Direction of evolution within Annelida and the definition of Pleistoannelida

The direction of evolution within Annelida has been a long and contentious debate since the middle of the last century. Based on functional morphology two hypotheses were predominantly discussed. The first hypothesis proposed an earthworm‐like ancestor and the evolution of a segmented coelom as an aid for peristaltic burrowing, whereas the second one postulated that the ancestral annelid resembled more an errant polychaete and the segmented coelom evolved as a requirement for blood transportation to the parapodia. A recent phylogenomic study of annelids as well as new fossil records of early annelids provide new insights to this debate, which will be discussed in this paper. Based on these data the ancestral annelid is most likely a mixture of sedentary and errant annelid features indicating an evolutionary origin of Annelida as a microphagous surface deposit‐feeder crawling upon and through soft‐bottom habitats. Hence, the evolution of iterative parapodia as locomotive devices most likely drove the evolution of segmentation in Annelida. Moreover, as Struck et al. (2011) did not provide a proper definition and name for their clade 1 this clade shall be named Pleistoannelida and is defined by the last common ancestor of Sedentaria and Errantia sensu Struck et al. (2011), and all the descendants of that ancestor.     

Developmental architecture of the nervous system in Themiste lageniformis (Sipuncula): New evidence from confocal laser scanning microscopy and gene expression

Sipuncula is a clade of unsegmented marine worms that are currently placed among the basal radiation of conspicuously segmented Annelida. Their new location provides a unique opportunity to reinvestigate the evolution and development of segmented body plans. Neural segmentation is clearly evident during ganglionic ventral nerve cord (VNC) formation across Sedentaria and Errantia, which includes the majority of annelids. However, recent studies show that some annelid taxa outside of Sedentaria and Errantia have a medullary cord, without ganglia, as adults. Importantly, neural development in these taxa is understudied and interpretation can vary widely. For example, reports in sipunculans range from no evidence of segmentation to vestigial segmentation as inferred from a few pairs of serially repeated neuronal cell bodies along the VNC. We investigated patterns of pan-neuronal, neuronal subtype, and axonal markers using immunohistochemistry and whole mount in situ hybridization (WMISH) during neural development in an indirect-developing sipunculan, Themiste lageniformis. Confocal imaging revealed two clusters of 5HT⁺ neurons, two pairs of FMRF⁺ neurons, and Tubulin⁺ peripheral neurites that appear to be serially positioned along the VNC, similar to other sipunculans, to other annelids, and to spiralian taxa outside of Annelida. WMISH of a synaptotagmin1 ortholog in T. lageniformis (Tl-syt1) showed expression throughout the centralized nervous system (CNS), including the VNC where it appears to correlate with mature 5HT⁺ and FMRF⁺ neurons. An ortholog of elav1 (Tl-elav1) showed expression in differentiated neurons of the CNS with continuous expression in the VNC, supporting evidence of a medullary cord, and refuting evidence of ontogenetic segmentation during formation of the nervous system. Thus, we conclude that sipunculans do not exhibit any signs of morphological segmentation during development.     

Unsegmented annelids? Possible origins of four lophotrochozoan worm taxa

In traditional classification schemes, the Annelida consists of the Polychaeta and the Clitellata (the latter including the Oligochaeta and Hirudinida). However, recent analyses suggest that annelids are much more diverse than traditionally believed, and that polychaetes are paraphyletic. Specifically, some lesser-known taxa (previously regarded as separate phyla) appear to fall within the annelid radiation. Abundant molecular, developmental, and morphological data show that the Siboglinidae, which includes the formerly recognized Pogonophora and Vestimentifera, are derived annelids; recent data from the Elongation Factor-1α (EF-1α) gene also suggest that echiurids are of annelid ancestry. Further, the phylogenetic origins of two other lesser-known groups of marine worms, the Myzostomida and Sipuncula, have recently been called into question. Whereas some authors advocate annelid affinities, others argue that these taxa do not fall within the annelid radiation. With advances in our understanding of annelid phylogeny, our perceptions of body plan evolution within the Metazoa are changing. The evolution of segmentation probably is more plastic than traditionally believed. However, as our understanding of organismal evolution is being revised, we are also forced to reconsider the specific characters being examined. Should segmentation be considered a developmental process or an ontological endpoint?     

