A brief review of holopelagic annelids

Annelids are one of the most successful major animal lineagesin terms of number of species and of habitats occupied. Despiteannelids being common in terrestrial, aquatic, and marine environments,only a limited number of lineages have evolved a holopelagicexistence. Most of these holopelagic lineages belong to Phyllodocida(nereidids, syllids, scale worms, and jawed worms) and moreparticularly often within the family Phyllodocidae. These wormsgenerally appear to retain many characteristics of adult annelids.Moreover, we provide molecular evidence showing that the well-knownalciopids are derived from within Phyllodocidae. In contrast,at least two lineages, Poeobius meseres/Flota flabelligera andprobably Chaetopterus pugaporcinus, are derived through paedomorphicprocesses acting on larvae from lineages that have sedentaryadult forms. Herein, we will briefly review the known diversityof holopelagic annelids with discussion of their evolutionaryorigins.     

Chimaeras and the origins of the holopelagic annelids Typhloscolecidae and Lopadorhynchidae: a reply to Struck & Halanych (2010)

Nygren, A. & Fredrik, P. (2010). Chimaeras and the origins of the holopelagic annelids Typhloscolecidae and Lopadorhynchidae: a reply to Struck & Halanych. —Zoologica Scripta, 40, 112–114.     

A scaleless scale worm: Molecular evidence for the phylogenetic placement of Pisione remota (Pisionidae, Annelida)

Pisionidae is a group of interstitial worms whose phylogentic affinities have been enigmatic. They have been allied to different Phyllodocida taxa. Although originally associated with Glyceridae and Phyllodocidae, they are more recently considered to be related to scale worms. Scale worms are a well-defined taxon, Aphroditiformia, within Annelida due to the unique possession of dorsal scales called elytra. Pisionidae lack elytra but they have been grouped with scale worms because they possess two pairs of jaws with venom glands, also found in Glyceridae. Determining the phylogenetic position of Pisionidae is important for understanding if features such as elytra and venomous jaws are evolutionarily labile in annelid history. Therefore, we explored 18S rDNA and Cytochrome c Oxidase subunit I data from several Aphroditiformia, Pisionidae, and other Phyllodocida to determine the phylogenetic placement of Pisionidae. Maximum likelihood and Bayesian inference of separate and combined data sets were conducted. All analyses support a derived position of Pisionidae within Aphroditiformia, close to Pholoidae and Sigalionidae. The loss of elytra in Pisionidae is probably due to adaptation for interstitial life. Furthermore, the results reject a monophyletic Aphroditoidea comprising Acoetidae, Aphroditidae, Eulepethidae and Polynoidae. Thus, the possession of only simple chaetae is either symplesiomorphic or convergent.     

Holopelagic Poeobius meseres ("Poeobiidae," Annelida) is derived from benthic flabelligerid worms

Phylogenetic relationships among and within the more than 70 recognized families of Annelida are poorly understood. In some cases, such as the monotypic Poeobiidae, derived morphology hinders the ability to find convincing synapomorphies that help elucidate evolutionary origins. In such cases, molecular data can be useful. Poeobiidae consists of the holopelagic polychaete Poeobius meseres, which is typically found in midwater depths off California. Morphologists have speculated that it is close to or within Flabelligeridae, but definitive evidence was lacking. Herein we use maximum likelihood phylogenetic reconstruction methods to examine the nuclear 18S rDNA (SSU) gene and the mitochondrial cytochrome b gene. Our results strongly support the hypothesis that P. meseres is a highly derived flabelligerid annelid closely related to Therochaeta. Thus, Poeobiidae is a junior synonym for Flabelligeridae. This result raises interesting questions about the evolution of the holopelagic P. meseres from a benthic ancestral flabelligerid.     

