隆肛蛙属Feirana种群的系统学与动物地理学研究

对隆肛蛙属的物种构成进行了订正，建立新属肛刺蛙属Yerana gen. nov.；订正后的隆肛蛙属现仅隶2种， 即隆肛蛙F. quadrana和太行隆肛蛙F. taihangnicus。运用形态学分析探讨了隆肛蛙属物种及种群的形态差异和分类关系，通过分子系统学研究探讨了隆肛蛙属物种及种群的分类和系统发育关系，运用动物地理学方法结合系统发育关系探讨了隆肛蛙属种群的地理分布格局成因与历史过程。主要结果和推论如下： 1．隆肛蛙属物种构成的订正及一新属建立 建立新属肛刺蛙属，将隆肛蛙属中的原叶氏隆肛蛙F. yei归隶新属肛刺蛙属并更名为叶氏肛刺蛙Y. yei，，新属建立的主要依据为：（1）雄性肛部隆起，肛孔下方有两个布满黑刺的大的白色球形隆起，具单咽下内声囊， 第一指具婚刺；（2）形态量度分析表明叶氏肛刺蛙与隆肛蛙和太行隆肛蛙的形态差异远大于后两者之间的差异；（3）叶氏肛刺蛙的分布区与隆肛蛙和太行隆肛蛙的分布区距离较远且呈隔离状态；（4）分子系统学研究资料（Jiang et al.,2005）证明叶氏肛刺蛙与隆肛蛙和太行隆肛蛙非单系发生；叶氏肛刺蛙在第二支中位于基部。因此，隆肛蛙属现仅隶2种，即隆肛蛙和太行隆肛蛙。 2．隆肛蛙属种群形态学研究 对隆肛蛙属中隆肛蛙和太行隆肛蛙的15个地理种群565只标本的28项形态性状进行了测量，运用典型判别分析法对其分析的结果表明：（1）太行隆肛蛙与隆肛蛙形态差异明显，支持其为不同的物种；（2）原隆肛蛙河南伏牛山种群和山西中条山种群应为太行隆肛蛙的地理种群；（3）隆肛蛙不同地理种群之间形态差异明显，其中四川安县种群、陕西周至种群和湖北利川种群与模式产地重庆巫山种群的差异可能达到了亚种或亚种以上分化水平。对隆肛蛙属量度分析的15个种群进行定性形态分析表明其分为三种形态型，对应隆肛蛙、过渡型和太行隆肛蛙，其变异特征主要为内跗褶、雄性肛部隆起及疣粒分布、第五趾外侧缘膜等，这与量度分析结果相似。 3．隆肛蛙属种群分子系统学研究 测定隆肛蛙属Feirana的2种19种群的线粒体12S rRNA和16S rRNA基因片段、ND2基因的DNA序列，比对后共计1953bps。（1）遗传多样性与距离分析：结果表明，隆肛蛙属种群具很高的遗传多样性，19个种群样品表现出19种单倍型（遗传多样性指数Hd＝1.0）； ND2基因的进化信息含量远高于12SrRNA和16SrRNA。隆肛蛙属2种群组内的种群间的遗传距离远小于两种群组间的距离，种群在不同基因上的遗传距离表现的关系与对应的系统树一致。（2）系统发育关系分析：结果表明，不同基因片断基于不同方法构建的隆肛蛙属种群系统发育树结构基本一致，基本表明隆肛蛙属种群为单系发生；它们在系统树中分为两大支，分别对应于隆肛蛙和太行隆肛蛙；支持中条山种群（沁水、历山和济源种群）和伏牛山种群（栾川和内乡种群）为太行隆肛蛙的地理种群，而原隆肛蛙秦岭中东段的部分种群（柞水、宁陕、长安大坝沟种群）也应为太行隆肛蛙的地理种群。（3）亚种分化分析：根据遗传距离分析和系统发育关系分析结果，并考虑形态上的差异情况以及地理分布信息，隆肛蛙所隶种群组可分为2亚种，即隆肛蛙指名亚种F. quadrana quadrana包括四川盆地东缘大巴山东段-巫山-武陵山北麓种群和秦岭中段（周至板房子和长安广货街）种群，他们在系统关系树上聚为一支；安县亚种F. quadrana anxianensis包括四川盆地西缘岷山东麓-龙门山-大巴山和秦岭西段的种群（安县、青川、文县、南江和凤县种群），他们在系统关系树上聚为一支。太行隆肛蛙所隶种群组也可分为2亚种，即太行隆肛蛙指名亚种F. taihangnicus taihangnicus包括中条山的种群（沁水、历山和济源种群）和中东秦岭的部分种群（柞水、长安大坝沟和宁陕种群），他们在系统关系树上聚为一支；太行隆肛蛙伏牛亚种F. taihangnicus funiuensis，为伏牛山地区的种群（栾川和内乡种群），他们在系统关系树上聚为一支。 4．隆肛蛙属种群动物地理学研究 隆肛蛙属19种群的分歧年代分析： 以长江巫山段和黄河三门峡段的形成历史时期为参考点，根据已测隆肛蛙属19种群及其外群包括N. pleski、P. yunnanesis、P. robertingeri、F. limnocharis的1953bps DNA序列构建分子钟，获得各支系的分歧年代。结果表明：①棘蛙族在70Ma左右开始其独立演化历程，这与Roelants et al.（2004）的分析结果～60±15Ma左右开始分化基本一致，后者印证了本文的分子钟。②隆肛蛙属的起始分化年代较早，隆肛蛙和太行隆肛蛙两种群组的最近祖先种群大概在46Ma～50Ma左右；隆肛蛙和太行隆肛蛙种群组内的种群分化年代相对两种群组间晚得多， 隆肛蛙种群组内两亚种分化起始年代约为10Ma左右，而太行隆肛蛙种群组内两亚种分化起始年代约为6Ma。 隆肛蛙属种群分布格局形成过程分析： ①隆肛蛙属的系统关系与地理分布格局密切相关，大部分系统分支分级与地理距离成正比；②隆肛蛙属最近祖先种群的分化中心可能位于秦岭中部地区， 隆肛蛙属的种群分布格局的形成表现为隔离分化与扩散相结合的机制，由隔离分化产生的隆肛蛙祖先种群主要从秦岭中部向西南方向扩散，后隔离分化为两亚种；太行隆肛蛙祖先种群向东北方向扩散也分化为两亚种。 隆肛蛙属种群分布区域地质历史的探讨：本文所建分子钟和种群分化方式印证了该区域的几次主要地质事件，包括岷山-龙门山-西秦岭等地区的快速差异隆起、第四纪冰期等。 The specific composition of the genus Feirana should be revised. A new genus Yerana gen. nov.（Ranidae：Dicroglossinae）was established based on morphological data-set and molecular phylogeny, as a result, only two species F. quadrana and F. taihangnicus are classified into Feirana now. Morphological differences and taxonomy of populations of Feirana were investigated based on morphological and morphometric data; phylogenetic relationships and taxonomy of populations of Feirana were elucidated using molecular data, and then the proceeding of the distribution pattern of populations of Feirana were discussed. The main results and conclusions and proposals were presented as following: 1. Revising of the specific composition of the genus