12 Phylogeny of the euglenoid loricate genera trachelomonas and strombomonas based on morphological and molecular data

Since the separation of the Trachelomonas subgroup “Saccatae” into a new genus, Strombomonas Deflandre (1930), there has been some question as to its validity. Deflandre's separation was based on morphological characteristics such as the shape of the lorica, lack of a distinctive collar, possession of a tailpiece, lack of ornamentation, and ability to aggregate particles on the lorica. Recent molecular analyses indicated that the loricate taxa were monophyletic, but few species have been sampled. The LSU rDNA from eleven Strombomonas and thirty‐eight Trachelomonas species was sequenced to evaluate the monophyly of the two genera. Bayesian and maximum‐likelihood analyses found one monophyletic clade for each genus. The Trachelomonas clade was weakly supported, but had five strongly supported subclades. Morphological characters, such as lorica development and pellicle strip reduction, also supported separation of the genera. Lorica development in Strombomonas occurred from the anterior of the cell to the posterior, forming a shroud over the protoplast whereas in Trachelomonas, a layer of mucilage was excreted over the entire protoplast, followed by creation of the collar at the anterior end. Taxa from both genera underwent exponential strip reduction at the anterior and posterior poles. In Strombomonas, only one reduction was visible in the anterior pole, while in most Trachelomonas species, two reductions were visible. Likewise, Strombomonas species possessed two whorls of strip reduction in their posterior end compared to a single whorl of strip reduction in Trachelomonas species. The combined morphological and molecular data support the retention of Trachelomonas and Strombomonas as separate genera.     

MORPHOLOGICAL SEPARATION OF THE EUGLENOID GENERA TRACHELOMONAS AND STROMBOMONAS (EUGLENOPHYTA) BASED ON LORICA DEVELOPMENT AND POSTERIOR STRIP REDUCTION1

Since the separation of the Trachelomonas Ehrenberg subgroup “Saccatae” into a new genus, Strombomonas Deflandre (1930) , there has been some question as to its validity. Deflandre's separation was based entirely on characteristics of the lorica, including the shape of the lorica, the lack of a distinctive collar, possession of a tailpiece, lack of ornamentation, and the ability of Strombomonas species to aggregate particles on the surface of the lorica. Recent molecular analyses indicated that the loricate taxa (Trachelomonas and Strombomonas) formed a single monophyletic clade; however, the phylogenetic relationship of Strombomonas to Trachelomonas remains unclear because only two Strombomonas taxa have been sequenced to date. In this study, we evaluated the monophyly of the loricate genera using two sets of morphological characters, lorica development and pellicle strip reduction. Lorica development in Strombomonas occurred from the anterior of the cell to the posterior, forming a shroud over the protoplast. In Trachelomonas, a layer of mucilage was excreted over the entire protoplast, followed by creation of the collar at the anterior end. Taxa from both genera underwent exponential strip reduction at the anterior and posterior poles. In Strombomonas only one reduction was visible in the anterior pole, whereas in most Trachelomonas species two reductions were visible. Likewise, Strombomonas species possessed two whorls of strip reduction in the posterior end compared with a single whorl of strip reduction in Trachelomonas species. These morphological characters support the separation of Trachelomonas and Strombomonas as distinct genera.     

Developmental studies on the loricate choanoflagellateStephanoeca diplocostata ellis

Summary Cells ofStephanoeca diplocostata comprise a colourless, flagellated, protoplast lodged in a lorica made of siliceous costae. The single anterior flagellum creates a water current from which bacteria and other food particles are filtered by the collar and ingested by linguiform pseudopodia that arise from the protoplast at the base of the collar. A waist divides the lorica into two chambers, the anterior of which contains three transverse and 17–20 longitudinal costae whereas the posterior chamber comprises two systems of spirally deflected costae and on some cells a pedicel at the hind end. Between 150–185 costal strips of similar length form the lorica. A thin investment covers the inner surface of the posterior chamber and lower part of the anterior chamber and joins with the protoplast at the level of the waist. Costal strips are produced within membrane-bounded vesicles in the peripheral cytoplasm and, although the origin of these vesicles is unknown, there is usually a close association with the Golgi apparatus. Once complete, strips are apparently released sideways through the plasmalemma into the cavity of the posterior lorica chamber. Later, bundles of strips are transferred to the top of the inner surface of the collar where they collectively form a horizontal ring. When sufficient strips to form a lorica have been accumulated at the top of the collar, cell division proceeds.     