A molecular phylogeny of annelids

We present parsimony analyses of annelids based on the largest taxon sample and most extensive molecular data set yet assembled, with two nuclear ribosomal genes (18S rDNA and the D1 region of 28S rDNA), one nuclear protein coding‐gene (Histone H3) and one mitochondrial ribosomal gene (16S rDNA) from 217 terminal taxa. Of these, 267 sequences are newly sequenced, and the remaining were obtained from GenBank. The included taxa are based on the criteria that the taxon must have 18S rDNA or at least two other loci. Our analyses show that 68% of annelid family ranked taxa represented by more than one taxon in our study are supported by a jackknife value > 50%. In spite of the size of our data set, the phylogenetic signal in the deepest part of the tree remains weak and the majority of the currently recognized major polychaete clades (except Amphinomida and Aphroditiformia) could not be recovered. Terbelliformia is monophyletic (with the exclusion of Pectinariidae, for which only 18S data were available), whereas members of taxa such as Phyllodocida, Cirratuliformia, Sabellida and Scolecida are scattered over the trees. Clitellata is monophyletic, although Dinophilidae should possibly be included, and Clitellata has a sister group within the polychaetes. One major problem is the current lack of knowledge on the closest relatives to annelids and the position of the annelid root. We suggest that the poor resolution in the basal parts of the trees presented here may be due to lack of signal connected to incomplete data sets both in terms of terminal and gene sampling, rapid radiation events and/or uneven evolutionary rates and long‐branch attraction. © The Willi Hennig Society 2006.     

Problems in polychaete systematics

Some of the intriguing issues in current polychaete systematics are reviewed. (1) The root of the `polychaete' tree. Currently there are two major hypotheses concerning the root position among polychaetes. One is based on rooting cladograms with outgroups such as Mollusca and result in simple-bodied taxa such as Opheliidae and Questidae forming a basal annelid grade along with Clitellata. Other hypotheses do not use outgroup rooting but involve scenarios on the evolution of the group and would place taxa in Aciculata as basal annelids, thus making Aciculata and Phyllodocida paraphyletic. Molecular sequence data has been of little help in resolving this issue thus far, largely due to limited taxon sampling. (2) Paraphyly. Owing, in part, to a tradition involving the emphasis on differences among taxa, and the application of Linnean ranks (e.g., family), paraphyly is undoubtedly a widespread phenomenon in polychaete systematics. An example of this has been proposed already for Spionidae. If the tree topology and rooting used by Blake & Arnofsky (1999) is correct, Spionidae is made paraphyletic by the recognition of the following four family-ranked taxa; Trochochaetidae, Poecilochaetidae, Longosomatidae and Uncispionidae. Another possible example is seen with Cirratulidae. A preliminary cladistic analysis shows that it is entirely possible that seven other taxa recognised as families may be nested within Cirratulidae. These include Acrocirridae, Ctenodrilidae, Fauveliopsidae, Flabelligeridae, Flotidae, Poeobiidae and Sternaspidae. (3) Problematic taxa. Apart from the problems exposed by the analysis of Cirratuliformia, the position of some of these groups, such as Aberranta, Alciopidae, Hesionides,Lopadorhynchidae, Microphthalmus,Nerillidae, Spinther,Tomopteridae and Sabellariidae, is discussed.     

Current status of annelid phylogeny

Annelida is an ecologically and morphologically diverse phylum within the Lophotrochozoa whose members occupy a wide range of environments and show diverse life styles. The phylogeny of this group comprising more than 17,000 species remained controversial for a long time. By using next-generation sequencing and phylogenomic analyses of huge data matrices, it was finally possible to reach a well-supported and resolved annelid backbone tree. Most annelid diversity is comprised in two reciprocal monophyletic groups, Sedentaria and Errantia, which are named after the predominant life style of their members. Errantia include Aciculata (Phyllodocida?+?Eunicida) and Protodriliformia, which is a taxon of interstitial polychaetes. Sedentaria comprise most of the polychaete families formerly classified as Canalipalpata or Scolecida, as well as the Clitellata. Six taxa branch as a basal grade outside of this major radiation: Oweniidae, Magelonidae, Chaetopteridae, Sipuncula, Amphinomida, and Lobatocerebrum. Oweniidae and Magelonidae form a monophyletic group which we name Palaeoannelida, which constitutes the sister taxon of the remaining annelids. The early splits of annelid phylogeny date back to the Cambrian. The new annelid phylogeny highlights the variability and lability of annelid body plans, and many instances of simplifications of body plan as adaptations to new life styles can be found. Therefore, annelids will be an appropriate model to understand major transitions in the evolution of Bilateria in general. Evolutionary developmental studies are one way to investigate macroevolutionary transition in annelids. We briefly summarize the state of developmental model organisms in Annelida and also propose new candidates on the background of the phylogeny.     

