A new genus of Nanaloricidae (Loricifera) from deep-sea sediments of volcanic origin in the Kilinailau Trench north of Papua New Guinea

A new genus and species of Nanaloricidae (Loricifera), Phoeniciloricus simplidigitatus, is described inhabiting fine sand covered by a layer of volcanic ash at a water depth of 1,813 m in the New Ireland Basin near the Kilinailau Trench (north of Papua New Guinea). The described specimen is a postlarva enclosed in a larval exuvium. This is the first report of a species belonging to the Nanaloricidae from the deep sea. This occurrence is surprising, because Nanaloricidae are typical inhabitants of coarse sands in the intertidal or littoral zone. Preference for these shallow water habitats is reflected in many morphological features which characterize the Nanaloricidae, and are not normally found in Loricifera inhabiting fine-grained, clayish, deep-sea bottoms. The postlarva of the new species is characterized by a long narrow mouth tube, an urn-shaped lorica divided into ten plates, and 13 small lorica spikes. Distinguishing features of the Higgins-larva include short spinose toes lacking mucros but having small and slightly enlarged bases, short scalids on the introvert, many thoracic plates arranged in 6–8 rows, numerous small papillate flosculi in the collar and caudal regions, and three pairs of filiform, short locomotory appendages on the ventral side. Some features of the new species, especially of the Higgins-larva, are discussed as adaptations to the deep-sea environment.Communicated by R.M. Kristensen     

Rugiloricus bacatus sp. nov. (Loricifera ‐Pliciloricidae) and a ghost‐larva with paedogenetic reproduction

Abstract

A new species of Loricifera, Rugiloricus bacatus sp. nov. is described together with the diagnoses of two other Rugiloricus species, Rugiloricus sp. nov. A and B, from the Faroe Bank (North Atlantic). Characteristic for all three species is the presence of a new type of reduced larva, a ghost‐larva. This type of reduced larva was discovered in 1986 by Jeanne Renaud‐Mornant, but it was with the Faroe Bank material that it was first discovered that the ghost‐larvae belonged to the phylum Loricifera. The ghost‐larvae are eitherfound inside late instar Higgins‐larvae, called penultimate Higgins‐larvae, or in the sediment. The three types of Higgins‐larvae from the Faroe Bank can be distinguished by characters such as anterior setae, posterior setae and toes. The adults of Rugiloricus bacatus sp. nov. are characterised by a prominent ruff resembling a pearl necklace; two of the eight clavoscalids are modified in the 1st row; the 2nd row of leg‐shaped scalids are very large and robust, and the 9th row with 30 beak‐like scalids alternating with 30 alternating plates. The postlarvae are free‐living and their scalids on the introvert are reduced to protoscalids. Postlarvae and adult stages have not been found for Rugiloricus sp. nov. A and B and therefore only diagnoses of these two species are presented here.     

Loricifera from the deep sea at the Galápagos Spreading Center,with a description of <Emphasis Type="Italic">Spinoloricus turbatio</Emphasis> gen. et sp. nov. (Nanaloricidae)

Specimens of a new species of Loricifera, Spinoloricus turbatio gen. et sp. nov., have been collected at the Galápagos Spreading Center (GSC) during the cruise SO 158, which is a part of the MEGAPRINT project. The new genus is positioned in the family Nanaloricidae together with the three already described genera Nanaloricus, Armorloricus and Phoeniciloricus. The postlarvae and adults of Spinoloricus turbatio gen. et sp. nov. are characterized by a mouth cone with eight oral ridges and basally with a cuticular reinforcement named mouth cone pleat; eighth row with 30 whip-like spinoscalids and 30 “alternating” plates; thorax with eight single and seven double trichoscalids, where the single trichoscalids are twice the length of the double ones, and the secondary appendage on the double trichoscalid is smooth whereas the others are serrated; lorica with eight cuticular plates with additional spikes in the anterior corners and intercalary plicae between the plates. Some of these genus-specific characters such as the mouth cone pleat, the “alternating” plates and the intercalary plicae have not been observed in Nanaloricidae before. The Higgins-larvae of Spinoloricus turbatio gen. et sp. nov. are characterized by six rectangular plates in the seventh row with two teeth, an indistinct honeycomb sculpture and long toes with little mucrones. The SO 158 cruise has yielded a minimum of ten new species of Loricifera out of only 42 specimens. These new species belong to two different orders, where one being new to science, and three different families. This result indicates a high diversity of loriciferans at the GSC. Nearly all the collected loriciferans are in a moulting stage, hence there is a new stage inside the present stage. This prolongation of life stages and the occurrence of multiple life stages inside each other are typical of deep-sea loriciferans. Here exemplified by the two postlarvae with adults inside, which is observed for the first time in Nanaloricidae.     