Feirana and establishment of a new genus The new genus Yerana, only containing the type species Y. yei, was established based on the following evidences: (1) In adult male, distinct up-heaved circular vesicle presents around the anal, and under anal there are two white balls on which black spines exist, black horny spines scatter on the upper side of first finger, and internal single subgular vocal sac presents; (2) there is obvious morphometric differences between Yerana and Feirana; (3) Yerana is distributed far from Feirana; (4) evidences of molecular phylogeny（Jiang et al.,2005）suggested that Yerana take a special phylogenetic clade which is different from other genus included in the tribe Paini. As a result, there are only two species in Feirana, i.e., F. quadrana and F. taihangnicus. 2. Morphological research of populations of Feirana Twenty-eight characters of 565 individuals of 15 populations of the genus Feirana were measured, the results of Canonical Discriminant analysis of the morphometric data-set indicated that: (1) there are very prominent differences between the two species F. quadrana and F. taihangnicus. The validity of species F. taihangnicus was approved here; (2) Mt. Funiu population and Mt. Zhongtiao population should belong to the species F. taihangnicus; (3) Obvious differences exist among 12 populations of F. quadrana, the differentiation among Zhouzhi population, Anxian population, Lichuan population, and Wushan population together with the others probably reach sub-specific or specific level. Result of morphological comparison between 15 different populations show that 3 morphological types are recogenized in according with F. quadrana, F. taihangnicus and intergradation, this result conform to the result of morphometric analysis. 3. Molecular phylogenetic study on populaions of Feirana Fragment of 12SrRNA and 16SrRNA genes, and ND2 gene of 19 populations of two species of Feirana were sequenced and aligned, from which 1953 bps were received. (1) analyses of genetic distance and hereditary diversity indicated that: genetic distance between populations in each group were less than distance between two groups of Feirana, 19 haplotypes were recognized from 19 samples of 19 populations, so the hereditary diversity of populations of Feirana was very high (Hd=1.0), phylogenetic information in ND2 gene is more than fragment sequence of 12SrRNA and 16SrRNA genes. (2) Result of molecular phylogeny indicate that the phylogenetic trees constructed using different methods based on different sequence data sets showed the revised genus Feirana is monophyletic since the 19 populations of Feirana were firstly clustered together as one large clade, which was further clustered into two major clades, corresponding to F. quadrana(GroupⅠ) and F. taihangnicus(GroupⅡ), respectively. So populations of Qinshui and Lishan in Mt. Zhongtiao, populations of Luanchuan and Neixiang in Mt. Funiu, and populations of Zhashui, Dabagou of Chang’an and Ningshan in eastern Mt. Qinling should belong to the species F. taihangnicus; (3) Subspecific