Scanning electron micrographs showing an overview and details of the trunk morphology of Condyloderes megastigma sp. nov. (A) Segments 1 and 2, lateral view. (B) Segments 3 and 4 in female specimen, ventral view. (C) Giant ventromedial sensory spot on segment 9. (D) Segments 7 to 9, ventral view. (E) Right sternal plate of segment 8. (F) Segments 9 to 11 in a female specimen (note the shape of the posterior segment margin of segment 10), ventral view. Abbreviations: gss, giant sensory spot; ld, laterodorsal sensory spot; ltas, lateral terminal accessory spine; lva, lateroventral acicular spine; lvc, lateroventral cuspidate spine; mda, middorsal acicular spine; ml, midlateral sensory spot; mts, midterminal spine; pl, placid; pv, paraventral sensory spots; sd, subdorsal sensory spot; sl, sublateral sensory spot; ss3, sensory spot type 3 (modified sensory spot); va, ventromedial appendage; vm, ventromedial sensory spots. 

Scanning electron micrographs showing an overview and details of the trunk morphology of Condyloderes megastigma sp. nov. (A) Segments 1 and 2, lateral view. (B) Segments 3 and 4 in female specimen, ventral view. (C) Giant ventromedial sensory spot on segment 9. (D) Segments 7 to 9, ventral view. (E) Right sternal plate of segment 8. (F) Segments 9 to 11 in a female specimen (note the shape of the posterior segment margin of segment 10), ventral view. Abbreviations: gss, giant sensory spot; ld, laterodorsal sensory spot; ltas, lateral terminal accessory spine; lva, lateroventral acicular spine; lvc, lateroventral cuspidate spine; mda, middorsal acicular spine; ml, midlateral sensory spot; mts, midterminal spine; pl, placid; pv, paraventral sensory spots; sd, subdorsal sensory spot; sl, sublateral sensory spot; ss3, sensory spot type 3 (modified sensory spot); va, ventromedial appendage; vm, ventromedial sensory spots. 

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A new kinorhynch species, Condyloderes megastigma sp. nov., is described from the Korea Strait. The new species is characterized by the presence of 16 placids with either eight, four, or two knobby projections, middorsal and lateroventral acicular spines on segments 1 to 9, lateroventral cuspidate spines on segment 2 in females only, but otherwise...

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Context 1
... 5C, 6D). Paradorsal sensory spots are not present. Elevated sensory spots are present in the lat- erodorsal, sublateral and ventromedial positions. A pair of unusual structures is present in the ventromedial positions (Figs. 2B, 5C, 6D, 6E). The structures are cone-shaped, approximately 8 μm long, and densely fringed throughout their length (see Fig. 6E for similar structure on segment 8). They may possibly be regarded enlarged tubules or papillae, but appear more like the distal part of a cuspidate spine. Due to the uncertainty in the nature of this new structure, we refer to them here with the neutral term "appendages". The ventromedial appendages are present in both sexes. Cutic- ...
Context 2
... 8 with middorsal and lateroventral acicular spines, and lateroventral cuspidate spines (Figs. 2A, 2B, 5C, 6D). A pair of non-elevated paradorsal sensory spots is present. Elevated sensory spots are also present in the lat- erodorsal and ventromedial positions. A pair of ventromedial appendages is present (Fig. 6E). Compared to those on the previous segment, these appendages are slightly longer, and are located slightly closer to the midsternal junction (Figs. 2B, 5C, 6D). Cuticular hairs and posterior segment margin appear as on the previous ...
Context 3
... spots (Figs. 2B, 5E, 6C, 6D). The giant sensory spots con- sist of numerous minute papillae that cover an area of 10 × 7 μm (Fig. 6C). Pores are not clearly visible. Cuticular hairs and posterior segment margin appear as on the previous segment. Segment 10 without spines or paradorsal sensory spots ( Fig. 2A, B). Elevated sensory spots are present in the sub- dorsal and lateroventral positions. The middorsal line is marked by a serrated ridge (Fig. 2A). The posterior ...
Context 4
... ( Fig. 2A, B). Elevated sensory spots are present in the sub- dorsal and lateroventral positions. The middorsal line is marked by a serrated ridge (Fig. 2A). The posterior segment margin is serrated as well and has no spicular fringe. The posterior margins of the sternal plates are straight in males, whereas they are broadly concave in females (Fig. 6F). Female gonopores are ...

