Morphology disentangles the systematics of a ubiquitous but elusive meiofaunal group (Kinorhyncha: Pycnophyidae)

Kinorhyncha is a group of benthic, microscopic animals distributed worldwide in marine sediments. The phylum is divided into two classes, Cyclorhagida and Allomalorhagida, congruent with the two major clades recovered in recent phylogenetic analyses. Allomalorhagida accommodates more than one‐third of the described species, most of them assigned to the family Pycnophyidae. All previous phylogenetic analyses of the phylum recovered the two genera within Pycnophyidae, Pycnophyes and Kinorhynchus, as paraphyletic and polyphyletic. A major problem in these studies was the lack of molecular data of most pycnophyids, due to the limited and highly localized distribution of most species, often in the Arctic and the deep‐sea. We here overcame the problem by adding a morphological partition with data for 79 Pycnophyidae species, 15 of them also represented by molecular data. Model‐based analyses yielded seven clades, which each was supported by several morphological apomorphies. Accordingly, Kinorhynchus is synonymized with Pycnophyes and six new genera are described for the remaining recovered clades: Leiocanthus gen. nov., Cristaphyes gen. nov., Higginsium gen. nov., Krakenella gen. nov., Setaphyes gen. nov. and Fujuriphyes gen. nov.     

MOLECULAR PHYLOGENY AND REVISION OF THE GENUS NETRIUM (ZYGNEMATOPHYCEAE,STREPTOPHYTA): NUCLEOTAENIUM GEN. NOV.1

Nuclear‐encoded SSU rDNA, chloroplast LSU rDNA, and rbcL genes were sequenced from 53 strains of conjugating green algae (Zygnematophyceae, Streptophyta) and used to analyze phylogenetic relationships in the traditional order Zygnematales. Analyses of a concatenated data set (5,220 nt) established 12 well‐supported clades in the order; seven of these constituted a superclade, termed “Zygnemataceae.” Together with genera (Zygnema, Mougeotia) traditionally placed in the family Zygnemataceae, the “Zygnemataceae” also included representatives of the genera Cylindrocystis and Mesotaenium, traditionally placed in the family Mesotaeniaceae. A synapomorphic amino acid replacement (codon 192, cysteine replaced by valine) in the LSU of RUBISCO characterized this superclade. The traditional genera Netrium, Cylindrocystis, and Mesotaenium were shown to be para‐ or polyphyletic, highlighting the inadequacy of phenotypic traits used to define these genera. Species of the traditional genus Netrium were resolved as three well‐supported clades each distinct in the number of chloroplasts per cell, their surface morphology (structure and arrangement of lamellae) and the position of the nucleus or nuclear behavior during cell division. Based on molecular phylogenetic analyses and synapomorphic phenotypic traits, the genus Netrium has been revised, and a new genus, Nucleotaenium gen. nov., was established. The genus Planotaenium, also formerly a part of Netrium, was identified as the sister group of the derived Roya/Desmidiales clade and thus occupies a key position in the evolutionary radiation leading to the most species‐rich group of streptophyte green algae.     

Toward Modern Classification of Eustigmatophytes,Including the Description of Neomonodaceae Fam. Nov. and Three New Genera1

The class Eustigmatophyceae includes mostly coccoid, freshwater algae, although some genera are common in terrestrial habitats and two are primarily marine. The formal classification of the class, developed decades ago, does not fit the diversity and phylogeny of the group as presently known and is in urgent need of revision. This study concerns a clade informally known as the Pseudellipsoidion group of the order Eustigmatales, which was initially known to comprise seven strains with oval to ellipsoidal cells, some bearing a stipe. We examined those strains as well as 10 new ones and obtained 18S rDNA and rbcL gene sequences. The results from phylogenetic analyses of the sequence data were integrated with morphological data of vegetative and motile cells. Monophyly of the Pseudellipsoidion group is supported in both 18S rDNA and rbcL trees. The group is formalized as the new family Neomonodaceae comprising, in addition to Pseudellipsoidion, three newly erected genera. By establishing Neomonodus gen. nov. (with type species Neomonodus ovalis comb. nov.), we finally resolve the intricate taxonomic history of a species originally described as Monodus ovalis and later moved to the genera Characiopsis and Pseudocharaciopsis. Characiopsiella gen. nov. (with the type species Characiopsiella minima comb. nov.) and Munda gen. nov. (with the type species Munda aquilonaris) are established to accommodate additional representatives of the polyphyletic genus Characiopsis. A morphological feature common to all examined Neomonodaceae is the absence of a pyrenoid in the chloroplasts, which discriminates them from other morphologically similar yet unrelated eustigmatophytes (including other Characiopsis-like species).     

