食肉目哺乳动物的系统发育学研究概述

追溯生物界不同生物类型的起源及进化关系,即重建生物类群的系统发育树是进化生物学领域中一个十分重要的内容。食肉目哺乳动物位于食物链顶端,很多成员不仅在我国野生动物保护工作中占有重要地位,而且还是研究动物适应性进化遗传机制的重要模式生物。因而,食肉目物种作为物种资源中的一个重要类群,其系统发育学一直是国内外研究的热门课题。构建可靠的食肉目分子系统树,无疑将具有重要的进化理论意义和保护生物学价值。鉴于目前食肉目各科间系统发育关系仍然处于“广泛争论”的状态,本文将针对食肉目科水平上的系统发育学研究进展,包括来自于形态学特征、细胞学及分子生物学方面的证据,做简要概述,并提出目前研究中存在的问题。这对今后食肉目系统发育方面的进一步研究工作具有指导意义,并为以该类群作为模式生物开展适应性进化研究奠定基础。     

食肉目哺乳动物的系统发育学研究概述

追溯生物界不同生物类型的起源及进化关系，即重建生物类群的系统发育树是进化生物学领域中一个十分重要的内容。食肉目哺乳动物位于食物链顶端，很多成员不仅在我国野生动物保护工作中占有重要地位，而且还是研究动物适应性进化遗传机制的重要模式生物。因而，食肉目物种作为物种资源中的一个重要类群，其系统发育学一直是国内外研究的热门课题。构建可靠的食肉目分子系统树，无疑将具有重要的进化理论意义和保护生物学价值。鉴于目前食肉目各科间系统发育关系仍然处于“广泛争论”的状态，本文将针对食肉目科水平上的系统发育学研究进展，包括来自于形态学特征、细胞学及分子生物学方面的证据，做简要概述，并提出目前研究中存在的问题。这对今后食肉目系统发育方面的进一步研究工作具有指导意义，并为以该类群作为模式生物开展适应性进化研究奠定基础。     

哺乳动物系统发育基因组学研究进展

哺乳动物是一类最进化并在地球上占主导地位的动物类群,重建其系统发育关系一直是分子系统学的研究热点。随着越来越多物种全基因组测序的完成,在基因组水平上探讨该类动物的系统发育关系与进化成为研究的热点。本文从全基因组序列,稀有基因组变异及染色体涂染等几个方面简要介绍了当前系统发育基因组学在现生哺乳动物分子系统学中的应用,综合已有的研究归纳整理了胎盘亚纲的总目及目间的系统发育关系,给出了胎盘动物19 个目的系统发育树。本文还分析了哺乳动物系统发育基因组学目前所面临的主要问题及未来的发展前景。     

中国兽类遗传与进化研究进展与展望

中国兽类物种丰富,且具有150个特有种。本文综述了60年来中国兽类遗传与进化的研究进展,内容涵盖系统发育关系重建、遗传多样性评估、种群遗传结构、适应性进化以及趋同进化的分子机制。本文重点概述了食肉目（大、小熊猫）、有蹄类、翼手目、灵长目、小型兽类以及海兽类等重要类群的研究进展,为中国兽类的物种保护提供了重要资料。另外,本文还对中国兽类遗传与进化研究未来的研究方向提出几点建议,包括运用各种组学技术、筛选新型遗传标记和候选基因（调控序列）、结合表观遗传学并借助进化发育生物学研究方法,以期全面深入地理解中国兽类分类地位、起源以及特异表型产生和独特适应的发育遗传学机制等,进而实现“天人合一”保护生物学的新理念和新愿景。     

雪豹保护遗传学和基因组学研究进展

雪豹(Panthera uncia)隶属于食肉目猫科豹属,是生活在青藏高原及其周边地区的旗舰物种。随着分子生物学和高通量测序技术的发展,雪豹保护遗传学和保护基因组学研究得到快速的发展,其中非损伤性遗传取样法显著推动了雪豹保护遗传学研究。本文综述了非损伤性遗传取样法在雪豹物种鉴定、个体识别和性别鉴定等研究中的应用,雪豹的系统发生地位、系统地理格局和种群遗传结构及其亚种争议、演化历史、适应性演化和基因组特征等保护遗传学和基因组学方面的研究现状和进展,并对雪豹保护遗传学和基因组学未来发展趋势进行了展望,以期促进雪豹保护生物学研究和保护对策的科学制定。     

