河北泥河湾"Pliopentalagus nihewanensis"分类地位的再思考

自20世纪60年代,欧亚大陆晚新生代地层中陆续发现了p3由5个主要褶沟组成的兔类化石,并先后建立了上新五褶兔(Pliopentalagus Gureev et Konkova,1964)和三裂齿兔(Tris- chizolagus Radulesco et Samson,1967)两属。近年,在中国安徽淮南大居山新生代晚期不同时代的洞穴、裂隙堆积物中发现了保存相当完整,数量颇多的上新五褶兔的材料。笔者在研究大居山新洞早上新世的上新五褶兔时, 将其与欧亚大陆该属内的已知种和三裂齿兔的各种进行了详细比较,发现产自河北省阳原县泥河湾盆地晚上新世稻地组的Pliopentalagus nihewanensis Cai,1989被指定为上新五褶兔值得商榷。Pliopentalagus nihewanensis的材料仅有1枚p3(GMC V 2008-1)和两枚中间颊齿(p4 或ml,GMC V 2008-2及m2,V 2008-3)。正型标本p3的中前褶沟(AR)中央、后外褶沟和后内褶沟后壁釉质层具有小褶曲,这些形态确实与上新五褶兔的特征有些相似。但褶曲的发育程度明显比上新五褶兔的弱;另外,其齿冠舌侧后缘向后内突出,后外褶沟和后内褶沟较平直并近与齿纵轴垂直,后外褶沟与后内褶沟深度相近等而与上新五褶兔也有所不同。褶沟的釉质层在两枚中间颊齿中较平直,与上新五褶兔显然也有较大的区别。就该p3而言,可以认为它属于一枚釉质构造极简单的上新五褶兔的前臼齿,也可以认为是釉质构造很复杂的三裂齿兔的p3。研究表明,在上新五褶兔和三裂齿兔的系统演化过程中,其颊齿褶沟釉质层的构造有越来越复杂的趋向。作为上新世较晚期的“Pliopentalagus nihewanensis”的p3,指定为上新五褶兔显然有悖于该属釉质构造在地史中越来越复杂的演化事实,而指定为三裂齿兔则正好与该属釉质构造的演化趋势相符(Averianov and Tesakov,1997)。另外,从共生动物群和动物的生活习性看,安徽淮南发现的上新五褶兔与大量现代东洋界特有的类型共生(如Tupaiidae、Platacanthomyidae等),上新五褶兔的后裔——现生的日本琉球奄美黑兔(Pentalagus)的栖息地为温暖、湿润的多山环境;而目前发现的三裂齿兔共生动物群所指示的是干凉的稀树草原环境,与上新五褶兔的生态环境有所不同。含“Pliopentalagus nihewanensis”的晚上新世稻地动物群主要由典型的古北界属种(如Mesosiphneus paratingi、Mimomys orientalis、Germanomys cf. G.weileri、Chardinomys nihewanicus及Ochotona spp.等)组成,而未见典型的东洋界成员。这种温带相对干旱的草原环境,显然与中国发现的上新五褶兔及其后裔的生存环境很不一样。因此,笔者认为“泥河湾上新五褶兔”应归入三裂齿兔属,正名泥河湾三裂齿兔Trischizolagus nihe—wanensis(Cai,1989)。种征订正为:p3具有较复杂的形态构造;中前褶沟具衍生的中央小褶曲, 后外褶沟和后内褶沟后壁釉质层褶曲相对发育。该种以这些基本特征而不同于欧亚大陆的已知种(如T.dumitrescuae、T.mirificus等),它可能代表该属的一个进步类型。

RECONSIDERATION OF THE GENERIC ASSIGNMENT OF "PLIOPENTALAGUS NIHEWANENSIS" FROM THE LATE PLIOCENE OF HEBEI, CHINA

"Pliopentalagus nihewanensis" was described by Cai (1989) , based on an isolated p3 and two isolated lower molariform teeth from the bottom of the "Nihewan beds" of Hebei Province, China. Detailed re-examination of the original specimens and comparisons with other species of the genus and Trischizolagus suggest: 1) the holotype (p3) alone can be interpreted as a species of the genus Pliopentalagus having p3 with least complicated enamel pattern. But, this hypothesis violates an evolutionary trend of the genus that the enamel pattern becomes more complicated through time. On the other hands, 2) the holotype can also be interpreted as a species of the genus Trischizolagus having p3 with most complicated enamel pattern. This hypothesis fits on an evolutionary trend of the genus that enamel pattern becomes more complicated through time ( Averianov and Tesakov, 1997) , 3) the associated fauna which is indicative of a dry and cool environment also suggests that the holotype may not belong to the genus Pliopentalagus. Therefore, we conclude that generic assignment of this taxon should be Trischizolagus, and T. nihewanensis is characterized by p3 with most complicated enamel pattern within the genus.