Symmetry of priapulids (Priapulida). 2. Symmetry of larvae

Larvae of priapulids are characterized by radial symmetry evident from both external and internal characters of the introvert and lorica. The bilaterality appears as a result of a combination of several radial symmetries: pentaradial symmetry of the teeth, octaradial symmetry of the primary scalids, 25-radial symmetry of scalids, biradial symmetry of the neck, and biradial and decaradial symmetry of the trunk. Internal radiality is exhibited by musculature and the circumpharyngeal nerve ring. Internal bilaterality is evident from the position of the ventral nerve cord and excretory elements. Externally, the bilaterality is determined by the position of the anal tubulus and two shortened midventral rows of scalids bordering the ventral nerve cord. The lorical elements define the biradial symmetry that is missing in adult priapulids. The radial symmetry of larvae is a secondary appearance considered an evolutionary adaptation to a lifestyle within the three-dimensional environment of the benthic sediment.     

Microscopic anatomy and ultrastructure of the introvert of Priapulus caudatus and P. tuberculatospinosus (Priapulida)

Introverts of Priapulus caudatus and P. tuberculatospinosus bear 25 rows of scalids, as well as 8 spines and scattered papillae in the region the circumoral lip. These, as well as the first ring of pharyngeal teeth in P. tuberculatospinosus, are sensory organs. Although superficially they differ between species, they are all characterized by apical and/or subapical openings which are located on tiny cuticular tubules. All sensory organs contain cilia bearing bipolar receptor cells. The 8 sensory spines situated between the circumoral area and the beginning of the scalids are ultrastructurally similar to the scalids. The introvert and pharyngeal body walls, and associated muscles are described. © 1994 Wiley-Liss, Inc.     

Successive reduction of the last instar larva of Loricifera, as evidenced by two new species of Pliciloricus from the Great Meteor Seamount (Atlantic Ocean)

Two new species of the genus Pliciloricus Higgins & Kristensen, 1986, are described from coarse sand at the plateau of the Great Meteor Seamount. Together with other species of the genus, Pliciloricus corvus sp. n. and Pliciloricus senicirrus sp. n., demonstrate that the last or seventh instar Higgins-larva can be more and more reduced.

Additionally, in the life cycle of P. corvus sp. n. a simplified adult is discovered which has a simplified body with persisting neck region and a sack-like trunk. The adult of P. corvus sp. n. is characterized, e.g., by 15 broad cross-walls in the clavoscalids, midventral spinoscalids of the second row transformed into a new type of double-organ, a secondary double-organ in the third row of scalids, type B spinoscalids of the fourth row ending in a grappling hook, a crescent-shaped ornamentation of the lorica margin with rounded gaps and protrusions, and a pair of longitudinal cuticular protrusions on the end cone. Distinguishing features of the first to sixth instar Higgins-larvae of P. corvus sp. n. are, e.g., setae of moderate length and long end spines of toes with double bulges. The seventh instar Higgins-larva of P. corvus sp. n. is not fully developed, e.g., the abdominal region is indistinct; the scalids, toes, and all five pairs of setae are clearly shorter than in the preceding instars of Higgins-larvae.

The adult of P. senicirrus sp. n. is characterized, e.g., by clavoscalids with many fine transversal striae, claw-like type B spinoscalids of the fourth row ventrally winged and with seven embedded needle-like teeth, edge of the lorica with 24 small protrusions, and 20 papillate flosculi at caudal end. The Higgins-larva of P. senicirrus sp. n. is characterized, e.g., by the wide and frill-like first section of the mouth cone, the second segment of spinoscalids with ventral serration and double tip, long anterosetae with strong spinules, and by the toes being nearly as long as the short lorica, to name a few. The most obvious feature is the extreme length of the three pairs of posterosetae. The posteroterminal setae are nearly as long as the whole body. The seventh instar Higgins-larva of P. senicirrus sp. n. is not reduced as in the case of P. corvus sp. n.     

