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The testis morphology, spermatogenesis, and the spermatozoon structure in two palaeonemerteans, Callinera sp. and Parahubrechtia sp., have been examined by light, laser confocal, and electron microscopy. Early spermatogenesis is quite similar between the two species. The similarities include the formation of the initial proacrosomal vesicle at the primary spermatocyte stage, the presence of electron-dense non-membranous substance around the centrioles in spermatogonia and spermatocytes, and the occurrence of lipid droplets at those stages. However, some differences in the process of acrosome formation during spermiogenesis were also found. In Callinera sp., a spherical acrosomal vesicle (rounded in the sections) transforms into a convex disk structure (arc-shaped in the longitudinal sections) in the developing posterior part of the spermatid and then migrates anteriorly; in Parahubrechtia sp., a rounded acrosomal vesicle migrates towards the apex of the cell where it is transformed into the arc-shaped structure. Both species have spermatozoa of a primitive type, i.e., with a small acrosomal complex, a rounded nucleus, a short mid-piece, and a posteriorly oriented flagellum. Nevertheless, some modifications in the sperm organization such as the asymmetric location of the acrosomal complex and mid-piece were recorded. It is possible that such modifications are synapomorphic for palaeonemerteans. In Callinera sp., the mid-piece contains 2–4 mitochondria, while Parahubrechtia sp. has only one ring-shaped but unclosed mitochondrion. The presence of several mitochondria in Callinera sp. may be regarded as a result of the so-called ‘progenetic spermatogenesis’.

From a biodiversity survey of a subterranean habitat near Sinan County, Guizhou Province, southwestern China, a new atyid shrimp of the genus Caridina H. Milne Edwards, 1837, C. sinanensissp. nov. was discovered. The new species can be separated from other congeners based on a combination of characters including depigmentation in body and reduction of eyes, small pigment spot at the centre of the cornea, the shape of rostrum and the endopod of the 1st male pleopod, and the relatively longer appendix interna on the appendix masculina of the 2nd pleopod. Mitochondrial COI and 16S rRNA gene sequences also support the establishment of the new species. Information on the habitat, and the levels of threat are discussed to guide the conservation of C. sinanensissp. nov.

The taxonomic identity of the palaeonemertean Cephalothrix linearis (Rathke, 1799) has been obscure for nearly two centuries, because its original description applies to almost any congeners, including Cephalothrix filiformis (Johnston 1828) and Cephalothrix rufifrons (Johnston, 1837), which occur commonly in the North Sea and adjacent waters. In this paper, I redescribe C. linearis based on two topotypes from Bergen, one herein designated as the neotype for C. linearis, because Rathke’s original material is not extant; I invoke Article 70.3.2 of the International Code of Zoological Nomenclature to fix Planaria linearis Rathke, 1799 as the type species of Cephalothrix Örsted, 1843 for the sake of stability. From the neotype, I determined sequences of the 28S rRNA, 16S rRNA, and cytochrome c oxidase subunit I (COI) genes. Using the COI sequence, I inferred the phylogenetic position of C. linearis along with 316 cephalotrichid sequences currently available in public databases. A tree-based species delimitation analysis detected 43 entities among them, with 34 in Cephalothrix and nine in eitherBalionemertes or Cephalotrichella. I apply valid species names to 12 of the 34 entities in Cephalothrix. I tabulated a total of 36 nominal species that are likely the members of the genus; the following five were excluded even though their specific names were originally combined with Cephalothrix: Cephalothrix armata Ulyanin, 1870 [Monostilifera, possibly Emplectonema gracile (Johnston, 1837)], Cephalothrix fragilis Bürger, 1892 [now Cephalotrichella signata (Hubrecht, 1879)], Cephalothrix signata Hubrecht, 1879 [now in Cephalotrichella], Cephalothrix unipunctata Parfitt, 1867 [now Tetrastemma melanocephalum (Johnston, 1837) (Monostilifera)], and Cephalothrix viridis Chapuis, 1886 [possibly Heteronemertea]. The five names cephalothrix Diesing, 1850 (as Borlasia cephalothrix), kroyeri Diesing, 1850 (as Cephalothrix kroyeri), linearis Diesing, 1850 (as Borlasia linearis), lineata Claparède, 1862 (as Cephalothrix lineata), and oerstedii Diesing, 1850 (as Cephalothrix oerstedii) aredeclared nomenclaturally unavailable.