MicroRNAs resolve an apparent conflict between annelid systematics and their fossil record

Both the monophyly and inter-relationships of the major annelid groups have remained uncertain, despite intensive research on both morphology and molecular sequences. Morphological cladistic analyses indicate that Annelida is monophyletic and consists of two monophyletic groups, the clitellates and polychaetes, whereas molecular phylogenetic analyses suggest that polychaetes are paraphyletic and that sipunculans are crown-group annelids. Both the monophyly of polychaetes and the placement of sipunculans within annelids are in conflict with the annelid fossil record—the former because Cambrian stem taxa are similar to modern polychaetes in possessing biramous parapodia, suggesting that clitellates are derived from polychaetes; the latter because although fossil sipunculans are known from the Early Cambrian, crown-group annelids do not appear until the latest Cambrian. Here we apply a different data source, the presence versus absence of specific microRNAs—genes that encode approximately 22 nucleotide non-coding regulatory RNAs—to the problem of annelid phylogenetics. We show that annelids are monophyletic with respect to sipunculans, and polychaetes are paraphyletic with respect to the clitellate Lumbricus, conclusions that are consistent with the fossil record. Further, sipunculans resolve as the sister group of the annelids, rooting the annelid tree, and revealing the polarity of the morphological change within this diverse lineage of animals.     

Polychaete phylogeny based on morphological data – a comparison of current attempts

Annelid phylogeny is one of the largest unresolved problems within the Metazoa. This is due to the enormous age of this taxon and also strongly influenced by the current discussion on the position of the Arthropoda, which traditionally is hypothesized to be the annelid sister taxon. Within the framework of recent discussions on the position of the Annelida, the ground pattern of this taxon is either a clitellate-like, parapodia-less dwelling organism or an organisms that resembles errant polychaetes in having parapodia and gills and probably being a predator. To solve this problem different attempts have been made in the past, cladistic analysis, scenario based plausibility considerations and a successive search for sister taxa base on isolated characters. These attempts are presented and critically discussed. There is at least strong support for the Annelida as wells as for several of its taxa above the level of traditional families; the monophyly of the Polychaeta, however, remains questionable. The term taxon is used here in the sense of group of things that share certain characteristics. Biological taxa are not necessarily monophyletic, although many of them turned out to be. In terms of phylogenetic systematics taxa should be monophyletic.     

Parasegmental appendage allocation in annelids and arthropods and the homology of parapodia and arthropodia

The new animal phylogeny disrupts the traditional taxon Articulata (uniting arthropods and annelids) and thus calls into question the homology of the body segments and appendages in the two groups. Recent work in the annelid Platynereis dumerilii has shown that although the set of genes involved in body segmentation is similar in the two groups, the body units of annelids correspond to arthropod parasegments not segments. This challenges traditional ideas about the homology of "segmental" organs in annelids and arthropods, including their appendages. Here I use the expression of engrailed, wingless and Distal-less in the arthropod Artemia franciscana to identify the parasegment boundary and the appendage primordia. I show that the early body organization including the appendage primordia is parasegmental and thus identical to the annelid organization and by deriving the different adult appendages from a common ground plan I suggest that annelid and arthropod appendages are homologous structures despite their different positions in the adult animals. This also has implications for the new animal phylogeny, because it suggests that Urprotostomia was not only parasegmented but also had parasegmental appendages similar to extant annelids, and that limb-less forms in the Protostomia are derived from limb-bearing forms.     

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.     

