙超科类群。鸭嘴龙超科由2个明确的共近裔性状支持:基翼突近水平伸展;肠骨骨板侧视较直。同样产自辽西义县组的锦州龙要比薄氏龙更进步一些。与其它的最近有关禽龙类的系统分析不同的是,本文保留禽龙科,包括Ouranosaurus、Mantellisaurus和Iguanodon,它们构成一单系类群,这一分支由2个明确的共近裔性状支持:骨化的轴上肌腱达第10颈椎;前足第I指的爪骨呈圆锥状。在禽龙科中Mantellisaurus和Ouranosaurus因为背椎和荐椎有较长的神经脊(大于相应椎体高度的2.5倍)而归为同一类群。鸭嘴龙超科与禽龙科互成姊妹群。
根据薄氏龙模式标本椎体的神经脊与神经弓全部愈合、身体长度、前后肢长度比、胫骨与股骨长度比、腕骨不愈合、以及前足第II与第III指的爪骨呈爪状等特征,推测薄氏龙可能为一个成年个体;体重约为0.2吨;可能以两足行走为主。薄氏龙的跖骨及前肢末端很短,这相似于其它用前肢支持身体重量的脊椎动物。因此推测薄氏龙可能与其它一些禽龙类相似也兼用四足行走;与其它的禽龙类相比,薄氏龙的奔跑速度可能较慢。另外,薄氏龙的前肢也可能更加灵活。
迄今发现最早的禽龙类化石产于英国的中侏罗统,亚洲地区在晚侏罗世没有禽龙类的分布。在辽西义县组中发现了德国提塘早期索伦霍芬层中的某些脊椎动物的相应的进步类群,这说明了亚洲大陆与欧洲大陆在晚侏罗世仍相互连接,之后才又相互裂解。而在欧洲晚白垩世发现的禽龙类的原始类群(Rhabdodon和Zalmoxes),可能也正是早白垩世这一阶段欧洲大陆与亚洲大陆之间的地理隔离的结果。原始的鸭嘴龙类起源于亚洲,并且在早白垩世的亚洲逐渐繁盛。在Aptian-Albian期亚洲大陆通过白岭海峡与北美相连,原始的鸭嘴龙类也在这一时期由亚洲迁徙至北美。
辽西义县组的地质时代长期存在较大争议,这主要是因为义县组所产各门类化石分别代表了的不同时代的演化特征。根据本文禽龙类系统发育的分析,薄氏龙可能生存于早白垩世Valanginian-Aptian期,综合参考原始禽龙类的古生物地理及义县组大康堡层生物群的总体特征,本文将薄氏龙的生存时代定为早白垩世中期,或略晚一些。
总之,义县薄氏龙的发现和研究对原始禽龙类及原始鸭嘴龙类的起源、演化、系统发育及古生物地理等方的研究都具有较重要意义。
Bolong yixianensis Wu, Godefroit and Hu, 2010 is a basal iguanodontian dinosaur, which was recently discovered from the Yixian Formation of western Liaoning, China. The holotype of Bolong yixianensis is virtually complete. It is the most complete known basal iguanodontian specimen found in China.
The synthetic biological age of the Yixianian Formation is determined as the Late Jurassic to the Early Cretaceous. And the Jurassic-Cretaceous boundary should be between the Jianshangou Bed and the Dakangpu Bed. Most of the isotopic ages of the volcanic rocks which have been obtained in recent years are different from the synthetic biological age of the Yixian Formation. However this problem could not be resolved in a short time.
The first detailed description of the holotype specimen of Bolong yixianensis is presented. And four autapomorphies of Bolong yixianensis are newly recognized from the postcranial skeleton. The revised diagnosis of Bolong yixianensis includes the following characters: depressed area at the junction between maxilla and lachrymal, corresponding to the emplacement of the antorbital fenestra in Iguanodon, Mantellisaurus and Ouranosaurus; caudal ramus of prefrontal forming a rostrocaudally depressed area above the orbital margin; rostrodorsal articular surface for the predentary occupying less than two-thirds of the height of the dentary and rostral tip of dentary therefore situated above the ventral third of the bone; primary ridge deflected distally on maxillary crowns; ulna and radius proportionally short and robust: ratio‘craniocaudal height of proximal part/length’= 0.28 for ulna and 0.33 for radius; proximal and distal ends of radius nearly symmetrically enlarged craniocaudally and triangular in medial and lateral views; postacetabular process of ilium dorsoventrally narrow (ratio‘length/maximal height’= 2); metatarsals are very short, ratio‘metatarsal III length/tibia length’= 0.18.