Phylogeny of split‐footed lacewings (Neuroptera,Nymphidae), with descriptions of new Cretaceous fossil species from China

A phylogeny of the lacewing family Nymphidae based on morphology and DNA sequences is presented including representatives of all living genera and selected fossil genera. Widely distributed Jurassic and Cretaceous genera gave rise to recent taxa now restricted to Australasia. Two previously defined clades (i.e. Nymphinae and Myiodactylinae) were recovered and reflect the diverging adult and larval morphology of members of these two subfamilies. From Chinese Cretaceous deposits, a new genus (Spilonymphes gen. nov.) is described with one new species, as well as new species described in the genera Baissoleon Makarkin and Sialium Westwood.     

MOLECULAR PHYLOGENY,ULTRASTRUCTURE, AND TAXONOMIC REVISION OF CHLOROGONIUM (CHLOROPHYTA): EMENDATION OF CHLOROGONIUM AND DESCRIPTION OF GUNGNIR GEN. NOV. AND RUSALKA GEN. NOV.1

We examined the molecular phylogeny and ultrastructure of Chlorogonium and related species to establish the natural taxonomy at the generic level. Phylogenetic analyses of 18S rRNA and RUBISCO LSU (rbcL) gene sequences revealed two separate clades of Chlorogonium from which Chlorogonium (Cg.) fusiforme Matv. was robustly separated. One clade comprised Cg. neglectum Pascher and Cg. kasakii Nozaki, whereas the other clade included the type species Cg. euchlorum (Ehrenb.) Ehrenb., Cg. elongatum (P. A. Dang.) Francé, and Cg. capillatum Nozaki, M. Watanabe et Aizawa. On the basis of unique ultrastructural characteristics, we described Gungnir Nakada gen. nov. comprising three species: G. neglectum (Pascher) Nakada comb. nov., G. mantoniae (H. Ettl) Nakada comb. nov., and G. kasakii (Nozaki) Nakada comb. nov. We also emended Chlorogonium as a monophyletic genus composed of Cg. euchlorum, Cg. elongatum, and Cg. capillatum. Because Cg. fusiforme was distinguished from the redefined Chlorogonium and Gungnir by the structure of its starch plate, which is associated with pyrenoids, we reclassified this species as Rusalka fusiformis (Matv.) Nakada gen. et comb. nov.     

Neomegamphopas kunduchii sp. nov. (Crust: Amphipoda) from East Africa, with a discussion of gnathopodal dominance in isaeid amphipods

A new species of Neomegamphopus Shoemaker, is described from East Africa. Ledoyerella gen. nov. is erected to include Lembos caputphotis Ledoyer and L. isochelatus Ledoyer, and Maragopsis gen. nov. to include Lemboides bidentata Ledoyer. It is suggested that the Aoridae sensu stricto are a distinct and easily recognizable family and should not be considered merely as polyphyletic offshoots of the Isaeidae. The hypothesis is put forward, that “neomegamphid” amphipods have been derived from ancestral isaeid stock which did not exhibit a marked secondary sexual development of the gnathopoda, while genera such as Aloiloi and Aorchoides are more recent derivatives of isaeids by a process of axial reversal of gnathopod dominance.     