Origins of holopelagic Typhloscolecidae and Lopadorhynchidae within Phyllodocidae (Phyllodocida,Annelida)

Struck, T. H. & Halanych, K. M. (2010). Origins of holopelagic Typhloscolecidae and Lopadorhynchidae within Phyllodocidae (Phyllodocida, Annelida).—Zoologica Scripta, 39, 269–275. Several distinct lineages of annelids have evolved holopelagic life styles. Unfortunately, our knowledge of the biology and evolution of most of these groups is limited. Typhloscolecidae and Lopadorhynchidae are two such examples of recognized families of holopelagic annelids about which little is known. Both groups have a limited number of known species (13 and 15, respectively) and are rarely discussed in the literature other than to note occurrence. Placing these groups in a phylogenetic context has been difficult due to their seemingly unique morphology, and, to the best of our knowledge, lack of molecular data. Nonetheless, previous authors have suspected that they are members of Phyllodocida and perhaps within Phyllodocidae. To test such hypotheses, we have employed nuclear 18S and 28S ribosomal data in both a Bayesian inference and a maximum‐likelihood framework. The resultant topology indicates that typhloscolecids and lopadorhynchs are sister taxa nested within Phyllodocidae near another holopelagic taxon, alciopids. Whereas posterior probabilities strongly support this placement, an AU hypothesis testing approach and bootstrap values are more equivocal but both still strongly suggest Phyllodocidae affiliations. Recognition of Typhloscolecidae and Lopadorhynchidae as annelid families is called into question.     

The origin of annelids

Annelids are a phylum of segmented bilaterian animals that have become important components of ecosystems spanning terrestrial realms to the deep sea. Annelids are remarkably diverse, possessing high taxonomic diversity and exceptional morphological disparity, and have evolved numerous feeding strategies and ecologies. Their interrelationships and evolution have been the source of much controversy over the past century with the composition of the annelid crown group, the relationship of major groups and the body plan of the ancestral annelid having undergone major recent revisions. There is a convincing body of molecular evidence that polychaetes form a paraphyletic grade and that clitellates are derived polychaetes. The earliest stem group annelids from Cambrian Lagerstätten are errant, epibenthic polychaetes, confirming that biramous parapodia, head appendages and diverse, simple chaetae are primitive for annelids. Current evidence from molecular clocks and the fossil record suggest that crown group annelids are a Late Cambrian – Ordovician radiation, with clitellates radiating in the Late Palaeozoic. Their body fossil record is largely confined to deposits showing exceptional preservation and is punctuated by the acquisition of hard parts in major groups. The discovery of an Ordovician fossil with soft tissues has shown that machaeridians are in fact a clade of crown polychaetes. They were in existence for more than 200 million years and possess unique calcitic dorsal armour, allowing their mode of life and phylogeny to be interpreted in the context of the annelid body plan. We identify a novel clade of machaeridians, the Cuniculepadida, which exhibit a series of adaptations for burrowing.     

The articulation of annelids

The aim of this paper is to assess the monophyly of the Annelida. Also, recent cladistic analyses of metazoan taxa, using a variety of data, have shown incongruities with regards to annelids and associated taxa that should be resolved. The Platyhelminthes is selected as the taxon to root our minimal length trees and polarise our characters in a parsimony analysis; ingroup taxa being Mollusca, Nemertea, Sipuncula, Echiura, Pogonophora, Vestimentifera, Euarthropoda, Onycho-phora, and the groups most commonly regarded as true ‘annelids’, the Clitellata and Polychaeta. We use 13 characters and a total of 33 states. This results in 18 minimal length trees of 23 steps. The consensus tree has the topology (Platyhelminthes (Nemertea (Sipuncula Mollusca (Echiura (Polychaeta (Vestimcntifera Pogonophora) Clitellata (Euarthropoda Onychophora)))))). The name Articulata is applied to the Clitellata, Euarthropoda, Onychophora, Pogonophora, Polychaeta, and Vestimentifera. The Vestimentifera is the sister group to, or more likely a clade within, the frenulate pogonophores, and the name Pogonophora is retained for this group. In half of the 18 minimal length trees, the traditionally formulated Annelida, i.e. Polychaeta and Clitellata, is paraphyletic if the Pogonophora are excluded. In the remaining minimal length trees, a monophyletic Annelida cannot be formulated. The name Annelida should not be used unless relationships within the Articulata are resolved to show it is a monophyletic taxon. The taxon name Articulata, originally formulated to include the Annelida and Arthropoda by Cuvier, is defined as the clade stemming from the first ancestor to show repetition of homologous body structures derived by teloblastic growth with a pygidial growth zone (segmentation) and longitudinal muscles broken into bands. The Articulata is considered, on current evidence, to consist of four monophyletic groups; the Arthropoda, Clitellata, Polychaeta, and Pogonophora, though the latter group may be a clade of polychaetes. If this is shown, the Pogonophora should revert to the original family name Lamellisabellidae Uschakov, 1933. An indented classification reflective of the cladistic pattern is provided. Other recent hypotheses about metazoan systematics arc analysed.     