Ultrastructure of the introvert and associated structures of the larvae of Halicryptus spinulosus (Priapulida)

Summary Based on new SEM and TEM observations, the arrangement and ultrastructure of the scalids and adhesive tubuli of the larvae of Halicryptus spinulosus are described. The scalids are arranged in 25 longitudinal rows. Transversally they form seven circlets which alternate. The first circlet consists of 8 and the second of 9 scalids. All following circlets consist of three subrings with 8+8+9 scalids each. New scalids develop in a sub frontal growing zone posterior to the second circlet. The ultrastructure of all different receptor cells and sense organs of the scalids is described. Whereas the receptor cell type 1 occurs in nearly all scalids, the verticalrootlet receptor is restricted to the first circlet, the basal receptor to the second and the coiled-cilium receptor to the third and fourth circlet. Based on these new data, the scalid arrangement and the equipment with different receptor cells can be postulated for the ground pattern of the priapulid larvae. Also the larval development, moulting and the structure of the cuticle are discussed in comparison with the Kinorhyncha and Loricifera.     

Introvert,mouth cone,and nervous system of Echinoderes capitatus (Kinorhyncha,Cyclorhagida) and implications for the phylogenetic relationships of Kinorhyncha

Summary The introvert, mouth cone, and nervous system of Echinoderes capitatus were examined by transmission and scanning electron microscopy. The introvert bears seven rings of primarily quincunxial sensory scalids, including type 1 and 2 spinoscalids as well as trichoscalids; the latter two types are additionally provided with glandular cells. The mouth cone bears one ring of decamerous sensory oral styles and three rings of quincunxial sensory pharyngeal styles. The intra- to basiepithelial, bilateral nervous system consists of a circumentric nerve ring in the introvert, a terminal and proximal nerve ring in the mouth cone, a ventral chain of ganglia, one in each trunk zonite, and a caudal ganglion. The introvert, the neck, and the trunk zonites are innervated from the forebrain; the mouth cone and the pharyngeal bulb are innervated from the hindbrain. The monophyly of the Kinorhyncha is based upon the following autapomorphic characters: (1) a mouth cone, (2) a neck with 16 placids, (3) a trunk with 11 zonites, (4) scalids of three types: type 1 and type 2 spinoscalids, and trichoscalids, (5) an anteriormost ring of ten type 1 spinoscalids (sensory organs divided into a basal and a terminal part), (6) a posteriormost ring of 14 trichoscalids (glandular sensory organs which are undivided), (7) rings in between the anteriormost and posteriormost are type 2 spinoscalids (glandular sensory organs divided into a basal and a terminal part), (8) a mouth cone with a terminal and a proximal nerve ring, (9) nine sensory oral styles with decamerous symmetry (the dorsal style is missing) and (10) three rings of sensory pharyngeal styles with, from anterior to posterior, ten, five, and five styles with quincunxial arrangement. The following characters are assumed to be autapomorphic for the taxon Nematoda+Gastrotricha+Kinorhyncha+Loricifera+Priapulida: (1) a basiepithelial circumentric brain and (2) a neuropileous nerve ring in a subterminal position. The following characters are assumed to be autapomorphic for the taxon Kinorhyncha+Loricifera+Priapulida: (1) a neuropileous nerve ring in a terminal position, (2) an introvert with scalids, (3) an eversible foregut and (4) tanycytes.The unpublished doctoral thesis of B. Neuhaus [1991 Zur Ultrastruktur, Postembryonalentwicklung und phylogenetischen Verwandtschaft der Kinorhyncha. PhD thesis. University of Götingen, Germany] was finished simultaneously with the completion of this study.     

