Phylogeny of the kinetoplastida: taxonomic problems and insights into the evolution of parasitism

To further investigate phylogeny of kinetoplastid protozoa, the sequences of small subunit (18S) ribosomal RNA of nine bodonid isolates and ten isolates of insect trypanosomatids have been determined. The root of the kinetoplastid tree was attached to the branch of Bodo designis and/or Cruzella marina. The suborder Trypanosomatina appeared as a monophyletic group, while the suborder Bodonina was paraphyletic. Among bodonid lineages, parasitic organisms were intermingled with free-living ones, implying multiple transitions to parasitism and supporting the 'vertebrate-first hypothesis'. The tree indicated that the genera Cryptobia and Bodo are artificial taxa. Separation of fish cryptobias and Trypanoplasma borreli as different genera was not supported. In trypanosomatids, the genera Leptomonas and Blastocrithidia were polyphyletic, similar to the genera Herpetomonas and Crithidia and in contrast to the monophyletic genera Trypanosoma and Phytomonas. This analysis has shown that the morphological classification of kinetoplastids does not in general reflect their genetic affinities and needs a revision.     

Systematics of the snake genera Stenophis and Lycodryas from Madagascar and the Comoros

Nagy, Z. T., Glaw, F. & Vences, M. (2010) Systematics of the snake genera Stenophis and Lycodryas from Madagascar and the Comoros. —Zoologica Scripta, 39, 426–435. Arboreal snakes belonging to the pseudoxyrhophiine genus Stenophis inhabit Madagascar but despite their spectacular appearance, surprisingly little is known about their natural history and systematics. Nonetheless, a close phylogenetic affinity of the genera Stenophis and Lycodryas (the latter genus currently includes a single species from the Comoros) has been hypothesized. Based on recent molecular genetic data, however, the monophyly of Stenophis was challenged. This study aimed at a systematic analysis and taxonomic revision of the genera Stenophis and Lycodryas. On the basis of new molecular genetic and morphological data and analyses, we propose to accommodate these snakes in three monophyletic genera: Lycodryas, Phisalixella and Parastenophis, and to consider Stenophis as a junior synonym of Lycodryas. In the new generic arrangement, the genera can also be well distinguished by morphological characters. On the specific level, Phisalixella tulearensis is resurrected and indications of further, undescribed taxa are revealed.     

Towards the phylogeny of the Curculionoidea (Coleoptera): Reconstructions from mitochondrial and nuclear ribosomal DNA sequences

With about 60,000 described species, Curculionoidea represent the most species-rich superfamily in the animal kingdom. The immense diversity apparently creates difficulties in the reconstruction of the phylogenetic relationships. Independent morphological studies have led to very different classifications. This study is based on molecular data from two independent molecular sources, the 16S and 18S rDNA. Sensitivity analyses were conducted for the sequence alignment (gap costs were varied) as well as the phylogenetic reconstruction algorithms and some of their parameters. The higher-level relationships reconstructed within Curculionoidea are sensitive to alignment and reconstruction method. Nemonychidae or Oxycorynidae+Belidae were found to be sister to all remaining Curculionoidea in many analyses. The 16S rDNA sequence data (obtained from 157 species) corroborate many tribes and genera as monophyletic. It is observed that the phylogenetic reconstruction of genera with specific genetic features such as polyploidy and parthenogenetic reproduction is difficult in weevils. The curculionid subfamily Lixinae appears monophyletic. A new monophylum consisting of Entiminae, Hyperinae, Cyclominae, Myllorhinus plus possibly the Cossoninae is distinguished and we call it Entiminae s.l. For most other subfamilies and families homoplasy concealed the phylogenetic signal (due to saturation of the 16S sequences), or the species sampling was insufficient, although our sampling scheme was rather broad. We observed that although data from one source can easily be misleading (16S) or hardly informative (18S), the combination of the two independent data sets can result in useful information for such a speciose group of organisms. Our study represents the most thorough analysis of molecular sequence data of the Curculionoidea to date and although the phylogenetic results appear less stable than expected, they reflect the information content of these sequence data realistically and thus contribute to the total knowledge about the phylogeny of the Curculionoidea.     

