A phylogeny of sparoid fishes (Perciformes, Percoidei) based on morphology

 The putative percoid superfamily Sparoidea includes the Nemipteridae, Lethrinidae, Sparidae, and Centracanthidae. Although a rigorous cladistic analysis has never been attempted, two hypotheses regarding relationships among these families have been proposed. One early noncladistic hypothesis considered the Sparidae to be intermediate between the more primitive Nemipteridae and the more derived Lethrinidae. A later nonformal phylogenetic treatment provided evidence for a close relationship between Sparidae and Centranthidae and suggested a closer affinity between the Nemipteridae and Lethrinidae. We examine 54 osteological, ligament, and squamation characters in representatives of all 45 genera of these families and 4 outgroup taxa. The results of our cladistic analysis are congruent with a cladistic interpretation of the earlier hypothesis, with strong support for the phyletic sequence Nemipteridae, Lethrinidae, Sparidae plus Centracanthidae, with placement of centracanthids unresolved with respect to sparid genera. Received: May 21, 2001 / Revised: October 26, 2001 / Accepted: November 19, 2001     

中国小蜂科属间系统发育关系分析(膜翅目:小蜂科)

利用Hennig86程序的nelsen合意和Phylip程序的多数规则合意2种支序分析方法,探讨了中国小蜂科的系统发育关系。基于中国21属的30个性状,计算得到2个合意树,其分类系统与传统分类系统基本保持一致。在进化关系和亲缘关系上表现为:小蜂属(Chalcis)、卡诺小蜂属(Conura)、大腿小蜂属(Brachymeria)和脊柄小蜂属(Epitranus)相对最为原始,而泰内小蜂属(Tainaniella)和背突小蜂属(Oxycoryphe)相对最为进化,前者和后者之间的亲缘关系最远;亲缘关系最近的有:泰内小蜂属(Tainaniella)和背突小蜂属(Oxycoryphe),泊卡小蜂属(Proconura)和日本小蜂属(Nipponochalcidia),小蜂属(Chalcis)和卡诺小蜂属(Conura)以及细尾小蜂属(Megalocolus)和三角小蜂属(Trigonura),它们分别构成姊妹群关系。     

Phylogenetic Analysis of the Family Taxodiaceae

In the present paper,both cladistic analysis and phenetic analysis were conducted to evaluate the phylogenetic relationships of the Taxodiaceae based on an extensive literature review and study of herbarium． In the cladistic analysis,the Sciadopityaceae was chosen as outgroup．The polarity of characters was determined mainly according to outgroup comparison,fossil evidence and generally accepted viewpoints of morphological evolution．By the result of compatibility analysis,character 2(leaf type),which possessed a much higher coefficient than others whether or not its polarity was altered,was deleted． Finally,a data matrix consisting of all the extant nine genera and 24 characters was analyzed using Maximal Same Step Method,Synthetic Method,Evolutionary Extremal Aggregation Method and Minimal Parallel Evolutionary Method,and four cladograms were generated,of which only the most parsimonious one (Fig．1)was presented for discussion. The cladogram shows that the Taxodiaceae are assorted along five lines of evolution： 1)Metasequoia;2)Sequoiadendron,Sequoia;3)Cryptomeria;4)Glyptostrobus and Taxodium;5)Cunninghamia,Athrotaxis and Taiwania． Ten genera(including Sciadopitys)and 59 characters were used in the phenetic analysis．The phenogram(Fig．2)indicates that Sciadopitys is a very distinct group with remote affinity to the other genera,and the Taxodiaceae are divided into four groups：1)Sequoia,Sequoiadendron;2)Athrotaxis,Cunninghamia and Taiwania;3)Cryptomeria,Glyptostrobus and Taxodium;4)Metasequoia． Based primarily on the result of cladistics,with reference to that of phenetics,the main conclusions were drawn as follows：(1)Generic relationships：Cryptomeria should be considered the most primitive genus in the extant groups of the Taxodiaceae. Glyptostrobus and Taxodium, close to Cryptomeria, are sister taxa and relatively primitive groups. Sequoiadendron and Sequoia are closely related and intermediate advanced. Metasequoia is a more or less isolated taxon, relatively close to Sequoiadendron and Sequoia. Cunninghamia. Athrotaxis and Taiwania might represent a single lineage and form a very advanced group, of which Taiwania may be the most specialized. (2) Systematic treatments: The authors support the viewpoint that Sciadopitys should be treated as an independent family, and suggest that the Taxodiaeae should be divided into five tribes. Systematic arrangements are as follows: Taxodiaceae Warming Trib. 1. Cryptomerieae Vierhapper Gen. 5. Sequoia Endl. Gen. 1. Cryptomeria D. Don Trib. 4. Metasequoieae Pilger et Melchior Trib. 2. Taxodieae Benth. et Hook. Gen. 6. Metasequoia Miki ex Hu et Cheng Gen. 2. Glyptostrobus Endl. Trib. 5. Cunninghamieae Zucc. Gen. 3. Taxodium Rich. Gen. 7. Cunninghamia R. Br. Trib. 3. Sequoieae Wettstein Gen. 8. Athrotaxis D. Don Gen. 4. Sequoiadendron Buchholz Gen. 9. Taiwania Hayata     