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Citations

... the Sea of Japan and adjacent waters (Neuhaus, 2013). (Higgins, 1966;Higgins and Korczynski, 1989;Adrianov, 1995;Grzelak and Sørensen, 2018;Sørensen and Grzelak, 2018;Yamasaki et al., 2018a), the Iberian Peninsula (Pardos et al., 1998;GªOrdóñez et al., 2008;Sørensen et al., 2010a;2012a;Sánchez et al., 2011;Neves et al., 2016), the Sea of Japan and adjacent waters (Adrianov, 1989;Higgins and Shirayama, 1990;Song and Chang, 2001;Adrianov et al., 2002;Lundbye et al., 2010;Sánchez et al., 2013;Sørensen et al., 2010a;2010b;2010c;2012a;2012b;2013;Yamasaki, 2016;Yamasaki and Kahijara, 2012;2020), or the north-western ...
... For this reason, the head is attached to the S1 and S2 through a complex net of muscles (Kristensen and Higgins, 1991;Altenburger, 2016). In addition, the S1 and S2 are completely developed from the first postembryonic stages with fully functional structures (Kozloff, 1972;Higgins, 1977a;1977b;Neuhaus, 1993;Sørensen et al., 2000;2010b;Schmidt-Rhaesa and Rothe, 2006;Neuhaus and Kegel, 2015), evidencing the tremendous importance of this body region for the kinorhynch life. Thus, the S1 and S2 would proportionally grow more slowly (compared to the general trunk growth) due to morpho-physiological restrictions, as an increase in its relative growth rate could lead to fatal head dysfunctions. ...
... At its anterior end, the mouth cone reveals 9 outer oral styles, which appear to be collapsed around the partly protruded inner foregut structures, but each style is separated from its neighbouring style. Each style is broader over about three quarters of its length and appears filamentous in the apical part (for comparison see Sørensen et al. 2010b, fig. 4A). ...
... In addition, the specimen reveals a sublateral cuspidate on segments 3, possibly 4 and 9 lacking in C. paradoxus, and the ventrolateral cuspidate spine on segment 5 is displaced towards the ventromedial position in comparison with C. paradoxus. Sublateral cuspidate spines do not occur in any other known species of Condyloderes (Table 7; Higgins 1969;Adrianov et al. 2002;Sørensen et al. 2010b;. The specimen is not described formally as a new species, because only a single specimen is available, many cuticular characters are hard to recognize if at all also because of the thin cuticle, the specimen is covered with detritus hampering observation, and the preparation is not too satisfying. ...
... Characters unique within genus are underlain with grey background. Measurements taken from Higgins (1969), Adrianov et al. (2002), , Sørensen et al. (2010b), , and this paper (Tables 2 6). ...
Article
The description of a new representative of the species-poor genus Condyloderes Higgins, 1969 from the Northeast Pacific (Alaska) is reported. The analyzed specimens of Condyloderes shirleyi sp. nov. showed a significant variation of numerous morphological characters, along with female-specific traits known also from other congeneric species. These findings stimulated the re-investigation of the type material of the six species of Condyloderes described so far, i.e., C. kurilensis Adrianov & Maiorova, 2016, C. megastigma Sørensen, Rho & Kim, 2010b, C. multispinosus (McIntyre, 1962) Higgins, 1969, C. paradoxus Higgins, 1969, C. setoensis Adrianov, Murakami & Shirayama, 2002, and C. storchi Higgins, 2004 in Martorelli & Higgins, 2004. Our study allowed to reveal various morphological novelties and to emend the diagnosis of these species and of the genus Condyloderes. Furthermore, our analysis led to synonymize C. megastigma with C. setoensis. The results of our investigation about the significant variation in C. shirleyi sp. nov. raise a wider question on species identity within Kinorhyncha, underscoring the necessity, if possible, to describe new species from a higher number of specimens and to concentrate on the morphological variation of the going-to-be-described species.
... coast of Argentina (San Jorge Gulf) [7]. Specimens of the fifth and more recently described species, C. megastigma Sorensen, Rho et Kim, 2010, were coll lected in the Korea Strait between Korea and Japan at the depth from 90 to 109 m [10]. The only previously known abyssal species of the Kinorhyncha from the NorthhWest Pacific is Pycnoo phyes abyssorum Adrianov et Maiorova, 2015, recently described from the locality at the abyssal plain near the KurillKamchatka Trench [1]. ...
... coast of Argentina (San Jorge Gulf) [7]. Specimens of the fifth and more recently described species, C. megastigma Sorensen, Rho et Kim, 2010, were coll lected in the Korea Strait between Korea and Japan at the depth from 90 to 109 m [10]. The only previously known abyssal species of the Kinorhyncha from the NorthhWest Pacific is Pycnoo phyes abyssorum Adrianov et Maiorova, 2015, recently described from the locality at the abyssal plain near the KurillKamchatka Trench [1]. ...
