The First Mitochondrial Genomes of Antlion (Neuroptera: Myrmeleontidae) and Split-footed Lacewing (Neuroptera: Nymphidae), with Phylogenetic Implications of Myrmeleontiformia

In the holometabolous insect order Neuroptera (lacewings), the cosmopolitan Myrmeleontidae (antlions) are the most species-rich family, while the closely related Nymphidae (split-footed lacewings) are a small endemic family from the Australian-Malesian region. Both families belong to the suborder Myrmeleontiformia, within which controversial hypotheses on the interfamilial phylogenetic relationships exist. Herein, we describe the complete mitochondrial (mt) genomes of an antlion (Myrmeleon immanis Walker, 1853) and a split-footed lacewing (Nymphes myrmeleonoides Leach, 1814), representing the first mt genomes for both families. These mt genomes are relatively small (respectively composed of 15,799 and 15,713 bp) compared to other lacewing mt genomes, and comprise 37 genes (13 protein coding genes, 22 tRNA genes and two rRNA genes). The arrangement of these two mt genomes is the same as in most derived Neuroptera mt genomes previously sequenced, specifically with a translocation of trnC. The start codons of all PCGs are started by ATN, with an exception of cox1, which is ACG in the M. immanis mt genome and TCG in N. myrmeleonoides. All tRNA genes have a typical clover-leaf structure of mitochondrial tRNA, with the exception of trnS1(AGN). The secondary structures of rrnL and rrnS are similar with those proposed insects and the domain I contains nine helices rather than eight helices, which is common within Neuroptera. A phylogenetic analysis based on the mt genomic data for all Neuropterida sequenced thus far, supports the monophyly of Myrmeleontiformia and the sister relationship between Ascalaphidae and Myrmeleontidae.     

The Association of the Stream Caddisfly Lasiocephala basalis (Kol.) (Trichoptera: Lepidostomatidae) with Wood

A population of the lepidostomatid caddisfly, Lasiocephala basalis, was studied in a third order stream, the Ohe, in Hesse, Germany. L. basalis showed an intimate association with wood debris in all of its aquatic life stages. Larvae exhibited a significant preference for organic substrates such as woody debris, leaves, and submerged alder roots. Gut content analysis revealed leaf material and wood fragments as the dominant larval food items classifying larvae as facultative xylophagous. Larval dominance on wood during spring corresponded to minimal leaf litter abundance. In winter, during periods of high discharge, the most important larval habitat was the submerged alder roots at the stream margin, thus acting as flow refugia. Most pupal cases were attached to submerged logs and roots. In addition, most of the egg masses were found on these two substrates. The role of L. basalis larvae in wood degradation is discussed.     

节肢动物线粒体基因组与系统发生重建

对mt基因组的比较研究是探讨节肢动物系统发生的有效手段之一。基因的排列和DNA序列可以为重建节肢动物的系统发生提供有用的信息。目前,已测定mt基因组全序列的节肢动物已增加到44种。归纳、总结了节肢动物mt基因组的基本特征、基因顺序、基因重排的发生和机制等。简要评述基于mt基因组的节肢动物系统发生研究。     

赤狐线粒体全基因组及系统发育分析

测定了赤狐的线粒体基因组全序列,总长度为16 723 bp,碱基组成为：31.3% A、26.1% C、14.8% G、27.8% T。和大多数哺乳动物一样,赤狐的线粒体全基因组包含13个蛋白质编码基因、2个核糖体RNA基因、22个转运RNA基因和1个控制区。除ND3基因起始密码子为不常见的ATT外,赤狐与北极狐、狼、家犬、郊狼的线粒体蛋白质编码遵循相同模式。在控制区的保守序列区段1和2之间发现一段较长的富含AC的随机重复序列。为了验证赤狐与其他犬科动物的系统发育关系,利用12个重链蛋白质编码基因,分别通过邻接法和最大简约法构建了系统发育树。结果表明：赤狐与北极狐是姐妹群,它们在犬科中都属于赤狐型分支,而灰狼、家犬和郊狼则属于狼型分支,与现有的系统进化研究结果一致。     