A new nemertean species of the genus Cephalothrix from intertidal calcareous red algae off the Vietnam coast is described based on histological sections, confocal laser scanning microscopy, and nucleotide sequences of three nuclear (18S rRNA, 28S rRNA, H3) and two mitochondrial (COI, 16S rRNA) DNA fragments. Cephalothrix suni sp. nov. is characterized by a unique body colour pattern consisting of transverse brown on ventral body side. A molecular phylogenetic analysis of the family Cephalotrichidae using a multigene approach has identified three clades for Cephalothrix: Cephalothrix, Procephalothrix, and interstitial cephalotrichids. The new species belongs to clade Procephalothrix sensu Chernyshev & Kajihara 2019 and a subclade of species with red, orange, or dark yellow anterior tip.

Kajihara, Hiroshi (2020): Three Species of Ribbon Worms (Nemertea) from Cebu, the Philippines. Species Diversity 25: 251-273, DOI: 10.12782/specdiv.25.251

Knowledge on the detailed body plan in Palaeonemertea (a group putatively considered to retain ancestral character states within Nemertea) is essential in understanding the morphological evolution among not only nemerteans but also the rest of bilaterians in general; however, such information has been scanty in previous literature. In this study, we examined the body wall musculature, the position of the nervous system, and the musculature of the proboscis apparatus in 14 species of Archinemertea (a palaeonemertean clade comprised of Cephalotrichidae and Cephalotrichellidae) using confocal laser scanning microscopy with phalloidin labeling as well as classical Mallory’s trichrome staining. Unexpectedly, all the archinemerteans possessed a diagonal muscle layer lying directly ‘above’—instead of ‘below’, as usually found in most vermiform animals—the body wall outer circular muscle layer, a character state that has never been reported among Bilateria except for some acoels and proseriates. We speculate that the aberrant position of the outer diagonal muscle layer might have been actualized as a result of a flip-flop in a genetic regulatory circuit that controls the body wall muscular morphogenesis, which must have happened independently among the bilaterian evolution at least once in Acoela (Xenacoelomorpha), Proseriata (Platyhelminthes), and Archinemertea (Nemertea), respectively. Based on available phylogenetic data and the results from the present observation, we reconstructed the evolution of morphological characters within Archinemertea and its subclades. The outer diagonal muscle layer is likely a synapomorphy for Archinemertea (Cephalotrichidae + Cephalotrichellidae). Members of Balionemertes and Cephalotrichella possess some unique states for palaeonemerteans in muscular morphology (muscular strand associated with lateral nerve cord; longitudinal muscles between brain and extraganglionic tissue; proboscis with outer longitudinal muscle strands). We propose that muscle fibers embedded in lateral nerve cords, termed myofibrillae, in hoplo- and heteronemerteans are homologous with the body wall longitudinal muscles near the lateral nerves surrounded by circular muscles.

6 pages, 3 figures, 2 tables, New data for Carinina ochracea Sundberg et al., 2009 are provided for the Iberian Peninsula, establishing the southernmost limit of its known distribution. This species was previously known from only two localities: the type locality in Tjärnö (Sweden) and Pouldohan (Brittany, France). The material examined here was obtained during a faunal survey in the Villaviciosa Estuary (Asturias, northern Iberian Peninsula). The identity of the new specimen was confirmed both by DNA barcoding and anatomical examination. The molecular divergence of all available sequences of this species for four molecular markers, cytochrome c oxidase subunit I (COI), 16S, 18S and 28S rDNA, is discussed. For COI, four polymorphic sites were found, indicating: 1) a nuclear pseudogene; 2) heteroplasmy; or 3) gene duplication of a region of the mitochondrial genome. Two previously overlooked morphological characters were found: the presence of a colour ring and a postfixation staining band (pigmented band), which is histologically characterized. This species is the 12th palaeonemertean and the 75th nemertean reported from Iberian waters, This study was supported by grant CGL2011-23306 from the Spanish government