A reevaluation of Wiwaxia and the polychaetes of the Burgess Shale

Wiwaxia corrugata and the indisputable polychaetes of the Middle Cambrian Burgess Shale, particularly Canadia spinosa, have figured prominently in recent hypotheses about the early evolution of polychaete annelids. Based on similarities between the sclerites of Wiwaxia and the notochaetae of Canadia with the broad notochaetae (paleae) of Recent chrysopetalid polychaetes, these two fossil taxa have been variously treated as closely related to the most highly derived stem forms of the polychaete (and annelid) crown group or as members of a specific, Recent subgroup within Polychaeta, the order Phyllodocida. Chrysopetalidae is a member of Phyllodocida, which is part of the major polychaete clade Aciculata; the latter two taxa are distinguished by four and six well defined autapomorphic characters, respectively. The best preserved or otherwise appropriate fossils of Wiwaxia corrugata, Canadia spinosa and the other polychaetes of the Burgess Shale have been studied in detail in order to determine whether they possess any characters that could support the homology of wiwaxiid sclerites, canadiid notochaetae and chrysopetalid paleae. Most of these fossil taxa have significant autapomorphies but the specific characters of the Aciculata and Phyllodocida are entirely absent. It is demonstrated that constraining cladograms in such a manner that wiwaxiid sclerites, canadiid notochaetae and chrysopetalid paleae are homologous leads to results that are markedly unparsimonious. Furthermore, Canadia and the other polychaetes of the Burgess Shale cannot be referred to any extant subgroup within the Polychaeta and cannot be used to polarize character evolution within the annelid crown group. Apart from its dubious sclerites, Wiwaxia has no characters that could indicate any close relationship with Polychaeta or Annelida.     

中国近海多毛纲底栖类群目与科水平的分类

多毛纲为环节动物重要类群, 具有重要的生态与经济意义。多毛纲的分类体系一直处于变化中。本文根据前人支序分类等研究成果, 综合《中国动物志》等资料, 整理了我国近海多毛纲底栖类群目与科的分类检索。根据Rouse和Fauchald (1997)的分类系统, 中国近海多毛纲的底栖类群可分为头节虫目(Scolecida)、海稚虫目(Spionida)、蛰龙介目(Terebellida)、缨鳃虫目(Sabellida)、叶须虫目(Phyllodocida)和矶沙蚕目(Eunicida)。基于头部、躯干部等附属物形态特征, 整理建立了中国近海多毛纲底栖类群上述六目至科的分类检索表, 考证了该分类方法, 总结了近年来我国近海多毛纲的研究和分子生物学对多毛纲系统演化的贡献。     

Phylogeny of embiopterans (Insecta)

A cladistic analysis of embiopterans, based on 157 species (representing 70% of the known genera) and 186 morphological characters, is presented, as well as a molecular analysis for 22 taxa using genes encoding 16S, 18S and 28S rDNA and COI. Species of all known families are included, except Andesembiidae Ross (specimens of which are in a private collection). The evidence presented supports the monophyly of four of the families (Australembiidae, Oligotomidae, Teratembiidae, and Anisembiidae). Notoligotomidae is paraphyletic and included within the Afro‐neotropical family Archembiidae (which is also paraphyletic). The genera Embia, Cleomia, Macrembia, and Dihybocercus (Embiidae) form, together with Australembiidae, a group strongly supported by morphology; the position of the remaining genera of Embiidae has two quite different resolutions. Almost 80% of the genera of Anisembiidae recently described appear as either paraphyletic or polyphyletic. Contrary to the opinion of other specialists, the major groups as well as the monophyly of some families are supported by features which have been ignored in classical approaches to the systematics of Embioptera, such as the ovipositor and cephalic and leg structures, characters with an almost perfect fit.     

