On the Innervation and Homologues of the Anterior End Appendages of the Eunicea (Polychaeta), with a Tentative Outline of the Fundamental Constitution of the Cephalic Nervous System of the Polychaetes

Earlier papers dealing with the architecture of the cephalic nervous system of the Eunicea were studied. Thereby, a re-investigation appeared necessary: the existing literature proved insufficient for detailed comparisons with other polychaete families and many earlier statements and conclusions turned out to be quite contradictory, especially as concerns the homologues of the anterior end appendages. In the present paper, the microanatomy of the brain and the innervation of, inter alia, the antennae, the nuchal organs and the alimentary canal of Hyalinoecia tubicola, Nothria conchylega, Eunice norvegica and E. pennata are described. The results are summarized in schematic diagrams and tables and compared with corresponding observations in other polychaete families. Thereby, some earlier opinions about the morphological value of the cephalic appendages of the Eunicea are strengthened while others are rejected. On the basis of the present observations and earlier results arrived at by the present author, and as an object for future discussion and criticism, a diagram of the fundamental constitution of the cephalic nervous system of the Polychaeta is tentatively outlined.     

On the Microanatomy of the Cephalic Nervous System of Nereidae (Polychaeta), with a Preliminary Discussion of Some Earlier Theories on the Segmentation of the Polychaete Brain

Abstract Earlier papers dealing with the microanatomy of the nereid brain have been studied. On this basis a re-investigation of the cephalic nervous system and of the innervation and homologues of the anterior end appendages of these animals appeared necessary: the existing literature proved insufficient for detailed comparisons with other polychaete families and many earlier statements were quite contradictory. In the present paper, the brain commissures and the innervation of, inter alia, the antennae and the palps of Neanthes virens and Nereis pelagica are described. Special attention was paid to the roots of the circum-oesophageal connectives and the ganglia in this part of the nervous system. The results, summarized in schematic diagrams and tables, are compared with corresponding observations in 14 other polychaete families. In a discussion of the architecture of the polychaete nervous system as a phylogenetic instrument, the supposed segmentation of the polychaete brain is questioned and the idea that the configuration of the polychaete nervous system offers support to the cyclomer theory is rejected. Other conclusions concerning the relationships within the Polychaeta are pointed out.     

Ultrastructure of nuchal organs in some marine polychaetes

Polychaetes normally possess one pair of nuchal organs at the posterior edge of the prostomium or peristomium. They have been regarded as chemosensory organs. The nuchal organs of four marine polychaete species with different habits were investigated by electron microscopy. Although the shapes of nuchal organs can vary greatly from simple ciliary bands (Scolelepis squamata, Spionidae) to retractile tongue-like, piston- or finger-shaped forms (Eteone longa, Anaitides mucosa, Phyllodocidae; Heteromastus filiformis, Capitellidae), the structural components, including the ciliated supporting cells, sensory cells, and nuchal epidermal cells, are essentially similar. The differences basically concern 1) the position of the sensory cells with relation to the ciliated supporting cells, 2) the location and structure of the nuchal nerve, and 3) the structure of the nuchal cuticle. The diverging nature of this modified cuticle is described and discussed in detail. Comparisons are made with the fine structure of nuchal organs of other polychaete species. Similarities of cellular components of nuchal organs are found not only in the four species studied here but also in all nuchal organs investigated so far. This is hypothesized to be due to the fact that the polychaete stem species already possessed nuchal organs with the respective cell types. Differences in the number and distribution of cellular components and in the overall shape of nuchal organs are thought to have evolved in correlation with the equipment of other cephalic appendages and with different habits and modes of nutrition.     

On the Innervation and Homologues of the Cephalic Appendages of the Aphroditacea (Polychaeta)

Abstract Earlier papers dealing with the innervation and homologues of the anterior end appendages of some aphroditacean families were re-studied. However valuable these earlier works may be, in some respects they proved to be insufficient for detailed comparisons with other polychaete families and some of their statements are quite contradictory. This prompted a re-investigation of the central-most parts of the nervous system of representatives of the families Polynoidae, Sigalionidae, Aphroditidae and Acoetidae. In the present paper, the brain commissures and the innervation of, inter alia, the antennae, the palps and the ommatophores (when present) in Lepidonotus squamatus, Harmothoe longisetis, Leanira (Sthenolepis) tetragona, Laetmonice producta benthaliana, and Panthalis oerstedi are described. Special attention is paid to the much-debated question about the presence or absence of a palp ganglion. The results, summarized in schematic diagrams, are compared with corresponding observations of the brains of, above all, the ‘spiomorphic’ polychaetes. Using the first Remanian criterion for identifying homologies, equivalents in the aphroditacean brain and the central nervous system of ‘sedentary’ families are proposed. By this a broader base is established for the discussion regarding the fundamental constitution of the anterior end of the polychaetes and the structure and homologues of their cephalic appendages.     