Evolution of blindness in scolopendromorph centipedes (Chilopoda: Scolopendromorpha): insight from an expanded sampling of molecular data

Relative to its diversity (34 genera, 700 species), Scolopendromorpha has been undersampled in molecular phylogenetic analyses compared with the other chilopod orders. Previous analyses based on morphology have not resolved several key controversies in systematics and evolutionary morphology unambiguously. Here we apply new molecular and morphological data to scolopendromorph phylogenetics, with a focus on the evolution of blindness. The taxonomic sample includes 19 genera, many lacking previous molecular data, and diverse, cosmopolitan genera of Scolopendridae are sampled by multiple species. Phylogenetic analysis with Direct Optimization used 94 morphological characters and ca. 4.5 kb of sequence data from two nuclear (18S and 28S rRNA) and two mitochondrial (16S rRNA and COI) loci. A single most‐parsimonious cladogram selected after sensitivity analyses resolves Scolopendromorpha as monophyletic, and divides it into a blind clade of three families (Plutoniumidae, Cryptopidae, Scolopocryptopidae) and its ocellate sister group, Scolopendridae. Some species‐rich, cosmopolitan genera (Cormocephalus, Otostigmus, Scolopendra) in Scolopendridae are non‐monophyletic, and in several instances (e.g. New and Old World Scolopendra) relationships are more congruent with geographical distributions than with traditional classifications. The tribe Asanadini is particularly subject to parameter‐sensitivity, nesting in the combined analysis within Scolopendrini but as sister to all other Scolopendrinae for molecular data alone. The total‐evidence tree unambiguously optimizes trunk segmentation: a 23‐segmented trunk has a single origin in the blind clade. © The Willi Hennig Society 2011.     

First phylogenetic analysis of the tribe Phyllotini (Rodentia: Sigmodontinae) combining morphological and molecular data

Previous phylogenetic analyses of the tribe Phyllotini, one of the largest components of the subfamily Sigmodontinae, have been based on a single source of evidence. In particular, morphological analyses were largely based on craniodental data, almost neglecting the potential phylogenetic information present in the postcranium. Despite the significant advances made in relation to the knowledge of phyllotine phylogeny in recent times, there are several unsolved issues that highlight the importance of a phylogenetic analysis that integrates multiple sources of evidence, including previously considered sources as well as new sources of data. We present here the first combined phylogenetic analysis (morphological and molecular) of phyllotines, which includes the widest taxon and character sampling to date. Our dataset includes 164 morphological characters, of which 83 are postcranial characters, plus 3561 molecular characters, scored for 52 species from 34 genera of Oryzomyalia. In this study 75 postcranial characters not previously considered in this group are thoroughly described, and their utility for solving the relationships within Phyllotini is evaluated by means of different complementary analyses. Phyllotini was retrieved as a monophyletic clade in the combined analysis, with a composition that matches that obtained in most other recent analyses. All genera of phyllotines were monophyletic and show high support values. Abrotrichini, Akodontini and Oryzomyini were also monophyletic. The inclusion of postcranial data appears to be of limited utility to solve the phylogenetic relationships within Phyllotini.     

New molecular data for tardigrade phylogeny, with the erection of Paramacrobiotus gen. nov.

Up to few years ago, the phylogenies of tardigrade taxa have been investigated using morphological data, but relationships within and between many taxa are still unresolved. Our aim has been to verify those relationships adding molecular analysis to morphological analysis, using nearly complete 18S ribosomal DNA gene sequences (five new) of 19 species, as well as cytochrome oxidase subunit 1 (COI) mitochondrial DNA gene sequences (15 new) from 20 species, from a total of seven families. The 18S rDNA tree was calculated by minimum evolution, maximum parsimony (MP) and maximum likelihood (ML) analyses. DNA sequences coding for COI were translated to amino acid sequences and a tree was also calculated by neighbour-joining, MP and ML analyses. For both trees (18S rDNA and COI) posterior probabilities were calculated by MrBayes. Prominent findings are as follows: the molecular data on Echiniscidae (Heterotardigrada) are in line with the phylogenetic relationships identifiable by morphological analysis. Among Eutardigrada, orders Apochela and Parachela are confirmed as sister groups. Ramazzottius (Hypsibiidae) results more related to Macrobiotidae than to the genera here considered of Hypsibiidae. Macrobiotidae and Macrobiotus result not monophyletic and confirm morphological data on the presence of at least two large groups within Macrobiotus. Using 18S rDNA and COI mtDNA genes, a new phylogenetic line has been identified within Macrobiotus , corresponding to the ' richtersi-areolatus group'. Moreover, cryptic species have been identified within the Macrobiotus ' richtersi group' and within Richtersius . Some evolutionary lines of tardigrades are confirmed, but others suggest taxonomic revision. In particular, the new genus Paramacrobiotus gen. n. has been identified, corresponding to the phylogenetic line represented by the ' richtersi-areolatus group'.     