Comparative mitochondrial and chloroplast genomics of a genetically distinct form of Sargassum contributing to recent “Golden Tides” in the Western Atlantic

Over the past 5 years, massive accumulations of holopelagic species of the brown macroalga Sargassum in coastal areas of the Caribbean have created “golden tides” that threaten local biodiversity and trigger economic losses associated with beach deterioration and impact on fisheries and tourism. In 2015, the first report identifying the cause of these extreme events implicated a rare form of the holopelagic species Sargassum natans (form VIII). However, since the first mention of S. natans VIII in the 1930s, based solely on morphological characters, no molecular data have confirmed this identification. We generated full‐length mitogenomes and partial chloroplast genomes of all representative holopelagic Sargassum species, S. fluitans III and S. natans I alongside the putatively rare S. natans VIII, to demonstrate small but consistent differences between S. natans I and VIII (7 bp differences out of the 34,727). Our comparative analyses also revealed that both S. natans I and S. natans VIII share a very close phylogenetic relationship with S. fluitans III (94‐ and 96‐bp differences of 34,727). We designed novel primers that amplified regions of the cox2 and cox3 marker genes with consistent polymorphic sites that enabled differentiation between the two S. natans forms (I and VIII) from each other and both from S. fluitans III in over 150 Sargassum samples including those from the 2014 golden tide event. Despite remarkable gene synteny and sequence conservation, the three Sargassum forms differ in morphology, ecology, and distribution patterns, warranting more extensive interrogation of holopelagic Sargassum genomes as a whole.     

Changes in pelagic Polychaete assemblages along the California current system

The composition, abundance, species richness and structural changes of the planktonic polychaete assemblages were analysed along a latitudinal transect in the California Current System (California, U.S.A. and Baja California, Mexico). The biological (species and abundance) and physical (temperature and salinity) data were analysed using Principal Component Analysis (PCA). The principal water masses in the survey area were determined. Twenty-four holoplanktonic species belonging to families Alciopidae, Iospilidae, Lopadorhynchidae, Tomopteridae and Typhloscolecidae were identified. Three clear species assemblages were discerned in the PCA results: 1. A `north group' (from Oregon-California border to San Francisco), with relatively high species richness (11) and the highest mean abundance (121 ind. per 500 m<sup>3</sup>) was characterised by Tomopteris septentrionalis, T. planktonis, Plotohelmis tenuis, and Travisiopis lobifera. California Current Water and Subtropical Central Water were present in the area occupied by this assemblage. 2. A `south group' (from off Bahía Magdalena to Cabo San Lucas), with the highest species richness (16), but low mean abundance (37.8 ind. per 500 m³); it included tropical affinity species, such Lopadorhynchus henseni, Tomopteris nationalis, and Travisiopsis dubia. In concordance Surface Equatorial Water was identified in this region. 3. A `transition group' (between the north and south regions) recorded the lowest mean abundance (2.3 ind. per 500 m³) and species richness (9). Only the California Current Water was detected in this area. The spatial pattern of species richness found along of this transect, was at least, partially due to the planktonic productivity distribution in the epipelagic region and the influence of several water masses coming from different directions.     

Position and Delineation of Chrysopetalidae and Hesionidae (Annelida,Polychaeta, Phyllodocida)

Previous studies suggest that the polychaete taxa Hesionidae and Chrysopetalidae may not represent separate groups, that Pilargidae constitute a subgroup within Hesionidae, and that Hesionides and Microphthalmus are highly derived hesionids. Phylogenetic systematic analyses of Phyllodocida and the subgroup Nereidiformia are presented in order to clarify the position and delineation of these taxa. The phyllodocida analysis includes 18 families representing the majority of the taxa in the group, is rooted with Onuphidae, and is based on 42 absent/present coded morphological characters, obtained mainly from literature. All 69 resulting shortest trees include the clade (Chrysopetalidae, Nereididae, Hesionidae), but with either Syllidae, Nautiliniellidae, Pilargidae or (Aphroditiformia, Pisionidae) as sister. In- and outgroup taxon selection for the Nereidiformia study is dictated by the outcome of Phyllodocida analysis, with scores based on examined species of two chrysopetalids, four hesionids, one nereid, one pilargid, one pisionid, one syllid, plus the putative hesionids Hesionides arenaria and Microphthalmus sp. It is based on 46 absent/present coded morphological characters. Two equally parsimonious trees indicate that chrysopetalids and hesionids are well delineated, that pilargids and hesionids are non-overlapping, and that Microphthalmus and Hesionides are not hesionids.     