differentiation. on the basis of genetic distance, phylogenetic trees and geographical distribution, F. quadrana should have two subspecies, i.e., F. quadrana qudadrana, consisting of the populations Guanghuojie of Chang’an and Zhouzhi in Mid-Mt. Qinling, populations in Wushan area and northern Mt. Wuling (Lichuan), and F. qudadrana anxianensis, consisting of the populations in eastern Mt. Ming shan-Mt. Longmen-western Mt. Daba-western Mt. Qinling (Anxian, Qingchuan, Wenxian, Nanjiang and Fengxian); F. taihangnicus should also has two subspecies, i.e., F. taihangnicus taihangnicus, consisting of the populations in Mt. Zhongtiao and eastern Mt. Qinling, and F. taihangnicus funiuensis, consisting of the populations in Mt. Funiu. 4. Zoogeography of populaions of Feirana Analysis for divergent time of 19 populations of Feirana: Using the dates of run-through of Wushan segment of Changjiang River as the time when the population of Lichuan started differentiated from the populations of Wushan and Shennongjia, and the dates of Sanmenxia segment of Yellow River as the time when the populations in Mt. Zhongtiao started differentiated from the population of Dabagou in Chang’an, molecular clock was established using sequences with 1953 bps of 19 populations of Feirana and outgroup including N. pleski, P. yunnanesis, P. robertingeri, F. limnocharis in order to estimate divergent time of all clades. Result of that indicated that: ① the tribe Paini started to evolve independently at about 70Ma when is in consistent with that estimated by Roelants et al.（2004）with result of about ～60±15Ma, they were corroborated by each other, this confirms the validity of this molecular clock; ② divergent time for speciation of Feriana is early, ancestral populations of F. quadrana and F. taihangnicus were found about 46Ma～50Ma; differentiation of populations within species is greatly late to the divergence of the two species, divergent time for F. quadrana is 10Ma and divergent time for F. taihangnicus is 6Ma. Proceeding of distribution pattern of Feirana. Phylogenetic relationships of populations of Feirana matched quite with distribution pattern of them, the relationships among clades showed in phylogenetic trees is direct ratio to geographical distance of them; the estimated date of speciation between two species of Feirana was as early as speciation of Paa yunnanesis and Nanara pleski; middle part of Mt. Qinling is the center of speciation of Feirana, combination of mult-events of dispersal and vicariance are probably the mechanism of speciation of Feirana, F. quadrana colonized the mid-Mt. Qinling and then differentiated into two subspecies in southwest direction, ancestral population of F. taihangnicus colonized the mid-Mt. Qinling and then differentiated into two subspecies in northeast direction. On geological history of the distribution of Feirana. According to molecular clock and speciation model of populations of Feirana, some geological events are confirmed, including special rise of Mt. Minshan- Mt. Longmen-western Mt. Qinling, glacial age.

