Ultrastructure of the body wall of three species of Grania (Annelida: Clitellata: Enchytraeidae)

De Wit P., Erséus C. and Gustavsson L.M. 2011. Ultrastructure of the body wall of three species of Grania (Annelida: Clitellata: Enchytraeidae). —Acta Zoologica (Stockholm) 92 : 1–11. The body wall of three species of Grania, including the cuticle, epidermis and the musculature, are studied using TEM. The cuticle is similar to previously studied enchytraeids, with an orthogonal grid pattern of collagen fibers. This pattern is also seen in Crassiclitellata, which has been suggested as the sister taxon of Enchytraeidae. Variation of epicuticular and fiber zone patterns seen in Naididae (formerly Tubificidae and Naididae) seem to be lacking in Enchytraeidae. The fiber thickness, however, varies between Grania species and may be a phylogenetically informative character. The epidermis consists of supporting cells, secretory cells and sensory cells. Basal cells, typical for Crassiclitellata, were not observed. The clitellum of Grania seems to consist of two types of gland cells, which develop from regular epidermal tissue. It is possible that more cell types exist in different regions of the clitellum, however. The body wall musculature is arranged somewhat differently from that of closely related taxa; this refers to the reduction of circular and outer, triangular longitudinal muscle fibers, while the inner, ribbon‐shaped longitudinal muscle fibers are well‐developed. A search was conducted for the cause of the peculiar green coloration of Grania galbina De Wit and Erséus 2007, and it was concluded that neither cyanobacteria nor epidermal pigment granules were present in the fixed material.     

New records of Adelodrilus (Oligochaeta,Tubificidae), with descriptions of two new species from the North-west Atlantic

Adelodrilus fimbriatus sp.n. is described from bathyal depths off the continental shelf of Eastern United States, A. cristatus sp.n. from sublittoral bottoms on this shelf (off Maryland and New Jersey). The two species are principally distinguished from their congeners by the morphology of their penial setae. New geographical records and morphological notes are given for A. pusillus Erséus (including the first find of Adelodrilus in the Mediterranean Sea), A. cooki Erséus, A. voraginus (Cook), and A. acochlearis Erséus & Loden. A pictorial key to all eleven species of Adelodrilus is provided.     

Taxonomy and Phylogeny of the Gutless Phallodrilinae (Oligochaeta,Tubificidae), with Descriptions of One New Genus and Twenty-Two New Species*

The gutless members of the subfamily Phallodrilinae are taxonomically revised and generically split off from their gut-bearing relatives. Inanidrilus Erséus, 1979, a much modified definition of which is presented, comprises the species leukodermatus (Giere, 1979) comb.n., speroi sp.n., fijiensis sp.n., bonomii sp.n., carterensis sp.n., gustavsoni sp.n., wasseri sp.n., scalprum sp.n., belizensis sp.n., aduncosetis sp.n., vacivus sp.n., triangulatus sp.n., ernesti sp.n., manae sp.n., falcifer Erséus & Baker, 1982, renaudae sp.n., extrmus (Erséus 1979) comb.n., mexicanus Erséus & Haker, 1982, and bulbosus Erséus, 1979. Olavius sp.n. is established to accommodate the species geniculatus (Erséus, 1981) comb.n., imperfectus sp.n., filithecatus (Erséus. 1981) comb.n. albidus (Jamieson, 1977) comb.n., propinquus sp.n., comorensis (Erséus. 1981) comb.n., cornuatus Davis, 1984, pellucidus sp.n., tenuissimus (Erséus, 1979) macer sp.n., caudatus (Erséus. 1979) comb.n., planus (Erséus, 1979) comb.n., clavatus (Erséus. 1981) comb.n., tantulus sp.n., longissimus (Giere, 1979) comb.n., alius sp.n., avisceralis (Erséus, 1981) comb.n., loisae sp.n., and hanssoni sp.n., Two subgenera are erected for tantulus—longissimus–alius (nominate subgenus Olavius) and avisceralis-loisae-hanssoni (Coralliodriloides subgen.n.), respectively. A tentative phylogeny of the gutless group, which appears monophyletic, is inferred on the basis of a partly computerized Wagner-tree analysis of all 38 species. According to this analysis. According to this analysis, Inanidrilus can be defined by one synapomorphy, more or less sickle-shaped penial setae, whereas Olavius can be discriminated by two such characters, (1) atrium comma-shaped/horizontal rather than erect and (2) copulatory sacs present.     