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.     

What is convergent/homoplastic in Pogonophora?*

Characters of Pogonophora, including Perviata (Frenulata and Scleronlinida) and Vestimentifera, are critically analysed and the characters re‐evaluated with respect to their reliability in determining relationships with Annelida‐Polychaeta in general and with Oweniidae in particular, as well as with Oligomera. In addition to several differentiations which possibly express parallelisms, only a few characters reflect trustworthy information based on the current state of knowledge. These, however, still include ‘sets’ of both Annelida‐Polychaeta features and Oligomera features; despite the more recent trend to regard the Pogonophora as annelids, no unequivocal preference can be given as to which of these ‘sets’ is convergent/homoplastic. The intra‐epithelial nervous system without true cerebral ganglia, typical of Epineuralia as well as Pogonophora and Oweniidae, is here considered to represent the paedomorphic peripheral‐epidermal larval system of Gastroneuralia in contrast to the ganglionated gastroneural adult system.     

A review of molecular markers used for Annelid phylogenetics

Annelida, one of the most successful animal phyla, exhibitsan amazing variety of morphological forms. Disparity betweensome of the forms is so great that until molecular tools wereused, some annelid lineages (for example, echiurids and pogonophorans)were not commonly recognized as belonging to the group. Althoughit is easy to assign annelids to a given family, understandingthe deeper relationships within the group has been difficult.The main working hypothesis for annelid phylogeny is based onmorphological cladistic analysis. However, the recent work usingmolecular tools has caused a revision of our view of annelidevolutionary history. For example, Scolecida and Palpata appearnot to be natural groups, and the phylogenetic positions ofsome aberrant taxa (for example, Siboglinidae, Poeobius, Pisione)have been determined. Herein, we discuss some of the main molecularmarkers that have been used to elucidate annelid phylogeny andthe contribution that such work is making to our understanding.A table highlighting the molecular literature and the genesused is included.     

Phylogenomics of Annelida revisited: a cladistic approach using genome‐wide expressed sequence tag data mining and examining the effects of missing data

We present phylogenomic analyses of the most comprehensive molecular character set compiled for Annelida and its constituent taxa, including over 347 000 aligned nucleotide sites for 39 taxa. The nucleotide data set was recovered using a pre‐existing amino acid data set of almost 48 000 aligned sites as a backbone for tBLASTn searches against NCBI. In addition, orthology determinations of the loci in the original amino acid data set were scrutinized using an All vs All Reciprocal Best Hit approach, employing BLASTp, and examining for statistical interdependency among the loci. This approach revealed considerable sequence redundancy among the loci in the original data set and a new data set was compiled, with the redundancy removed. The newly compiled nucleotide data set, the original amino acid data set, and the new reduced amino acid data set were subjected to parsimony analyses and two forms of bootstrap resampling. The last‐named data set also was analysed using a maximum‐likelihood approach. There were two main objectives to these analyses: (i) to examine the general topology, including support, resulting from the analyses of the new data sets and (ii) to assess the consistency of the branching patterns across optimality criteria by comparison with previous probabilistic approaches. The phylogenetic hypotheses resulting from analyses of the three data sets are largely unsupported, reflecting the continued difficulty of finding numerous, reliable, and suitable loci for a group as ancient as Annelida. Resulting parsimonious hypotheses disagree, in some respects, with the previous probabilistic approaches; Sedentaria and, in most cases, Errantia are not supported as monophyletic groups but Pleistoannelida is recovered as a (unsupported) monophyletic group in one of the three parsimony analyses as well as the likelihood analysis. In addition, we performed missing data titration studies to estimate the impact of missing data on overall support and support for specific clades.     