The proboscis and rhynchocoel musculature of 56 nemertean species was studied using phalloidin labelling and confocal laser scanning microscopy. Six types of muscle layers are found in the anterior proboscis of the nemerteans: inner circular, inner diagonal, inner longitudinal, outer diagonal, outer circular, and outer longitudinal. Only the inner circular and inner longitudinal muscle layers are present in all the nemerteans studied. Ten types of arrangement of the proboscis musculature are described. Three primary types ('palaeotype', 'heterotype', and 'hoplotype') correspond to the three nemertean supergroups (Palaeonemertea, Heteronemertea, and Hoplonemertea). The evolutionary transformations of the initial 'palaeotype' proboscis proceeded in two primary ways: increasing bilateral symmetry (Callinera, Hubrechtella, and most of Heteronemertea) and increasing polyradial symmetry (Baseodiscidae, Oxypolellinae, and Hoplonemertea). The musculature of the middle portion of the proboscis differs among the three groups with armature: Palaeonemertea (genus Callinera), Polystilifera, and Monostilifera. The musculature of the stylet apparatus of the monostiliferous nemerteans is more complicated than that of the polystiliferous nemerteans, and consists of four muscle components--basal and anterior sphincters, radial and longitudinal musculature. Among the studied monostiliferans, the different components of the stylet musculature are developed to varying degrees. In addition, data on the structure of the rhynchocoel with interwoven musculature are provided. The taxonomic significance and phylogenetic interpretation of the proboscis and rhynchocoel musculature is discussed.

The structure of pseudocnidae of 16 species of Palaeonemertea clade Archinemertea (= Cephalotrichida s.l.) was investigated with confocal laser, scanning, and transmission electron microscopy (TEM). All species of the genus Cephalothrix possess two kinds of pseudocnidae, large and small. Only one type of pseudocnida is present in Balionemertes and Cephalotrichella. TEM revealed variation in the ultrastructure of large and small pseudocnidae of four species of Cephalothrix. Pseudocnidae of Balionemertes, Cephalotrichella, and Cephalothrix differ in substructure: in Balionemertes and Cephalotrichella the medulla is located in the basal half of the pseudocnidae with а precore layer situated in the apical half, whereas in Cephalothrix spp. and other palaeonemerteans the medulla surrounds a precore layer. Our results confirm the division of archinemerteans into Cephalotrichidae (with genus Cephalothrix) and Cephalotrichellidae (with genera Cephalotrichella and Balionemertes). The synapomorphy of Cephalotrichidae is pseudocnida dimorphism and the synapomorphies of Cephalotrichellidae are the position of the pseudocnidae on epithelial ridges and the distinct organization of pseudocnida layers, specifically the relative position of the medulla and precore layers. The pseudocnida lateral process, one or more of which is present in most species observed, is a probable synapomorphy of the clade Archinemertea. This is the first application of pseudocnida features to distinguish super-generic nemertean taxa and the results suggest that pseudocnidae provide a useful source of characters for nemertean systematics.

A checklist of benthic ribbon worm species from the Caribbean coast of Colombia is presented, including synonyms, distributions, a photographic record, and the main morphologic characters of each species for a rapid identification. This is the first research focused broadly on nemerteans in Colombia. 54 specimens of nemerteans were hand-collected from the rocky littoral of two different localities, and identified according to personal experience and specialist literature. 13 species were found; of which 11 represent new records for the country. These species belong to eight different traditionally used families: Tubulanidae, Valenciniidae, Lineidae, Amphiporidae, Cratenemertidae, Emplectonematidae, Drepanophoridae and Ototyphlonemertidae. The most common and abundant species was Dushia atra. The biodiversity of nemerteans in Colombia seems to overlap with the nemertean fauna from Florida and Brazil, explained by the convergence of the North Brazil Current, Guiana Current, Caribbean Currents and the Panama-Colombia Contracurrent in the sampled region. The results of this work suggest that the Caribbean coast of Colombia is a region with a high diversity of nemerteans, and provide important taxonomic data for environmental assessments and future biological research.

