中国子午沙鼠形态地理变异及亚种分类研究
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摘要
子午沙鼠(Meriones meridianus Pallas,1773)(英文名:midday gerbil)隶属于啮齿目(Rodentia),仓鼠科(Cricetidae),沙鼠属(Meriones)。国外主要分布于蒙古、哈萨克斯坦、塔吉克斯坦、伊朗和阿富汗等中亚国家,向西一直可达里海(Caspian Sea)沿岸地区。国内主要分布于内蒙古锡林郭勒的浑善达克沙地以西,河北西北部,山西中北部,河南西北部,陕西秦岭以北,甘肃和宁夏全境,青海东北部和柴达木盆地,以及新疆的全境。垂直分布可从海拔-150m的吐鲁番盆地,上升到昆仑山脉北坡海拔3200m的高山荒漠地区。子午沙鼠由于分布很广,目前为止全世界描述过的亚种共有10多个。据文献记载,子午沙鼠在国内共有7个亚种,分别为:(1)内蒙古亚种(M. m. psammophilus Miline-Edwards,1871).(2)阿勒泰亚种(M. m. buechneri Thomas,1909).(3)木垒亚种(M.m. muleiensis Wang,1981)、(4)伊犁亚种(M. m. penicilliger Heptner,1933)、(5)叶氏亚种(M.m.jei Wang,1964)、(6)叶城亚种(M.m. cryptorhinus Blanford,1875).(7)塔里木亚种(M. m. lepturus Buchner,1889)。
     为了弄清楚中国子午沙鼠的形态地理变异规律,本文以2008年7月~2010年10月采自内蒙古、宁夏、甘肃、青海和新疆5省区25个采样点的339只成年子午沙鼠为材料,通过偏相关分析以及多元线性回归分析分析了子午沙鼠的体重(BM)、体长(BL)、尾长(TL)、耳长(EL)、后足长(HFL)、相对尾长(RTL)、相对耳长(REL)、相对后足长(RHFL)8个身体量衡度,颅全长(GLS)、后头宽(MB)、鼻骨长(GLN)、相对后头宽(RMB)、相对鼻骨长(RGLN).颅体积(SS)、听泡体积(SAB)、相对听泡体积(RSAB)8个头骨量度,心脏干重(HE)、肺脏干重(LU)、肝脏干重(LI)、肾脏干重(KI)、脾脏干重(SP)、相对心脏干重(RHE)、相对肺脏干重(RLU)、相对肝脏干重(RLI)、相对肾脏干重(RKI)、相对脾脏干重(RSP)10个内脏衡度与纬度(Latitude)和海拔(Altitude)2个地理因子以及年均温(TEM)、无霜期(FFP)、相对湿度(HUM)、降雨量(PRC)、日照时数(SUN)和年均风速(WIN)6个气候因子之间的变异规律,得出以下主要结论:
     偏相关分析分析了子午沙鼠的8个身体量衡度、8个头骨量度和10个内脏衡度随2个地理因子的变异规律,结果显示:(1)对于子午沙鼠的身体量衡度来说,耳长、尾长、后足长、相对耳长和相对尾长与纬度显著负相关;体重、体长和后足长与海拔显著正相关,尾长、相对耳长和相对尾长与海拔显著负相关;(2)对于子午沙鼠的头骨量度来说,相对后头宽与纬度显著正相关,鼻骨长、相对鼻骨长、听泡体积和相对听泡体积与纬度显著负相关;颅全长、鼻骨长、相对后头宽、相对鼻骨长、颅体积、听泡体积和相对听泡体积与海拔显著正相关;(3)对于子午沙鼠的内脏衡度来说,脾脏干重、相对肝脏干重和相对脾脏干重与纬度显著正相关,肺脏干重和相对肺脏干重与纬度显著负相关;心脏干重、肺脏干重、肝脏干重、肾脏于重、脾脏干重(雄性)和相对肝脏干重与海拔显著正相关。
     多元线性回归分析分析了子午沙鼠的8个身体量衡度、8个头骨量度和10个内脏衡度随6个气候因子的变异规律,结果显示:(1)对于子午沙鼠的身体量衡度来说,体重、体长和相对后足长的变化主要是与无霜期的变化有关;耳长和相对耳长的变化主要是与同照时数的变化有关;尾长和相对尾长的变化主要是与年均风速的变化有关;后足长的变化主要是与相对湿度的变化有关;(2)对于子午沙鼠的头骨量度来说,颅全长、鼻骨长、相对后头宽、相对鼻骨长、听泡体积和相对听泡体积的变化主要是与无霜期的变化有关;后头宽的变化主要是与降雨量的变化有关;颅体积的变化主要是与相对湿度的变化有关;(3)对于子午沙鼠的内脏衡度来说,心脏干重(雌性)、脾脏干重(雄性)和相对心脏干重(雌性)的变化主要是与年均温的变化有关;心脏干重(雄性)、肺脏干重、肝脏干重、肾脏干重和相对肺脏干重的变化主要是与无霜期的变化有关;相对心脏干重(雄性)的变化主要是与降雨量的变化有关;相对肾脏干重的变化主要是与日照时数的变化有关;相对脾脏干重(雌性)的变化主要是与相对湿度的变化有关。