系统发育多样性在植物区系研究与生物多样性保护中的应用

利用系统发育关系从进化角度理解物种的组成、分布格局对可为有效保护和合理利用生物多样性提供科学依据。以支长为基础的系统发育多样性是最基本的测度指数,由此衍生出的几十个参数加大了选择难度。根据具体研究的问题选择合适的指标,是目前最为可行的方案,已经在植物系统发育区系研究和生物多样性保护中取得了一系列研究成果。DNA序列迅速积累特别是DNA条形码项目在全球的大力发展,提供了海量的标准化数据,为构建解决能力较好的宏系统发育树提供了可能。但是围绕系统发育多样性开展的相关研究还需要更多的物种分布、环境因子、气候等基础数据,同时对其在生态系统功能上的表征作用需要进一步验证。     

基于DNA条形码评估西双版纳国家级自然保护区对樟科植物进化历史的保护

为评估西双版纳国家级自然保护区对樟科这一重要植物类群进化潜力的保护情况, 揭示将物种进化历史纳入生物多样性保护评估的重要性, 本研究通过对西双版纳地区长期的野外调查并查阅标本记录与文献资料, 整理出该地区樟科13属121种物种的具体分布信息, 以植物条形码ITS序列作为分子标记构建了反映整个西双版纳地区樟科植物系统发育关系的系统发育树。我们以此为基础, 从物种层面分析了各物种的进化特异性(evolutionary distinctiveness, ED), 从区域层面分析了自然保护区内、外以及32个行政乡镇的系统发育多样性(phylogenetic diversity, PD), 并结合物种丰富度(species richness, SR)与物种濒危等级, 综合探讨了西双版纳国家级自然保护区对樟科植物进化历史的保护情况。研究发现, 西双版纳国家级自然保护区仅拥有整个西双版纳地区54.5%的樟科物种数, 却保护了该地区樟科植物约88.8%的进化历史, 没有被列入保护范围但却拥有高系统发育多样性的区域有打洛镇、易武乡等。就物种而言, 进化特异性相对较高的19个物种中, 有5种(26.3%)在自然保护区内没有分布; 濒危等级高的54个物种中, 有20种(37.0%)在自然保护区没有分布, 同时拥有高进化特异性和濒危等级的物种仅有1种不在保护区内分布。结果表明, 虽然西双版纳国家级自然保护区对樟科这一植物类群的系统发育多样性以及高保护价值物种的保护较好, 但仍有部分重要樟科植物的进化历史没有涵盖在现有自然保护区范围内; 按照传统方法设定的自然保护区虽能在一定程度上保护樟科物种的进化历史, 但仍然存在与标准化系统发育多样性保护策略相矛盾的地方。因此, 今后在建立自然保护区时, 应将系统发育多样性考虑在内, 以保护生物多样性应对环境变化的潜力。     

猫科动物

前言从各个角度看来,食肉目堪称为哺乳纲中最重要的目,而猫科则在食肉目中占有首要地位。猫科动物种类繁多,分布广泛,与人类的利害矛盾也多而且深。动物学教科书中把猫科当做一项重点,诚然有理。这篇稿除了讲述猫科的分类和分布(特别注重在我国的种类和分布)外,并有重点地说明本科动物的形态和生态,以充实授课的内容。     

系统发育研究中“长枝吸引”现象概述

系统发育研究(phylogeny)不仅有助于重建地球所有生物体的进化历史, 而且还可以揭示进化生物学领域中的一些基本问题。清晰了解各生物物种进化历程及不同物种之间的进化关系, 是进一步研究和探索生物学其他学科的基础。但是现今广泛应用的所有系统发育分析方法都存在一定的局限性, 在一定程度上不能有效消除各种误差, 从而不能客观地处理和分析数据, 也就不能成功重建生物进化历程, 真实反映物种进化关系。系统发育研究中, “长枝吸引” (Long-branch Attraction, LBA)假象是最为困扰研究者的问题。文章从“长枝吸引”问题的产生原由、检测方法以及消除策略等多个方面进行详尽概述, 并通过列举典型实例, 阐述了解决“长枝吸引”问题的途径。     