Phylogenetic relationships within the South American fish family Anostomidae (Teleostei,Ostariophysi, Characiformes)

Analysis of a morphological dataset containing 152 parsimony‐informative characters yielded the first phylogenetic reconstruction spanning the South American characiform family Anostomidae. The reconstruction included 46 ingroup species representing all anostomid genera and subgenera. Outgroup comparisons included members of the sister group to the Anostomidae (the Chilodontidae) as well as members of the families Curimatidae, Characidae, Citharinidae, Distichodontidae, Hemiodontidae, Parodontidae and Prochilodontidae. The results supported a clade containing Anostomus, Gnathodolus, Pseudanos, Sartor and Synaptolaemus (the subfamily Anostominae sensu Winterbottom) albeit with a somewhat different set of relationships among the species within these genera. Anostomus as previously recognized was found to be paraphyletic and is split herein into two monophyletic components, a restricted Anostomus and the new genus Petulanos gen. nov. , described herein. Laemolyta appeared as sister to the clade containing Anostomus, Gnathodolus, Petulanos, Pseudanos, Sartor and Synaptolaemus. Rhytiodus and Schizodon together formed a well‐supported clade that was, in turn, sister to the clade containing Anostomus, Gnathodolus, Laemolyta, Petulanos, Pseudanos, Sartor and Synaptolaemus. Anostomoides was sister to the clade formed by these nine genera. Leporinus as currently defined was not found to be monophyletic, although certain clades within that genus were supported, including the species with subterminal mouths in the former subgenus Hypomasticus which we recognize herein as a genus. Abramites nested in Leporinus, and Leporellus was found to be the most basal anostomid genus. The presence of cis‐ and trans‐Andean species in Abramites, Leporellus, Leporinus and Schizodon, all relatively basal genera, suggests that much of the diversification of anostomid species pre‐dates the uplift of the Andean Cordilleras circa 11.8 million years ago. Several important morphological shifts in anostomid evolution are illustrated and discussed, including instances of convergence and reversal. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 70–210.     

Distinguishing features of immature stages of Panchaetothripinae (Thysanoptera: Thripidae) known in New Zealand

Diagnostic morphological characters of the juvenile Panchaetothripinae in New Zealand are illustrated. Keys developed enable colonies with only immature stages to be identified without needing to rear adults. Live larvae or larvae in ethanol are distinguished by the presence of expanded tips of body setae (Parthenothrips dracaenae), the absence of setae at the abdomen tip (Hercinothrips bicinctus), setae at abdomen tip not longer than abdominal tip width (Heliothrips haemorrhoidalis) and abdominal tip setae longer than abdominal tip width (Sigmothrips aotearoana, endemic species). The presence or absence of spine-like setae on abdominal segments 9 and 10, and the number and length of setae on the wing buds, enable identification of pupae. Abdominal spine-like setae were on the prepupa and pupa of H. bicinctus and S. aotearoana, species that pupate off the plant, and are probably defensive structures. This is the first record of spine-like setae on segment 10 of terebrantian pupae.     

Diversity in the section Compressa of the genus Chaetoceros (Bacillariophyceae), with description of two new species from Chinese warm waters

Monoclonal strains of Chaetoceros species were collected from Chinese warm waters. Vegetative cells and resting spores were examined using light microscopy as well as scanning and transmission electron microscopy. Combined morphological and DNA sequence data from the hypervariable D1–D3 region of the nuclear ribosomal large subunit showed the presence of two new species within the section Compressa, herein described as Chaetoceros bifurcatus sp. nov. and C. millipedarius sp. nov. Both species possessed features typical of the section Compressa, but C. bifurcatus was characterized by the heavy intercalary setae fusing and extending together for a distance before diverging. The heavy setae were not visually contorted, lacking poroids and spines, in contrast with how the section Compressa has been defined. Chaetoceros millipedarius was characterized by a horizontal stagger of ordinary intercalary setae when the chain was seen in broad girdle view, making the chain look like a millipede. The two new species were supported by molecular phylogenetic analyses with C. bifurcatus sister to C. contortus var. ornatus, while C. millipedarius was sister to C. contortus var. contortus. Based on the morphological features exhibited in C. bifurcatus, the diagnosis of the section Compressa was emended to exclude “contorted” heavy setae and compressed valve faces.     