... Only two species, C. kurilensis sp. nov. in females and C. megastigma in both sexes, have ventromedial appendages (tubules) on trunk segments 7–8 and large rounddshaped areas of densely arranged cuticular spinelets on trunk segment 9. In C. megastigma, these areas were interpreted as giant sensory spots [10]. Con trary to this species, females of C. kurilensis sp. ...
Article
A new species of centroderid kinorhynchs, Condyloderes kurilensis sp. nov., collected from muddy sediment at the abyssal plain near the Kuril-Kamchatka Trench, North-West Pacific, is described and illustrated using light and electron microscopy. Previously, members of the genus Condyloderes were found only in nearly shallow water enviroments—C. multispinosus (North Sea), C. paradoxus (Bengal Bay), C. setoensis (Tanabe Bay, Japan), C. storchi (Argentina coast), and C. megastigma (Korea Strait). The new species is characterized by the presence of lateroventral cuspidate spines only on trunk segment 8 in both sexes, paired ventromedial appendages (tubules) on trunk segments 7 and 8 in females only, and dorsolateral spines on trunk segment 10 in males only. Condyloderes kurilensis sp. nov. constitutes the sixth species of the genus Condyloderes described so far and the only second abyssal species of the Kinorhyncha described from the North-West Pacific. A key to the six species of the genus Condyloderes is proposed.
... Current biogeographical evidence, however, challenges this hypothesis because the coasts of Japan and Korea have been intensively surveyed for kinorhynchs from deep sea sediments to the intertidal zone, and so far E. ohtsukai has only been reported from the type locality (e.g. Sørensen et al., 2010Sørensen et al., , 2012aSørensen et al., , 2013Thomsen et al., 2013;Yamasaki, 2015;Yamasaki and Fujimoto, 2014;Yamasaki and Kajihara, 2012). ...
... This trait is autapomorphic for the genus. Interestingly though, the outer oral styles in species of Condyloderes appear to be, if not lost or fused, then at least strongly reduced (Fig 4D) [47], but it is uncertain whether or not there is an evolutionary link between the lack of outer oral styles in Campyloderes and the reduction in Condyloderes. In the morphological analysis of the present study, these two conditions were coded as different stages in the same character, but the analysis did not unambiguously support the existence of a transformation series. ...
... 5. Placids with either (a) broad midventral placid + two narrow ventrolateral placids + other placids alternatingly narrow and broad, or (b) very broad midventral placid + other placids alternatingly narrow and broadcondition (a) applies to species of Centroderes and Condyloderes [47]- [49], whereas condition (b) applies to Campyloderes exclusively [45], [46]. Hence, this trait does not support Campyloderes as more closely related with neither Centroderes nor Condyloderes. ...
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The phylogeny of Kinorhyncha was analyzed using morphology and the molecular loci 18S rRNA and 28S rRNA. The different datasets were analyzed separately and in combination , using maximum likelihood and Bayesian Inference. Bayesian inference of molecular sequence data in combination with morphology supported the division of Kinorhyncha into two major clades: Cyclorhagida comb. nov. and Allomalorhagida nom. nov. The latter clade represents a new kinorhynch class, and accommodates Dracoderes, Franciscideres, a yet undescribed genus which is closely related with Franciscideres, and the traditional homalor-hagid genera. Homalorhagid monophyly was not supported by any analyses with molecular sequence data included. Analysis of the combined molecular and morphological data furthermore supported a cyclorhagid clade which included all traditional cyclorhagid taxa, except Dracoderes that no longer should be considered a cyclorhagid genus. Accordingly, Cyclorhagida is divided into three main lineages: Echinoderidae, Campyloderidae, and a large clade, 'Kentrorhagata', which except for species of Campyloderes, includes all species with a midterminal spine present in adult individuals. Maximum likelihood analysis of the combined datasets produced a rather unresolved tree that was not regarded in the following discussion. Results of the analyses with only molecular sequence data included were incongruent at different points. However, common for all analyses was the support of several major clades, i.e., Campyloderidae, Kentrorhagata, Echinoderidae, Dracoderi-dae, Pycnophyidae, and a clade with Paracentrophyes+ New Genus and Franciscideres(in those analyses where the latter was included). All molecular analyses including 18S rRNA sequence data furthermore supported monophyly of Allomalorhagida. Cyclorhagid mono-phyly was only supported in analyses of combined 18S rRNA and 28S rRNA (both ML and BI), and only in a restricted dataset where taxa with incomplete information from 28S rRNA had been omitted. Analysis of the morphological data produced results that were similar with those from the combined molecular and morphological analysis. E.g., the PLOS ONE |
... Cuspidate spines are found in species of several cyclorhagid genera, including Antygomonas, Condyloderes, Semnoderes, Sphenoderes, Triodontoderes, Wollunquaderes andZelinkaderes (see e.g. Higgins, 1969, 1990;Nebelsick, 1990;Bauer-Nebelsick, 1995Adrianov et al., 2002;Sørensen, 2007;Sørensen et al., , 2010bSørensen et al., , 2010cSørensen & Thormar, 2010). Of course the position of spines differ between the species of the genera, but in all species, cuspidate spines appear consistently on segment 5, and only Sphenoderes poseidon lacks cuspidate spines on segment 8 (Sørensen et al., 2010b). ...