克氏光唇鱼线粒体基因组测定及光唇鱼属的系统发育分析

根据侧条光唇鱼(Acrossocheilus parallens)线粒体基因(mt DNA)序列设计引物,采用引物步移和PCR扩增产物测序,获得了克氏光唇鱼(A.kreyenbergii)的mt DNA全序列。结构分析表明,克氏光唇鱼mt DNA为首尾闭合的环状基因,全长16 596 bp,编码37个基因,包括13个蛋白编码基因、22个t RNA基因、2个r RNA基因和两段非编码区(D-loop和轻链复制起点OL),碱基组成具有明显的A+T偏好和反G偏倚现象。13个蛋白编码基因中,除COⅠ的起始密码子是GTG,其余基因的起始密码子均为ATG;终止密码子包括完全的终止密码子TAA(38.46%)和TAG(7.69%),不完全的终止密码子TA(15.38%)和T(38.46%)。在D-loop区的811~837 bp区间发现了一段"TA"短串联重复序列。从蛋白编码基因所包含的信息量、变异位点和变异率看,ND5、ND4、COⅠ和ND2最适合作为光唇鱼属种间系统发育分析的分子标记。采用贝叶斯法利用13个蛋白编码基因所构建的系统发育树显示,光唇鱼属和白甲鱼属(Onychostoma)的24种鱼类各自没有聚为单系群,相互间不能明确区分。     

线粒体DNA序列分析在鳞石蛾科(昆虫纲：毛翅目)成、幼虫联系中的应用

水生昆虫幼虫的种类鉴定是影响水质生物监测准确性的重要因素 ,我国这方面的形态学鉴定资料极其缺乏。本研究测定和分析了 4种鳞石蛾成虫及其疑似幼虫的mtDNA COⅠ、Ⅱ及tRNA基因序列 ,发现鳞石蛾成虫与其疑似幼虫的序列歧异度均 <1% ,属于种内差异 ,由此确认了 4种供试幼虫与其对应成虫的联系。研究表明利用线粒体DNA细胞色素氧化酶基因序列分析鉴定鳞石蛾类幼虫是可行的 ,为加快水生昆虫的幼虫分类研究提供了新途径。     

Some characteristics of the trichoptera distribution in the Mesozoic of Eurasia (Insecta: Trichoptera)

The stratigraphic distribution and peculiarities of burial of caddis cases from 293 Mesozoic localities in Eurasia are analyzed. An attempt is made to analyze and interpret the co-occurrence of trichopteran cases and adults in the major localities. The main stages in the evolution of the building behavior of trichopteran larvae in the Mesozoic have been elucidated.     

湾鳄(Crocodylus porosus)线粒体基因组全序列结构分析与鳄类系统发生

该文测序了湾鳄的线粒体基因组全序列,全长为16,917bp。湾鳄mtDNA结构与其他脊椎动物相似,由22个tRNA,2个rRNA和13个蛋白质编码基因及1个非编码的控制区(D-loop)所组成。除NADH6和tRNAGln、tRNAAla、tRNAAsn、tRNACys、tRNATyr、tRNASer(UCN)、tRNAGlu、tRNAPro在L-链上编码之外,其余基因均在H-链编码。基因排列顺序与已测序的鳄类一致,这显示了鳄类线粒体基因排列顺序上的保守性。但鳄类线粒体基因排列顺序与脊椎动物的典型排列方式相比,有较大的差异,尤其是tRNAPhe基因的重排、tRNASer-tRNAHis-tRNALeu基因族的排列方式等。湾鳄mtDNA和已测序的鳄类一样,缺失轻链复制起始点(OLR)。基于17种鳄mtDNA控制区保守区,采用PAUP4.0最大简约法(Maximumparsimony,MP)构建MP树,邻接法(Neighbor-joiningmethod,NJ)构建NJ树,结果显示:食鱼鳄(Gavialisgangeticus)和假食鱼鳄(Tomistomaschlegelii)聚为一支后再与鳄科(Crocodylidae)的其他物种形成姐妹群,这与基于食鱼鳄和假食鱼鳄的线粒体全序列的分析结果一致,支持将食鱼鳄并入鳄科的观点。结果还支持非洲窄吻鳄(Crocodyluscataphractus)与鳄属(Crocodylus)构成姐妹群,可以单独划分为属的观点。     

Monoaminhaltige Strukturen im Zentralnervensystem der Trichoptera (Insecta) Teil II