Early Palaeogene Louisinidae (Macroscelidea,Mammalia), their relationships and north European diversity

Seventeen species from the Palaeocene and Early Eocene of northern Europe, of which 12 are new, are described belonging to the extinct macroscelidean family Louisinidae, raised here from subfamily rank. These species belong to nine genera, of which five are new. The new genera are Walbeckodon , Berrulestes , Gigarton , Thryptodon , and Prolouisina . The new species are Walbeckodon krumbiegeli , Walbeckodon girardi , Paschatherium levei , Berrulestes phelizoni , Berrulestes pellouini , Berrulestes poirieri , Gigarton meyeri , Gigarton sigogneauae , Gigarton louisi , Thryptodon brailloni , Louisina marci , and Teilhardimys brisswalteri . Prolouisina is erected for ‘Louisina’atavella Russell, 1964. Cladistic analysis was undertaken to understand the relationships within the Louisinidae and between them and the North American family Apheliscidae, in which they had earlier been included as a subfamily. Louisinidae are shown to be sister group to a clade consisting of Apheliscidae plus Amphilemuridae and part of a paraphyletic and polyphyletic Adapisoricidae, all of which are tentatively considered to be stem members of the order Macroscelidea. The most primitive macroscelidid, Chambius, from the Early Eocene of northern Africa is nested within Apheliscidae when postcranial characters were included, but in a majority of cases within the Louisinidae when postcranial characters were excluded. Most species from northern Europe became extinct at the end of the Palaeocene, although the genus Paschatherium survived for much of the Early Eocene and Teilhardimys survived into the earliest Eocene. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 856–936.     

Immunohistochemical (cLSM) and ultrastructural analysis of the central nervous system and sense organs in Aeolosoma hemprichi (Annelida, Aeolosomatidae)

The Aeolosomatidae are very small limnetic or terrestrial annelids of apparently simple organisation and uncertain phylogenetic position. They have been placed either at the base of the Clitellata, as a highly derived taxon within the Clitellata closely related to the Naididae, or as their sister group within the „Polychaeta”. A combined immunohistochemical (cLSM) and ultrastructural investigation of the central nervous system and the sense organs in Aeolosoma hemprichi was undertaken to look for characters which might support one of these theories. The position of the brain within the prostomium and the organisation of the ventral nerve cord, with its intraepithelial paired longitudinal nerves lying far apart from each other and the presence of a median longitudinal nerve, are atypical for the Clitellata and clearly differ from the situation found in Naididae. Moreover, the circumoesophageal connectives are bifurcated and enter the brain as dorsal and ventral roots; this arrangement is unknown in Clitellata, in which these connectives are unbranched. An ultrastructural analysis of the ciliated pits located laterally in the furrow between prostomium and peristomium in A. hemprichi and other Aeolosomatidae show that they are in fact nuchal organs. Such presumed chemosensory organs are typical of „Polychaeta” and absent in all Clitellata. Two pairs of ciliary sense organs are present in the prostomium in front of the brain of A. hemprichi. Although similarly organised sensory structures occur in many species of the Clitellata, they differ in position and certain ultrastructural features and are known from other Annelida as well. These results clearly support the exclusion of the Aeolosomatidae from the Clitellata and do not provide any evidence for a sister-group relationship between these two taxa. Accepted: 8 February 2000     

Osteology of the South Asian Genus Psilorhynchus McClelland, 1839 (Teleostei: Ostariophysi: Psilorhynchidae), with investigation of its phylogenetic relationships within the order Cypriniformes