Morphology and life cycle of a new loriciferan from the Atlantic coast of Florida with an emended diagnosis and life cycle of Nanaloricidae (Loricifera)

Abstract. A new interstitial loriciferan, Nanaloricus gwenae sp. nov., is described from coarse-sand and shell-hash habitats ( Amphioxus sand), at 15–17 m depth, 6–7 miles off the coast of Fort Pierce, FL, USA. The new species is very closely related to the type species Nanaloricus mysticus found off the coast of Roscoff, France in nearly the same kind of sediment ( Dentalium sand). All life stages (Higgins larva, postlarva, and adult) in the life cycle of the new species were found through the 10-year investigation, leading to the conclusion that all species of Nanaloricidae have only a sexual reproductive cycle. The adult of the new species can be distinguished by the different body shape, the lorical spikes and sculpture, and the shape of the scalids. The postlarva has stronger longitudinal ridges on the lorical plates and the Higgins larva has toes with smaller mucrones than those of N. mysticus . An emended definition of the Nanaloricidae is provided based on the new formula of the number of rows and shape of the scalids on the introvert. The associated meiofauna found in the subtidal sand or shell hash is both abundant and diverse. Tardigrades, gastrotrichs, and kinorhynchs are among the most common associates. N . gwenae sp. nov. is represented by only eight specimens collected during a 10-year period of sampling this habitat.     

Two new species of the genus Pliciloricus (Loricifera, Pliciloricidae) from the Faroe Bank, North Atlantic

Five cruises to the Faroe Bank in the northeast Atlantic revealed a rich loriciferan fauna inhabiting shell gravel to fine carbonate sand. This paper describes two new species of Pliciloricus found at depths of 120-260 m. Currently, only six species of this genus have been described from various localities around the world, but several new species are currently being described. The diagnosis of the genus Pliciloricus is emended to include the two new taxa. The first of the two new species Pliciloricus leocaudatus sp. n. is characterized by having four new kinds of scalids in the second row and an additional row of alternating plates in the eighth row. The second species Pliciloricus shukeri sp. n. is characterized in having a secondary double organ in the third row. The morphology of the two species, including the new structures, has been investigated using LM and scanning electron microscopy. Conclusively, the scalid pattern of the introvert of Pliciloricus has been re-evaluated, due to the finding of the new structures. Additionally, a discussion of the life cycle of the genus Pliciloricus is given, since the postlarval stage is reduced in both new species.     

Structure of the nervous system in Tubiluchus troglodytes (Priapulida)

Abstract. The nervous system of the meiobenthic priapulid species Tubiluchus troglodytes is described by immunohistochemistry and confocal laser scanning microscopy. The brain is circumpharyngeal, consisting of a central ring of neuropil and both anterior and posterior somata. From the brain emerges a ventral nerve cord, which shows ganglion-like swellings in the neck and caudal region. The introvert includes longitudinal neurite bundles running below and between the rows of scalids, with a small cluster of sensory cells under each scalid. In the body wall of the neck and trunk region, longitudinal and circular neurite bundles are present in an orthogonal pattern. The tail is innervated from the caudal swelling of the ventral nerve cord; it also includes longitudinal and circular bundles in an orthogonal pattern. The pharynx has a reticulated system of neurite bundles running between the pharyngeal teeth and fimbrillae. Below each tooth and fimbrilus is a ganglion-like cluster of somata. The intestine is surrounded by a nerve net. The data on the nervous system are compared within other priapulids and with other species of Scalidophora (Kinorhyncha and Loricifera).     

Scanning and Transmission Electron Microscopical Observations on the Larvae of Priapulus caudatus (Priapulida)

Abstract Scanning electron microscopical studies revealed four distinct morphological larva-types (not instars) of Priapulus caudatus whose lorica-length measured 82–860 μm. The smallest of the larvae are round to oval, have 20 longitudinal ridges, a series of transverse ridges, and have two pairs of laterally situated tubuli near the posterior limits. The second larva-type is dorsoventrally compressed, has a single dorsal and single unsculptured ventral plate each with a prominent midridge near the anterior limits, three infolded lateral plates with a pair of tubuli near the posterior limits. The third and fourth larva-types remain dorsoventrally flattened; the third larva-type has less pronounced sculpturing than the fourth larva-type and is smaller; two pairs of tubuli are situated as in the previous stage. The number and arrangement of scalids on the introvert and pharyngeal teeth differ according to each of the four larva-types and are described. The ultrastructure of all organ systems is described. All scalids are sensilla equipped with ciliated receptor cells. The cuticle is similar to the cuticle in the adult and differs fundamentally from the cuticle of larval Halicryptus. The trunk is richly supplied with sensory structures. Trunk tubuli contain secretory cells. Posteriorly, a gland complex was found which presumably is the precursor of the equivalent gland in postlarvae and adults. The implications of the first larva-type's structural similarity to the larva of Tubiluchus are discussed.     