     本文还以大沙鼠为外群,以25个采样点子午沙鼠的体重(BM)、体长(BL)、尾长(TL)、耳长(EL)、后足长(HFL)、相对尾长(RTL)、相对耳长(REL)、相对后足长(RHFL)8个身体量衡度和颅全长(GLS)、颅基长(BL)、鼻骨长(GLN)、腭长(MPL)、齿隙(DL)、颧宽(ZB)、眶间宽(OB)、后头宽(MB)、听泡长(LAB)、听泡宽(WAB)、上颊齿列长(LUCR)、下颊齿列长(LLCR)、相对颅基长(RBL)、相对鼻骨长(RGLN)、相对腭长(RMPL)、相对齿隙(RDL)、相对颧宽(RZB)、相对眶间宽(ROB)、相对后头宽(RMB)、相对听泡长(RLAB)、相对听泡宽(RWAB)、相对上颊齿列长(RLUCR)、相对下颊齿列长(RLLCR)23个头骨量度共31个变量的平均值进行了系统聚类分析,并结合以前分子生物学方面的文献资料对国内分布的7个子午沙鼠亚种进行了分类讨论,得出以下主要结论:
     基于8个身体量衡度和23个头骨量度共31个形态变量的系统聚类分析树状图以及Cytb基因序列构建的NJ系统树和MP系统树的结果显示:(1)子午沙鼠叶城亚种(M.m.cryptorhinus)和子午沙鼠塔里木亚种(M. m.lepturus)均不成立,是内蒙古亚种(M.m. psammophilus)的同物异名,应归入子午沙鼠内蒙古亚种之内,其主要特征是:体型较大,平均重约60g左右,体长约为120mm。尾长较短,约占体长的90%。毛色深棕带红,几乎不显沙灰色调。尾毛长且蓬松,尾端毛束黑色或较淡。国内主要分布于内蒙古浑善达克沙地以西,经河北西北部,山西中部和北部,陕西秦岭以北,甘肃,宁夏全境,青海省东北部和柴达木盆地,新疆东部,直到新疆南部的塔里木盆地西缘;(2)子午沙鼠木垒亚种(M.m. muleiensis)和子午沙鼠叶氏亚种(M.m.jei)也不成立,是伊犁亚种(M.m. penicilliger)的同物异名,应归入子午沙鼠伊犁亚种之内,其主要特征是:体型较小,平均重约45g左右,体长约为110mm。尾长较长,约占体长的100%。毛色沙黄带灰,呈现较浓的沙灰色调。尾毛短且稀疏,尾端毛束黑色或较浓。国内分布于天山南麓的吐鲁番盆地,天山北麓的古尔班通古特沙漠,往西经艾比湖附近的乌苏沙漠,直至伊犁河谷地;(3)子午沙鼠阿勒泰亚种(M. m. buechneri)由于没有获得足够的标本进行形态变量的系统聚类分析,故暂时无法讨论其亚种的分类地位是否成立,但分子生物学方面的证据目前来看是支持的。
Midday gerbils(Meriones meridianus Pallas,1773) belong to the Rodentia, Cricetidae, Meriones. They are common in sandy areas in Mongolia, Kazakstan, Iran and Afghanistan, and can be found to the west as far as the Caspian Sea coast. In China midday gerbils are mainly distributed in midwest of Inner Mongolia, north of Shanxi and Shanxi, Ningxia, Gansu, Qinghai and Xinjiang province. They can be found from150meters below the sea level to3200meters above sea level. As a highly adaptable species, more than10subspecies so far are recorded. According to the literatures, there are7subspecies in China. They are (1) M. m. psammophilus Miline-Edwards,1871;(2) M. m. buechneri Thomas,1909;(3) M. m. muleiensis Wang,1981;(4) M. m. penicilliger Heptner,1933;(5) M. m.jei Wang,1964;(6) M. m. cryptorhinus Blanford,1875;(7) M. m. lepturus Buchner,1889, respectively.