Members of the genus Psilorhynchus are small benthic fishes, commonly referred to as torrent minnows, which inhabit the fast to swift flowing water bodies of the Indo‐Burma region and the Western Ghats of Peninsular India. Despite being described scientifically in the mid 18^th century, the morphology of Psilorhynchus remains poorly known and its phylogenetic placement within the order Cypriniformes is a matter of considerable debate. In this paper the osteology of Psilorhynchus sucatio is described and illustrated in detail. Notes and/or illustrations on the osteology of 12 other species of Psilorhynchus are also provided for the first time. A phylogenetic investigation of the position of Psilorhynchus within the order Cypriniformes is also conducted. Analysis of 127 morphological characters scored for 52 ingroup taxa (including 12 species of Psilorhynchus) and four outgroup taxa resulted in 14 equally parsimonious cladograms (287 steps long; consitency index, CI = 0.48; retention index, RI = 0.88). Psilorhynchus is recovered as the sister group to the family Cyprinidae, and is regarded as a member of the superfamily Cyprinoidea, which forms the sister group to the Cobitoidea (including all other cypriniform families). The sistergroup relationship between Psilorhynchus and Cyprinidae is supported by eight derived characters (five of which are homoplastic within the order Cypriniformes). The monophyly of Psilorhynchus is supported by 16 derived characters (eight of which are homoplastic within Cypriniformes). Three species groups of Psilorhynchus are proposed, the Psilorhynchus balitora group (including P. amplicephalus, P. balitora, P. breviminor, P. nepalensis, P. rahmani, P. pavimentatus, and P. brachyrhynchus), the Psilorhynchus gracilis group (including P. gracilis, P. melissa, P. robustus, and P. tenura), and the Psilorhynchus homaloptera group (including P. arunachalensis, P. homaloptera, P. microphthalmus, and P. pseudecheneis). The continued use of the family group name Psilorhynchidae is recommended. Comments on the interrelationships of the Cypriniformes are also provided. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011.     

Phylogenetics,character evolution,and distribution patterns of the greenbriers,Smilacaceae (Liliales), a near‐cosmopolitan family of monocots

Smilacaceae, composed of Smilax and Heterosmilax, are a cosmopolitan family of > 200 species of mostly climbing monocots with alternate leaves characterized by reticulate venation, a pair of petiolar tendrils and usually prickly stems. Although there has been a long history of studying Smilax since Linnaeus named the genus in 1753, the phylogenetic history of this dioecious family remains unclear. Here we present results based on nuclear ribosomal internal transcribed spacer (nrITS) and plastid matK and rpl16 intron DNA sequence data from 125 taxa of Smilacaceae. Our taxon sampling covers all sections of Smilax and Heterosmilax and major distribution zones of the family; species from Ripogonaceae and Philesiaceae are used as outgroups. Our molecular analysis indicates that phylogenetic relationships largely contradict the traditional morphological classification of the family, instead showing a conspicuous geographical pattern among the species clades. The previously recognized genus Heterosmilax was found to be embedded in Smilax. Species in the family are separated into primarily New World and Old World clades, except for a single species lineage, Smilax aspera, that is sister to the remaining species of the family, but with poor statistical support. Ancestral character state reconstructions and examination of distribution patterns among the clades provide important information for future taxonomic revisions and historical biogeography of the group. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 535–548.     

An unusual new archosauriform from the Middle–Late Triassic of southern Brazil and the monophyly of Doswelliidae

Until now the Doswelliidae was considered a monospecific family including Doswellia kaltenbachi from the Late Triassic of North America. The phylogenetic position of this taxon remained enigmatic until recently, when a sister‐group relationship with the Proterochampsidae was suggested. In the present contribution we describe the new doswelliid species Archeopelta arborensis gen. et sp. nov. from the Middle–Late Triassic of Brazil. A cladistic analysis recovered Archeopelta, Doswellia, and Tarjadia within a monophyletic group of basal archosauriforms, the Doswelliidae. The monophyly of this family is supported by the presence of osteoderm ornamentation that is coarse, incised, and composed of regular pits and the presence of an unornamented anterior articular lamina. Archeopelta is more closely related to Doswellia than to other archosauriforms by the presence of basipterygoid processes anterolaterally orientated, dorsal centra with a convex surface, width of the neural arch plus ribs of the first primordial sacral that are three times the length of the neural arch, and iliac blade laterally deflected, with strongly convex dorsal margin, and a length less than three times its height. The phylogenetic analysis indicates that Doswellidae is the closest large monophyletic entity to Archosauria, which achieved a wide palaeolatitudinal distribution during the late Middle and Late Triassic time span. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 839–871.     