We compared the anatomy of the holotype of the palaeonemertean Cephalothrix simula ( Iwata, 1952 ) with that of the holotypes of Cephalothrix hongkongiensis Sundberg, Gibson and Olsson, 2003 and Cephalothrix fasciculus ( Iwata, 1952 ), as well as additional specimens from Fukue (type locality of C. simula) and Hiroshima, Japan. While there was no major morphological discordance between these specimens, we found discrepancies between the actual morphology and some statements in the original description of C. simula with respect to supposedly species-specific characters. Our observation indicates that these three species cannot be discriminated by the anatomical characters so far used to distinguish congeners. For objectivity of scientific names, topogenetypes of the mitochondrial cytochrome c oxidase subunit I (COI) sequences are designated for C. simula, C. hongkongiensis, and C. fasciculus. Analysis of COI sequence showed that the Hiroshima population can be identified as C. simula, which has been found in previous studies from Trieste, Italy, and also from both the Mediterranean and Atlantic coasts of the Iberian Peninsula, indicating an artificial introduction via (1) ballast water, (2) ship-fouling communities, or (3) the commercially cultured oyster Crassostrea gigas ( Thunberg, 1793 ) brought from Japan to France in 1970s. Cephalothrix simula is known to be toxic, as it contains large amounts of tetrodotoxin (TTX). We report here that the grass puffer Takifugu niphobles ( Jordan and Snyder, 1901 )-also known to contain TTX- consumes C. simula. We suggest that the puffer may be able to accumulate TTX by eating C. simula.

Tubulanus mawsoni (Wheeler 1940) comb. nov. is redescribed based on the material collected during the Spanish Antarctic expedition Bentart-03 through the Bellingshausen Sea (Antarctica). Previously belonging to the genus Carinina Hubrecht 1885, the species is transferred to the genus Tubulanus based on the situation of the brain and lateral nerves between the basement membrane and the outer circular muscle layer, the presence of muscle crosses in the body wall, the position of the blood and excretory systems, the presence of proboscis nerves and the structure of the cerebral sensory organs, the rhynchocoel wall and the rhynchodaeal and buccal nerves. Microphotographs of the different regions of the body and the proboscis structure are provided for the first time. Based on the ambiguities between Tubulanus diagnosis and the species descriptions, as well as the possible paraphyletic status of the genus, the need for new morphological and molecular studies is suggested.http://zoobank.org/urn:lsid:zoobank.org...

Carinoma hamanako sp. nov., the ninth member of the genus, is described based on three specimens collected in Lake Hamana, Shizuoka Prefecture, Pacific coast of Honshu, Japan. The new species can be distinguished from its congeners by the blind-ending rhynchocoelic villar vessels, the lateral rhynchocoel vessels rejoining the lateral vessels, the unbranched excretory canals, the situation of the posterior part of the mid-dorsal nerve within the longitudinal muscle layer of the body wall, and the absence of a vascular plexus in the foregut region. A phylogenetic analysis based on the nuclear 28S rRNA and mitochondrial cytochrome c oxidase subunit I genes shows that C. hamanako sp. nov. forms a clade with two congeners, C. mutabile Griffin, 1898 and C. tremaphoros Thompson, 1900, thus supporting the generic placement of the new species.

Callinera emiliae sp. nov., the ninth member of the genus, is described based on three specimens collected in Dumaguete, Negros Island, Republic of the Philippines. The new species can be distinguished from its congeners by the following characteristics: lateral sensory organs present; sub-epidermal glandular cells absent; blood vascular system without a ventral cephalic connective; nervous system with two dorsal cerebral commissures; and foregut nerves fused to form a ganglion in front of the mouth. In living specimens, epidermal constrictions were observed in the intestinal region; the presence of intestinal sphincters was confirmed in sectioned material and these correspond with the epidermal constrictions.