On the Anatomy of the Central Nervous System of Phyllodocidae (Polychaeta) and the Phylogeny of Phyllodocid Genera: a New Alternative

A reinvestigation of the central nervous system of the Phyllodocidae appeared necessary as the existing literature proved insufficient for detailed comparisons with other polychaete families. Many earlier opinions turned out to be quite contradictory, especially as concerns the morphological value of the anterior end appendages. In the present paper the microanatomy of the brain and the anterior ventral cord, as well as the innervation of, inter alia, the cephalic appendages, the nuchal organs and the alimentary canal of Eulalia viridis (main object of study), Eumida sanguinea, Pterocirrus macroceros, Sige fusigera, Eteone picta, E. foliosa, Phyllodoce groenlandica, P. longipes, P. mucosa, Paranaitis wahlbergi, Notophyllum foliosum and Nereiphylla lutea are described. The results are summarized in schematic diagrams and compared with corresponding observations on other polychaete families. Thereby, some earlier opinions about the nature of the cephalic appendages of the Phyllodocidae are strengthened while others are rejected. Most significantly, it is concluded that the so-called ventrolateral antennae of phyllodocids are homologous with the palps of other polychaetes and the median antenna found in certain genera is homologous with the nuchal papilla found in most other genera of the family. These partly new or revised characters and character states formed the basis for a reconsideration of earlier ideas about the phylogeny of the Phyllodocidae: a new alternative.     

Identifying the sister group to the bees: a molecular phylogeny of Aculeata with an emphasis on the superfamily Apoidea

Debevec, AH., Cardinal, S & Danforth, BN. Identifying the sister group to the bees: a molecular phylogeny of Aculeata with an emphasis on the superfamily Apoidea. —Zoologica Scripta, 41, 527–535. The hymenopteran superfamily Apoidea includes the bees (Anthophila) as well as four predatory wasp families (Heterogynaidae, Ampulicidae, Sphecidae and Crabronidae) collectively referred to as the “sphecoid” or “apoid” wasps. The most widely cited studies suggest that bees are sister to the wasp family Crabronidae, but alternative hypotheses have been proposed based on both morphological and molecular data. We combined DNA sequence data from previously published studies and newly generated data for four nuclear genes (28S, long‐wavelength rhodopsin, elongation factor‐1α and wingless) to identify the likely sister group to the bees. Analysis of our four‐gene data set by maximum likelihood and Bayesian methods indicates that bees most likely arise from within a paraphyletic Crabronidae. Possible sister groups to the bees include Philanthinae, Pemphredoninae or Philanthinae + Pemphredoninae. We used Bayesian methods to explore the robustness of our results. Bayes Factor tests strongly rejected the hypotheses of crabronid monophyly as well as placement of Heterogynaidae within Crabronidae. Our results were also stable to alternative rootings of the bees. These findings provide additional support for the hypothesis that bees arise from within Crabronidae, rather than being sister to Crabronidae, thus altering our understanding of bee ancestry and evolutionary history.     

Additional insights into phylogenetic relationships of the Class Ophiuroidea (Echinodermata) from rRNA gene sequences

Ophiuroids are important benthic marine invertebrates with an unstable taxonomic history. Recent phylogenetic and morphological investigations have promoted major departures from established taxonomy. As such, additional insights into evolutionary relationships of ophiuroids are valuable. We analyzed ribosomal sequence data from a mitochondrial gene (16S rRNA) and a nuclear gene (18S rRNA) from 39 ophiuroids representing 14 of the 18 currently accepted families. Main findings from our study include support for a polyphyletic Ophiomyxidae, paraphyletic Amphiuridae, monophyletic Euryalida, a sister relationship between the Ophiactidae and Ophiotrichidae, and a clade comprised of the Amphiuridae and Amphilepididae. Relationships within the families Gorgonocephalidae, Ophiuridae, Ophiodermatidae and Ophiomyxidae are discussed.     