Phylogeny of Eastern North American Coreopsis (Asteraceae-Coreopsideae): insights from nuclear and plastid sequences, and comments on character evolution

A molecular phylogenetic study of eastern North American Coreopsis and representatives of other genera of tribe Coreopsideae was conducted using combined sequences from nuclear ITS and two plastid regions (matK, rpl16). A total of 25-30 species has been recognized in five sections of Coreopsis in eastern North America. Based on morphological characters, these taxa have generally been considered a monophyletic group. Our well-resolved phylogeny supports the monophyly of sections that have been recognized in Coreopsis, but the sections collectively do not comprise a monophyletic group because species of north temperate Bidens occur within one of the two major Coreopsis clades. The most notable departure of present results from prior views of relationships among sections is the lack of a sister group relationship between sections Calliopsis and Eublepharis; the shared presence of four-lobed disk floret corollas had been used to support a close relationship between these two sections. Relationships within sections show both similarities and differences with the results of previous studies based primarily on morphological characters. Mapping of morphological characters used taxonomically in Coreopsis and related genera onto the phylogeny indicates that the evolution of these characters has been complex, and this compromises their value for defining monophyletic groups. Examples include the annual habit, alternate leaves, winged fruits, red or brown basal spots on the yellow ligules, and four-lobed disk floret corollas.     

New taxa refresh the phylogeny and classification of pleurostomatid ciliates (Ciliophora,Litostomatea)

A high diversity of pleurostomatid ciliates has been discovered in the last decade, and their systematics needs to be improved in the light of new findings concerning their morphology and molecular phylogeny. In this work, a new genus, Protolitonotus gen. n., and two new species, Protolitonotus magnus sp. n. and Protolitonotus longus sp. n., were studied. Furthermore, 19 novel nucleotide sequences of SSU rDNA, LSU rDNA and ITS1‐5.8S‐ITS2 were collected to determine the phylogenetic relationships and systematic positions of the pleurostomatid ciliates in this study. Based on both molecular and morphological data, the results demonstrated that: (i) as disclosed by the sequence analysis of SSU rDNA, LSU rDNA and ITS1‐5.8S‐ITS2, Protolitonotus gen. n. is sister to all other pleurostomatids and thus represents an independent lineage and a separate family, Protolitonotidae fam. n., which is defined by the presence of a semi‐suture formed by the right somatic kineties near the dorsal margin of the body; (ii) the families Litonotidae and Kentrophyllidae are both monophyletic based on both SSU rDNA and LSU rDNA sequences, whereas Amphileptidae are non‐monophyletic in trees inferred from SSU rDNA sequences; and (iii) the genera Loxophyllum and Kentrophyllum are both monophyletic, whereas Litonotus is non‐monophyletic based on SSU rDNA analyses. ITS1‐5.8S‐ITS2 sequence data were used for the phylogenetic analyses of pleurostomatids for the first time; however, species relationships were less well resolved than in the SSU rDNA and LSU rDNA trees. In addition, a major revision to the classification of the order Pleurostomatida is suggested and a key to its families and genera is provided.     

Phylogenetic relationships of the Sparidae (Teleostei: Percoidei) and implications for convergent trophic evolution

A phylogenetic analysis of the majority of sparid genera and representatives of the sparoid families Centracanthidae, Lethrinidae and Nemipteridae is presented using 87 predominately osteological characters. The Sparidae constitute a monophyletic grouping, with the inclusion of the centracanthid Spicara smaris , which nests deep within the ingroup. The phylogeny was then used to investigate agreement with the most recent molecular study, taxonomic stability of subfamilial classification and the evolution of feeding strategies. Results show that the incongruence between morphological and molecular data appears largely to be an artifact of errors in rooting. However, there appears to be real and substantial conflict between the molecular tree and the morphological data, which is not attributable to the different positions of the least congruent taxa. The data support the molecular hypothesis that none of the subfamilial classification, based on dentition and trophic specialization, is monophyletic, and should be rejected pending further taxonomic revision. The phylogeny supports multiple independent origins of trophic types and it is suggested that the evolutionary plasticity of the oral teeth of sparids has been fundamental to the adaptive radiation of this family compared to their closest allies. ©2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 269–301.     