Based on specimens collected subtidally (similar to 10 m in depth) in Tomioka Bay, Japan, we describe the palaeonemertean Tubulanus tamias sp. nov., which differs from all its congeners in body coloration. In molecular phylogenetic analyses based on partial sequences of the nuclear 18S and 28S rRNA genes and histone H3, as well as the mitochondrial 16S rRNA and cytochrome c oxidase subunit I genes, among selected palaeonemerteans, T. tamias nested with part of the congeners in Tubulanus, while the genus as currently diagnosed appears to be non-monophyletic. Molecular cloning detected polymorphism in 28S rDNA sequences in a single individual of T. tamias, indicating incomplete concerted evolution of multiple copies. Tubulanus tamias is peculiar among tubulanids in having 9-10 pigment-cup eyes in the epidermis on either side of the head anterior to the cerebral sensory organs, and remarkably there are two types of eyes. The anterior 8-9 pairs of eyes, becoming larger from anterior to posterior, are completely embedded in the epidermis and proximally abutting the basement membrane; each pigment cup contains bundle of up to seven, rod-shaped structure that resemble a rhabdomeric photoreceptor cell. In contrast, the posterior-most pair of eyes, larger than most of the anterior ones, have an optical cavity filled with long cilia and opening to the exterior, thus appearing to have ciliary-type photoreceptor cells. The size and arrangement of the eyes indicate that the posterior-most pair of eyes are the remnant of the larval (or juvenile) eyes.

16 pages, 4 figures, 2 tables, A literature-based taxonomic catalogue of nemerteans (phylum Nemertea) from Spain and Portugal is provided, listing 75 species (12 Palaeonemertea, 24 Pilidiophora, and 39 Hoplonemertea) belonging to 34 genera. This is a low species number compared with the approximately 400 species listed in Europe. This lack of knowledge is mainly due to the low number of researchers interested in the phylum and the well-known taxonomic difficulties of its study. Geographic records are indicated for each species, and for some, comments are included on certain biological and taxonomic aspects, This research has benefited from funds from the Instituto de Estudios Ceutíes, Ceuta, Spain. The study was also supported by the Franklin Institute, Universidad de Alcalá

Callinera monensis sp. n., from Derbyhaven Harbour in the Islc of Man, is described and illustrated. The anatomy of the species is compared and contrasted with that of the other known members of the genus, Callinera buergeri Bcrgcndal and C. buergeri f. grandis Bergendal, from Scandinavian waters;f. grandis is afforded separate specific status as C. grandis Bcrgcndal.

Sundberg, Per, Chernyshev, Alexey. V., Kajihara, Hiroshi, Kånneby, Tobias, Strand, Malin (2009): Character-matrix based descriptions of two new nemertean (Nemertea) species. Zoological Journal of the Linnean Society 157 (2): 264-294, DOI: 10.1111/j.1096-3642.2008.00514.x, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2008.00514.x

Kajihara, Hiroshi (2006): Four palaeonemerteans (Nemertea: Anopla) from a tidal flat in middle Honshu, Japan. Zootaxa 1163: 1-47, DOI: 10.5281/zenodo.2645302, {"references":["Bergendal, D. (1900a) Studier ofver nemertiner. I. Callinera burgeri Bergendal, en representant for ett afvikande slagte bland Palaonemertinerna. Lunds Universitets Arsskrift, 36, 1-47.","Bergendal, D. (1900b) Till kannedomen om de nordiska Nemertinerna. Ofversigt af Kongliga Vetenskaps-Akademiens Forhandlingar, 57, 581-602.","Bergendal, D. (1900c) Uber ein Paar sehr eigenthumliche nordische Nemertinen. Zoologischer Anzeiger, 23, 313-328.","Bergendal, D. (1901) Studier ofver nemertiner. I. Callinera burgeri Bergendal, en representant for ett afvikande slagte bland Palaonemertinerna. Lunds Universitets Arsskrift, 37, 49-116.","Bergendal, D. (1902a) Zur Kenntnis der nordischen Nemertinen. 2. Eine der construierten Urnemertine entsprechende Palaeonemerine aus dem Meere der schwedischen Westkuste. 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Two new palaeonemerteans, Carinoma patriciaesp. nov., and Hubrechtella queenslandicasp. nov., from the Great Barrier Reef province of Australia, are described and illustrated. The systematic relationships of these nemerteans are discussed and emended diagnoses are given for the genera Carinoma (Carinomidae) and Hubrechtella (Hubrechtidae).

On the basis of the axial relationship between the larva and adult, it is possible to divide the larvae of the members of the Heteronemertea and a species of the Palaeonemertea into five types, forming a sequential series in which the adult axis of each advances by an angle of 45° from that immediately preceding it. Phylogenetic consideration of the Nemertea has been undertaken in the light of evidence provided by the axiality of the embryos and the developmental data hitherto recorded.