Zusammenfassung Mit Hilfe der fluoreszenzmikroskopischen Methode nach Falck und Hillarp wurden die monoaminhaltigen Strukturen im Zentralnervensystem einiger Trichopteren untersucht (vgl. Klemm, 1968). Im Protocerebrum können vier Gruppen von katecholamin-haltigen Zellkörpern unterschieden werden. Eine weitere unregelmäßig darstellbare Gruppe von globulösen Perikarya liegt im Lobus opticus. Fluoreszierende Varikositäten durchsetzen locker das Cerebralganglion und sind in folgenden Neuropilstrukturen konzentriert: Medulla, Lobula, Corpus centrale, Nodulus, Corpus ventrale, - und -Lobus und Lobus communis. Letzterer verbindet den - und den -Lobus mit dem nicht fluoreszierenden Pedunculus. In der Lamina, in der accessorischen Medulla, in der Pons cerebralis, im Tuberculum opticum, im Tractus olfactorio-globularis, im Pedunculus, im Stielglomerulus und in den Globulizellen ließen sich keine Katecholamine nachweisen. Zwischen dem monoaminhaltigen Lobus communis und dem nicht fluoreszierenden Pedunculus besteht eine scharfe Grenze. Fluoreszierende variköse Fasern und einzelne fluoreszierende Perikarya befinden sich im Deuto- und Tritocerebrum. Zwei fluoreszierende Bahnen können im Cerebralganglion unterschieden werden: 1. Stratum caudale, 2. Tractus ventralis. Letzterer beginnt mit seiner Pars anterior und posterior im dorsalen fluoreszierenden Neuropil des Protocerebrum. Die beiden Teile laufen frontal und caudal um den Zentralkörper herum und vereinigen sich unterhalb des Zentralkörpers in der Pars communis. Von hier aus zieht der paarige Tractus ventralis bis in das Tritocerebrum. Die Frage einer Homologisierung dieser Bahnen wird diskutiert.Ähnlich wie im Ganglion suboesophageale sind in den drei thorakalen und fünf separaten abdominalen Ganglien zwei Paar fluoreszierender Zellkörper vorhanden, in denen Dopamin mikrospektrofluorimetrisch festgestellt werden konnte. Ein Paar dieser Zellen liegt caudoventral, das andere dorso-median bis dorso-caudal. Ihre Zellfortsätze und Abzweigungen werden beschrieben. In einzelnen Fällen konnte noch ein zweites dorsales Paar fluoreszierender Perikarya sichtbar gemacht werden. Das letzte Ganglion der Bauchkette setzt sich aus zwei bis drei Gangliomeren zusammen. Die Anzahl ihrer fluoreszierenden Perikarya ist reduziert. Das Neuropil der Ganglien im Bauchmark ist von monoaminhaltigen Fasern durchsetzt, wobei sich in der dorsalen Hälfte mehr fluoreszierendes Neuropil befindet als in der ventralen. Lateral in den Ganglien sind die monoaminhaltigen varikösen Fasern vorwiegend dorsoventrad angeordnet. Im medianen Teil laufen sie in Längsrichtung, verzweigen sich und setzen sich in die Konnektive fort und verbinden die katecholaminhaltigen Neuropilbereiche der einzelnen Ganglien miteinander.

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Monoamine-containing structures in the central nervous system in Trichoptera (Insecta). Part II

Summary The distribution of monoamine-containing structures in the central nervous system of Trichoptera has been investigated with the histochemical fluorescence method of Falck and Hillarp (cf. also Klemm, 1968). In the protocerebrum, four groups of catecholamine containing perikarya can be distinguished. Another group of irregularly appearing cell bodies is situated in the optical lobe. Fluorescent varicosities occur scattered throughout the cerebral ganglion, being concentrated in the following neuropile areas: In the medulla, the lobula, the corpus centrale, the noduli, the corpora ventralia, and the - and -lobes, as well as in the lobus communis where the - and -lobes join the non-fluorescent pedunculus. In the lamina, the accessory medulla, the pons cerebralis, the tuberculum opticum, the tractus olfactorio-globularis, the pedunculus including its glomerular region, and the globuli-cells, no catecholamines are detectable. There is a sharp borderline between the monoamine-containing lobus communis and the non-fluorescent pedunculus. Fluorescent varicose fibres and single fluorescent perikarya occur in the Deuto- and Tritocerebrum. Two fluorescent tracts can be distinguished in the cerebral ganglion: 1. Stratum caudale. 2. Tractus ventralis. The pars anterior and pars posterior of the tractus ventralis begin in the dorsal fluorescent neuropile of the cerebral ganglion and join underneath the central body into their pars communis. From this, it can be traced as tractus ventralis into the tritocerebrum. The possible homologies of these tracts are discussed. Similar to the ganglion suboesophageale, the three thoracal and the five separate abdominal ganglia contain two paires of fluorescent cell bodies. Microspectrofluorometrically a content of dopamine in these cells could be established. One pair lies caudoventrally. The two other cell bodies are situated dorso-medially to dorso-caudally; their position varies notably, especially in the abdominal ganglia. The cell process and its arborisations are described. Occasionally a second dorsal cellpair could be observed. The last ganglion of the abdominal chain is composed of at least two gangliomeres with a reduced number of fluorescent perikarya. The neuropile of the thoracal- and abdominal ganglia is penetrated by monoamine-containing fibres, with a predominance in the dorsal half. In the ganglia, the fluorescent varicose fibres are mainly oriented dorso-ventrally in the lateral part and longitudinally in the medial part, where they branch and continue into the connectives. In this way, the catecholamine-containing neuropiles of adjacent ganglia are connected to each other.