On the Microanatomy of the Brain and the Innervation and Homologues of the Cephalic Appendages of Hesionidae and Syllidae (Polychaeta)

Abstract Earlier papers dealing with the anatomy of the hesionids and syllids were studied. Thereby it was found that information about the structure of the central nervous system was meagre. As a result, the anterior end appendages, especially the large, laterloventral ones of the Syllidae, have been differently interpreted. This prompted a re-investigation. The circum-oesophageal connectives, the brain commissures and the innervation of the alimentary canal and the cephalic appendages of a number of hesionid and syllid species were studied. The results, summarized in schematic diagrams, were compared with corresponding observations in other polychaete families. Among other things, it was concluded that not only the latero-ventral cephalic appendages of the hesionids but also those of the syllids are homologous with the palps of the nereids and of many other “errant” and “sedentary” families.     

On the Microanatomy of the Supraoesophageal Ganglion of Some Amphinomids (Polychaeta Errantia), with Further Discussion of the Innervation and Homologues of the Polychaete Palps

Earlier papers dealing with the anatomy of the amphinomid brain were studied. Discrepancies in the studied works prompted a re-investigation of the central nervous system of these animals. In this paper the brain commissures, the clusters of ganglion cells and the innervation of the antennae, the palps and the nuchal organs of Eurythoe complanata, Amphinome rostrata, Pherecardia striata, and Notopygos variabilis are described in some detail. The brain of Hermodice carunculata is studied more cursorily. The results, summarized in schematic diagrams, are compared with corresponding observations of the brains of ‘spiomorphic’ and ‘serpulimorphic’ polychaetes and to information given by earlier authors about structures in the Nereidiformia and Aphroditiformia. Using the major Remanian criteria for identifying homologies, equivalents in the amphinomid brain and the central nervous systems of some ‘sedentary’ and ‘errant’ families are proposed. As a consequence, some earlier opinions about the morphological value of the cephalic appendages of the amphinomids are strengthened while others are rejected. The necessity of further investigations of the polycheaete nervous system is emphasized.     

On the anatomy of the central nervous system and the morphological value of the anterior end appendages of Ampharetidae, Pectinariidae and Terebellidae (Polychaeta)

Earlier papers dealing with the anatomy of the central nervous system of ampharetids, pectinariids and terebellids were studied. On this basis a re‐investigation appeared necessary: statements in the literature about the structure of the brain and the innervation of the appendages of the anterior end were incomplete and contradictory. In the present paper, the brain, the circum‐oesophageal connectives and the innervation of, inter alia, the tentacular membrane (including the dorsal ridge), the buccal tentacles, the alimentary canal, the nuchal organs and the branchiae (when present) of Amphicteis gunneri, Anobothrus gracilis, Melinna cf. cristata, Pectinaria auricoma, P. belgica, P. koreni, Petta pusilla, Pista cristata, Eupolymnia nebulosa, Thelepus cincinnatus and Polycirrus medusa are described. The results are summarized in schematic diagrams and compared with each other and with the central nervous system of other polychaetes. It is concluded that the ampharetids, the pectinariids and the terebellids bear no antennae and no palps and that their buccal tentacles belong to the alimentary canal. It is emphasized that all attempts to range their cephalic nervous system into previously proposed common and general schemes of ‘the polychaete nervous system’ seem totally fruitless.     

