Activity of digestive enzymes of thick-billed murre (Uria lomvia) and common murre (U. aalga) invaded by cestodes

Activities of digestive enzymes (proteases, carbohydrases, acid and alkaline phosphatases) are determined in intestinal mucosa of the thick-billed and common murres Comparative analysis of the obtained results is performed for non-infected and for birds infested by cestodes. It has been established that at invasion by cestode Alcataenia armillaris (Cestoda: Tetrabothriidae), activities of carbohydrase and alkaline phosphatase in intestinal mucosa of the thick-billed murre decreases. Parasitizing of cestodes Tetrabothrius jaegerskieldi (Cestoda: Tetrabothriidae) in intestine of the common murre induces a decrease of saccharase activity. There is studied kinetics of desorption of enzymes from digestive-transport surfaces of the bird intestine. Peculiarities of firmness of enzyme fixation are established on the surface of intestinal mucosa of invaded murres. According to the obtained data, a decrease of the carbohydrase activities in intestine of infested murres is likely to be due to absorption of a part of enzymes hydrolyzing carbohydrates on the surface of cestodes.     

Activity of digestive enzymes of thick-billed guillemot (<Emphasis Type="Italic">Uria lomvia</Emphasis>) and common guillemot (<Emphasis Type="Italic">U. aalge</Emphasis>) invaded by cestodes

Activities of digestive enzymes (proteases, carbohydrases, acid and alkaline phosphatases) are determined in intestinal mucosa of the thick-billed and common guillemots. Comparative analysis of the obtained results is performed for non-infested and for birds infested by cestodes. It has been established that at infestation by cestode Alcataenia armillaris (Cestoda: Tetrabothriidae), activities of carbohydrase and alkaline phosphatase in intestinal mucosa of the thick-billed guillemot decreases. Parasitizing of cestodes Tetrabothrius jaegerskieldi (Cestoda: Tetrabothriidae) in intestine induces a decrease of saccharase activity. There is studied kinetics of desorption of enzymes from digestive-transport surfaces of the bird intestine. Peculiarities of firmness of enzyme fixation are established on the surface of intestinal mucosa of invaded guillemots. According to the obtained data, a decrease of the carbohydrase activities in intestine of infested guillemots is likely to be due to absorption of a part of enzymes hydrolyzing carbohydrates on the surface of cestodes.     

Phylogenetic relationships of phrynosomatid lizards based on nuclear and mitochondrial data,and a revised phylogeny for Sceloporus

Phrynosomatid lizards are among the most common and diverse groups of reptiles in western North America, Mexico, and Central America. Phrynosomatidae includes 136 species in 10 genera. Phrynosomatids are used as model systems in many research programs in evolution and ecology, and much of this research has been undertaken in a comparative phylogenetic framework. However, relationships among many phrynosomatid genera are poorly supported and in conflict between recent studies. Further, previous studies based on mitochondrial DNA sequences suggested that the most species-rich genus (Sceloporus) is possibly paraphyletic with respect to as many as four other genera (Petrosaurus, Sator, Urosaurus, and Uta). Here, we collect new sequence data from five nuclear genes and combine them with published data from one additional nuclear gene and five mitochondrial gene regions. We compare trees from nuclear and mitochondrial data from 37 phrynosomatid taxa, including a “species tree” (from BEST) for the nuclear data. We also present a phylogeny for 122 phrynosomatid species based on maximum likelihood analysis of the combined data, which provides a strongly-supported hypothesis for relationships among most phrynosomatid genera and includes most phrynosomatid species. Our results strongly support the monophyly of Sceloporus (including Sator) and many of the relationships within it. We present a new classification for phrynosomatid lizards and the genus Sceloporus, and offer a new tree with branch lengths for use in comparative studies.     

The activities of digestive enzymes as a determinant factor in the localization of <Emphasis Type="Italic">Tetrabothrius erostris</Emphasis> (Loennberg) (Cestoda: Tetrabothriidae) in the intestine of the herring gull <Emphasis Type="Italic">Larus argentatus</Emphasis> Pontoppidan

We determined the parameters of invasion and localization of Tetrabothrius erostris (Loennberg) (Cestoda: Tetrabothriidae) in the intestine of herring gulls Larus argentatus Pontoppidan (adults and nestlings). Adult herring gulls have T. erostris parasites in the proximal and middle segment of their small intestine. Nestlings have T. erostris in the middle segment of their small intestine. High activities of the digestive enzymes (proteases and glycosidaeses) in these segments of the intestine were found. Apparently, this factor, as well as the physiological peculiarities of the development of T. erostris, determine the localization of tetrabothriids in the intestine of herring gulls.     