Die vorliegende Arbeit wurde unterstützt durch The Swedish Medical Research Council (No. B70-14X-56-06 und B70-14X-05).     

Covariation of Mitochondrial Genome Size with Gene Lengths: Evidence for Gene Length Reduction During Mitochondrial Evolution

Reduction of genome size and gene shortening have been observed in a number of parasitic and mutualistic intracellular symbionts. Reduction of coding capacity is also a unifying principle in the evolutionary history of mitochondria, but little is known about the evolution of gene length in mitochondria. The genes for cytochrome c oxidase subunits I–III, cytochrome b, and the large and small subunit rRNAs are, with very few exceptions, always found on the mitochondrial genome. These resident mitochondrial genes can therefore be used to test whether the reduction in gene lengths observed in a number of intracellular symbionts is also seen in mitochondria. Here we show that resident mitochondrial gene products are shorter than their corresponding counterparts in -proteobacteria and, furthermore, that the reduction of mitochondrial genome size is correlated with a reduction in the length of the corresponding resident gene products. We show that relative genomic AT content, which has been identified as a factor influencing gene lengths in other systems, cannot explain gene length/genome size covariance observed in mitochondria. Our data are therefore in agreement with the idea that gene length evolves as a consequence of selection for smaller genomes, either to avoid accumulation of deleterious mutations or triggered by selection for a replication advantage.     

New records of polycentropodid caddisflies from Korea (Insecta: Trichoptera)

Four species of the family Polycentropodidae –Plectrocnemia baculifera, P. kusnezovi, Paranyctiophylax hjangsanchonus and Polyplectropus nocturnus– are recorded from Korea. Polyplectropus nocturnus is newly recorded from the Korean peninsula. The other three species are recorded from the Republic of Korea for the first time.     

Complete Mitochondrial Genome Sequences of the South American and the Australian Lungfish: Testing of the Phylogenetic Performance of Mitochondrial Data Sets for Phylogenetic Problems in Tetrapod Relationships

We determined the complete nucleotide sequences (16403 and 16572 base pairs, respectively) of the mitochondrial genomes of the South American lungfish, Lepidosiren paradoxa, and the Australian lungfish, Neoceratodus forsteri (Sarcopterygii, Dipnoi). The mitochondrial DNA sequences were established in an effort to resolve the debated evolutionary positions of the lungfish and the coelacanth relative to land vertebrates. Previous molecular phylogenetic studies based on complete mtDNA sequences, including only the African lungfish, Protopterus dolloi, sequence were able to strongly reject the traditional textbook hypothesis that coelacanths are the closest relatives of land vertebrates. However, these studies were unable to statistically significantly distinguish between the two remaining scenarios: lungfish as the closest relatives to land vertebrates and lungfish and coelacanths jointly as their sister group (Cao et al. 1998; Zardoya et al. 1998; Zardoya and Meyer 1997a). Lungfish, coelacanths, and the fish ancestors of the tetrapod lineage all originated within a short time window of about 20 million years, back in the early Devonian (about 380 to 400 million years ago). This short divergence time makes the determination of the phylogenetic relationships among these three lineages difficult. In this study, we attempted to break the long evolutionary branch of lungfish, in an effort to better resolve the phylogenetic relationships among the three extant sarcopterygian lineages. The gene order of the mitochondrial genomes of the South American and Australian lungfish conforms to the consensus gene order among gnathostome vertebrates. The phylogenetic analyses of the complete set of mitochondrial proteins (without ND6) suggest that the lungfish are the closest relatives of the tetrapods, although the support in favor of this scenario is not statistically significant. The two other smaller data sets (tRNA and rRNA genes) give inconsistent results depending on the different reconstruction methods applied and cannot significantly rule out any of the three alternative hypotheses. Nuclear protein-coding genes, which might be better phylogenetic markers for this question, support the lungfish–tetrapod sister-group relationship (Brinkmann et al. 2004).This article contains online supplementary material.Reviewing Editor: Dr. Rafael Zardoya     