Molecular phylogeny of the hexagrammid fishes using a multi-locus approach

Ideally, organisms are grouped into monophyletic assemblages reflecting their evolutionary histories. Single (molecular) markers can reflect the evolutionary history of the marker, rather than the species in question, therefore, phylogenetic relationships should be inferred from adequate sampling of characters. Because the use of multiple loci greatly improves the resolving power of the molecular assay, we constructed a molecular phylogeny of the family Hexagrammidae based on six loci, including two mitochondrial and four nuclear loci. The resulting molecular phylogeny, from the combined data, was significantly different from the morphological topology suggested by Shinohara [Memoirs of the Faculty of Fisheries, Hokkaido University 41 (1994) 1]. Our data support a monophyletic assemblage for the genera Hexagrammos and Pleurogrammus. However, other taxa traditionally included in the family Hexagrammidae did not form a monophyletic assemblage. The monotypic genus Ophiodon was more closely associated with cottids than with other hexagrammids. Our data concur with the morphological topology in that the genera Zaniolepis and Oxylebius formed a monophyletic clade, which was distinct and basal to the remaining hexagrammids, seven cottids and one agonid.     

Phylogeny of Nerillidae (Polychaeta, Annelida) as inferred from combined 18S rDNA and morphological data

A phylogeny of the meiofaunal polychaete family Nerillidae based on morphological, molecular and combined data is presented here. The data sets comprise nearly complete sequences of 18S rDNA and 40 morphological characters of 17 taxa. Sequences were analyzed simultaneously with the morphological data by direct optimization in the program POY, with a variety of parameter sets (costs of gaps: transversions: transitions). Three outgroups were selected from the major polychaete group Aciculata and one from Scolecida. The 13 nerillid species from 11 genera were monophyletic in all analyses with very high support, and three new apomorphies for Nerillidae are identified. The topology of the ingroup varied according to the various parameter settings. Reducing the number of outgroups to one decreased the variance among the phylogenetic hypotheses. The congruence among these was tested and a parameter set, with equal weights (222) and extension gap weighted 1, yielded minimum incongruence (ILD). Several terminal clades of the combined analysis were highly supported, as well as the position of Leptonerilla prospera as sister terminal to the other nerillids. The evolution of morphological characters such as segment numbers, chaetae, appendages and ciliation are traced and discussed. A regressive pathway within Nerillidae is indicated for several characters, however, generally implying several convergent losses. Numerous genera are shown to require revision. © The Willi Hennig Society 2005.     

Testing morphology-based hypotheses of phylogenetic relationships in Parmeliaceae (Ascomycota) using three ribosomal markers and the nuclear RPB1 gene

Parmeliaceae is the largest family of lichen-forming fungi with more than 2000 species and includes taxa with different growth forms. Morphology was widely employed to distinguish groups within this large, cosmopolitan family. In this study we test these morphology-based groupings using DNA sequence data from three nuclear and one mitochondrial marker from 120 taxa that include 59 genera and represent the morphological and chemical diversity in this lineage. Parmeliaceae is strongly supported as monophyletic and six well-supported main clades can be distinguished within the family. The relationships among them remain unresolved. The clades largely agree with the morphology-based groupings and only the placement of four of the genera studied is rejected by molecular data, while four other genera belong to clades previously unrecognised. The classification of these previously misplaced genera, however, has already been questioned by some authors based on morphological evidence. These results support morphological characters as important for the identification of monophyletic clades within Parmeliaceae.     