Ultrastructure of introvert and pharynx of Halicryptus spinulosus (priapulida)

The introvert of Halicryptus spinulosus bears three kinds of sensilla: buccal papillae, ordinary scalids, and dentoscalids. They are all characterized by bipolar monociliary receptor cells. The former two have apical openings at which the sensory cilia are in close contact with the ambient sea water. The pharyngeal teeth are composed of slender epithelial cells the tips of which are devoid of organelles and a thick cuticle. The anatomy of the muscle arrangement of the pharynx is described. Glands occur at the junction of the pharynx and midgut.     

Recent Priapulidae and their Early Cambrian ancestors: comparisons and evolutionary significance

Although priapulid worms form a relatively small phylum in present-day marine environments, they were important animals in Cambrian endobenthic communities. Two Early Cambrian priapulids, namely Xiaoheiqingella peculiaris and Yunnanpriapulus halteroformis nov. gen., nov. sp. from the Maotianshan Shale Lagerstätte of SW China are revised and described. Several key-features of the body plan of Recent Priapulidae are recognized in these two forms: 1) the four-fold body division (introvert, neck, trunk, and caudal appendage); 2) the well-developed introvert armed with ca. 25 longitudinal rows of scalids; 3) the caudal appendage; 4) the pharyngeal teeth arranged in a pentagonal disposition (Xiaoheiqingella); 5) the ventral nerve cord present in Yunnanpriapulus. This morphology indicates close evolutionary relationships with modern priapulids. Xiaoheiqingella and Yunnanpriapulus nov. gen. are tentatively placed within the recent family Priapulidae. The Priapulidae lineage may therefore have a remote origin (Early Cambrian) much older than was previously assumed (Priapulites; Late Carboniferous). The functional morphology of Xiaoheiqingella and Yunnanpriapulus nov. gen. suggests that these two worms were chiefly carnivorous with possible occasional mud-eating habits.     

Morphology and ultrastructure of the sensory appendages of a kinorhynch introvert

The structure and arrangement of appendages (scalids) on the head of the homalorhagid kinorhynch Kinorhynchus phyllotropis Brown & Higgins, 1983 are named, described and illustrated. In adults of this species, seven rings of external scalids are separated by segment boundary structures from the oral styles and three rings of internal scalids. All of these appendages contain ciliated receptor cells which pass to pores at the scalid tips, and all but the two anterior rings are jointed. All of these appendages are radially arranged in multiples of five, and closely associated with the ten lobed circumoral nerve ring. The most posterior scalid ring consists of fourteen trichoscalids, of which six are longer than the other eight. The six longer trichoscalids are arranged in bilateral symmetry, two to each of the three facets of the body. Similarities between scalid arrangement in Kinorhyncha and Loricifera are discussed.     

A new type of loriciferan larva (Shira larva) from the deep sea of Shatsky Rise,Pacific Ocean

Loricifera is a phylum of minute animals that live exclusively in marine sediments. A total of 33 species have been described so far in this phylum; however, several more are already known from preliminary observations. Loriciferans are characterised by a complex life cycle, which involves a succession of several adult and larval stages. Here, we describe a new type of loriciferan larval stage: the Shira larva. The gross morphology of this larva is generally similar to that of the most prominent larval type of Loricifera, the so-called Higgins larva. However, the Shira larva possesses a number of unique features, namely (1) a single pair of anteroventral setae is present in the most anterior region of the abdomen, (2) the bases of the anteroventral setae are very large and swollen, (3) the thorax and abdomen are thinner than the introvert and (4) the abdominal region is divided into five sub-regions. Accordingly, we described the new species, Tenuiloricus shirayamai gen. nov. et sp. nov. (incertae sedis). The new findings are discussed from a comparative perspective with the Higgins larva as well as with the fossil of a putative loriciferan larval stage from the Middle Cambrian.     