Ecomorphological diversification among South American spiny rats (Rodentia; Echimyidae): a phylogenetic and chronological approach

The phylogeny of South American spiny rats (Rodentia; Echimyidae) was studied using the exon 28 of the von Willebrand Factor nuclear gene (vWF). Sequences were analysed separately and in combination with a mitochondrial dataset (cyt b, 12S and 16S rRNAs) used in previous publications. The basal polytomy of echimyids was partially resolved and unexpected intergeneric clades were recovered. Thus, the intimate nested position of Myocastor within echimyids is evidenced. A well-supported clade is identified, including all the arboreal genera, and a group formed by Myocastor, Thrichomys, and Proechimys+Hoplomys. The clustering of Euryzygomatomys+Clyomys with Trinomys is also suggested. On the opposite, the phylogenetic position of Capromys as well as the relationships among arboreal genera remain unclear. Molecular divergence times were estimated using a Bayesian relaxed molecular clock and suggest a Middle Miocene origin for most of modern genera. The ecomorphological diversification of echimyids is discussed in the light of these new results and past environmental modifications in South America.     

Evolution of mitochondrial and ribosomal gene sequences in anophelinae (Diptera: Culicidae): implications for phylogeny reconstruction

In this study, two mitochondrial genes, cyt b and ND5, and the D2 expansion segment of the 28S nuclear ribosomal gene were used to reconstruct a phylogeny of the mosquito subfamily Anophelinae. The ingroup consisted of all three genera of Anophelinae and five of six subgenera of Anopheles. Six genera of Culicinae were used as the outgroup. Extreme conservation at the protein level coupled with rapid saturation of synonymous positions probably accounted for the lack of meaningful phylogenetic signal in the cyt b gene. In contrast, abundant variation at all codon positions of the ND5 gene allowed recovery of the basal and most of the recent relationships. Phylogenetic analysis of D2 produced results consistent with those of ND5. Combined analysis indicated well-supported monophyletic Anophelinae (with Chagasia basal), Anopheles + Bironella, and subgeneric clades within the genus Anopheles. Moreover, subgenera Nyssorhynchus and Kerteszia were supported as a monophyletic lineage. The Kishino-Hasegawa test could not reject the monophyly of Anopheles, whereas the recently proposed hypothesis of close affinity of Bironella to the subgenus Anopheles was rejected by the analyses of ND5 and combined data sets. The lack of resolution of Bironella and Anopheles clades, or basal relationships among subgeneric clades within Anopheles, suggests their rapid diversification. Recovery of relationships consistent with morphology and previous molecular studies provides evidence of substantial phylogenetic signal in D2 and ND5 genes at levels of divergence from closely related species to subfamily in mosquitoes.     

Phylogenetic relationships among microgastrine braconid wasp genera based on data from the 16S, COI and 28S genes and morphology

Abstract Phylogenetic relationships among the genera of the large braconid wasp subfamily Microgastrinae were explored using DNA sequence data from the mitochondrial large ribosomal subunit (16S), nuclear large ribosomal subunit (28S) and mitochondrial cytochrome oxidase (COI) genes, along with morphological characters, both new and from previous studies. The taxonomic history of this group of wasps is reviewed, along with a critique of previous phylogenetic studies on the group. Molecular data were sampled from forty-six species representing twenty-six genera of microgastrines, plus three species representing the close outgroup taxa Cardiochilinae and Miracinae. Some 2300 base pairs of aligned sequence were obtained per taxon from the three genes. In addition, fifty-three morphological characters were coded for all known genera, including two undescribed genera, except Semionis Nixon (known from only a single male type specimen). Relationships among several groups of genera are clarified and challenge some major assumptions made in earlier classifications. In particular, it is clear that dependence on one or a few major morphological character systems oversimplifies relationships, and can lead to misleading results. Despite the large amount of data analysed, basal divergences within the subfamily remain poorly resolved and essentially unsupported in any rigorous statistical sense.     