RED SNOW CAUSED BY A NEW SPECIES OF TRACHELOMONAS12

Trachelomonas kolii sp. nov., a member of the Euglenophyta, was the causative agent of red snow near Mt. Bachelor, Oregon. The cells are uniflagellate or without flagella and, in general, exhibit the characteristics of the genus Trachelomonas. The lorica, when present, is transparent, elongate, reticulate, and has a funnel-shaped collar at the anterior end. The protoplast is completely filled with red pigment. Red spherical resting stages that appear similar to the resting stages of the ubiquitous snow alga Chlamydomonas nivalis (Bauer) Wille are described.     

囊裸藻属与陀螺藻属囊壳微细结构及元素组成比较

对采自自然水体中囊裸藻属(Trachelomonas)的8种和陀螺藻属(Strombomonas)的两种藻类的囊壳微细结构和元素组成进行了比较分析,并探讨了这两个属的系统演化关系。结果表明：组成囊壳的矿质元素主要是铁和硅,在陀螺藻和表面粗糙的囊裸藻中,硅的含量较高,铁的含量相对较低;在大多数囊裸藻中,尤其是表面致密、光滑的种类中,铁的含量较高,硅的含量较低,矿质元素的组成与囊壳的颜色无直接关系。建议将陀螺藻属(Strombomonas)撤消,原陀螺藻属的种作为囊裸藻属(Trachelomonas)中的一个种组(Group),它们是裸藻科中最原始的具囊壳的类群。     

Reconciling cladistic and genetic analyses in choreotrichid ciliates (Ciliophora, Spirotricha, Oligotrichea)

Fifty-six features of halteriid, oligotrichid, and choreotrichid ciliates are cladistically analysed, including an updated hypothesis about the evolution of the somatic ciliary patterns. Based on its morphology, Lynnella clusters with Parastrombidinopsis, Parastrombidium, and Strombidinopsis, while it is basal to the other choreotrichids in the molecular phylogenies. The two clusters of Favella species in small subunit rRNA gene trees are supported by morphological features, justifying a separation at genus and family level. The genus Favella has a smooth lorica surface and a somatic ciliary pattern comprising a left and lateral ciliary field as well as two dorsal kineties and a monokinetidal ventral kinety abutting on the right ciliary field. The new genus Schmidingerella n. gen., established for the second Favella cluster, groups with Metacylis and Rhabdonella in the molecular trees. It differs from Favella in (i) a lorica wall with reticulate surface ridges and minute openings and (ii) a ventral kinety that is distinctly apart from the right ciliary field and composed of a monokinetidal anterior and a dikinetidal posterior portion. The genera Codonaria, Codonella, and Codonellopsis are affiliated with the family Dictyocystidae, whose diagnosis is improved to include the lorica sac.     

中国具囊壳裸藻类的新种类

本文报道了在我国首次发现的１３个具囊壳裸藻类的新种类,在它们之中隶属于囊裸藻属的有５个新变种;属于陀螺藻属的有８个新变种。     

Kakoeca antarctica gen. et sp.n., a loricate choanoflagellate (Acanthoecidae, Choanoflagellida) from Antarctic sea ice with a unique protoplast suspensory membrane

A new genus and species of loricate choanoflagellate, Kakoeca aniarctica Buck & Marchant gen. et sp.n. grown in rough culture from an Antarctic sea ice innoculum is described. This organism has a distinctive lorica morphology consisting of more than 200 costal strips arranged in transverse and longitudinal costae that arc perpendicular to one another in the posterior portion of the lorica. The transverse costae show declination with respect to the lorica axis in the anterior part of the lorica. The cell is suspended in the lorica by a robust protoplast suspensory membrane. This membrane blocks water flow from the posterior of the lorica necessitating water entry through the side of the lorica, an area where the maximum sized apertures in the lorica are found. Terminology (lorica lining and protoplast suspensory) is suggested for the two types of lorica membranes which have been found associated with loricas.     