Molecular phylogeny,analysis of character evolution,and submersible collections enable a new classification of a diverse group of gobies (Teleostei: Gobiidae: Nes subgroup), including nine new species and four new genera

The Nes subgroup of the Gobiosomatini (Teleostei: Gobiiformes: Gobiidae) is an ecologically diverse clade of fishes endemic to the tropical western Atlantic and eastern Pacific oceans. It has been suggested that morphological characters in gobies tend to evolve via reduction and loss associated with miniaturization, and this, coupled with the parallel evolution of adaptations to similar microhabitats, may lead to homoplasy and ultimately obscure our ability to discern phylogenetic relationships using morphological characters alone. This may be particularly true for the Nes subgroup of gobies, where several genera that are diagnosed by ‘reductive characters’ have been shown to be polyphyletic. Here we present the most comprehensive phylogeny to date of the Nes subgroup using mitochondrial and nuclear sequence data. We then evaluate the congruence between the distribution of morphological characters and our molecular tree using maximum‐likelihood ancestral state reconstruction, and test for phylogenetic signal in characters using Pagel's λ tree transformations (Nature, 401 , 1999 and 877). Our results indicate that all of the characters previously used to diagnose genera of the Nes subgroup display some degree of homoplasy with respect to our molecular tree; however, many characters display considerable phylogenetic signal and thus may be useful in diagnosing genera when used in combination with other characters. We present a new classification for the group in which all genera are monophyletic and in most cases diagnosed by combinations of morphological characters. The new classification includes four new genera and nine new species described here, many of which were collected from rarely sampled deep Caribbean reefs using manned submersibles. The group now contains 38 species in the genera Carrigobius gen. nov., Chriolepis, Eleotrica, Gobulus, Gymneleotris, Nes, Paedovaricus gen. nov., Pinnichthys gen. nov., Psilotris, and Varicus. Lastly, we provide a key to all named species of the Nes subgroup along with photographs and illustrations to aid in identification.     

Phylogeny of the deep‐sea cirripede family Scalpellidae (Crustacea,Thoracica) based on shell capitular plate morphology

A cladistic analysis of 23 extant species of the deep‐sea pedunculate cirripede family Scalpellidae was undertaken, based on 61 shell plate characters, and taking the Jurassic–Cretaceous scalpellomorph genus Cretiscalpellum as an out‐group. The consensus tree shows progressive morphological change from basal to more derived taxa, but a derived group is marked by major morphological innovation, including 27 character state changes that permit subdivision of the family into two sharply demarcated clades – the more basal group is here placed within a redefined Scalpellinae (Arcoscalpellum, A rcuatoscalpellum gen. nov. , D iotascalpellum gen. nov. , Graviscalpellum, R egioscalpellum gen. nov. , and Scalpellum), and a more derived group named A migdoscalpellinae subfam. nov. that shows numerous progressive trends in morphology, permitting the recognition of three genera (Amigdoscalpellum, Catherinum, and Weltnerium). The phylogeny is independently supported by a recently published multiple DNA marker‐based molecular phylogeny. The more basal Scalpellinae appeared in the Aptian (Early Cretaceous, 120 Mya), and derived Amigdoscalpellinae were already present by the Campanian (Late Cretaceous, 78 Mya), represented by C atherinum anglicum sp. nov. and Amigdoscalpellum bellulum from the UK Chalk. Specialized receptacles to accommodate dwarf males in the apical interior of the scutum evolved at least three times during the history of the scalpellids. © 2015 The Linnean Society of London     

New bugs of the family Archegocimicidae (Hemiptera: Heteroptera) from the Upper Jurassic of Mongolia

Two new genera and five new species of Archegocimicidae are described from the Upper Jurassic of the Shar-Teg locality in Mongolia: Shartegocimex rasnitsyni, gen. et sp. nov., S. distans, sp. nov., Shartegocorpus paranotalis, gen. et sp. nov., Saldonia formosa, sp. nov., S. insolita, sp. nov.     

Two new genera,Caveana gen. nov. and Triviola gen. nov., and two new Torodora species from Thailand (Lepidoptera: Lecithoceridae)

Two new genera, Caveana, gen. nov. and Triviola, gen. nov., belonging to the subfamily Torodorinae (Lecithoceridae) are described, based on Caveana diemseoki, sp. nov. and Triviola puiensis, sp. nov. At the same time, two new Torodora species, T. karismata, sp. nov. and T. youwon, sp. nov., are also described from Thailand. Images of adults and genitalia are given.     