Organization of the Mitochondrial Genome of Mantis Shrimp Pseudosquilla ciliata (Crustacea: Stomatopoda)

We determined the nearly complete mitochondrial genome of Pseudosquilla ciliata (Crustacea, Stomatopoda), including all protein-coding genes and all but one of the transfer RNAs. There were no gene rearrangements relative to the pattern shared by crustaceans and hexapods. Phylogenetic analysis using concatenated amino acid sequences of the mitochondrial protein-coding genes confirmed a basal position of Stomatopoda among Eumalacostraca. Pancrustacean relationships based on mitogenomic data were analyzed and are discussed in relation to crustacean and hexapod monophyly and hexapod affinities to crustacean subtaxa.     

The Mitochondrial Genome of Raphanus sativus and Gene Evolution of Cruciferous Mitochondrial Types

To explore the mitochondrial genes of the Cruciferae family, the mitochondrial genome of Raphanus sativus (sat) was sequenced and annotated. The circular mitochondrial genome of sat is 239,723 bp and includes 33 protein-coding genes, three rRNA genes and 17 tRNA genes. The mitochondrial genome also contains a pair of large repeat sequences 5.9 kb in length, which may mediate genome reorga-nization into two sub-genomic circles, with predicted sizes of 124.8 kb and 115.0 kb, respectively. Furthermore, gene evolution of mitochondrial genomes within the Cruciferae family was analyzed using sat mitochondrial type (mitotype), together with six other re-ported mitotypes. The cruciferous mitochondrial genomes have maintained almost the same set of functional genes. Compared with Cycas taitungensis (a representative gymnosperm), the mitochondrial genomes of the Cruciferae have lost nine protein-coding genes and seven mitochondrial-like tRNA genes, but acquired six chloroplast-like tRNAs. Among the Cruciferae, to maintain the same set of genes that are necessary for mitochondrial function, the exons of the genes have changed at the lowest rates, as indicated by the numbers of single nucleotide polymorphisms. The open reading frames (ORFs) of unknown function in the cruciferous genomes are not conserved. Evolutionary events, such as mutations, genome reorganizations and sequence insertions or deletions (indels), have resulted in the non- conserved ORFs in the cruciferous mitochondrial genomes, which is becoming significantly different among mitotypes. This work represents the first phylogenic explanation of the evolution of genes of known function in the Cruciferae family. It revealed significant variation in ORFs and the causes of such variation.     

Ultrastructure and central projections of extraocular photoreceptors in caddiesflies (Insecta: Trichoptera)

Summary Retained larval eyes (stemmata) were studied in the imagines of three species of Trichoptera: Phrygania grandis, Agrypnia varia, and Trichostegia minor. At the light-microscopic level the stemmata of all three species appeared to represent different stages of reduction with respect to size, shape and number of lenses. However, in all three species electron-microscopic studies showed units with monolayered rhabdoms, each formed by four retinula cells. By use of immunocytochemistry the presence of S-antigen was demonstrated in the retinula cells and their axons. This method also revealed the central projections of the axons of the retinula cells, which were found (i) to terminate either in the lamina accessoria or (ii) to penetrate this area to join the fibers of the outer chiasma of the optic lobes and then terminate in the medulla accessoria. The lamina accessoria and the medulla accessoria are the assumed remnants of the larval optic lobes. It is suggested that the imaginal stemmata might still be functioning photoreceptors.     