PHYLOGENY OF THE EUGLENOPHYTES: ANALYSIS USING SSU RDNA AND ULTRASTRUCTURAL CHARACTERS

The use of both molecular and morphological data to determine relationships among the euglenoids is vital for a complete understanding of their phylogeny, and the development of an accurate taxonomy. Analyses of the SSU (18S) rDNA from 12 euglenoid genera have resulted in tree topologies that are in agreement with many defining morphological characters. The euglenoid lineage is formed by phagotrophic euglenoids at its base, followed by the divergence of phototrophs that in-turn gave rise to osmotrophs. The photosynthetic lineage is anchored by euglenoids with two emergent flagella, Eutreptia and Eutreptiella , while the remainder of the lineage is composed of euglenoids with a single emergent flagellum. Among the photosynthetic euglenoids with a single emergent flagellum those that secrete mucilaginous stalks, Colacium , or form a lorica, Trachelomonas and Strombomonas , are closely associated. The remaining photosynthetic genera Euglena , Phacus , and Lepocinclis are intermixed with each other and the osmotrophic genera Astasia , and Khawkinea. Hence, they are not monophyletic, sensu Hennig. To reinforce molecular phylogenies, a robust morphological character database is necessary. For taxa with complex internal structures complete serial reconstruction is required. Serial reconstruction of the flagellar and feeding apparatuses in Ploeotia costata illustrate this necessity. Originally described as having both an MTR (Type I) and a Type II feeding apparatus, reconstruction has shown P. costata to have a single, Type II, feeding apparatus. Moreover, the Type II now appears to be an autapomorphy for Ploeotia species, while euglenoid feeding apparatuses, in toto, appear to form a continuum of structural types.     

PHYLOGENY OF THE EUGLENOID LORICATE GENERA TRACHELOMONAS AND STROMBOMONAS (EUGLENOPHYTA) INFERRED FROM NUCLEAR SSU AND LSU rDNA1

Previous studies using the nuclear SSU rDNA and partial LSU rDNA have demonstrated that the euglenoid loricate taxa form a monophyletic clade within the photosynthetic euglenoid lineage. It was unclear, however, whether the loricate genera Trachelomonas and Strombomonas were monophyletic. In order to determine the relationships among the loricate taxa, SSU and LSU nuclear rDNA sequences were obtained for eight Strombomonas and 25 Trachelomonas strains and combined in a multigene phylogenetic analysis. Conserved regions of the aligned data set were used to generate maximum‐likelihood (ML) and Bayesian phylogenies. Both methods recovered a strongly supported monophyletic loricate clade with Strombomonas and Trachelomonas species separated into two sister clades. Taxa in the genus Strombomonas sorted into three subclades. Within the genus Trachelomonas, five strongly supported subclades were recovered in all analyses. Key morphological features could be attributed to each of the subclades, with the major separation being that all of the spine‐bearing taxa were located in two sister subclades, while the more rounded, spineless taxa formed the remaining three subclades. The separation of genera and subclades was supported by 42 distinct molecular signatures (33 in Trachelomonas and nine in Strombomonas). The morphological and molecular data supported the retention of Trachelomonas and Strombomonas as separate loricate genera.     

Molecular evolution of southern North American Cyprinidae (Actinopterygii), with the description of the new genus Tampichthys from central Mexico

Most of the recognized species of the genus Dionda inhabit drainages of the Gulf of Mexico from central Mexico to central Texas, USA, and have been considered a monophyletic group based on morphological, osteological, and allozyme investigations. Phylogenetic relationships of 15 species of Dionda and 34 species from closely related genera were inferred from one mitochondrial (cytb) and three nuclear gene sequences (S7, Rhodopsin, Rag1) totaling 4487 nucleotides. Separate analyses of all four genes yield congruent phylogenies; however the 15 putative species of Dionda evaluated were never recovered as a monophyletic group when species from nine related genera were included in the analyses. Among the ingroup taxa, one well-supported and highly divergent clade is consistently recognized and consists of six recognized and three undescribed northern species currently recognized in the genus Dionda. These nine species inhabit present or past tributaries of the Rio Grande basin of northern Mexico and southern USA, and were recovered as a basal clade in all analyses. Another large, also strongly supported clade, consisting of seven genera, include five southern recognized species currently in the genus Dionda, forming the sister group to the Codoma clade. These five species comprise the "Southern Dionda clade" and inhabit headwaters of the Pánuco-Tamesí drainage and some adjacent coastal rivers in the Tampico Embayment. The consistent and repeated identification of eight different clades recovered in most of the separate gene analyses strongly supports a division of the non-natural genus Dionda. A new genus, Tampichthys, is proposed for the clade of species endemic to east-central Mexico and formerly in Dionda. Tampichthys and the putative monotypic genus Codoma are more related to Mexican species of the genera Cyprinella and Notropis than to other species referred to Dionda sensu stricto.     