Annelid phylogeny and the status of Sipuncula and Echiura

Background  

Annelida comprises an ancient and ecologically important animal phylum with over 16,500 described species and members are the dominant macrofauna of the deep sea. Traditionally, two major groups are distinguished: Clitellata (including earthworms, leeches) and "Polychaeta" (mostly marine worms). Recent analyses of molecular data suggest that Annelida may include other taxa once considered separate phyla (i.e., Echiura, and Sipuncula) and that Clitellata are derived annelids, thus rendering "Polychaeta" paraphyletic; however, this contradicts classification schemes of annelids developed from recent analyses of morphological characters. Given that deep-level evolutionary relationships of Annelida are poorly understood, we have analyzed comprehensive datasets based on nuclear and mitochondrial genes, and have applied rigorous testing of alternative hypotheses so that we can move towards the robust reconstruction of annelid history needed to interpret animal body plan evolution.     

Cellular and muscular growth patterns during sipunculan development

Sipuncula is a lophotrochozoan taxon with annelid affinities, albeit lacking segmentation of the adult body. Here, we present data on cell proliferation and myogenesis during development of three sipunculan species, Phascolosoma agassizii, Thysanocardia nigra, and Themiste pyroides. The first anlagen of the circular body wall muscles appear simultaneously and not subsequently as in the annelids. At the same time, the rudiments of four longitudinal retractor muscles appear. This supports the notion that four introvert retractors were part of the ancestral sipunculan bodyplan. The longitudinal muscle fibers form a pattern of densely arranged fibers around the retractor muscles, indicating that the latter evolved from modified longitudinal body wall muscles. For a short time interval, the distribution of S-phase mitotic cells shows a metameric pattern in the developing ventral nerve cord during the pelagosphera stage. This pattern disappears close to metamorphic competence. Our findings are congruent with data on sipunculan neurogenesis, as well as with recent molecular analyses that place Sipuncula within Annelida, and thus strongly support a segmental ancestry of Sipuncula.     

The phylogenetic position of the Clitellata and the Echiura - on the problematic assessment of absent characters*

Absent characters (negative characters) are difficult to assess and their correct interpretation as symplesiomorphies, synapomorphies or convergencies (homoplasies) is one of the greatest challenges in phylogenetic systematics. Different phylogenetic assessments often result in contradictory phylogenetic hypotheses, in which the direction of evolutionary changes is diametrically opposed. Especially in deciding between primary (plesiomorphic) and secondary (apomorphic) absence, false conclusions may be reached if only the outgroup comparison and the principle of parsimony are employed without attempting any biological evaluation or interpretation of characters. For example, in the higher‐level systematization of the Annelida and related taxa different assessments of absent characters have led to conflicting hypotheses about the phylogenetic relationships and the ground pattern of the annelid stem species. Varying phylogenetic interpretations regarding the absence of the chemosensory nuchal organs in the clitellates and their presence in polychaetes initiated a controversy that produced two alternative phylogenetic hypotheses: (1) the Clitellata are highly derived Annelida related to a subtaxon within the, in this case, paraphyletic ‘Polychaeta’ or (2) the Clitellata are comparatively primitive Annelida representing the sister group of a monophyletic taxon Polychaeta. In the former, the absence of nuchal organs in the Clitellata is regarded as a secondary character, in the latter as primary. As most Clitellata are either limnetic or terrestrial, we must ask which characters are plesiomorphies, taken from their marine stem species without changes. In addition to a thorough investigation and evaluation of clitellate characters, a promising approach to these questions is to look for such characters in limnetic and terrestrial annelids clearly not belonging to the Clitellata. A similar problem applies to the evaluation of the position of the Echiura, which lack both segmentation and nuchal organs. Evidence is presented that in both taxa these absent characters represent derived, apomorphic character states. The consequences for their phylogenetic position and the questionable monophyly of the Polychaeta are discussed. The conclusion drawn from morphological character assessments is in accordance with recently published hypotheses based on molecular data.     