Revision of the genusgirella (girellidae) from east asia

The taxonomy of three species ofGirella from East Asian waters,G. punctata Gray, 1835,G. leonina (Richardson, 1846) andG. mezina Jordan & Starks, 1907, is reviewed and intraspecific (individual and ontogenic) variations detailed.Girella mezina is characterized by a very wide mouth and thick upper lip, the soft-rayed portion of the anal fin high and round, dorsal profile of the head abruptly slanting in front of the eyes in adults, a transverse yellow band on the body in life, and 3–4 rows of teeth along the outer jaw margins, the central cusp of each tooth being wider than the lateral cusps in adults.Girella punctata is characterized by usually 2 rows of teeth along the outer jaw margins, usually 7 transverse series of scales between the lateral line and median spinous portion of the dorsal fin (TRac), and usually 52–55 pored lateral line scales (LLp). The species is variable in body and caudal fin shape, extent of squamation on the opercular region, and number and position of dark spots on the scales. It also exhibits ontogenetic variation in the number of tooth-rows.Girella leonina is characterized by a conspicuously black opercular flap, essentially a single row of teeth along the outer jaw margins, usually 10–11 TRac and usually 59–64 LLp. Intraspecific variations were evident in the mouth position, scale condition and body color after death, and in the number of pores of cephalic lateral line canals. The holotype ofG. punctata, previously known only from a figure, is described for the first time.Girella melanichthys is synonymized underG. punctata with a lectotype designated for the former.     

Ultrastructural observations of the larva ofTubiluchus corallicola (Priapulida)

Larvae ofTubiluchus corallicola van der Land 1968 were investigated by scanning and transmission electron microscopy. The scalids are sensory organs, each has a bipolar receptor cell with a single apical cilium similar to the scalid in the adult. Muscle cells of the larva are more differentiated than previously reported for other Priapulida; the larval arrangement of circular and longitudinal musculature differs from that of the adult, and a diaphragm is reported for the first time in Priapulida. The diaphragm may function in hydrostatic control of eversion and inversion of the introvert and mouth cone. The functional morphology of these two structures is discussed and contrasted with the Kinorhyncha.     

Life Cycle and Morphology of a Cambrian Stem-Lineage Loriciferan

Cycloneuralians form a rich and diverse element within Cambrian assemblages of exceptionally preserved fossils. Most resemble priapulid worms whereas other Cycloneuralia (Nematoda, Nematomorpha, Kinorhyncha, Loricifera), well known at the present day, have little or no fossil record. First reports of Sirilorica Peel, 2010 from the lower Cambrian Sirius Passet fauna of North Greenland described a tubular lorica covering the abdomen and part of a well developed introvert with a circlet of 6 grasping denticles near the lorica. The introvert is now known to terminate in a narrow mouth tube, while a conical anal field is also developed. Broad muscular bands between the plates in the lorica indicate that it was capable of movement by rhythmic expansion and contraction of the lorica. Sirilorica is regarded as a macrobenthic member of the stem-lineage of the miniaturised, interstitial, present day Loricifera. Like loriciferans, Sirilorica is now known to have grown by moulting. Evidence of the life cycle of Sirilorica is described, including a large post-larval stage and probably an initial larva similar to that of the middle Cambrian fossil Orstenoloricusshergoldii .     

Morphology,Morphogenesis, and Molecular Phylogeny of Paraparentocirrus sibillinensis n. gen., n. sp., a “Stylonychine Oxytrichidae” (Ciliophora,Hypotrichida) Without Transverse Cirri

A terrestrial oxytrichid ciliate Paraparentocirrus sibillinensis n. gen., n. sp., which was found in soil samples of a beech forest stand within the National Park of Sibillini Mountains, Italy, was investigated using live observation and protargol impregnation. The morphology of interphase, morphogenesis, and molecular phylogeny inferred from SSU rDNA sequences of this ciliate were studied. Paraparentocirrus n. gen., is mainly characterized by a semirigid body, an undulating membrane in the Oxytricha pattern, six fronto‐ventral (FV) rows, the absence of transverse cirri, one right and one left row of marginal cirri, four dorsal kineties, two dorsomarginal rows, and caudal cirri at the end of dorsal kinety 4. During morphogenesis, oral primordia develop through the proliferation of basal bodies from some cirri of FV rows 4 and 5, and FV row 6 takes part in the anlagen formation of the proter. The dorsal morphogenesis was typical of oxytrichids, with simple fragmentation of dorsal kinety 3, and the dorsomarginal rows developed from the right marginal row. Phylogenetic analyses based on the SSU rDNA sequences support the classification of this new genus in the stylonychines.