On the phylogeny of the Phyllodocidae (Polychaeta Annelida): an alternative

The phylogeny of the Phyllodocidae (Polychaeta Annelida) is examined in an analysis which includes previously unnoticed characters of the nervous system. Concentrating on four problems of importance in understanding the phylogeny of the family, a study of the nervous system demonstrated that: 1. A median antenna, such as that present in species of Eulalia, is not homologous to the nuchal papilla of, for example, Eteone and Phyllodoce; 2. Nuchal organs, presumably found in all phyllodocids, are basically homologous, but four separate types may be recognized; 3. The anterior concentration of the first segments (particularly their tentacular cirri), which is evident in most phyllodocids, is accompanied by different alterations of the ventral nervous system. The polarities of these changes are not self-evident; 4. A study of the ventral nervous system of Eteone reveals that the first apparent segment, bearing two pairs of tentacular cirri, must be regarded as homologous to the second segment of all other phyllodocids. Furthermore, this genus descended from forms in which a segment was present anterior to the first apparent segment, the first two segments possibly bearing tightly packed tentacular cirri. In connection with other characters, this information was used to produce a cladogram, which has some surprising features. The most unexpected result is the prediction that Eteone is the sister group of a taxon containing the genera Paranaitis, Chaetoparia, and Phyllodoce. The results are discussed and compared to those of other investigators.     

Ultrastructure of the extensively developed nuchal organs of Laonice bahusiensis (Annelida: Canalipalpata: Spionidae)

The nuchal organs of annelid Laonice bahusiensis (Spionidae) from northern Europe have been studied using scanning and transmission electron microscopy. L. bahusiensis is the first spionid species in which extensively developed, continuous nuchal organs are described. The nuchal organs of this genus are the longest known among polychaete annelids. They consist of paired double bands extending from the prostomium on a mid‐dorsal caruncle for about 24–30 setigers. Their microanatomy corresponds to the general structural plan of nuchal organs: there are ciliated supporting cells and bipolar sensory cells with sensory cilia traversing an olfactory chamber. The organs are overlaid by a secondary paving‐stone‐like cover and innervated by one pair of longitudinally elongated nuchal nerves. These findings clearly favor the hypothesis that the paired, extensively developed ciliated structures found in some Spionidae are homologous with the prostomial nuchal organs characteristic of polychaete annelids. J. Morphol. 2010. © 2009 Wiley‐Liss, Inc.     

Ultrastructural investigations on the cephalic and metameric nuchal organs of Spio cf. filicornis (Polychaeta, Spionidae)

The nuchal organs of Spio cf. filicornis from northern Europe have been studied by scanning and transmission electron microscopy. Spio cf. filicornis is the first species in which metameric nuchal organs are described. The nuchal organs consist of a distinct cephalic nuchal complex followed by metameric structures for a variable number of chaetigers. Their microanatomy corresponds to the general structural plan of nuchal organs: these are ciliated supporting cells and bipolar sensory cells with sensory cilia traversing an olfactory chamber. The organs are overlaid by a secondary paving-stone-like cover and innervated by longitudinally elongated paired nuchal nerves. The findings clearly favour the hypothesis that the paired metameric ciliated structures found in some Spionidae are in fact homologous with the prostomial nuchal organs characteristic of Polychaeta.     

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.     

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.     

An early Cambrian chelicerate from the Emu Bay Shale,South Australia

The Emu Bay Shale Lagerstätte (Cambrian Series 2, Stage 4) occurs on the north coast of Kangaroo Island, South Australia. Over 50 species are known from here, including trilobites and non‐biomineralized arthropods, palaeoscolecids, a lobopodian, a polychaete, vetulicolians, nectocaridids, hyoliths, brachiopods, sponges and chancelloriids. A new chelicerate, Wisangocaris barbarahardyae gen. et sp. nov., is described herein, based on a collection of some 270 specimens. It is up to 60 mm long, with the length of the cephalic shield comprising about 30% that of the exoskeleton. The cephalic margin has three pairs of bilaterally‐symmetrical small triangular spines. A pair of small eyes is placed well forwards on the ventral margin of the cephalic shield. The trunk comprises 11 segments that increase in length while narrowing posteriorly, each possibly bearing a pair of biramous appendages; the most posterior segment is almost square whereas the others are transversely elongated. The spatulate telson is proportionately longer than in taxa such as Sanctacaris, Utahcaris and Leanchoilia. Up to eight (?four pairs) of 3 mm‐long elements bearing alternating inward‐curving short and long spines beneath the cephalic shield are interpreted as basipodal gnathobases, part of a complex feeding apparatus. A well‐developed gut includes a stomach within the cephalic shield; it extends to the base of the telson. In a few specimens there are shell fragments within the gut, including those of the trilobite Estaingia bilobata (the most common species in the biota); these fragments have sharp margins and extend across the gut lumen. The species may have been a predator or a scavenger, ingesting material already broken up by a larger predator/scavenger. The morphology of this taxon shares many overall body features with Sanctacaris, and some with Sidneyia, particularly its gnathobasic complex. These chelicerate affinities are corroborated by phylogenetic analyses.     