A Phylogenetic Analysis of Families in the Hamamelidae

A cladistic analysis of the families in the Hamamelidae is made in the present paper. As a monophyletic group, the subclass Hamamelidae includes 19 families, namely, the Trochodendraceae, Tetracentraceae, Cercidiphyllaceae, Eupteleaceae, Eucommiaceae, Hamamelidaceae (incl. Rhodoleiaceae and Altingiaceae), Platanaceae, Daphniphyllaceae, Balanopaceae, Didymelaceae, Myrothamnaceae, Buxaceae, Simmondsiaceae, Casuarinaceae, Fagaceae (incl. Nothofagaceae), Betulaceae, Myricaceae, Rhoipteleaceae and Juglandaceae. The Magnoliaceae was selected for outgroup comparison after careful consideration. Thirty-two informative character states were used in this study. Three principles, namely, outgroup comparison, fossil evidence and generally accepted viewpoints of morphological evolution, were used for polarization of the characters. An incompatible number concept was first introduced to evaluate the reliable degree of polarization of the characters and, by this method, the polarization of the three character states was corrected. A data matrix was constructed by the 19 ingroup families and 32 character states. The data matrix was analysed with the Minimal Parallel Evolutionary Method, Maximal Same Step Method (Xu 1989), and Synthetic Method. Three cladograms were constructed and a parsimonious cladogram (Length= 131)was used as the base for discussing the systematic relationships of families in the Hamamelidae. According to the cladogram, the earlist group differented in the subclass Hamamelidae consists of two vesselless wood families, the Trochodendraceae and Tetracentraceae. This result supports the concept proposed by Takhtajan (1987)and Cronquist (1981, 1988)that the Trochodendrales is probably a primitive taxon in the Hamamelidae. As in a clade group, the Cercidiphyllaceae, Eucommiaceae, Balanopaceae and Didymelaceae originated apparently later than the Trochodendrales. The Cercidiphyllaceae diverged earlier in this group, which implies that this family and the Trochodendrales form a primitive group in the subclass. The Cercidiphyllaceae is either placed in Hamamelidales (Cronquist 1981, Thorne 1983), or treated as an independent order (Takhtajan 1987).This analysis suggests that the Cercidiphyllaceae is a relatively isolated taxon, far from the Hamamelidaceae but close to the Trochodendrales in relation. The Eucommiaceae and Didymelaceae are both isolated families and considered as two distinct orders (Takhtajan 1987, Cronquist 1981, 1988).The Balanopaceae is included in the Fagales (Cronquist 1981, 1988) or Pittosporales (Thorne 1983), or treated as a distinct order Balanopales (Takhtajan 1987 ).Obviously the Balanopaceae and Eucommiaceae are not closely related because of the sole synapomorphy (placentation).In fact these four families are more or less isolated taxa and it is probably more reasonable to treat them as independent orders. Cronquist ( 1981, 1988) places the Eupteleaceae, Platanaceae and Myrothamnaceae in the Hamamelidales, while Takhtajan (1987)puts Hamamelidaceae and Platanaceae into the Hamamelidales and treats the Eupteleaceae and Myrothamnaceae as two independent monofamilial orders. These three families are grouped by more synapomorphies (palmateveined, serrate or lobate leaves, deciduous and anemophilous plants)which may indicate their close phylogenetical affinity. A core group of the Hamamelidae includes ten families, among which the Hamamelidaceae originated earlier than the others, so that it is a relatively primitive family. The Betulaceae, Fagaceae and Myricaceae differentiated later than the Hamamelidaceae. The former two are very closely related, and thus thought to be two neighbouring orders by Takhtajan (1987)or included in the Fagales by Cronquist (1981, 1988)and Thorne (1983). The Myricaceae and Fagaceae are connected in the cladogram by only a single synapomorphy (endosperm absent), and therefore the close relationship does not exist between them. The Buxaceae, Simmondsiaceae and Daphniphyllaceae form an advanced group, in which they are weakly linked with each other by only one synapomorphy (pollen grains＜25μm). The Daphniphyllaceae is closely related to the Simmondsiaceae, but the Buxaceae is rather isolated. The Rhoipteleaceae and Juglandaceae share a number of synapomorphies (axile placentation, endosperm absent, embryo larger, fruit indehiscent) , forming a highly specialized group. The opinion that the Juglandales is composed of the Juglandaceae and Rhoipteleaceae(Cronquist 1981; 1988, Lu et Zhang 1990)is confirmed by this analysis. A contrary point of view, which treated them as two distinct orders by Takhtajan (1987), apparently could not be accepted. The Casuarinaceae was regarded as the primitive angiosperm (Engler 1893), but in fact it is a highly reduced and specialized group. It is united with Rhoipteleaceae and Juglandaceae by four synapomorphies, i. e. placentation type, endosperm absent, embryo large and fruit indehiscent. However, the family presents six autapomorphies, and thus the position of the Casuarinaceae as an advanced family is confirmed by this analysis. Finally a strict consensus tree, which represents the phylogenetic relationships of thefamilies in the Hamamelidae, was given as a result of the analysis.     