贝叶斯推论及其在百合目分子系统学中的应用

贝叶斯推论作为进化生物学的最新进展,在适用复杂模型、大型数据集、计算速度和结果容易解释等方面明显优于其它算法。本文简要介绍了贝叶斯推论原理及其在分子进化和系统发育研究中的重要性,并使用该方法对百合目主要类群的系统发育关系进行了重建。结果显示,百合目rbcL基因最适合的DNA进化模型为GTR I G,贝叶斯法与距离法和最大简约法构建的系统发育树拓扑结构相似,没有显著差异,但是分辨率和支持率明显比后者高。贝叶斯分析结果显示,百合目内划分的7个科,除Smilacaceae科外,其余各科均为高后验概率(PP=1·0)支持的单系类群;文中作者还对各科间的系统关系进行了探讨。     

Evolution of Acoustic Communication Signals of Mammals: Friendly Close-Range Vocalizations in Felidae (Carnivora)

The distribution of the three friendly close-range vocalization types known in the Felidae was plotted on a recently published phylogeny of the cat family (Felidae) based on sequence comparisons of two mitochondrial DNA genes and other molecular and biochemical characters, with extrapolated divergence ages of its various lineages. It was found to be congruent with this phylogeny. One of the sound types is likely to be present in 30 species of the family (documented in 22 so far), another is present in 4, and the third in 2 species only; these sound types represent a phylogenetic transformation series. The latter two vocalization types also differ considerably from the first in the mode of sound production. From this, evolutionary conservatism over a long epoch for the one widespread vocalization type can be inferred, and less conservatism in the type present in four species, while the emergence of the least common type is evidence of relatively considerable and rapid evolutionary change. Thus, acoustic communication signals in a group of taxa can evolve at considerably different rates, and for a specific character this rate can differ between different lineages of that group. The ultimate causes of the evolutionary stability or of the subsequent relatively rapid change in sound structure and mode of sound production in these felid vocalizations are unknown.     

Phylogenetic Autocorrelation Analysis of Extinction Risks and the Loss of Evolutionary History in Felidae (Carnivora: Mammalia)

Phylogeny provides a natural measurement of biodiversity by allowing the computation of indexes that express the amount of phylogenetic diversity that are, in principle, independent of species counts and provides an objective measurement to evaluate the amount of biodiversity lost under the ongoing extinction crisis. In this note, we analyzed patterns of phylogenetic autocorrelation in extinction risks in Felidae (Mammalia: Carnivora) and estimated, using simulation procedures, the amount of phylogenetic diversity loss if k species are preserved in this clade. The simulations showed that loss of phylogenetic diversity based on the IUCN list for extinction threats in Felidae is within the expected values based on the simulated random model, a result also confirmed by the absence of phylogenetic autocorrelation in extinction risks, indicating that extinction threats are randomly distributed across the phylogeny. So, we confirm that loosing species will not necessarily generate a direct proportional loss of phylogenetic information and, consequently, that alternative measures of biodiversity could be used to establish conservation priorities under the common restriction of resources.     

猫科动物的分类——分子系统学框架和化石证据

猫科动物的演化关系传统上一直是富有争议的,这很大程度上造成了猫科在属一级的分类众说纷纭。新兴的分子生物学技术为我们理解猫科动物的系统发育提供了新的视角,本文回顾了近些年来的猫科分子系统学研究,以Johnson等（2006）和李钢等（2016）的研究结果为核心,同时结合确切的化石记录进行佐证,推断了猫科的演化历史。我们发现,分子系统学提出的现代猫科动物于晚中新世开始适应辐射、先后分化出了8个支系等观点,与化石证据基本吻合;但是分子系统学对部分支系起源的推断与化石证据存在差异。结合化石记录的分析,我们认为现代猫科8个支系中除狞猫支系和虎猫支系以外,其余均最有可能起源于亚洲,现代猫科动物在演化过程中至少经历了30次洲际迁徙。结合演化历史和形态学研究,我们遵循邱占祥等（2004）的观点,将现存的猫科动物全部归于猫亚科（Felinae）,并将现代猫亚科动物划分为15个属40个物种。     

Molecular Phylogeny of Mitochondrial Cytochrome b and 12S rRNA Sequences in the Felidae: Ocelot and Domestic Cat Lineages

Molecular phylogeny of the cat family Felidae is derived using two mitochondrial genes, cytochrome b and 12S rRNA. Phylogenetic methods of weighted maximum parsimony and minimum evolution estimated by neighbor-joining are employed to reconstruct topologies among 20 extant felid species. Sequence analyses of 363 bp of cytochrome b and 376 bp of the 12S rRNA genes yielded average pair-wise similarity values between felids ranging from 94 to 99% and from 85 to 99%, respectively. Phylogenetic reconstruction supports more recent, intralineage associations but fails to completely resolve interlineage relationships. Both genes produce a monophyletic group ofFelisspecies but vary in the placement of the pallas cat. The ocelot lineage represents an early divergence within the Felidae, with strong associations between ocelot and margay, Geoffroy's cat and kodkod, and pampas cat and tigrina. Implications of the relative recency of felid evolution, presence of ancestral polymorphisms, and influence of outgroups in placement of the topological root are discussed.     