A MOLECULAR ANALYSIS OF THE EUGLENOPHYTES USING SSU RDNA

Almost since the creation of the genus Euglena (Ehrenberg), the taxa assigned to it have been separated, split apart, and reorganized into new genera based on morphological relationships, resulting in the creation of the genera Phacus (Dujardin) , Lepocinclis (Perty) , Astasia (Pringsheim), and Khawkinea ( Jahn and McKibben) based on intuitive methods. In an effort to assess the validity of these genera, we have used small subunit (SSU) rDNA data to generate a phylogenetic framework for these genera, with particular attention to the genus Euglena . Using the conserved sequence areas, we performed a phylogenetic analysis using parsimony, maximum likelihood, and distance methods. These different criteria have resulted in trees of the same topology. The euglenoid clade was composed of phagotrophic euglenoids at the base, which gave rise to phototrophs that in turn gave rise to osmotrophs. Among the photosynthetic taxa, the biflagellate form diverged prior to the uniflagellate form. Additionally, the need for a revision in the taxonomy of some of these genera was demonstrated. Currently, taxa from the photosynthetic genera Euglena, Phacus, and Lepocinclis do not form monophyletic clades, but are intermixed with each other as well as with the osmotrophic taxa, Astasia and Khawkinea.     

常用核基因序列在双翅目昆虫系统学中的研究进展

双翅目昆虫分为长角亚目和短角亚目,前者主要类群包括蚊、蠓、蛉和蚋,后者主要类群为虻类和蝇类。国内外学者对双翅目昆虫形态分类和分子系统发育关系研究均较多。本文整理总结了几种主要核基因在双翅目昆虫进化和系统发育关系的研究资料,结果显示:双翅目的单系性得到了众多形态学、生物学和分子数据的支持,多数系统发育研究认为传统的长角亚目为并系,短角亚目是一个单系,其主要类群舞虻下目、环裂类、有缝组和有瓣蝇类均为单系,但非环裂类、无缝组为并系,无瓣类可能为并系;基本搞清了有重要医学意义和与环境关系密切的类群,特别是有瓣蝇类各科类群分类系统的进化关系;双翅目昆虫发生辐射进化的三个分支节点时间即:低等双翅目(蚊类)2.2亿年、低等短角亚目(虻类)1.8亿年、有缝组(蝇类)6500万年;大量双翅目昆虫自然生命史历经吸血性、植食性和寄生性,有2.6亿年以上的演化历程。从相关核基因研究中总结出:18SrRNA、28SrRNA和CAD基因能很好的解决高级阶元从目到属的系统发育问题;EF-1ɑ基因和White基因更适合从科到属水平的分类阶元;ITS基因一般应用在从属到种水平的低级分类阶元,并被广泛应用到双翅目昆虫分子系统学研究中。     

Revisiting Ross and Sims (1971): toward a molecular phylogeny of the Biddulphiaceae and Eupodiscaceae (Bacillariophyceae)

The ocellate and pseudocellate diatoms in the Eupodiscaceae and Biddulphiaceae (respectively) are common inhabitants of the marine littoral (and plankton zone) with a rich fossil history making them important components of marine stratigraphic studies and good candidates for molecular dating work. These diatoms are important for un‐derstanding the phylogeny of the diatoms as a whole, as molecular phylogenies have blurred the traditional distinction between the pennate and multipolar non‐pennate diatoms. However, the convoluted taxonomic history of these groups has the potential to disrupt both stratigraphic and molecular dating studies. Although efforts have been made to examine frustule morphology of several ocellate and pseudocellate diatoms and develop a morphological scheme to define genera, very little work has been done to determine how these groups are interrelated. In this study, we use nuclear and chloroplast molecular markers to construct a phylogeny of a diverse sampling of Eupodiscaceae and Biddulphiaceae taxa. The ocellus‐bearing taxa (Eupodiscaceae) are monophyletic, and thus the ocellus may be a useful character in delimiting the Eupodiscaceae, the Biddulphiaceae are polyphyletic and scattered across a number of lineages of multipolar non‐pennate diatoms. Hypothesis testing aimed at assessing the likeliness of several morphology based hypotheses against the molecular data highlights uncertainty in both types of data. We present evidence that there are monophyletic genera within both the Biddulphiaceae and Eupodiscaceae, and recommend the taxa within the Odontella mobilensis/sinensis/regia clade be transferred to a new genus: Trieres Ashworth & Theriot.     