龙胆属的系统发育分析

本文运用支序分类的原理和方法,对龙胆科龙胆属的属下等级进行了重新归类和系统发育分析。龙胆属是一个单系群,以3项近裔共性为归类依据。性状分析作了性状同源性分析和性状极性分析。性状极化主要以外类群比较、性状相关性及染色体资料为依据,其它方法,如生物重演律原则、地理递进原则以及孢粉形态等也被结合使用。分析结果,双蝴蝶属和蔓龙胆属被选择为外类群,71个性状被选择作为建立数据矩阵的基本资料。使用PAUP程序对矩阵进行了运算,得到4个最简约的谱系分支图,它们均具一致性系数0.637,支序长度为160步,f-比值范围为0.179～0.189,其中具最低f-比值的图被选作为类群归类和讨论亲缘关系的基础。在支序图上龙胆属归为15个组;其中5个组又划分为系,共包括23个系,其余组为单型组,故共有33个属下类群。一个严格的一致性谱系分支图总结了所有的一致点,从而支持了支序分析的结果。     

小车蝗属Oedaleus Fieber昆虫的种间系统发育关系支序分析(直翅目:斑翅蝗科)

在已有比较形态学研究的基础上 ,本文选择了 1 9个性状 ,利用 PAUP程序 ,以支序分析探讨了小车蝗属昆虫的种间系统发育关系。结果红胫小车蝗 ( 1 )与黄胫小车蝗 ( 2 )的亲缘关系很近 ,是 1对姐妹群 ,亚洲小车蝗 ( 3)、黑条小车蝗 ( 4 )、红翅小车蝗( 5)、隆叉小车蝗 ( 6)这 4个种之间的相互关系比较近 ,它们的亲缘关系可表示为 :( ( ( 3,6) 4) 5) ,而台湾小车蝗 ( 7)与其它种类的亲缘关系均较远。     

中国灵猫科的支序系统学分析

选取分布于中国境内的8属9种灵猫科动物的88个骨骼性状、31个外部形态性状和1个行为性状,运用替代外群法分别以赤狐(犬科狐属)和青鼬(鼬科貂属)共同或单独作为外群进行支序分析,得出10个支序图,其步长(TL)为：106～136,一致性指数(CI)为0．581—0．660,保留指数(RI)为0．610—0．714。经合意分析得到4个相似的支序图,其TL为41—136,CI为0．581—0．732,RI为0．610—0．818,结果支持：①斑灵狸、大灵猫、小灵猫构成一个单系群,与传统分类一致(均属灵猫亚科);②椰子狸、花面狸、熊狸构成一个单系群,与传统分类一致(均属长尾狸亚科);③红颊獴、食蟹獴与灵猫科其他种为姐妹群关系(Bootstrap检验支持率100％),建议獴类为一亚科;④长颌带狸拥有较多自近裔性状如三个门齿孔,是一个高度特化的种类,其系统地位有待进一步研究。     