Genetic diversity and phylogenetic position of the subclass Astomatia (Ciliophora) based on a sampling of six genera from West African oligochaetes (Glossoscolecidae, Megascolecidae), including description of the new genus Paraclausilocola n. gen

To more confidently assess phylogenetic relationships among astome ciliates, we obtained small subunit (SSU) rRNA sequences from nine species distributed in six genera and three families: Almophrya bivacuolata, Eudrilophrya complanata, Metaracoelophrya sp. 1, Metaracoelophrya sp. 2, Metaracoelophrya intermedia, Metaradiophrya sp., Njinella prolifera, Paraclausilocola constricta n. gen., n. sp., and Paraclausilocola elongata n. sp. The two new species in the proposed new clausilocolid genus Paraclausilocola n. gen. are astomes with no attachment apparatus, two files of contractile vacuoles, and an arc-like anterior suture that has differentiations of thigmotactic ciliature on the anterior ends of the left kineties of the upper surface. Phylogenetic analyses were undertaken using neighbor-joining, Bayesian inference, maximum likelihood, and maximum parsimony. The nine species of astomes formed a strongly supported clade, showing the subclass Astomatia to be monophyletic and a weakly supported sister clade to the scuticociliates. There were two strongly supported clades within the astomes. However, genera assigned to the same family were found in different clades, and genera assigned to the same order were found in both clades. Thus, astome taxa appear to be paraphyletic when morphology is used to assign species to genera.     

Morphology,morphogenesis, and molecular phylogeny of a new marine urostylid ciliate (Ciliophora,Stichotrichia) from the South China Sea,and a brief overview of the convergent evolution of the midventral pattern within the Spirotrichea

The morphology, infraciliature, morphogenetic events, and small subunit ribosomal RNA (SSU rRNA) gene‐based phylogeny of a stichotrich ciliate, Bergeriella ovata gen. et sp. nov. , found in coastal waters off Daya Bay, Guangdong, South China, were investigated. The new taxon has an atypical midventral pattern, and is characterized by several rather unusual morphological features: frontal cirri forming an irregular tricorona; one nonmigratory row located right of the midventral rows, which is morphogenetically derived from the posteriormost fronto‐ventral‐transverse (FVT) streak; conspicuously enlarged postoral ventral cirri, whereas the left ventral cirri are delicate and arranged in oblique rows. Based on both the morphological and morphogenetical data, a new family, Bergeriellidae fam. nov. , is proposed. In the SSU rRNA phylogenetic trees, B. ovata gen. et sp. nov. groups with the classic urostylids (urostylids s.s.) with high nodal support. Our phylogenetic analyses have revealed at least seven separate clades for midventral pattern‐bearing taxa (e.g. Pattersoniella, Neokeronopsis, Rigidothrix, urostylids s.s., Uroleptidae, Holostichidae + Psammomitridae, and Pseudoamphisiellidae), suggesting that the order Urostylida Jankowski, 1979 sensu Lynn, 2008 is a polyphyletic assemblage, and that the convergent evolution of the urostylid midventral pattern might have occurred more times among the spirotrichs than has previously been recognized. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 697–710.     

Molecular phylogenetic analysis of Podostemaceae: implications for taxonomy of major groups