A phylogenetic analysis was carried out. One hundred and twenty six characters of 26 taxa of Iguanodontia are employed in the phylogenetic analysis. This analysis shows that Bolong is the most basal Hadrosauroidea; Two synapomorphies supporting the Hadrosauroidea: basipterygoid process extending nearly horizontally; ischium shaft nearly straight. Jinzhousaurus, also from the Yixian Formation of western Liaoning Province, is more advanced than Bolong. Contrary to other recent phylogenetic analyses here retain a monophyletic Iguanodontidae family, including Iguanodon, Mantellisaurus and Ouranosaurus. This clade is characterized by the following autapomorphies: the ossified epaxial tendons extend cranially from cervical 10; and the pollex is spike-like and conical. Within Iguanodontidae, Mantellisaurus and Ouranosaurus are united by the presence of elongated neural spines on the posterior dorsals and sacrals, more than 2.5 times centrum height. Iguanodontidae is the sister group of Hadrosauroidea.
The complete closure of the sutures between the neural arches and centra throughout the vertebral column, the body length, the ratio between the forelimb length and hindlimb length, the ratio between the tibia length and femur length, the carpals were not coossified, the unguals on manus digits II and III are claw-shaped, and the other characters of Bolong suggest that the holotype specimen of Bolong is an adult individual when it died, its body mass is about 0.2 tonne, and Bolong spent considerable periods of time walking or running bipedally. However, the metatarsals and the distal portion of the forelimbs are particularly short in Bolong, like other vertebrates using their forelimbs for weight supporting. So, Bolong might have been a facultive quadruped, as in many other iguanodontians, and it probably has an inferior running ability compared to other iguanodonitans. Additionally, the forelimbs of Bolong are probably more agile.
The earliest basal Iguanodontia is known from the Middle Jurassic in England. Iguanodontia are absent from Late Jurassic localities in Asia. Some vertebrates which were discovered from Late Jurassic Solnhofen Bed in Germany also have their phylogenetic affinites in Yixian Formation. This indecates that Asian and European continents were connected in the Late Juarassic, and the primitive iguanodontians (Rhabdodon and Zalmoxes) lived in Europe during the Late Cretaceous, perhaps just caused by the vicariant event which separated Asian and European continents in the Early Cretaceous. The Hadrosauroidea was origined from Asia, and they immigrated to North America during the Late Early Cretaceous.
There's a lot of debate on the age of Yixian Formation. The reason is different taxa of Yixian Biota displays different ages. Although based on the current phylogenetic analysis, the geological age of Bolong is presumbly between Valanginian and Aptian. Here the geological age of Bolong is determined as the middle Early Cretaceous or later, based on the paleobiogeographical distribution of basal Iguanodontia and the synthetic biological age of the Dakangup Bed of Yixian Formation.
In sum, the discovery and study of Bolong have great value on the study of the origin, evolution, phylogeny and paleobiogeography of basal Iguanodontia and basal Hadrosauroidea.