Phylogenetic relationships of glyptosternoid fishes (Siluriformes: Sisoridae) inferred from mitochondrial cytochrome b gene sequences

To explore phylogenetic relationships among glyptosternoid fishes, we determined nucleotide sequences of the complete mitochondrial cytochrome b gene region (1138 base pair). Thirteen species of glyptosternoid fishes and six species of non-glyptosternoids represent 10 sisorid genera were examined. Molecular phylogenetic trees were constructed using the maximum parsimony, minimum evolution, maximum likelihood, and Bayesian methods. Bayesian and maximum likelihood analyses support the monophyly of glyptosternoids, but our hypothesis of internal relationships differs from previous hypothesis. Results indicated that glyptosternoid is a monophyletic group and genera Glyptosternum and Exostoma are two basal species having a primitive position among it. Genera Euchiloglanis and Pareuchiloglanis form a sister-group. Then they form a sister-group with Pseudexostoma plus Oreoglanis. Our result also found that Pareuchiloglanis anteanalis might be considered as the synonyms of Parechiloglanis sinensis, and genus Euchiloglanis might have only one valid species, Euchiloglanis davidi.     

Phylogenetic relationships among the lepidopteran and trichopteran suborders (Insecta) from the embryological standpoint1

Adopting the cladistic method in comparative embryology, 27 embryonic characters were analyzed to reconstruct the phylogenetic relationships among the lepidopteran and trichopteran suborders, viz., Annulipalpia, Integripalpia, Zeugloptera, Dacnonypha, Exoporia, Monotrysia, and Ditrysia. The resultant cladogram is basically coincident with that proposed by Kristensen . The order Trichoptera and Lepidoptera constitute a monophyletic group on the basis of one synapomorphy, the presence of well developed silk glands in embryos. The Trichoptera are separable from the Lepidoptera by the states of four characters. The Trichoptera, as a whole, are quite homogenous, and embryological data provide no rational ground for the separation of this order into the Annulipalpia and Integripalpia at a subordinal level. On the contrary, the embryonic development of the Lepidoptera becomes divergent from the most primitive condition to a specialized one according to suborders. The Zeugloptera are the sister group of all other Lepidoptera, because they share two apomorphies with the latter. The Dacnonypha are the most primitive next to the Zeugloptera, and have a sister-group relationship with the Exoporia + (Monotrysia + Ditrysia), being held together with the latter by five synapomorphies. Although the Exoporia retain almost as many plesiomorphic characters as the Dacnonypha, they have a sister-group relationship with the Monotrysia + Ditrysia, as inferred on the basis of two synapomorphies. The Monotrysia and Ditrysia have a sistergroup relationship, and are the most advanced groups in the Lepidoptera judging from their shared acquisition of many apomorphic characters.     

鲤科鲌亚科鱼类的系统发育及动物地理学

以雅罗鱼亚科为外群,采用分支系统学的原理和方法对鲌亚科17个属(须鳊属除外)进行了系统发育和动物地理学的分析.结果表明,(1)鲌亚科这17个属为一个单系群,并由4个较小的单系群组成;(2)鲌亚科现在的地理分布格局除源自离散事件外,主要源于扩散事件.鲌亚科的特征演化还表明,东亚地区地质历史的反转演化和平行演化可能在鲌亚科系统发育过程中起了重要的作用.     