Myoanatomy of Myzostoma cirriferum (Annelida,Myzostomida): Implications for the evolution of the myzostomid body plan

Studies of rare genomic marker systems suggest that Myzostomida are a subgroup of Annelida and phylogenomic analyses indicate an early divergence of this taxon within annelids. However, adult myzostomids show a highly specialized body plan, which lacks typical annelid features, such as external body annulation, coelomic cavities with metanephridia, and segmental ganglia of the nervous system. The putative loss of these features might be due to the parasitic/symbiotic lifestyle of myzostomids associated with echinoderms. In contrast, the larval anatomy and adult locomotory system resemble those of annelids. To clarify whether the myoanatomy of myzostomids reflects their relationship to annelids, we analyzed the distribution of f‐actin, a common component of muscle fibers, in specimens of Myzostoma cirriferum using phalloidin‐rhodamine labeling in conjunction with confocal laser‐scanning microscopy. Our data reveal that the musculature of the myzostomid body comprises an outer circular layer, an inner longitudinal layer, numerous dorsoventral muscles, and prominent muscles of the parapodial complex. These features correspond well with the common organization of the muscular system in Annelida. In contrast to other annelids, however, several elements of the muscular system in M. cirriferum, including the musculature of the body wall, and the parapodial flexor muscles, exhibit radial symmetry overlaying a bilateral body plan. These findings are in line with the annelid affinity of myzostomids and suggest that the apparent partial radial symmetry of M. cirriferum arose secondarily in this species. Based on our data, we provide a scenario on the rearrangements of muscle fibers that might have taken place in the lineage leading to this species. J. Morphol., 2013. © 2012 Wiley Periodicals, Inc.     

Reconciling taxonomy and phylogeny in the bristleworm family Eunicidae (polychaete,Annelida)

Eunicid annelids inhabit diverse marine habitats worldwide, have ecological and economic importance and have been pictured in the news as giant predator worms. They compose a traditional stable taxon recently supported as monophyletic but characterized by plesiomorphies. Most genera within the family have been recovered as paraphyletic in previous studies. We present a phylogenetic hypothesis for eunicid based on molecular (COI, 16S rDNA, 18S rDNA) and morphological data (213 characters), including an explicit attempt to account for serial homology. Eunicidae as well as monophyletic genera Marphysa sensu stricto and Lysidice is redefined based on synapomorphies. Nematonereis is synonymized to Lysidice. Leodice and Nicidion are resurrected to name monophyletic groups including species previously included in Eunice and Marphysa sensu lato. Traditional diagnostic characters such as the absence/presence of peristomial cirri, lateral antennae and branchiae are homoplasies and not informative at the generic level. Different coding of traditional characters (i.e. articulation of prostomial appendages) and novel characters of prostomial features and regionalization of the body support the monophyly of the family and genera level clades. Thus, the phylogenetic hypothesis presented here and the evolution of characters provided background information for taxonomic changes yielding evolutionary meaningful classification and diagnoses for the family and genera.     

Worms by number

This paper investigates alternation patterns in length, shape and orientation of dorsal cirri (fleshy segmental appendages) of phyllodocidans, a large group of polychaete worms (Annelida). We document the alternation patterns in several families of Phyllodocida (Syllidae, Hesionidae, Sigalionidae, Polynoidae, Aphroditidae and Acoetidae) and identify the simple mathematical rule bases that describe the progression of these sequences. Two fundamentally different binary alternation patterns were found on the first four segments: 1011 for nereidiform families and 1010 for aphroditiform families. The alternation pattern in all aphroditiform families matches a simple one-dimensional cellular automaton and that for Syllidae (nereidiform) matches the Fibonacci string sequence. Hesionidae (nereidiform) showed the greatest variation in alternation patterns, but all corresponded to various known substitution rules. Comparison of binary patterns of the first 22 segments using a distance measure supports the current ideas on phylogeny within Phyllodocida. These results suggest that gene(s) involved in post-larval segmental growth employ a switching sequence that corresponds to simple mathematical substitution rules.