黄脸油葫芦线粒体基因组:一种新的基因排列方式

采用长距PCR扩增及保守引物步移法测定并注释了黄脸油葫芦（Teleogryllus emma）线粒体基因组全序列。结果表明,黄脸油葫芦的线粒体基因组全长15 660 bp,A+T含量为73.1%。谷氨酸、色氨酸及天冬酰胺的转运RNA基因由N链编码,形成了直翅目中的第三种基因排列顺序,其余结构与其它螽亚目昆虫的线粒体结构一致。基因间隔序列共计73 bp,间隔长度从1—24 bp不等;有14对基因间存在共54 bp重叠,重叠碱基数在1—11 bp之间。13个蛋白质编码基因中12个基因（除COⅠ基因外）的起始密码为标准的ATN组成,COI基因的起始密码子为TTA。有10个基因在基因3＇端能找到完全的TAA或TAG终止密码子,而有三个基因（COII,ND5和ND4）终止密码子为不完整的T。除tRNASer（AGN）外,其余21个tRNA基因的二级结构均属典型的三叶草结构。黄脸油葫芦940bp的A+T富集区中存在一个被认为与复制起始有关的保守的二级结构,该结构不仅存在于直翅目昆虫中,而且也存在于双翅目、鳞翅目和膜翅目中,但是未见于昆虫纲的早期分化类群——弹尾目中。     

New species and new records of Apataniidae (Insecta: Trichoptera) from east central China

Three new species of Apataniidae from China, Apatania protracta sp. n., Apatidelia paramartynovi sp. n., and Moropsyche dawuensis sp. n. are described and illustrated. We also report Apatania semicircularis Leng and Yang, 1998 for the first time from Hubei Province.

Apatania protracta: http://zoobank.org/urn:zoobank.org:pub:8317EB90-9316-4F7B-94C6-65938A6F2873

Apatidelia paramartynovi: http://zoobank.org/urn:zoobank.org:pub:8741CE6F-B339-4842-BD8C-04BE8B21A222

Moropsyche dawuensis: http://zoobank.org/urn:zoobank.org:pub:B5CB815D-59B5-401F-9308-9EE24A9BBAF7     

鳙的线粒体基因组核苷酸全序列分析

对采集自我国长江的鳙的线粒体DNA全序列进行了测定.结果表明,鳙的线粒体DNA全长为166221 bp,其碱基因组成为A=31.6%;C=27.1%;G=16.0%;T=25.3%,A+T含量为56.9%.鳙线粒体基因组的排列、结构和组成与其它鲤科鱼类相似,包括37个基因,即13个蛋白质编码基因,2个rRNA基因,22个tRNA基因和一个非编码控制区(D-loop).在13个蛋白编码基因中,除ND6由轻链编码外,其余12个基因均由重链编码.COI基因的起始密码子为GTG,而其它12个蛋白编码基因的起始密码子均为ATG.     

齿缘摄龟线粒体基因组全序列与系统进化

采用常规PCR扩增并测序获得了齿缘摄龟(Cyclemys dentata)线粒体DNA(mtDNA)全序列,并研究了其基因组结构特点;根据20种龟的线粒体基因组重链蛋白编码基因序列,分别利用最大简约法(MP)、最大似然法(ML)和贝叶斯法(Bayesian)构建系统进化树,探讨这些龟鳖物种之间的系统进化关系。结果显示,齿缘摄龟线粒体基因组全序列长为16489 bp(GenBank登录号为JX455823),A+T含量为61.51%,编码37个基因,包括13个蛋白质编码基因,2个rRNA,22个tRNA和1个控制区(Dloop),基因组成与其他龟鳖类动物相似;非编码区D-loop长973 bp,包含1个中央保守区(CD),2个扩展终止结合序列区(ETASs),3个保守盒(CSBs);构建的MP树、ML树和Bayesian树的拓扑结构相似,闭壳龟属7种龟聚为一枝,拟水龟属6种龟聚为一枝,齿缘摄龟与黑桥摄龟聚在一枝,3种进化树均支持这些龟鳖物种现有的分类学地位。     

基于线粒体基因组的藏马高原适应及系统发育分析

研究测定了西藏那曲（4,500m）、云南中甸（3,300m）、云南德钦（3,300m）地区3匹藏马线粒体全基因组序列。3个地区的藏马线粒体基因组全长以及结构均与韩国济州岛的马类似,但比瑞典马线粒体基因组短。藏马基因组在DNA序列上的两两相似性达993％。通过对线粒体蛋白编码区的分析发现,NADH6基因的蛋白序列在三匹藏马中均表现快速进化的现象。这表明NADH6基因在藏马高原适应进化过程中扮演着重要角色。此外,利用7匹藏马的D—loop区域序列以及与其亲缘关系较近的马的序列首次构建的藏马的系统发育树显示,那曲藏马与中甸、德钦藏马属于不同的分支,且存在较大的遗传多样性,表明藏马可能为多地区起源。