A MOLECULAR PHYLOGENY OF THE FELIDAE: IMMUNOLOGICAL DISTANCE

The phylogenetic distances between 34 of the 37 extant species of Felidae were estimated using albumin immunological distances (AID). Albumins from ten cat species were used to prepare antisera in rabbits. A consensus phylogeny was constructed from a matrix of reciprocal AID measurements using four distinct phylogenetic algorithms. A series of one-way measurements using the ten index antisera and those 24 species for which albumins were available (but antisera were not), permitted addition of these “species' limbs” to the previously derived phylogenetic trees. The major conclusions of the derived topology were: 1) the earliest branch of the feline radiation occurred approximately 12 million years B.P. and led to the small South American cats (ocelot, margay, Geoffroy's cat, etc.); 2) the second branching occurred 8-10 million years B.P. and included the close relatives of the domestic cat (wildcats, jungle cat, sand cat, and black-footed cat) plus Pallas's cat; 3) the third lineage which began to radiate 4-6 million years B.P. was the pantherine lineage, which included several early branches (cheetah, serval, clouded leopard, golden cats, and puma) and a very recent (2 million years B.P.) split between the lynxes and the modern great cats (Panthera). The topology of the Felidae derived from albumin immunological distance is highly consistent with the karyological disposition of these species, as well as with the fossil record of this family. Because of the recent divergence of this group, the presented data set and the derived topology contain certain unresolved phylogenetic relationships which are so indicated.     

Morphological integration in the carnivoran skull

The correlated evolution of traits may be a principal factor in morphological evolution, but it is typically studied in genetic or developmental systems. Most studies examining phenotypic trait correlations, through analysis of morphological integration, consider only few taxa, with limited ability to test hypotheses of the influence of trait integration on morphological variation and diversity. The few comparative studies in less inclusive groups have yielded varying relationships of integration to the key factors of phylogeny and diet. In this paper, I present analyses of cranial morphological integration in 30 species from the mammalian order Carnivora, spanning eight extant families and a wide range of ecological and morphological diversity. Fifty-five cranial landmarks were captured through three-dimensional digitization of 15-22 specimens for each species. Using a node-based phylogenetic distance matrix, a significant correlation was found between similarity in patterns of integration and phylogenetic relatedness within Felidae (cats) and Canidae (dogs), but not within more inclusive clades, when size-related variation was removed. When size was included, significant correlations were found across all Caniformia, Musteloidea, Mustelidae, and Felidae. There was a significant correlation between phylogeny and morphological integration only within the higher-level clade Feliformia (cats, civets, mongooses, and hyaenas) when a branch-length-based phylogenetic distance matrix was analyzed, with and without size. In contrast, diet was significantly correlated with similarity in morphological integration in arctoid carnivorans (bears, raccoons, and weasels), but had no significant relationship with integration in feliforms or canids. These results support the proposition that evolutionary history is correlated with cranial integration across large clades, although in some smaller clades diet also exerts significant influence on the correlated evolution of traits.     

Phylogenetic assessment of introns and SINEs within the Y chromosome using the cat family felidae as a species tree

The cat family Felidae was used as a species tree to assess the phylogenetic performance of genes, and their embedded SINE elements, within the nonrecombining region of the Y chromosome (NRY). Genomic segments from single-copy X-Y homologs SMCY, UBE1Y, and ZFY (3,604 bp) were amplified in 36 species of cat. These genes are located within the X-degenerate region of the NRY and are thought to be molecular "fossils" that ceased conventional recombination with the X chromosome early within the placental mammal evolution. The pattern and tempo of evolution at these three genes is significant in light of the recent, rapid evolution of the family over approximately 12 Myr and provides exceptional support for each of the eight recognized felid lineages, as well as clear diagnostic substitutions identifying nearly all species. Bootstrap support and Bayesian posterior probabilities are uniformly high for defining each of the eight monophyletic lineages. Further, the preferential use of specific target-site motifs facilitating SINE insertion is empirically supported by sequence analyses of SINEs embedded within the three genes. Target-site insertion is thought to explain the contradiction between intron phylogeny and results of the SMCY SINE phylogeny that unites distantly related species. Overall, our data suggest X-degenerate genes within the NRY are singularly powerful markers and offer a valuable patrilineal perspective in species evolution.     