Effects of starvation on larval growth, survival and metamorphosis of Ivory shell Babylonia formosae habei Altena et al., 1981 (Neogastropoda : Buccinidae)

The impact of starvation on larvae of Ivory shell Babylonia formosae habei was studied in a laboratory experiment. Newly hatched veligers showed considerable tolerance to starvation due to their endogenous yolk material, and time to the point-of-no-return (PNR; the threshold point during starvation after which larvae can longer metamorphose even if food is provided) was calculated to be 104.5 h. However, starvation still affected larval growth, survival, and metamorphosis. Mean shell length of larvae increased 49.77 mum day(-1) for nonstarved, but only 11.13 mum day (-1) for larvae starved for 108 h. After larvae began feeding, their growth rates rapidly recovered to the level of the nonstarved following short periods of starvation (less than 48 h), but were inhibited and unable to ever reach the level of the nonstarved when being starved beyond 48 h. Percent metamorphosis was 53.75% for the nonstarved, but all larvae died before 10 days for those starved for 108 h. Starvation not only affected larval time to reach metamorphosis, but also caused the delay in the time to metamorphosis. For the nonstarved, larvae took only 11.5 days to reach spontaneous metamorphosis, but they took 20 days to reach spontaneous metamorphosis when starved for 96 h, and this duration of delayed metamorphosis reached 8.5 days. Furthermore, the importance of yolk material for maintaining larval survival of B. formosae habei during starvation periods is also discussed. (C) 2004 Elsevier B.V. All rights reserved.

晚中新世以来中国北方风成沉积的磁性地层学和沉积学研究

The past two decades have witnessed an unprecedented growth of interest in the palaeoenvironmental significance of the Pleistocene loess deposits in northern China. However, it is only several years ago that the Tertiary red clay sequence underlying Pleistocene loess attracted much attention. One of the major advances in recent studies of eolian deposits on the Loess Plateau is the verification of the eolian origin for the Tertiary red clay sediments. The evidence of the eolian origin for the red clay is mainly from geochemical and sedimentological studies. However, sedimentological studies of the red clay deposits are still few compared with those of the overlying loess sediments. To date, the red clay sections located near Xifeng, Baoji, Lantian, Jiaxian, and Lingtai have been studied, with an emphasis on magnetostratigraphy. These sections have a basal age ranging from ～4.3 Ma to ～7.0 Ma. The thickness of the sections varies significantly, depending perhaps on the development of local geomorphological conditions and the drainage system. Although the stratigraphy of the red clay sections has been recorded in some detail, correlation of the red clay sequences has not yet been undertaken. Geological records (Sun J. et al., 1998) have shown that during glacial periods of the Quaternary the deserts in northem China were greatly expanded compared with modern desert distribution. During interglacial periods, desert areas contracted and retreated mostly to northwestern China because of the increase in inland penetration of monsoonal precipitation. According to pedogenic characteristics of the red clay deposits, the climatic conditions of the Loess Plateau is warmer and wetter generally in the Neogene than in the late Pleistocene. Panicle analyses show that grain size distribution of the red clay sequence is similar to that of the paleosols in the Pleistocene loess record, thus implying a relatively remote provenance of the red clay materials. However, the quantitative or semiquantitative estimates of the distance from the source region to the Loess Plateau during the red clay development remains to be investigated. In this study, magnetostratigraphic and sedimentological studies are