Redefinition and phylogenetic relationships of the family Pinguipedidae (Teleostei: Perciformes)

The phylogenetic relationships of the family Pinguipedidae plus the genus Cheimarrichthys von Haast, 1874, were studied to redefine the family. Based on a phylogenetic analysis using derived characters belonging to 21 transformation series, accepting the monophyly of Pinguipedidae plus Cheimarrichthys provisionally for the operational procedure of the analysis, it was hypothesized that Cheimarrichthys is not closely related to Parapercis Bleeker, 1863, although these genera had been considered as having a close affinity. Although the Pinguipedidae and Cheimarrichthys share nine derived characters, it was determined that these characters are also found in other trachinoid families. In addition, several families, such as the Leptoscopidae, Uranoscopidae, and Trichodontidae, have many more derived characters in common with Cheimarrichthys than the pinguipedids have with that genus. The conclusion drawn here is that it is not parsimonious to recognize the monophyly of the Pinguipedidae and Cheimarrichthys based on these nine derived characters, and that these characters are not compelling enough to link these groups. The redefined Pinguipedidae is proposed to include the following five genera: Parapercis, Prolatilus Gill, 1865, Pinguipes Cuvier, 1829, Pseudopercis de Miranda-Ribeiro, 1903, and Kochichthys Kamohara, 1960. Cheimarrichthys, excluded from the Pinguipedidae, is put into its own family, Cheimarrichthyidae.     

Molecular phylogeny of the stromateoid fishes (Teleostei: Perciformes) inferred from mitochondrial DNA sequences and compared with morphology-based hypotheses

The phylogenetic relationships among 21 species of stromateoid fishes, representing five families and 13 genera, were reconstructed using 3263bp of mitochondrial DNA sequences, including the posterior half of the 16S rRNA and entire COI and Cytb genes. The resultant molecular phylogenies were compared with previous phylogenetic hypotheses inferred from morphological characters. Molecular phylogenetic trees were constructed using the maximum parsimony, maximum likelihood, and Bayesian methods. All three methods resulted in well-resolved trees with most nodes being supported by moderate to high support values. In contrast to previous morphological analyses, which resulted in non-monophyly of Centrolophidae, all three methods utilized for the present molecular analyses supported the monophyly of Centrolophidae, as well as the reciprocal monophyly of the other stromateoid families, previous morphological hypotheses being rejected by the Templeton and Shimodaira-Hasegawa tests. In addition, the three methods indicated a sister-group relationship between Ariommatidae and Nomeidae. The position of Tetragonuridae was, however, incongruent between the MP method and the ML and Bayesian methods, being placed in the most basal position of Stromateoidei in the former, but occupying a sister relationship to Stromateidae in the latter. Comparison of the molecular phylogenies to previous morphological hypotheses suggested that evolutionary changes in morphological characters have not occurred equally among the stromateoid lineages, the evolution of the centrolophids not having been accompanied by appreciable morphological changes, whereas other stromateoids have undergone considerable morphological changes during their evolutionary history. The molecular phylogenies also shed some light on the evolutionary pattern of the pharyngeal sac, two of the four types of sac corresponding to two main lineages of Stromateoidei. Some taxonomic implications were also discussed.     

蝽科Dalpada属群的系统发育分析(半翅目)

作者对Halyini族中Dalpada属群的9个属级阶元的代表属作了形态学研究,选取头、翅、腹部、雌、雄外生殖器以及受精囊等方面的18项形态特征,运用PAUP程序进行支序分析,得出该属群的属级关系支序图,并对该属群的系统发育进行了讨论。     

Phylogenetic relationships among atheriniform fishes (Teleostei: Atherinomorpha)

We present evidence from adult and larval morphology for the monophyly and relationships of Atheriniformes, using other atherinomorphs, mugilids and acanthomorph fishes as outgroups. Atheriniformes is diagnosed by ten characters (larval: short preanal length, single mid-dorsal row of melanophores; adult: vomerine ventral face concave, long Al muscle tendon to lacrimal, two anterior infraorbital bones, pelvic-rib ligament, pelvic medial plate not extended to anterior end, and second dorsal-fin spine flexible). We recognize six families within the order, the hierarchical relationships among which are: (Atherinopsidae (Notocheiridae (Melanotaeniidae (Atherionidae (Phallostethidae, Atherinidae))))). Other major conclusions include: (1) Atherinopsidae (Menidiinae, Atherinopsinae) is diagnosed by 20 characters (e.g. ethmomaxillary ligament attached to palatine dorsal process, ventral postcleithrum with two dorsal rami); (2) Melanotaeniidae (Bedotiinae (Melanotaeniinae (Telmatherinini, Pseudomugilini))) is diagnosed by six characters (e.g. absence of second dorsal-fin spine, sexual dimorphism in body colour and median-fin development, greater body depth); (3) Dentatherina is in Phallostethidae; (4) Atherinidae (Atherinomorinae (Craterocephalinae, Atherininae)) is diagnosed by three characters (lacrimal notch, ventral postcleithrum between first and second pleural ribs, pelvic ventral spine); (5) Atherinidae and Phallostethidae form the Atherinoidea clade diagnosed by seven characters (e.g. interopercle dorsal process absent, dorsal wings of urohyal absent, ventral postcleithrum laminar, pelvic medial plate extended to anterior end, presence of anal plate). Bedotia, Rhodes , and melanotaeniines are shown to be derived within atheriniforms rather than the plesiomorphic sister groups to a paraphyletic 'atherinoid' group. We also demonstrate that groups traditionally placed in Atherinidae (Menidiinae, Atherininae, Atherioninae, etc.) comprise a paraphyletic assemblage.     