The river‐weed family Podostemaceae (c. 300 species in c. 54 genera) shows a number of morphological innovations to be adapted to its unusual aquatic habitat, and its unique or rare bauplan features have been reflected in the traditional (i.e. non‐molecular) classification recognizing numerous monotypic or oligospecific genera. The infrasubfamilial relationships of many genera remained unclear. The present study used molecular phylogenetic analysis of matK sequences for 657 samples (c. 132 species/c. 43 genera). The family was traditionally divided into three subfamilies (Podostemoideae, Tristichoideae and Weddellinoideae). American Podostemoideae were shown to be polyphyletic and divided into four clades, i.e. Ceratolacis, Diamantina, Podostemum and all other genera. Among the podostemoid clades, Diamantina was the first branching clade and a clade comprising Mourera and the Apinagia subclade was then sister to the remainder of the New World and Old World Podostemoideae with low statistic supports. The Old World Podostemoideae comprised four monophyletic clades, i.e. two African clades, one Madagascan clade and one Asian clade, although the relationships among these clades and American Ceratolacis and Podostemum were poorly resolved. African Podostemoideae were polyphyletic, with Saxicolella pro parte being weakly supported as sister to the remaining Old World Podostemoideae plus Ceratolacis and Podostemum. In contrast to the American and African clades, monophyly of four Asian subclades was well supported. Plants of Tristicha (Tristichoideae) and of Weddellina (Weddellinoideae), which are currently treated as monospecific, had great matK differentiation equivalent to at least interspecific variation. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 169 , 461–492.     

Phylogeny and taxonomy of European funnel‐web spiders of the Tegenaria−Malthonica complex (Araneae: Agelenidae) based upon morphological and molecular data

The taxonomy and systematics of European house spiders, currently constituting the ill‐defined Tegenaria?Malthonica complex (including Aterigena) in the family Agelenidae, are revised. In Europe four monophyletic genera and 81 species are defined. One genus, Eratigena gen. nov. , and seven species are described as new; at species level 17 new synonyms and 20 new combinations are proposed, and the original combination of 14 species is reinstated. Five species could not be placed (incertae sedis) because of insufficient material and one taxon is regarded as ‘nomen dubium’. On the basis of a detailed morphological assessment, 88 characters were chosen for a cladistic analysis. Phylogenetically informative characters include mostly spination patterns as well as spinneret and genital structures. In addition to morphology, three gene sections [cytochrome c oxidase subunit 1 (CO1), nicotinamide adenine dinucleotide dehydrogenase subunit 1 (NADH1) 28S] were analysed. Morphological and molecular analyses were performed individually and in combination applying maximum parsimony and Bayesian tree search methods. In all resulting trees Malthonica and Tegenaria in their present composition are either polyphyletic or paraphyletic. Consequently, we redefined the two genera and erected a new genus, Eratigena gen. nov. Identification keys are provided for the European agelenid genera as well as for the European species of Tegenaria and Eratigena gen. nov. The genera and most of the constituent species are described and illustrated. The new classification has also been applied to some extra European members of the Tegenaria‐Malthonica complex resulting in additional three new synonyms, seven reversals to the original combination, and four new combinations. © 2013 The Linnean Society of London     

A NEW LOOK AT AN ANCIENT ORDER: GENERIC REVISION OF THE BANGIALES (RHODOPHYTA)1

The red algal order Bangiales has been revised as a result of detailed regional studies and the development of expert local knowledge of Bangiales floras, followed by collaborative global analyses based on wide taxon sampling and molecular analyses. Combined analyses of the nuclear SSU rRNA gene and the plastid RUBISCO LSU (rbcL) gene for 157 Bangiales taxa have been conducted. Fifteen genera of Bangiales, seven filamentous and eight foliose, are recognized. This classification includes five newly described and two resurrected genera. This revision constitutes a major change in understanding relationships and evolution in this order. The genus Porphyra is now restricted to five described species and a number of undescribed species. Other foliose taxa previously placed in Porphyra are now recognized to belong to the genera Boreophyllum gen. nov., Clymene gen. nov., Fuscifolium gen. nov., Lysithea gen. nov., Miuraea gen. nov., Pyropia, and Wildemania. Four of the seven filamentous genera recognized in our analyses already have generic names (Bangia, Dione, Minerva, and Pseudobangia), and are all currently monotypic. The unnamed filamentous genera are clearly composed of multiple species, and few of these species have names. Further research is required: the genus to which the marine taxon Bangia fuscopurpurea belongs is not known, and there are also a large number of species previously described as Porphyra for which nuclear SSU ribosomal RNA (nrSSU) or rbcL sequence data should be obtained so that they can be assigned to the appropriate genus.     