本文首次对义县薄氏龙模式标本的头骨和头后骨骼进行了详细的描述,新发现了义县薄氏龙的4个自近裔性状,并重新修订了义县薄氏龙的鉴定特征,其特征如下:上颌骨与泪骨之间,在相对于Iguanodon、Mantellisaurus和Ouranosaurus眶前孔的位置发育有一凹坑;前额骨后部在眼眶之上发育有一前后延长的凹陷;齿骨背侧的前齿骨关节面高度不足齿骨高度的2/3,齿骨前尖位于齿骨自下向上1/3高处;上颌骨齿齿冠主脊向后弯曲;尺骨和桡骨相对短粗,尺骨与桡骨的近端宽度与骨干长度的比值分别为0.28和0.33;桡骨近端和远端前后几乎对称膨大,外、内侧视呈三角形;肠骨后髋臼突腹背向窄(长/高=2);跖骨非常短,第III跖骨与胫骨长度比为0.18。
运用分支系统学研究方法,本文对禽龙类的32个类群的126个特征进行了系统发育分析。分析结果表明义县薄氏龙为最原始的鸭嘴龙超科类群。鸭嘴龙超科由2个明确的共近裔性状支持:基翼突近水平伸展;肠骨骨板侧视较直。同样产自辽西义县组的锦州龙要比薄氏龙更进步一些。与其它的最近有关禽龙类的系统分析不同的是,本文保留禽龙科,包括Ouranosaurus、Mantellisaurus和Iguanodon,它们构成一单系类群,这一分支由2个明确的共近裔性状支持:骨化的轴上肌腱达第10颈椎;前足第I指的爪骨呈圆锥状。在禽龙科中Mantellisaurus和Ouranosaurus因为背椎和荐椎有较长的神经脊(大于相应椎体高度的2.5倍)而归为同一类群。鸭嘴龙超科与禽龙科互成姊妹群。
根据薄氏龙模式标本椎体的神经脊与神经弓全部愈合、身体长度、前后肢长度比、胫骨与股骨长度比、腕骨不愈合、以及前足第II与第III指的爪骨呈爪状等特征,推测薄氏龙可能为一个成年个体;体重约为0.2吨;可能以两足行走为主。薄氏龙的跖骨及前肢末端很短,这相似于其它用前肢支持身体重量的脊椎动物。因此推测薄氏龙可能与其它一些禽龙类相似也兼用四足行走;与其它的禽龙类相比,薄氏龙的奔跑速度可能较慢。另外,薄氏龙的前肢也可能更加灵活。
迄今发现最早的禽龙类化石产于英国的中侏罗统,亚洲地区在晚侏罗世没有禽龙类的分布。在辽西义县组中发现了德国提塘早期索伦霍芬层中的某些脊椎动物的相应的进步类群,这说明了亚洲大陆与欧洲大陆在晚侏罗世仍相互连接,之后才又相互裂解。而在欧洲晚白垩世发现的禽龙类的原始类群(Rhabdodon和Zalmoxes),可能也正是早白垩世这一阶段欧洲大陆与亚洲大陆之间的地理隔离的结果。原始的鸭嘴龙类起源于亚洲,并且在早白垩世的亚洲逐渐繁盛。在Aptian-Albian期亚洲大陆通过白岭海峡与北美相连,原始的鸭嘴龙类也在这一时期由亚洲迁徙至北美。
辽西义县组的地质时代长期存在较大争议,这主要是因为义县组所产各门类化石分别代表了的不同时代的演化特征。根据本文禽龙类系统发育的分析,薄氏龙可能生存于早白垩世Valanginian-Aptian期,综合参考原始禽龙类的古生物地理及义县组大康堡层生物群的总体特征,本文将薄氏龙的生存时代定为早白垩世中期,或略晚一些。
总之,义县薄氏龙的发现和研究对原始禽龙类及原始鸭嘴龙类的起源、演化、系统发育及古生物地理等方的研究都具有较重要意义。
Bolong yixianensis Wu, Godefroit and Hu, 2010 is a basal iguanodontian dinosaur, which was recently discovered from the Yixian Formation of western Liaoning, China. The holotype of Bolong yixianensis is virtually complete. It is the most complete known basal iguanodontian specimen found in China.
The synthetic biological age of the Yixianian Formation is determined as the Late Jurassic to the Early Cretaceous. And the Jurassic-Cretaceous boundary should be between the Jianshangou Bed and the Dakangpu Bed. Most of the isotopic ages of the volcanic rocks which have been obtained in recent years are different from the synthetic biological age of the Yixian Formation. However this problem could not be resolved in a short time.