A global phylogeny of apple snails: Gondwanan origin, generic relationships, and the influence of outgroup choice (Caenogastropoda: Ampullariidae)

Apple snails (Ampullariidae) are a diverse family of pantropical freshwater snails and an important evolutionary link to the common ancestor of the largest group of living gastropods, the Caenogastropoda. A clear understanding of relationships within the Ampullariidae, and identification of their sister taxon, is therefore important for interpreting gastropod evolution in general. Unfortunately, the overall pattern has been clouded by confused systematics within the family and equivocal results regarding the family's sister group relationships. To clarify the relationships among ampullariid genera and to evaluate the influence of including or excluding possible sister taxa, we used data from five genes, three nuclear and two mitochondrial, from representatives of all nine extant ampullariid genera, and species of Viviparidae, Cyclophoridae, and Campanilidae, to reconstruct the phylogeny of apple snails, and determine their affinities to these possible sister groups. The results obtained indicate that the Old and New World ampullariids are reciprocally monophyletic with probable Gondwanan origins. All four Old World genera, Afropomus, Saulea, Pila, and Lanistes, were recovered as monophyletic, but only Asolene, Felipponea, and Pomella were monophyletic among the five New World genera, with Marisa paraphyletic and Pomacea polyphyletic. Estimates of divergence times among New World taxa suggest that diversification began shortly after the separation of Africa and South America and has probably been influenced by hydrogeological events over the last 90 Myr. The sister group of the Ampullariidae remains unresolved, but analyses omitting certain outgroup taxa suggest the need for dense taxonomic sampling to increase phylogenetic accuracy within the ingroup. The results obtained also indicate that defining the sister group of the Ampullariidae and clarifying relationships among basal caenogastropods will require increased taxon sampling within these four families, and synthesis of both morphological and molecular data. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 61–76.     

Physical gills in Elmidae (Coleoptera: Byrrhoidea): Structure and evolutionary pattern of plastron in Stenelmis and related genera

Many species within Elmidae (Coleoptera: Byrrhoidea) have plastrons composed of flattened setae. However, some genera display fine plastrons on the epicuticle, called plastron hairs. In Japanese elmids, members of the genera Stenelmis, Ordobrevia, Nomuraelmis and Leptelmis bear ventral plastron hairs. Based on a maximum likelihood tree including most Japanese genera within Elmidae, we found that these genera are monophyletic and that plastron hairs are a derived character in Elmidae. We also found that the genus Graphelmis bears jigsaw puzzle‐like plastron scales with plastron hair‐like projections, and is sister to the group with plastron hairs.     

Phylogenetic relationships within the superfamily Desmodoroidea (Nematoda: Desmodorida), with descriptions of two new and one known species