基于16S rDNA和28S rDNA D2基因序列与形态特征联合分析的中国角顶叶蝉亚科系统发育研究（半翅目： 叶蝉科）

首次在国内利用28S rDNA D2区段和16S rDNA基因序列,结合50个形态特征对角顶叶蝉亚科(Deltocephalinae)［半翅目（Hemiptera）: 叶蝉科（Cicadellidae）］19个属进行系统发育分析研究。从无水乙醇浸泡保存的标本中提取基因组DNA并扩增了19个内群和1种外群Typhlocybinae［半翅目(Hemiptera): 叶蝉科(Cicadellidae)］种类的28S rDNA D2基因片段并测序,同时扩增了16S rDNA基因片段并测序11条,采用了GenBank中1个种类的16S rDNA同源序列。采用PAUP*4.0和MrBayes3.0两个分析软件和3种建树方法,利用同源28S D2 rDNA和16S rDNA两个基因序列与形态特征结合进行系统发育分析研究。分析结果表明,二叉叶蝉族Macrostelini是一个单系,并在角顶叶蝉亚科的系统发育中处于基部的位置,是内群中最原始的族;角顶叶蝉族Deltocephalini中除了纹翅叶蝉属Nakaharanus,其余各属构成单系;殃叶蝉族Euscelini内属的归属比较混乱,可能是一个并系群,属间差异有待进一步研究。隆额叶蝉族Paralimnini与顶带叶蝉族Athysanini是姐妹群。带叶蝉属Scaphoideus与纹翅叶蝉属Nakaharanus是姐妹群,二者与木叶蝉属Phlogotettix的关系最近,三者构成一个单系,建议将三者归为带叶蝉族Scaphoideini。研究结果还表明,小眼叶蝉族Xestocephalini和Balcluthini的系统发育位置不明,有待进一步研究。     

雪雀属系统发育关系的研究（雀形目：文鸟科)

雷雀属Montifringilla7种36个形态特征作为分类单元,进行支序分析。以麻雀属Passer、石雀属Petronia部分种类作为比较外群,讨论雪雀属的分类地位及其与近缘属麻雀属、石雀属系统发育关系;分析了雪雀属内种间的系统发育。支序分析的结果表明雪雀属为单系群,建议将雪雀属划分成两个亚属,讨论了雪雀属种类在进货上的位置,在3个近缘属的系统发育关系中,麻雀属类群较原始,雪雀属类群较进化,石雀属在进货中介于二之间。     

Phylogenetic analysis of the Amphitritinae (Polycnaeta: Terebellidae)

The phylogenetic relationships of the Amphitritinae (Polychaeta: Terebellidae) were studied using parsimony analysis of 22 external morphological characters. To choose outgroups to polarize the characters, I carried out a preliminary analysis of the relationships of the four terebellid subfamilies and the Trichobranchidae. The single most parsimonious tree from the analysis supports monophyly of the Terebellidae by the presence of ventral glandular shields. However, this character is homoplasious within the Terebellomorpha, and further evaluation of the Terebellidae is recommended. Artacama and Thelepus were chosen as outgroups for the analysis of amphitritine genera. The generic level analysis yielded seven equally parsimonious trees, which are consistent in their topologies except for the relationships among seven genera in one large clade. In all trees, Artacama is the sister taxon to a large clade within the Amphitritinae; the Artacaminae is therefore synonymized with the Amphitritinae, which is diagnosed by the presence of double rows of uncini. Within the Amphitritinae, the status of several monotypic genera is questioned; plesiomorphic character states indicated by the analysis are discussed. The results presented are offered as working hypotheses of the relationships among amphitritine genera. The large number of homoplasies indicated by the analysis emphasizes the need to further evaluate these hypotheses using additional characters. With a robust phylogenetic hypothesis of amphitritine relationships, a re-classification of the group based on apomorphic character states can be undertaken, and questions regarding the evolution of morphological characters, reproductive modes, or biogeographical patterns can be properly addressed.     

Phylogeny and biogeography of ice crawlers (Insecta: Grylloblattodea) based on six molecular loci: designating conservation status for Grylloblattodea species

Ice crawlers (Insecta: Grylloblattodea) are rarely encountered insects that consist of five genera representing 26 species from North America and Asia. Asian grylloblattids are the most diverse, but North American ice crawlers (genus Grylloblatta) are known for their adaptation to cold conditions. Phylogenetic relationships among grylloblattid species and genera are not known. Six genes were sampled in 35 individuals for 18S rRNA, 28S rRNA, histone 3, 12S rRNA, 16S rRNA, and cytochrome oxidase II from 21 populations of Grylloblatta, three populations from Japan (genus Galloisiana), and three populations from Russia (genus Grylloblattina). Phylogenetic analysis of these data with two mantophasmid outgroups in POY supported monophyletic genera, with Grylloblatta as sister to Grylloblattina. Grylloblatta was shown to contain two major lineages: a clade in Northern California and Oregon and a clade in Washington and Oregon. One new species and six candidate species are proposed. IUCN Red List Conservation Criteria were implemented to designate conservation status for each lineage.     