Description of a new species of Parasesarma (Crustacea; Decapoda; Brachyura; Sesarmidae) from the Persian Gulf, based on morphological and genetic characteristics

The genera Parasesarma and Perisesarma are common inhabitants of mangroves and estuaries of the Indo-West Pacific. These, overall similar, genera of sesarmid crabs can be told apart by the presence or absence of an anterolateral tooth, but recent and ongoing studies show that they are not reciprocally monophyletic. In recent years, detailed morphological studies have revealed a surprising number of undescribed new species within these genera. In the present paper, we describe a new species of the genus Parasesarma from the relatively poorly studied Persian Gulf. So far, these specimens had been referred to as Parasesarma plicatum (Latreille, 1803), but morphological and genetic characteristics clearly allow separation of the two species. Consequently, the species from the Persian Gulf is here described as new and morphological and mitochondrial genetic characters are provided that allow recognition of the species.     

Revised classification and evolution of leucotrichiine microcaddisflies (Trichoptera: Hydroptilidae) based on morphological and molecular data

Hydroptilidae constitute the most diverse caddisfly family, with over 2000 species known from every habitable continent. Leucotrichiinae are exclusively New World microcaddisflies, currently including over 200 species and 17 genera. Phylogenetic analyses of Leucotrichiinae relationships based on 114 morphological characters and 2451 molecular characters from DNA sequences were conducted. DNA sequences analysed were from one mitochondrial gene, cytochrome oxidase I (653 bp), and four nuclear genes, carbamoylphosphate synthase (802 bp), elongation factor 1α (352 bp), histone 3 (308 bp) and 28S rDNA (336 bp). The morphological matrix included 94 taxa (with representatives of all included genera) and the molecular matrix included 62 taxa. Individual and combined datasets were analysed under parsimony and Bayesian inference. In addition, a relaxed molecular clock divergence time estimate was conducted to determine the age of the subfamily and major lineages. All Bayesian inference analyses strongly suggest a monophyletic Leucotrichiinae, which initially diverged at approximately 124 Ma into two monophyletic groups of genera. These groups are herein elevated to tribal status, Alisotrichiini trib.n. and Leucotrichiini Flint sensu n. Several genera of Leucotrichiini were not recovered as monophyletic clades and some nomenclatural changes are proposed to reflect their phylogenetic history. These include the synonymy of Abtrichia with Peltopsyche; transfer of Betrichia hamulifera to Costatrichia; transfer of Betrichia alibrachia and Costatrichia falsa to Leucotrichia; and transfer of Costatrichia fluminensis to Acostatrichia. Additionally, Tupiniquintrichia gen.n. is described to include Peltopsyche maclachlani and Leucotrichia procera. According to our results, crown diversifications of both Alisotrichiini trib.n. (～80 Ma) and Leucotrichiini sensu n. (～103 Ma) occurred after complete separation of South America from Africa. Current distributions of most leucotrichiine genera are probably a result of migration from South America towards the north using the proto‐Caribbean archipelago (100 to 49 Ma). This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:FB6A3385-323D-41AF-B4BE-E19A393A493C .     

Phylogenetic systematics of leaffishes (Teleostei: Polycentridae,Nandidae)

The Asian (nandid) and Afro‐Neotropical (polycentrid) leaffishes represent two superficially similar, but historically poorly diagnosed families – a situation resulting in a convoluted systematic history. Here, and including for the first time in a molecular study all leaffish genera, we generate a hypothesis of the phylogenetic history of both groups. We analyse a multilocus molecular data set encompassing 257 acanthomorph taxa, carry out a survey and assessment of selected osteological characters for the polycentrid leaffishes and also provide a reanalysis of previously published morphological data. Our results confirm: (1) that the Polycentridae and Nandidae are only remotely related, and hence, the classic leaffishes are diphyletic; (2) that the Polycentridae is monophyletic, with new skeletal synapomorphies being congruent with molecular data in placing the enigmatic Afronandus – a taxon that thus far has never been included in any molecular study – as sistergroup to the remaining genera; (3) the monophyly of the Nandidae + Badidae and their inclusion into a larger monophyletic group – along with the Pristolepididae, Anabantoidei and Channoidei – comprising the Labyrinthici sensu Rosen & Patterson. We also review the morphological and molecular evidence for both the conflicting placement of Pristolepis and the putative sistergroup relationship between the labyrinth fishes (Anabantoidei) and snakeheads (Channoidei).     

13 Viral control of phytoplankton

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.