Article
A new genus and species of Kinorhyncha, Franciscideres kalenesos gen. et sp. nov., is described from tidal and subtidal sandy habitats in Brazil. The new genus and species is characterized by an extremely flexible trunk without pachycycli that appears perfectly circular in cross-section, segments 1, 2 and 11 consisting of closed rings and 3 to 10 of single, bent plates with midventral articulations, a neck without placids that resembles an additional segment, densely packed scale-like, cuticular hairs, and a terminal segment with a middorsal spine and two sets of lateral terminal spines, but no midterminal spine. Phylogenetic analyses of 18S rRNA of Franciscideres kalenesos gen. et sp. nov. and 47 other kinorhynch ingroup taxa suggest that the new genus is a basal homalorhagid, whereas comparison of morphological characters indicates affinities between the new genus and the peculiar cyclorhagid Cateria.http://zoobank.org/urn:lsid:zoobank.org:pub:64E29D97-DE1D-4511-8683-C969DD2EED43
... Sørensen et al. 2010aSørensen et al. , 2012a. Exceptions are Polacanthoderes martinezi, Zelinkaderes klepali, and Z. brightae that have more scalids in even numbered sections (see Bauer-Nebelsick 1995;Sørensen et al. 2007;Sørensen 2008), and Condyloderes megastigma, Triodontoderes anulap and Z. floridensis that have the same number of scalids in all sections (see Higgins 1990;Sørensen and Rho 2009;Sørensen et al. 2010c). Also the differences of scalid numbers in the posterior introvert rings have been reported previously. ...
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Full-text available
Meristoderes is the most recently described kinorhynch genus and has until now only accommodated its type species M. macracanthus from the Mediterranean and M. galatheae from the Solomon Islands in Southeast Asia. The present contribution is an account on the genus based on samples from the ongoing exploration of the kinorhynch fauna in coastal and subtidal waters around in the Korean Peninsula. The samples yielded no less than five new species of Meristoderes, of which four are formally described herein, whereas only diagnostic notes are provided for the fifth. The descriptions are supplemented with a key to species of the genus, and morphological notes on the organization of cephalic structures and cuticular trunk structures. New morphological findings include intrageneric variation in the composition of the terminal segment, which may have either one unpaired or two paired tergal plates. The data also show that the differentiation of a tergal and sternal plate in segment 2 varies greatly between the species. Results of the present study indicate that the genus probably holds a significant, unrecovered biodiversity, but the data also shed light on potential future taxonomic problems among genera of Echinoderidae and stress the importance of identifying additional diagnostic traits to define Meristoderes.
... 158; ssp, sensory spot; st, sternal plate; su, suture; TL, total length; tm, transversal muscle; tp, tergal plate; tr, trichoscalid; TVG, TV-grab; vl, ventrolateral; vl5, (length of) ventrolateral spine of segment; vm, ventromedial; ZMB, followed by catalogue number of expeditions with RV Sonne and RV Polarstern revealed plenty of material of Kinorhyncha (Fahrbach and Gerdes, 1997; Arntz and Gutt, 1999; Hoernle et al., 2003; Werner et al., 2000 Werner et al., , 2009 Werner et al., , 2010 Werner, 2002; Werner and Hauff, 2007; Sørensen, 2008). New species and genera of Kinorhyncha have been described in recent years with the aim of searching for characters for a phylogenetic analysis of Kinorhyncha (Neuhaus, 2004; Neuhaus and Blasche, 2006; Sørensen et al., 2007, 2010a Sørensen, 2008; Sørensen and Rho, 2009; Sørensen and Thormar, 2010; Herranz et al., in press). Among the genera studied, Campyloderes seemed to represent an easy target, because it consists of three species and one subspecies only, namely C. vanhoeffeni Zelinka, 1913, C. vanhoeffeni var. ...