Molecular systematics of sponges (Porifera)

The first application of molecular systematics to sponges was in the 1980s, using allozyme divergence to dis-criminate between conspecific and congeneric sponge populations. Since this time, a fairly large database has been accumulated and, although the first findings seemed to indicate that sponge species were genetically more divergent than those of other marine invertebrates, a recent review of the available dataset indicates that levels of interspecific gene identities in most sponges fall within the normal range found between species of other invertebrates. Nevertheless, some sponge genera have species that are extremely divergent from each other, suggesting a possible polyphyly of these genera. In the 1990s, molecular studies comparing sequences of ribosomal RNA have been used to reappraise the phylogenetic relationships among sponge genera, families, orders and classes. Both the 18S small subunit and the 28S large subunit rRNA genes have been sequenced (41 complete or partial and 75 partial sequences, respectively). Sequences of 18S rRNA show good support for Porifera being true Metazoa, but they are not informative for resolving relationships among genera, families or orders. 28S rRNA domains D1 and D2 appear to be more informative for the terminal nodes and provide resolution for internal topologies in sufficiently closely related species, but the deep nodes between orders or classes cannot be resolved using this molecule. Recently, a more conserved gene, Hsp70, has been used to try to resolve the relationships in the deep nodes. Metazoan monophyly is very well supported. Nevertheless, the divergence between the three classes of Porifera, as well as the divergence between Porifera, Cnidaria and Ctenophora, is not resolved. Research is in progress using other genes such as those of the homeodomain, the tyrosine kinase domain, and those coding for the aggregation factor. For the moment the dataset for these genes is too restricted to resolve the phylogenetic relationships of these phyla. However, whichever the genes, the phylogenies obtained suggest that Porifera could be paraphyletic and that the phylogenetic relationships of most of the families and orders of the Demospongiae have to be reassessed. The Calcarea and Hexactinellida are still to be studied at the molecular level.     

Comprehensive molecular phylogeny of the sub-family Dipterocarpoideae (Dipterocarpaceae) based on chloroplast DNA sequences

Dipterocarpoideae, the largest sub-family of well-known plant family Dipterocarpaceae, dominates in South Asian rain forests. Although several previous studies addressed the phylogeny of the Dipterocarpaceae family, relationships among many of its genera from the Dipterocarpoideae sub-family are still not well understood. In particular, little is known about the relationships of the genera Vateriopsis, Stemonoporus, Vateria and inconsistence remains between phylogenetic results and taxonomic classifications of Shorea and Hopea species. We studied molecular phylogeny of the sub-family Dipterocarpoideae using the trnL-trnF spacer, trnL intron and the matK gene sequences of chloroplast DNA (cpDNA). This study is the first comprehensive phylogeny reconstruction for the sub-family Dipterocarpoideae based on cpDNA, as it includes most genera (14) and a large number of species (79) with most species endemic to Sri Lanka, as well as one species from Seychelles and one species from the genus Monotes from Madagascar. Phylogenetic trees were constructed using the Neighbor Joining (NJ) and Maximum Likelihood (ML) methods using combined set of sequences including all three cpDNA regions. The topologies of the NJ and ML trees were to a certain extent, consistent with the current taxonomy of Dipterocarpoideae based on morphology and with previous molecular phylogenies based on cpDNA. Furthermore, our results provided new evidence regarding the relationships of the following genera: Vateriopsis and Stemonoporus and about the validity of the previous morphology based classifications of Shorea species. In addition, the topology of our trees was consistent with the classification of Shorea species proposed by Maury (1978), Maury-Lechon (1979) and Symington (1943). Finally, our results provided evidence for the affinity of the genus Monotes to Asian Dipterocarpoideae rather than to Tiliaceae and indicated that it is a good candidate for outgroup species for future studies of the former sub-family.     