Phylogenetic relationships within mammalian order Carnivora indicated by sequences of two nuclear DNA genes

Phylogenetic relationships among 37 living species of order Carnivora spanning a relatively broad range of divergence times and taxonomic levels were examined using nuclear sequence data from exon 1 of the IRBP gene (approximately 1.3 kb) and first intron of the TTR gene (approximately 1 kb). These data were used to analyze carnivoran phylogeny at the family and generic level as well as the interspecific relationships within recently derived Felidae. Phylogenetic results using a combined IRBP+TTR dataset strongly supported within the superfamily Califormia, the red panda as the closest lineage to procyonid-mustelid (i.e., Musteloidea) clade followed by pinnipeds (Otariidae and Phocidae), Ursidae (including the giant panda), and Canidae. Four feliform families, namely the monophyletic Herpestidae, Hyaenidae, and Felidae, as well as the paraphyletic Viverridae were consistently recovered convincingly. The utilities of these two gene segments for the phylogenetic analyses were extensively explored and both were found to be fairly informative for higher-group associations within the order Carnivora, but not for those of low level divergence at the species level. Therefore, there is a need to find additional genetic markers with more rapid mutation rates that would be diagnostic at deciphering relatively recent relationships within the Carnivora.     

Evolution and systematics of the feliform Carnivora

Recent studies have improved our knowledge about the evolution and phylogeny of feliform taxa. Detailed study on new fossil remains of extinct feliform nimravides allows a new hypothesis concerning interrelationships within this family. Many factors indicate lack of sister relationships of Nimravinae and Barbourofelinae. However, only further investigations may bring full acceptance of this hypothesis. The paraphyly of Viverridae has been resolved by excluding the taxa Nandinia, Prionodon and Cryptoprocta and Fossa, which today are placed basally to all remaining Feliformia (family Nandiniidae), as sister taxon of Felidae (family Prionodontidae) and as Malagasy Carnivora lineage basal to hyaenid–herpestid clade, respectively. Still, incongruence among results concerning the systematic position of these taxa exhibits the necessity of further investigation. Detailed study revealed inconsistencies within genet and genet-like taxa phylogeny, which have still to be resolved. Malagasy Carnivora belong to a separate lineage, which originated from herpestid–hyaenid ancestors and colonised Madagascar during a single colonisation event. However, interrelationships among Malagasy Carnivora are poorly resolved. The situation of the social mongooses clade was resolved by including ethologic data to phylogenetic analyses; however, there is little information on solitary mongooses, which have a paraphyletic status today. Felid morphology and taxonomic revision attained during recent years show greater evolutionary differentiation. Nevertheless, no clear taxonomy has been achieved. New investigating methods are required. In the hyaenid family, which includes only four living species, some investigations related to the ecomorphological evolutionary path have been performed. The comparisons of fossil and subfossil remains with modern feliforms, combined with recent molecular methods, may improve our knowledge.     

中国陆生哺乳类的易绝灭特征

本研究首先收集了我国367个陆生哺乳类物种的4类基本特征数据:个体大小、繁殖力、利用和濒危等级。从类群和特征两个角度出发,用随机检验和回归分析的方法,检验(1)受威胁物种在各科中是否是随机分布的?如果不是,哪些科含有比随机分布更多或更少的受威胁物种?不同类群受威胁概率与人类的利用有何关系?(2)物种受威胁程度与个体大小和繁殖力有何关系?人类的利用是否对受威胁程度与个体大小和繁殖力的关系产生了重要影响?结果表明:受威胁物种在各科中的分布不是随机的,猴科、牛科和猫科含比随机分布更多的受威胁物种,而鼠科和科则相反;不同类群受威胁概率与被利用率成正比;物种受威胁程度相对于繁殖力和身体大小也不是随机的;当控制系统发育关系的影响后,物种受威胁程度随个体增大和繁殖力降低而增大;多元回归分析显示,只有个体大小与物种受威胁程度相关,这一点与其它的研究不同;个体大小与被利用率成正比。过度利用对我国哺乳类多样性受威胁模式产生了重要影响