Central nervous system and sense organs, with special reference to photoreceptor-like sensory elements, in Polygordius appendiculatus (Annelida), an interstitial polychaete with uncertain phylogenetic affinities

Abstract. The phylogenetic position of Polygordius is still pending; relationships with either Opheliidae or with Saccocirrus are the most favored hypotheses. The present study of Polygordius appendiculatus was designed to look for morphological characters supporting either of these two hypotheses. The homology of the anterior appendages, and the structure of the central nervous system and nuchal organ all required clarification; we also examined whether photoreceptor‐like sense organs exist in adults. From their innervation pattern, it is likely that the anterior appendages represent palps. They lack structures typical of palps in Canalipalpata, such as musculature and coelomic cavities, which would be expected in the case of a saccocirrid relationship. Thirteen photoreceptor‐like sense organs were found in front of the brain, the only structures resembling photoreceptors in adults of P. appendiculatus. These multicellular sense organs comprise a supportive cell and several sensory cells enclosing an extracellular cavity. There are three different types of sensory cells: one rhabdomeric and two ciliary. These sensory cells are combined differently into three forms of sense organ: the most frequent uses all three types of sensory cells, the second possesses one rhabdomeric and one ciliary cell type, and the third has two types of ciliary sensory cells. Whereas similar sensory cells are frequently found in various polychaetes, their combination in one sensory organ is unique to Polygordius and is considered to represent an autapomorphy. The nuchal organs exhibit features typical of polychaetes; there are no specific features in common with Saccocirrus. Instead, the covering structures show obvious similarities to Opheliidae, as can also be found in the central nervous system. Altogether, the current observations do not contradict a relationship with opheliids but provide no evidence of a relationship with Saccocirrus as has been found in certain molecular analyses, and thus currently leave the phylogenetic position of Polygordius unresolved.     

A conserved genetic mechanism specifies deutocerebral appendage identity in insects and arachnids

The segmental architecture of the arthropod head is one of the most controversial topics in the evolutionary developmental biology of arthropods. The deutocerebral (second) segment of the head is putatively homologous across Arthropoda, as inferred from the segmental distribution of the tripartite brain and the absence of Hox gene expression of this anterior-most, appendage-bearing segment. While this homology statement implies a putative common mechanism for differentiation of deutocerebral appendages across arthropods, experimental data for deutocerebral appendage fate specification are limited to winged insects. Mandibulates (hexapods, crustaceans and myriapods) bear a characteristic pair of antennae on the deutocerebral segment, whereas chelicerates (e.g. spiders, scorpions, harvestmen) bear the eponymous chelicerae. In such hexapods as the fruit fly, Drosophila melanogaster, and the cricket, Gryllus bimaculatus, cephalic appendages are differentiated from the thoracic appendages (legs) by the activity of the appendage patterning gene homothorax (hth). Here we show that embryonic RNA interference against hth in the harvestman Phalangium opilio results in homeonotic chelicera-to-leg transformations, and also in some cases pedipalp-to-leg transformations. In more strongly affected embryos, adjacent appendages undergo fusion and/or truncation, and legs display proximal defects, suggesting conservation of additional functions of hth in patterning the antero-posterior and proximo-distal appendage axes. Expression signal of anterior Hox genes labial, proboscipedia and Deformed is diminished, but not absent, in hth RNAi embryos, consistent with results previously obtained with the insect G. bimaculatus. Our results substantiate a deep homology across arthropods of the mechanism whereby cephalic appendages are differentiated from locomotory appendages.     

Ultrastructure of the nuchal organs in the polychaete Platynereis dumerilii (Annelida,Nereididae)