Phylogeny and classification of the Culicidae (Diptera)

The generic relationships and higher classification of the family Culicidae are examined on the basis of a phylogenetic analysis. New and traditional morphological characters studied and compared throughout the Culicidae resulted in the acquisition of character data relative to the majority of species within each genus. Polymorphisms and morphological observations are discussed and additional information and illustrations are provided for the majority of characters and their character states. The analysis of seventy-three adult, pupal and fourth-instar larval characters coded for the thirty-eight currently recognized genera of mosquitoes resulted in relationships and groupings which differ significantly from traditional hypotheses. The analysis supports the monophyly of the subfamily Anophelinae and the tribes Culicini and Sabethini. The Anophelinae form the most basal clade of the family. The results indicate that Aedini is a paraphyletic assemblage with respect to the Mansoniini, each of which is monophyletic in itself. The Aedini + Mansoniini form a sister group to the Culicini + Sabethini, with the Aedini and the Culicini placed in ancestral relationships to the Mansoniini and the Sabethini, respectively. Based on the topography of generic relationships among more 'generalized' mosquitoes, the boundaries and relationships of the tribes Aedeomyiini, Uranotaeniini, Ficalbiini, Hodgesiini, Orthopodomyiini and Culisetini appear to be problematic. Relationships between genera of the tribe Aedini are generally poorly resolved due to a significant amount of polymorphism, especially within the genus Aedes as currently defined. There is no support for separate subfamily recognition for the genus Toxorhynchites , which is downgraded to tribal status as a result of the analysis. The results are discussed in relation to previous hypotheses based on subjective inference and cladistic analyses.     

Phylogenetic Analysis of the Diplomonadida (Wenyon, 1926) Brugerolle, 1975: Evidence for Heterochrony in Protozoa and Against Giardia lamblia as a "Missing Link"

ABSTRACT. A suite of 23 ultrastructural characters was used in a phylogenetic analysis of the protozoan order Diplomonadida. A single most parsimonious solution was found, with a length of 38 transformations and a consistency index of 0.84. The cladogram supports previous hypotheses of the relationships of the genera in the suborder Diplomonadina, as well as the inclusion of the genera Enteromonas and Trimitus in the order. Heterochrony is suggested in the change to binary axial symmetry, as hypermorphosis resulting from delayed cytokinesis in the ancestor. Hypotheses regarding a pivotal position for Giardia lamblia in the evolution of eukaryotes are inconsistent with the phylogeny proposed here.     

Phylogenetic analysis of the Diplomonadida (Wenyon, 1926) Brugerolle, 1975: evidence for heterochrony in protozoa and against Giardia lamblia as a "missing link".

A suite of 23 ultrastructural characters was used in a phylogenetic analysis of the protozoan order Diplomonadida. A single most parsimonious solution was found, with a length of 38 transformations and a consistency index of 0.84. The cladogram supports previous hypotheses of the relationships of the genera in the suborder Diplomonadina, as well as the inclusion of the genera Enteromonas and Trimitus in the order. Heterochrony is suggested in the change to binary axial symmetry, as hypermorphosis resulting from delayed cytokinesis in the ancestor. Hypotheses regarding a pivotal position for Giardia lamblia in the evolution of eukaryotes are inconsistent with the phylogeny proposed here.     

Evolutionary relationships among the protostrongylidae (Nematoda: Metastrongyloidea) as inferred from morphological characters, with consideration of parasite-host coevolution.

The phylogeny of nematodes in the family Protostrongylidae (Nematoda: Metastrongyloidea) was reconstructed by cladistic analysis of 28 binary and multistate characters derived from comparative morphology. Analyses were hierarchical, and examined (1) relationships among genera, including 13 ingroup taxa and Metastrongylidae as an outgroup (single tree, 78 steps, consistency index [CI] = 0.705); and (2) relationships among genera and species groups, including 21 ingroup taxa and Metastrongylus apri as an outgroup (single tree, 76 steps, CI = 0.582). In the species-level tree, Protostrongylidae was divided into 2 major clades, 1 containing the subfamilies Muelleriinae (including the recently described Umingmtakstrangylus pallikuukensis), Elaphostrongylinae, and the Varestrongylinae (excluding Pneumocaulus kadenazii). Varestrongylus was paraphyletic as it included Pneumostrongylus calcaratus. The second major clade consisted of a paraphyletic group containing Protostrongylus spp. and Spiculocaulus leuckarti and, basal to this subclade, several other individual protostrongylid lineages. The various subclades generally correspond to the subfamilial divisions of the Protostrongylidae. The Neostrongylinae, however, is not supported as Neostrongylus and Orthostrongylus are not sister groups. Based on a large number of hypothesized synapomorphies, the elaphostrongylines appear to be a highly derived group of protostrongylids, a feature potentially correlated with their habitat localization in muscular and nervous tissues. The generic-level tree retained most of the primary structure revealed among the species but excluded the varestrongylines from the Muelleriinae + Elaphostrongylinae subclade. Artiodactyles of the family Cervidae are considered basal hosts for protostrongylids; secondary colonization in Caprini, Rupicaprini, and among lagomorphs is postulated.     