Ultrastructure of pigmented adult eyes in errant polychaetes (Annelida): implications for annelid evolution

The evolution of photoreceptor cells and eyes in Metazoa is far from being resolved, although recent developmental and structural studies have provided strong evidence for a common origin of photoreceptor cells and existence of sister cell types already in early metazoans. These sister cell types are ciliary and rhabdomeric photoreceptor cells, depending on which part of each cell is involved in photoreception proper. However, a crucial point in eye evolution is how the enormous structural diversity of photoreceptor cells and visual systems developed, given the general molecular conservation of the photoreceptor cells. One example of this diversity can be observed in Annelida. Within the polychaetes the errant forms, taxon Aciculata, constitute the only group possessing true multicellular eyes in the adult stage. Thus far these organs have been investigated only in taxa of Phyllodocida, a subgroup of Aciculata. Data on Eunicida and Amphinomida as well as certain phyllodocidan taxa had been lacking. The ultrastructure of these adult eyes was investigated in various species of errant polychaetes, belonging to Amphinomidae, Eunicidae and Hesionidae, to elucidate whether they provide any phylogenetic clues regarding either the evolution of visual systems in Annelida or lophotrochozoan phylogeny in general. These eyes are composed of numerous supportive pigment cells and rhabdomeric photoreceptor cells and sometimes additional cell types. As a rule the pigment and rhabdomeric cell types form a continuous epithelium in which the two types intermingle. Presence of granules with shading pigment in sensory cells is a common feature but is apparently restricted to a taxon comprising Phyllodocida and Eunicida s. str. Very likely a lens-like structure does not belong to the ground pattern of annelid eyes, despite its presence in Phyllodocida. These lens-like structures are formed by secretions or cellular processes of the pigment cells. In many species the eye cup communicates with the exterior via a small cuticularized canal. This canal is interpreted as a rudiment due to the mode of formation in the epidermis. With respect to current phylogenetic hypotheses, these multicellular eyes have either been developed in the stem species of a taxon Aciculata nested within the polychaetes or have been evolved in the stem lineage of Annelida. Similarities to gastropod eyes are interpreted as convergent and not as indication of common origin. Except for the photoreceptor cells proper, the structure of the adult eyes in polychaetes most likely does not help to resolve lophotrochozoan phylogeny.     

Functionally diverse front-end desaturases are widespread in the phylum Annelida

Aquatic single-cell organisms have long been believed to be unique primary producers of omega-3 long-chain (≥C₂₀) polyunsaturated fatty acids (ω3 LC-PUFA). Multiple invertebrates including annelids have been discovered to possess methyl-end desaturases enabling key steps in the de novo synthesis of ω3 LC-PUFA, and thus potentially contributing to their production in the ocean. Along methyl-end desaturases, the repertoire and function of further LC-PUFA biosynthesising enzymes is largely missing in Annelida. In this study we examined the front-end desaturase gene repertoire across the phylum Annelida, from Polychaeta and Clitellata, major classes of annelids comprising most annelid diversity. We further characterised the functions of the encoded enzymes in selected representative species by using a heterologous expression system based in yeast, demonstrating that functions of Annelida front-end desaturases have highly diversified during their expansion in both terrestrial and aquatic ecosystems. We concluded that annelids possess at least two front-end desaturases with Δ5 and Δ6Δ8 desaturase regioselectivities, enabling all the desaturation reactions required to convert the C₁₈ precursors into the physiologically relevant LC-PUFA such as eicosapentaenoic and arachidonic acids, but not docosahexaenoic acid. Such a gene complement is conserved across the different taxonomic groups within Annelida.