Abstract. We examined the nuchal organs of adults of the nereidid polychaete Platynereis dumerilii by means of scanning and transmission electron microscopy. The most prominent features of the nuchal organs are paired ciliary bands located dorsolaterally at the posterior margin of the prostomium. They are composed of primary sensory cells and multiciliated supporting cells, both covered by a thin cuticle. The supporting cells have motile cilia that penetrate the cuticle and are responsible for the movement of water. Subapically, they have a narrowed neck region; the spaces between the neck regions of these supporting cells comprise the olfactory chamber. The dendrites of the sensory cells give rise to a single modified cilium that crosses the olfactory chamber; numerous thin microvillus-like processes, presumably extending from the sensory cells, also traverse the olfactory chamber. At the periphery of the ciliated epithelium runs a large nervous process between the ciliated supporting cells. It consists of smaller bundles of sensory dendrites that unite to form the nuchal nerve, which leaves the ciliated epithelium basally and runs toward the posterior part of the brain, where the perikarya of the sensory cells are located in clusters. The ciliated epithelium of the nuchal organs is surrounded by non-ciliated, peripheral epidermal cells. Those immediately adjacent to the ciliated supporting cells have a granular cuticle; those further away have a smooth cuticle. The nuchal organs of epitokous individuals of P. dumerilii are similar to those described previously in other species of polychaetes and are a useful model for understanding the development of nuchal organs in polychaetes.     

Expression of Distal-less, dachshund, and optomotor blind in Neanthes arenaceodentata (Annelida, Nereididae) does not support homology of appendage-forming mechanisms across the Bilateria

The similarity in the genetic regulation of arthropod and vertebrate appendage formation has been interpreted as the product of a plesiomorphic gene network that was primitively involved in bilaterian appendage development and co-opted to build appendages (in modern phyla) that are not historically related as structures. Data from lophotrochozoans are needed to clarify the pervasiveness of plesiomorphic appendage-forming mechanisms. We assayed the expression of three arthropod and vertebrate limb gene orthologs, Distal-less (Dll), dachshund (dac), and optomotor blind (omb), in direct-developing juveniles of the polychaete Neanthes arenaceodentata. Parapodial Dll expression marks pre-morphogenetic notopodia and neuropodia, becoming restricted to the bases of notopodial cirri and to ventral portions of neuropodia. In outgrowing cephalic appendages, Dll activity is primarily restricted to proximal domains. Dll expression is also prominent in the brain. dac expression occurs in the brain, nerve cord ganglia, a pair of pharyngeal ganglia, presumed interneurons linking a pair of segmental nerves, and in newly differentiating mesoderm. Domains of omb expression include the brain, nerve cord ganglia, one pair of anterior cirri, presumed precursors of dorsal musculature, and the same pharyngeal ganglia and presumed interneurons that express dac. Contrary to their roles in outgrowing arthropod and vertebrate appendages, Dll, dac, and omb lack comparable expression in Neanthes appendages, implying independent evolution of annelid appendage development. We infer that parapodia and arthropodia are not structurally or mechanistically homologous (but their primordia might be), that Dll's ancestral bilaterian function was in sensory and central nervous system differentiation, and that locomotory appendages possibly evolved from sensory outgrowths.     

Morphology and ultrastructure of the anterior end of Diplocirrus longisetosus Marenzeller, 1890 (Flabelligeridae, Polychaeta, Annelida)

The morphology and ultrastructure of the sedentary polychaete Diplocirrus longisetosus Marenzeller, 1890, collected from the White Sea, were studied using dissection, histological methods, light microscopy, and both scanning and transmission electron microscopy. The prostomium and peristomium carry a pair of palps, eight branchiae, a pair of nuchal organs and two nephridiopores, ciliated folds and the mouth. The prostomium, peristomium and the first chaetigerous segment with all appendages comprise the so-called siphon complex. The mouth leads to a pharyngeal organ that is closed ventrally and composed of a ventral muscle bulb adjoined dorsally by two folds projecting into the pharyngeal lumen. These parts are connected and enveloped by the longitudinal investing muscle. No tongue-like organ is present. The nervous system of the siphonal part comprises the brain, the circum-oesophageal connectives and the ganglia of the peristomium and first chaetigerous segment.     

A new genus of Platycopioida (Copepoda) from a marine cave on Bermuda

Nanocopia minuta was collected from an inland marine cave on Bermuda. This new genus is reminiscent of Platycopia in its cephalic appendages and in the 5th legs of the male. The 1st leg has a 3-segmented exopod and a 1-segmented endopod. The other legs show reductions with no armature along the inner margin of the exopods and with 2-segmented endopods in the female. Only 2nd and 3rd legs bear two outer spines on the first exopodal segment.Owing to similarities in sexual characters, mouthparts, and modifications of the swimming legs, Nanocopia and Platycopia are considered more closely related to each other than either is to Antrisocopia.Platycopioida now contains three genera of which two are found only in Roadside Cave. The order has retained several primitive characters and seems to have separated early from the gymnoplean stem.