混合营养型马勒姆杯棕鞭藻的重描述及其系统发育分析

马勒姆杯棕鞭藻(Poterioochromonas malhamensis)是典型的混合营养型单细胞鞭毛藻, 因个体微小且形态多变, 该鞭毛藻的正确鉴定十分困难, 且至今针对该物种一直没有详细全面的形态描述。从小球藻的大规模培养中分离出一株捕食性鞭毛藻P. malhamensis CMBB008, 利用荧光染色、扫描和透射电镜等方法, 对其形态学特征进行了详尽的描述, 并获取该物种的SSU rDNA和rbcL基因序列用于探讨该物种的系统发育地位。文章在前人形态描述的基础上, 详细记录了囊壳和叶绿体的形态: 囊壳柄长18.3—47.5 μm, 杯体宽8.5—11.3 μm, 深6.3—10.7 μm; 叶绿体1—2个, 为中间桥接的二裂片状。文章还结合光镜和电镜观察首次报道了该物种的硅质胞囊, 发现其具有独特的三层领状结构和顶端孔塞结构, 并揭示了其主要细胞器的超微结构及重要代谢产物, 如油滴和金藻昆布多糖液泡等。通过不同种株间的比较研究发现, 作为Poterioochromonas属重要形态特征的囊壳和叶绿体, 由于形态变异性高, 不适合用于P. malhamensis的鉴定, 而胞囊形态很可能作为Poterioochromonas种间区分的重要形态特征。分子系统发育分析显示Poterioochromonas所有株系聚为一大支, 表明该属是单系类群。P. malhamensis CMBB008位于模式株P. malhamensis SAG933-1a所在的大分支内。Poterioochromonas支系可分为P. malhamensis和P. stipitata两个姊妹群, P. malhamensis支系可进一步分为两个姊妹支, 其中CCMP3181可能并不是P. malhamensis。研究补充和完善了P. malhamensis的形态和分子数据, 并从形态学和分子系统发育学角度为Poterioochromonas属的分类研究提供重要的参考信息。     

Phylogeny and taxonomic revision of plastid-containing euglenophytes based on SSU rDNA sequence comparisons and synapomorphic signatures in the SSU rRNA secondary structure

Sequence comparisons and a revised classification of the Euglenophyceae were based on 92 new SSU rDNA sequences obtained from strains of Euglena, Astasia, Phacus, Trachelomonas, Colacium, Cryptoglena, Lepocinclis, Eutreptia, Eutreptiella and Tetreutreptia. Sequence data also provided molecular signatures for taxa from genus to class level in the SSU rRNA secondary structure, revealed by a novel approach (search for non-homoplasious synapomorphies) and used for taxonomic diagnoses. Photosynthetic euglenoids and secondary heterotrophs formed a clade, designated as Euglenophyceae (emend.) with two orders: Euglenales and Eutreptiales. The mostly marine Eutreptiales (Eutreptia, Eutreptiella; not Distigma) comprised taxa with two or four emergent flagella (the quadriflagellate Tetreutreptia was integrated within Eutreptiella). The Euglenales (freshwater genera with < or = one emergent flagellum) formed nine clades and two individual branches (single strains); however, only two clades were congruent with traditional genera: Trachelomonas (incl. Strombomonas) and Colacium. Euglena was polyphyletic and diverged into four independent clades (intermixed with Astasia, Khawkinea and Lepocinclis) and two individual branches (e.g. E. polymorpha). Phacus was also subdivided into Phacus s. str. and two combined lineages (mixed with Lepocinclis spp. or Cryptoglena). In consequence, Euglena (s. str.), Phacus and other genera were emended and one lineage (mixed Phacus/Lepocinclis-clade) was recognized as the previously neglected genus Monomorphina Mereschkowsky (1877). The sister clade of Phacus s. str. (mixed Euglena/Lepocinclis-clade) was identified as Lepocinclis Perty (emended).     

Phylogeny of the tribe Colletieae, Rhamnaceae

A cladistic analysis based on 63 morphological characters was carried out on the tribe Colletieae including two presumed closely related genera, Geanothus and Noltea as outgroupS. In addition to a parsimony analysis of the equally weighted characters, analyses investigating the effects of character weighting, removal of a presumed hybrid species as well as the impact of uncertainly scored characters were undertaken. In all analyses Noltea was placed as sister group to a well supported monophyletic Colletieae. Nineteen different in group topologies were found, with the additional analyses mainly supporting two of them. Within the Colletieae a basal dichotomy divides Trevoa and Retanilla from the remainder of the tribe. While the Trevoa-Retanilla clade is fully resolved, the second lacks detailed resolution. Within this clade the Colletia species from a well supported monophyletic group, while monophyly of the disjunct genus Discaria could not be confirmed.     