Three nematode species of the superfamily Desmodoroidea Filipjev, 1922, were isolated from beach sediments in Wellington, New Zealand, for morphological and molecular analyses. Two of these species, D esmodorella verscheldei sp. nov. and D racograllus ngakei sp. nov. , were new to science and are described herein. Epsilonema rugatum Lorenzen, 1973, comb. nov. , which was originally described from New Zealand material as a subspecies of Epsilonema dentatum from Chile, is redescribed and elevated to the rank of species based on cuticular ornamentation. The phylogenetic relationships amongst the three Desmodoroidea families are investigated based on new and existing sequences of the D2 and D3 expansions segments of large subunit (LSU) 28S rRNA gene and small subunit (SSU) of 18S rDNA gene. Our analyses suggest that the Draconematidae is a sister taxon to the Desmodorinae and Spiriniinae, with the Draconematidae forming a monophyletic crown group and the Desmodorinae and Spiriniinae forming a paraphyletic stem group. Phylogenetic relationships between the Epsilonematidae and Stilbonematinae, however, could not be determined with certainty. The SSU and D2‐D3 of LSU consensus trees indicate that the morphological resemblance between the Draconematidae and Epsilonematidae, which are both characterized by swollen pharyngeal body regions and mid‐posterior body regions with specialized setae, reflects distinct and independently evolved adaptations to their unusual mode of locomotion, with differences in the structure and distribution of specialized setae between the two families also consistent with convergent evolution. We show that the family Desmodoridae and superfamily Desmodoroidea as currently defined are not monophyletic. It was not possible to determine whether the Prodesmodorinae are more closely related to the Desmodoroidea or Microlaimoidea, although it is clear that they do not belong to the Desmodoridae. The single Molgolaiminae sequence available formed a distinct clade together with the superfamily Microlaimoidea, and should therefore be placed with the latter. Clarifying the phylogenetic relationships within the Desmodoroidea will require greater focus on the Pseudonchinae, Molgolaiminae, and Epsilonematidae, for which no or very few sequences are available at present. © 2016 The Linnean Society of London     

A reanalysis of Parabuthus (Scorpiones: Buthidae) phylogeny with descriptions of two new Parabuthus species endemic to the Central Namib gravel plains,Namibia

Two new thick‐tail scorpions in the genus Parabuthus Pocock, 1890 are described from the gravel plains of the Central Namib Desert, Namibia: Parabuthus glabrimanus sp. nov. ; Parabuthus setiventer sp. nov. The two new species occupy discrete distributional ranges, allopatric with the closely related species Parabuthus gracilis Lamoral, 1979 and Parabuthus nanus Lamoral, 1979. The distributions of the four species are mapped and a key provided for their identification. Revised diagnoses are provided for P. gracilis and P. nanus. The two new species are added to a previously published morphological character matrix for Parabuthus species and their phylogenetic positions determined in a reanalysis of Parabuthus phylogeny. Parabuthus setiventer sp. nov. is found to be the sister species of P. nanus, whereas P. glabrimanus sp. nov. is sister to a monophyletic group comprising P. gracilis, P. nanus, and P. setiventer sp. nov. The discovery of two new scorpion species endemic to the Central Namib gravel plains contributes to a growing body of evidence that this barren and desolate region is a hotspot of arachnid species richness and endemism. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159 , 673–710.     

The phylogenetic position of the Aeolosomatidae and Parergodrilidae, two enigmatic oligochaete-like taxa of the 'Polychaeta', based on molecular data from 18S rDNA sequences

The Aeolosomatidae and the Parergodrilidae are meiofaunal Annelida showing different combinations of clitellate‐like and non‐clitellate character states. Their phylogenetic positions and their systematic status within the Annelida are still in debate. Here we attempt to infer their systematic position using 18S rDNA sequences of the aeolosomatid Aeolosoma sp. and the parergodrilid Stygocapitella subterranea and several other meiofaunal taxa such as the Dinophilidae, Polygordiidae and Saccocirridae. The data matrix was complemented by sequences from several annelid, arthropod and molluscan species. After evaluation of the phylogenetic signal the data set was analysed with maximum‐parsimony, distance and maximum‐likelihood algorithms. Sequences from selected arthropods or molluscs were chosen for outgroup comparison. The resolution of the resulting phylogenies is discussed in comparison to previous studies. The results do not unequivocally support a sister‐group relationship of Aeolosoma sp. and the Clitellata. Instead, depending on the algorithms applied, Aeolosoma clusters in various clades within the polychaetes, for instance, together with eunicidan species, the Dinophilidae, Harmothoë impar or Nereis limbata. The position of Aeolosoma sp. thus cannot be resolved on the basis of the data available. S. subterranea always falls close to a cluster comprising Scoloplos armiger, Questa paucibranchiata and Magelona mirabilis, all of which were resolved as not closely related to both Aeolosoma sp. and the Clitellata. Therefore, convergent evolution of clitellate‐like characters in S. subterranea and hence in the Parergodrilidae is suggested by our phylogenetic analysis. Moreover, the Clitellata form a monophyletic clade within the paraphyletic polychaetes.