PHYLOGENY OF EUGLENOPHYCEAE BASED ON SMALL SUBUNIT rDNA SEQUENCES: TAXONOMIC IMPLICATIONS1

Small subunit rDNA sequences of 42 taxa belonging to 10 genera were used to infer phylogenetic relationships among euglenoids. Members of the phototrophic genera Euglena, Phacus, Lepocinclis, Colacium, Trachelomonas, and Strombomonas plus the osmotrophs Astasia longa, Khawkinea quartana, and Hyalophacus ocellatus were included. Six major clades were found in most trees using multiple methods. The utility of Bayesian analyses in resolving these clades is demonstrated. The genus Phacus was polyphyletic with taxa sorting into two main clades. The two clades correlated with overall morphology and corresponded in large part to the previously defined sections, Pleur‐ aspis Pochmann and Proterophacus Pochmann. Euglena was also polyphyletic and split into two clades. In Bayesian analyses species with less plastic pellicles and small disk‐like chloroplasts diverged at the base of the tree. They grouped into a single clade which included the two Lepocinclis spp., which also are rigid and bear similar chloroplasts. The metabolic Euglena species with larger plastids bearing pyrenoids and paramylon caps arose near the top of the tree. The loricates Strombomonas and Trachelomonas formed two well‐ supported, but paraphyletic, clades. The strong support for the individual clades confirmed the value of using lorica features as taxonomic criteria. The separation of the osmotrophic species A. longa, K. quartana, and H. ocellatus into different clades suggested that the loss of the photosynthetic ability has occurred multiple times.     

New radiolaria of the family Prunobrachidae from the uppermost Cretaceous of the eastern Polar Urals

New radiolarian genera and species from Upper Cretaceous siliceous clays of borehole 22 (Polar Urals) are described. The family Prunobrachidae Pessagno, 1975 is reviewed. The new genus Pseudobrachium gen. nov. has a dumbbell skeleton without a patagium and Spinibrachium gen. nov. has apophysis-like spines and a patagium in the central part of the skeleton. Five new species, Spinibrachium amoni, Prunobrachium kozlovae, P. boreale, Pseudobrachium gracile, and P. trilobatum, are described; they are of great importance for paleogeography and stratigraphic correlation of Campanian deposits.     

New Jurassic Cercopoidea from China and their evolutionary significance (Insecta: Hemiptera)

Abstract: A new family, Sinoalidae Wang and Szwedo fam. nov., is described from the Middle Jurassic Daohugou Biota (Inner Mongolia, China). Two new genera with three new species (Sinoala parallelivena Wang and Szwedo gen. et sp. nov., Jiania crebra Wang and Szwedo gen. et sp. nov., and Jiania gracila Wang and Szwedo sp. nov.) are erected based on well‐preserved whole‐bodied specimens. The morphological characters of Procercopidae and the new family are given, and a key to the species of Sinoalidae is presented. The genera Luanpingia Hong, 1983 and Huabeicercopis Hong, 1983 are revised and attributed to Sinoalidae. The genera Mesocercopis Hong, 1983 and Sinotettegarcta Hong, 1986 are considered junior synonyms of Anthoscytina Hong, 1983. Taxa assigned to Sinoalidae fam. nov. cannot be attributed to any previously known family, but share some general (plesiomorphic) characters with ancient Clypeata, for example, postclypeus distinctly swollen, transversely wrinkled, antenna with a flagellum of a few elongate segments, frons with median ocellus and lateral ocelli on crown. Moreover, the new family is closely related to Procercopidae (Cercopoidea) based on a pronotum with a median incision at the hind margin; tegmen slender, partly punctuate, basal portion of Sc not exceeding apex of basal cell, and hind legs with lateral spines. It has a mixture of ancestral characters shared with Hylicelloidea and Jurassic Procercopidae and some derived characters. Therefore, it is tentatively placed in Cercopoidea. A preliminary phylogram of Clypeata is presented based on the combination of fossil and morphological data.