conducted at two thick red clay sequences-Jingchuan and Lingtai section. The objectives of these studies are focused on further sedimentological evidence for the eolian origin of the red clay, correlation of red clay sequences, provenance of the red clay, and the palaeoclimate reconstruction in the Neogene. Paleomagnetic studies show that the Jingchuan red clay has a basal age of 8.0 Ma, which is 1 million years older than the previously studied Lingtai section. The Lingtai red clay sequence was divided into five units on the basis of pedogenica characteristics (Ding et al., 1999a). The Jingchuan red clay sequence, however, can be lithologically divided into six units according to field observations. The upper five units of the Jingchuan red clay can generally correlate well with the five units of the Lingtai red clay. Comparison of magnetic susceptibility and color reflectance records of four red clay sections suggests that the Lingtai red clay sequence can be the type-section of the Neogene red clay deposits in northern China. Pleistocene loess and modem dust deposits have a unimodal grain-size distribution. The red clay sediments at Jingchuan and Lingtai also have a unimodal grain-size distribution especially similar to the paleosols in the Pleistocene loess record. Sedimentological studies of a north-south transect of loess deposits above S2 on the Loess Plateau show that loess deposits had distinct temporal and spatial sedimentary differentiation. The characteristics of such sedimentary differentiation can be well presented in a triangular diagram of normalized median grain size, normalized skewness, and normalized kurtosis. The triangular diagrams of the red clay-loess sequence at Lingtai and Jingchuan indicate that loess-paleosol-red clay may be transported and sorted by the same agent wind, thus extending the eolian record in the Loess Plateau from 2.6 Ma back to about 8.0 Ma. It has been recognized that during the last glacial maximum (LGM) the deserts in northern China had a distribution similar to the present, whereas during the Holocene Optimum the deserts retreated to the area west of the Helan Mountains. Advance-retreat cycles of the deserts will lead to changes in the distance of the Loess Plateau to the dust source regions, thereby controlling changes in grain size of the loess deposited in a specific site. To observe spatial changes in sedimentological characteristics of loess during the last glacial-interglacial cycle, the texture of loess was measured along the north-south transect of the Loess Plateau. Since the southern margin of the Mu Us desert during the LGM is already known, several models of grain size parameters versus the minimum distance from the source region to depositional areas were developed. According to these semiquantitative models, the minimum distance from the source region to Lingtai and Jingchuan areas is about 600 km during the Neogene. Therefore the estimated provenance of the Tertiary red clay deposits is the areas now occupied by the Badain Jaran desert and arid regions west of it. The ratio of the free iron to total iron concentration attests to being a good proxy indicator for the summer monsoon evolution. The Lingtai Fe_20_3 ratio record shows high values over three time intervals: 4.8-4.1 Ma, 3.4-2.6 Ma, and during the interglacial periods of the past 0.5 Ma. The increase in summer monsoon intensity over the three intervals also coincides with the well-developed soil characteristics. It is therefore concluded that the East-Asia summer monsoon has experienced a non-linear evolution since the late Miocene. In general, the East Asia summer monsoon was stronger in Neogene than in Quaternary and the strongest East Asia summer monsoon may occur between 4.1 and 4.8 Ma. The relatively small ice volume and high global temperature may be responsible for the strong summer monsoon during the early Pliocene.