Isoenzyme diversity and phylogenetic relationships among the American beans of the genus Vigna Savi (Fabaceae)

Electrophoretic variation of 13 isoenzyme systems encoded by 25 gene loci has been studied among the 12 American bean species of the genus Vigna, subgenera Sigmoidotropis and Lasiospron, in comparison with the two pantropical species Vigna luteola and V. vexillata of the subgenera Vigna and Plectrotropis, respectively. Of the 202 electromorphs (putative allozymes) discovered, 106 were phylogenetically informative, i.e. shared by at least two taxa, and 96 were taxon-specific. Parsimony, neighbour-joining and UPGMA analyses of the isozyme data as unordered presence–absence characters revealed the division of the American Vigna species into six clades which mostly correspond to the grouping of species into subgenera and sections in the traditional morphology-based systems, except in the placement of species V. gentryi and V. caracalla. The type species of the section Caracallae, V. caracalla, appears on the cladogram as basally paraphyletic in the subgenus Sigmoidotropis, but apart from the V. linearis group of the same section which forms a separate subclade. Vigna gentryi is not linked to other species of the section Sigmoidotropis, but appears in a distant subclade as a sister species to V. adenantha s. l. of the section Leptospron. The subgenus Lasiospron is sister to the pantropical V. vexillata of the African subgenus Plectrotropis.     

Phylogenetic relationships within the Retiolitidae (Graptolithina) and a new genus, Cometograptus

A new retiolitid genus Cometograptus, restricted to the lundgreni Biozone (Late Wenlock), is characterized by a combination of primitive features such as strong transverse rods and ancora sleeve lists with seams facing out, with advanced features such as a long sicula, central, free nema and pustulose bandages. Cometograptus links two Subfamilies of the Family Retiolitidae. The Subfamily Retiolitinae is divided into redefined lineages: Pseudoretiolites, Retiolites, Paraplectograptus, whereas the Subfamily Plectograptinae is divisible into the Sokolovograptus (questionable), Gothograptus, Spinograptus, Plectograptus lineages, and Cometograptus. The Sokolovograptus group, the first retiolitids bearing ancora sleeve lists with seams facing out, did not survive the lundgreni crisis. The only survivors were Upper Wenlock retiolitids of the Gothograptus lineage. The new genus Cometograptus includes Cometograptus koreni n.gen. and n.sp., Cometograptus tomczyki n.gen. and n.sp., as well as the previously described Cometograptus nevadensis, Cometograptus marsupium and Cometograptus apoxys.     

寄生红蜡蚧的扁角跳小蜂属6个种(膜翅目:跳小蜂科)的系统发育支序分析

选用寄生红蜡蚧的扁角跳小蜂属(Anicetus)6个种的14个形态特征和2个其他特征(寄主种数和是否寄生软蚧)作为内群性状,以柯氏花翅跳小蜂(Microterys clauseni)作为外群,运用PAUP(Version 4．0b10)程序做了支序分析。所得支序树状图有如下参数：树长=28;一致性指数=0．6786;稳定性指数=0.4375。6种扁角跳小蜂中,亲缘关系表示为：(A．annulatus,((A．rubensi,(A．beneficus,A．rarisetus)),(A．howardi,A.ohgushii)))。软蚧扁角跳小蜂(A.annulatus)与其他5个种的亲缘关系相对较远,与传统分类学观点一致。