A new genus and species of bothriurid scorpion from the Brandberg Massif, Namibia, with a reanalysis of bothriurid phylogeny and a discussion of the phylogenetic position of Lisposoma Lawrence

Abstract. Brandbergia haringtoni , a new genus and species of bothriurid scorpion, is described from the Brandberg Massif, northwestern Namibia. A cladistic analysis, based on seventy-four morphological characters scored for thirty-one exemplar species representing all genera of Bothriuridae, and one genus from each of the six remaining families of Scorpionoidea, was conducted to test the phylogenetic placement of the new genus and whether it affects the internal relationships of Bothriuridae. The available data demonstrate, under a range of weighting regimes, that the new genus is the most basal bothriurid, and confirm the scheme of relationships among the remaining bothriurid genera that was recovered in a previous analysis: ( Brandbergia ( Lisposoma ( Thestylus ( Phoniocercus ( Tehuankea ( Cercophonius  +  Urophonius ) ( Bothriurus  +  Brachistosternus  + Orobothriurus  +  Centromachetes ( Timogenes  +  Vachonia ))))))). On the basis of this evidence, Lourenço's recent proposal of family Lisposomidae for Lisposoma is rejected and Lisposomidae is synonymized with Bothriuridae. The implications of the phylogeny for understanding the biogeography of Bothriuridae are discussed.     

Molecular phylogenetics of the family Cyprinidae (Actinopterygii: Cypriniformes) as evidenced by sequence variation in the first intron of S7 ribosomal protein-coding gene: further evidence from a nuclear gene of the systematic chaos in the family

The family Cyprinidae is the largest freshwater fish group in the world, including over 200 genera and 2100 species. The phylogenetic relationships of major clades within this family are simply poorly understood, largely because of the overwhelming diversity of the group; however, several investigators have advanced different hypotheses of relationships that pre- and post-date the use of shared-derived characters as advocated through phylogenetic systematics. As expected, most previous investigations used morphological characters. Recently, mitochondrial DNA (mtDNA) sequences and combined morphological and mtDNA investigations have been used to explore and advance our understanding of species relationships and test monophyletic groupings. Limitations of these studies include limited taxon sampling and a strict reliance upon maternally inherited mtDNA variation. The present study is the first endeavor to recover the phylogenetic relationships of the 12 previously recognized monophyletic subfamilies within the Cyprinidae using newly sequenced nuclear DNA (nDNA) for over 50 species representing members of the different previously hypothesized subfamily and family groupings within the Cyprinidae and from other cypriniform families as outgroup taxa. Hypothesized phylogenetic relationships are constructed using maximum parsimony and Basyesian analyses of 1042 sites, of which 971 sites were variable and 790 were phylogenetically informative. Using other appropriate cypriniform taxa of the families Catostomidae (Myxocyprinus asiaticus), Gyrinocheilidae (Gyrinocheilus aymonieri), and Balitoridae (Nemacheilus sp. and Beaufortia kweichowensis) as outgroups, the Cyprinidae is resolved as a monophyletic group. Within the family the genera Raiamas, Barilius, Danio, and Rasbora, representing many of the tropical cyprinids, represent basal members of the family. All other species can be classified into variably supported and resolved monophyletic lineages, depending upon analysis, that are consistent with or correspond to Barbini and Leuciscini. The Barbini includes taxa traditionally aligned with the subfamily Cyprininae sensu previous morphological revisionary studies by Howes (Barbinae, Labeoninae, Cyprininae and Schizothoracinae). The Leuciscini includes six other subfamilies that are mainly divided into three separate lineages. The relationships among genera and subfamilies are discussed as well as the possible origins of major lineages.