Developmental studies on the loricate choanoflagellateStephanoeca diplocostata Ellis

Summary Cell division inStephanoeca diplocostata follows the accumulation of a large number of costal strips in horizontal bundles at the top of the parent collar. Prior to nuclear division the flagellum is lost and the protoplast is large and rectangular. Nuclear division takes place whilst the protoplast undergoes vigorous metabolic movements and subsequent cytokinesis is achieved by equatorial constriction. The anterior of the two daughter protoplasts is the juvenile and is inverted with respect to the sister which remains attached to the parent lorica. The two protoplasts are joined by a cytoplasmic strand that consists of two threads both of which are initially attached to the daughter protoplasts at one side of the collar. Cell separation involves elongation of the strand and after each thread has broken contact with one of the daughter cells the two threads slide over each other until the juvenile is released. The juvenile takes the accumulation of supernumerary strips as it leaves the parent lorica and after release of the juvenile the strips are mobilised to form a new lorica. The collar tentacles of the parent are thought to play a significant role in the movement of strips during division and certain selected tentacles on the juvenile are associated with lorica assembly. Cell separation takes between 9–12 minutes and lorica assembly by the juvenile 2–3 minutes.     

Phylogeny and classification of Marantaceae

Relationships of Marantaceae were estimated from nucleotide sequence variation in the rps16 intron (plastid DNA) and from morphological characters. Fifty-nine species (21 genera) formed the ingroup, and 12 species (12 genera) of other Zingiberales formed the outgroup. There is no support for the traditional subdivision of Marantaceae into a triovulate and a uniovulate tribe or the informal groups previously proposed. The so-called Donax group forms a paraphyletic grade that is basal within Marantaceae. Thalia appears as the distal branch of this grade, but its position is not supported in jackknife analysis. The so-called Calathea group is monophyletic in all shortest trees but not supported with greater than 50% jackknife. The genus Calathea appears to be paraphyletic. The Maranta and Phrynium groups are clearly polyphyletic. Maranta, Koernickanthe , and genera of the Mymsma group, all neotropical, form a strongly supported monophyletic group. The sister of this group is the palaeotropical genus Halopegia. Koernickanthe is nested within Maranta , as this genus is traditionally circumscribed. The African genera Ataenidia and Marantochloa form a strongly supported clade in which Ataenidia is the sister group to Marantochloa . Based on phylogeny it is concluded that Africa, in spite of being much poorer in species, is the most likely ancestral area of Marantaceae     

基于线粒体ND1和COI基因序列探讨锯眼蝶亚科主要类群的系统发生关系

测定了分布于中国的锯眼蝶亚科4族10属共20个种的线粒体ND1和COI基因的部分序列,结合从GenBank中获得的4个国外种类的同源序列,以凤蝶科的迪洛尔娟凤蝶(Allancatria deyrolle)、丝带凤蝶(Sericinus montela),以及娟蝶科的西猛娟蝶(Parnassius simonius)为外类群,通过邻接法、最大简约法、最大似然法和贝叶斯法重建了分子系统树,分析了该亚科内主要类群的系统发生关系。分析结果表明:帻眼蝶族和锯眼蝶族具有较近的亲缘关系;黛眼蝶族不是单系群,该族中的黛眼蝶属、荫眼蝶属与眉眼蝶族具有较近的亲缘关系,带眼蝶属、藏眼蝶属、毛眼蝶属和帕眼蝶属聚合为一个独立的支系,其中带眼蝶属和藏眼蝶属在所有的分析方法中均以100%的置信度(BP=100%,PP=1.00)相聚合,建议将它们合并为一属。     

A preliminary study on relationships among selected Australian members of the tribe Spilomelini (Lepidoptera: Crambidae: Pyraustinae)

A preliminary study was conducted on phylogenetic relationships among some selected genera of the Australian Spilomelini, focusing on relationships among the Australian Glyphodes group (Glyphodes Guenée, 1854, Dysallacta Lederer, 1863, Talanga Moore, 1885 and Agrioglypta Meyrick, 1932) and the 17 genera which are morphologically similar to it. Representatives of three genera of the Pyarustini were used as outgroups. Cladistic analysis of morphological data from the adult moths produced 10 equally MP trees (length = 221, CI=0.294, and RI=0.608). The clade formed by the 21 selected genera of the Australian Spilomelini had low bootstrap support even though a good apomorphy supported the monophyly of this group, namely, a strong, bilobed praecinctorium of abdominal tympanal organs. The analysis showed that the Glyphodes group is not monophyletic because the genus Chrysothyridia Snellen appears within it in the 10 MP trees. The concept of the Glyphodes group should be expanded to include Chrysothyridia and also the Synclera Zeller and Didymostoma (Walker) since the Synclera + Didymostoma clade, as the hypothesised sister group of the Glyphodes group, is not sufficiently supported as a separate monophyletic group. The analysis also showed that genus Glyphodes is not a monophyletic group, while Metallarcha Meyrick is a monophyletic group.