     In order to find out the rule of geographic variation on morphological characters of midday gerbils in China,339adult individuals of M. meridianus were measured, which were trapped by the rattrap at25sites from July of2008to October of2010in Inner Mongolia, Ningxia, Gansu, Qinghai and Xinjiang province, northwest of China. Partial correlation analysis and multiple linear regression analysis were used to analyze the variation between midday gerbil's body mass (BM), body length (BL), tail length (TL), ear length (EL), hind foot length (HFL), relative tail length (RTL), relative ear length (REL), relative hind foot length (RHFL)8body mass and measurements, greatest length of skull (GLS), mastoid breadth (MB), greatest length of the nasals (GLN), relative mastoid breadth (RMB), relative greatest length of the nasals (RGLN), skull size (SS), size of auditory bulla (SAB), relative size of auditory bulla (RSAB)8skull measurements and heart dry mass (HE), lung dry mass (LU), liver dry mass (LI), kidneys dry mass (KI), spleen dry mass (SP), relative heart dry mass (RHE), relative lung dry mass (RLU), relative liver dry mass (RLI), relative kidneys dry mass (RKI), relative spleen dry mass (RSP)10organ masses with latitude and altitude2geographical factors, and annual mean temperature (TEM), frost-free period (FFP), relative humidity (HUM), precipitation (PRC), annual sunshine hours (SUN), annual average wind speed (WIN)6climatic factors. This study got the following conclusion:
     Partial correlation analysis were used to analyze the relationship between midday gerbil's8body mass and measurements,8skull measurements and10organ masses with2geographical factors. The results showed that:(1) For the midday gerbil's8body mass and measurements, ear length, tail length, hind foot length, relative ear length and relative tail length were significantly negatively correlated with latitude, body mass, body length and hind foot length were significantly positively correlated with altitude, tail length, relative ear length and relative tail length were significantly negatively correlated with altitude;(2) For the midday gerbil's8skull measurements, relative mastoid breadth was significantly positively correlated with latitude, greatest length of the nasals, relative greatest length of the nasals, size of auditory bulla and relative size of auditory bulla were significantly negatively correlated with latitude, greatest length of skull, greatest length of the nasals, relative mastoid breadth, relative greatest length of the nasals, skull size, size of auditory bulla and relative size of auditory bulla were significantly positively correlated with altitude;(3) For the midday gerbil's10organ masses, spleen dry mass, relative liver dry mass, and relative spleen dry mass were significantly positively correlated with latitude, lung dry mass and relative lung dry mass were significantly negatively correlated with latitude; heart dry mass, lung dry mass, liver dry mass, kidneys dry mass, spleen dry mass (male) and relative liver dry mass were significantly positively correlated with altitude.
     Multiple linear regression analysis were used to analyze the relationship between midday gerbil's8body mass and measurements,8skull measurements and10organ masses with6climatic factors. The results showed that:(1) For the midday gerbil's8body mass and measurements, the change of body mass, body length and relative hind foot length were mainly caused by the change of frost-free period, the change of ear length and relative ear length were mainly caused by the change of annual sunshine hours, the change of tail length and relative tail length were mainly caused by the change of annual average wind speed, the change of hind foot length was mainly caused by the change of relative humidity;(2) For the midday gerbil's8skull measurements, the change of greatest length of skull, greatest length of the nasals, relative mastoid breadth, relative greatest length of the nasals, size of auditory bulla and relative size of auditory bulla were mainly caused by the change of frost-free period, the change of mastoid breadth was mainly caused by the change of precipitation; the change of skull size was mainly caused by the change of relative humidity;(3) For the midday gerbil's10organ masses, the change of heart dry mass (female), spleen dry mass (male) and relative heart dry mass (female) were mainly caused by the change of annual mean temperature, the change of heart dry mass (male), lung dry mass, liver dry mass, kidneys dry mass and relative lung dry mass were mainly caused by the change of frost-free period, the change of relative heart dry mass (male) was mainly caused by the change of precipitation, the change of relative kidneys dry mass was mainly caused by the change of annual sunshine hours, the change of relative spleen dry mass (female) was mainly caused by the change of relative humidity.
     In this study, hierarchical cluster analysis were used to cluster31variable mean measurements of midday gerbil from the25sites, including8body mass and measurements and23skull measurements, and great gerbil (Rhombomys opimus) as outer cluster. At the same time, some literatures of molecular biology of midday gerbils were reviewed, and we discussed taxonomy of7subspecies in China.
     We banded together the Phylogenetic tree analyses (NJ method and MP method) composed of mitochondrial cytochrome-6gene. The results demonstrated that:(1) M. m. cryptorhinus and M. m. lepturus are all belong to M. m. psammophilus. Diagnosis: they are bigger, with the body weight averaging60g, and the length of head and body about120mm. Tail is slightly shorter than the head and body length, averaging90%of the latter. Upper surface with hairs is thus of a dark brown hue, almost not showing gray in colour. Tail is thickly and fluffy, and the tail tuft black or light. Distribution: midwest of Inner Mongolia, north of Shanxi and Shaanxi, Ningxia, Gansu, Qinghai, east and south of Xinjiang;(2) M. m. muleiensis and M. m. jei are all belong to M. m. penicilliger. Diagnosis:smaller, with the body weight averaging45g, and the length of head and body about110mm. Tail is almost as same long as the head and body length, averaging100%of the latter. Upper surface with hairs being thus of a grayish-buff hue, almost show dark gray in colour. Tail thinly and closely, the tail tuft is black or dark. Distribution:Tianshan Mounts areas of Xinjiang, including Turpan Basin, Gurbantonggut Desert, Usu Desert and Yili River Valley;(3) As there were not enough specimens to carry on hierarchical cluster analysis, we were not sure of the subspecies classification of M. m. buechneri, but the evidence of molecular biology supported this conclusion.
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