The first detailed description of the holotype specimen of Bolong yixianensis is presented. And four autapomorphies of Bolong yixianensis are newly recognized from the postcranial skeleton. The revised diagnosis of Bolong yixianensis includes the following characters: depressed area at the junction between maxilla and lachrymal, corresponding to the emplacement of the antorbital fenestra in Iguanodon, Mantellisaurus and Ouranosaurus; caudal ramus of prefrontal forming a rostrocaudally depressed area above the orbital margin; rostrodorsal articular surface for the predentary occupying less than two-thirds of the height of the dentary and rostral tip of dentary therefore situated above the ventral third of the bone; primary ridge deflected distally on maxillary crowns; ulna and radius proportionally short and robust: ratio‘craniocaudal height of proximal part/length’= 0.28 for ulna and 0.33 for radius; proximal and distal ends of radius nearly symmetrically enlarged craniocaudally and triangular in medial and lateral views; postacetabular process of ilium dorsoventrally narrow (ratio‘length/maximal height’= 2); metatarsals are very short, ratio‘metatarsal III length/tibia length’= 0.18.
A phylogenetic analysis was carried out. One hundred and twenty six characters of 26 taxa of Iguanodontia are employed in the phylogenetic analysis. This analysis shows that Bolong is the most basal Hadrosauroidea; Two synapomorphies supporting the Hadrosauroidea: basipterygoid process extending nearly horizontally; ischium shaft nearly straight. Jinzhousaurus, also from the Yixian Formation of western Liaoning Province, is more advanced than Bolong. Contrary to other recent phylogenetic analyses here retain a monophyletic Iguanodontidae family, including Iguanodon, Mantellisaurus and Ouranosaurus. This clade is characterized by the following autapomorphies: the ossified epaxial tendons extend cranially from cervical 10; and the pollex is spike-like and conical. Within Iguanodontidae, Mantellisaurus and Ouranosaurus are united by the presence of elongated neural spines on the posterior dorsals and sacrals, more than 2.5 times centrum height. Iguanodontidae is the sister group of Hadrosauroidea.
The complete closure of the sutures between the neural arches and centra throughout the vertebral column, the body length, the ratio between the forelimb length and hindlimb length, the ratio between the tibia length and femur length, the carpals were not coossified, the unguals on manus digits II and III are claw-shaped, and the other characters of Bolong suggest that the holotype specimen of Bolong is an adult individual when it died, its body mass is about 0.2 tonne, and Bolong spent considerable periods of time walking or running bipedally. However, the metatarsals and the distal portion of the forelimbs are particularly short in Bolong, like other vertebrates using their forelimbs for weight supporting. So, Bolong might have been a facultive quadruped, as in many other iguanodontians, and it probably has an inferior running ability compared to other iguanodonitans. Additionally, the forelimbs of Bolong are probably more agile.
The earliest basal Iguanodontia is known from the Middle Jurassic in England. Iguanodontia are absent from Late Jurassic localities in Asia. Some vertebrates which were discovered from Late Jurassic Solnhofen Bed in Germany also have their phylogenetic affinites in Yixian Formation. This indecates that Asian and European continents were connected in the Late Juarassic, and the primitive iguanodontians (Rhabdodon and Zalmoxes) lived in Europe during the Late Cretaceous, perhaps just caused by the vicariant event which separated Asian and European continents in the Early Cretaceous. The Hadrosauroidea was origined from Asia, and they immigrated to North America during the Late Early Cretaceous.
There's a lot of debate on the age of Yixian Formation. The reason is different taxa of Yixian Biota displays different ages. Although based on the current phylogenetic analysis, the geological age of Bolong is presumbly between Valanginian and Aptian. Here the geological age of Bolong is determined as the middle Early Cretaceous or later, based on the paleobiogeographical distribution of basal Iguanodontia and the synthetic biological age of the Dakangup Bed of Yixian Formation.
In sum, the discovery and study of Bolong have great value on the study of the origin, evolution, phylogeny and paleobiogeography of basal Iguanodontia and basal Hadrosauroidea.
引文
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