... Therefore, these characters have virtually not been available for species identification. In more recent publications, Pardos, Higgins, Sørensen, Neuhaus and coworkers summarize data about the arrangement of spines, sensory spots, gland cell outlets, and papillae in a table with reference positions introduced by Pardos et al. (1998) (see also Neuhaus and Blasche, 2006; Sørensen, 2006; Sørensen et al., 2000, 2007, 2010a,b; Sørensen and Thormar, 2010). Information about intraspecific variation of characters in Kinorhyncha such as asymmetrical distribution of sensory spots is available for very few species, namely for juvenile and adult Pycnophyes kielensis Zelinka, 1928 and P. dentatus Reinhard, 1881 and for juvenile Paracentrophyes praedictus Higgins, 1983 (see Neuhaus, 1993). ...
... Consequently, the proposed complete fusion of outer oral styles has to be regarded as an autapomorphic character of Campyloderes. Primary scalids with internal septa are not known for any kinorhynch except specimens of Campyloderes (Higgins, 1969Higgins, , 1983Higgins, , 1990 Brown, 1989; Neuhaus, 1993 Neuhaus, , 1995 Neuhaus, , 2004 BauerNebelsick, 1995, 1996 Adrianov and Malakhov, 1999; Sørensen, 2007; Sørensen et al., 2007, 2010a Sørensen and Rho, 2009; this paper). The chambering of scalids is regarded as an autapomorphy of species of Campyloderes. ...
Article
Altogether, 103 adult and 23 juvenile specimens of Campyloderes from 33 locations both in the deep sea and on the continental shelf all over the world were studied by light microscopy (97 specimens) and scanning electron microscopy (28 specimens). Especially from the Faroe Island, the Central American East Pacific Ocean and from the area east and northeast of New Zealand, enough specimens are available to study the regional variation of characters. Specimens both from these regional areas and worldwide reveal a significant morphological variation, especially in the distribution of sensory spots, gland cell outlets, and papillae, whereas characters conventionally used for species identification, such as spine pattern do not vary much. Overlapping character patterns do not allow identification of different species and to discriminate the current populations from previously described species. We conclude that the morphological variation results from ongoing species formation processes. We also report observations that two adult life history stages may exist in Campyloderes. The character set in the ground pattern of Campyloderes is presented.
... Positional terminology follows that proposed by Pardos et al. (1998) with the addition of a paraventral position (Sørensen et al., 2010) and an amended specification of the divisions between the positions proposed for the genus Echinoderes (Thormar and Sørensen, 2010), which can also be applied to Meristoderes gen. nov. ...
... Positional terminology follows that proposed by Pardos et al. (1998) with the addition of a paraventral position (Sørensen et al., 2010) and an amended specification of the divisions between the positions proposed for the genus Echinoderes (Thormar and Sørensen, 2010), which can also be applied to Meristoderes gen. nov. ...
Article
A new kinorhynch genus, Meristoderes gen. nov., and two new species from Spain and the Solomon Islands, respectively, are described. The new genus is distinguished from all other genera by the first segment consisting of a closed cuticular ring, and the second segment having partial tergosternal junctions, and a superficial midventral fold. This is a new cuticular configuration that may shed light into the phylogenetic relationships of echinoderid kinorhynchs. Meristoderes macracanthus gen. et sp. nov. from the Mediterranean coast of Spain is recognised by the presence of middorsal spines on segments 4, 6 and 8, ventrolateral tubules on segment 2, lateroventral tubules on segment 5, lateroventral spines on segments 6–9, lateral accessory tubules on segment 8, one pair of laterodorsal tubules on segment 10. Meristoderes galatheae sp. nov. from the Solomon Islands is recognized by having a middorsal spine on segment 4 only, ventrolateral tubules on segment 2, lateroventral tubules on segment 5, lateroventral spines on segments 6–9, lateral accessory tubules on segment 8 and subdorsal tubules on segment 10. Both species have a pattern of paraventral perforation site clusters on segments 3–9, with conspicuously long bracteate hairs from the posteriormost perforations sites on the segments 3–7 and 3–6, respectively.The new genus Meristoderes gen. nov. is included into the family Echinoderidae Bütschli, 1876 and appears closely related with the genera Cephalorhyncha Adrianov, 1999 and Echinoderes Claparède, 1863. The new information it provides is discussed to clarify the internal phylogeny of Echinoderidae. The terminology for cuticular characters in the overlapping area between consecutive segments is also standardized.
... However, ongoing studies show that the Korean kinorhynch fauna is far from being well-described. Various non-echinoderid new species from Korea have recently been described in a series of papers (Sørensen et al. 2010a(Sørensen et al. , 2010b(Sørensen et al. , 2010c, and the next step in this survey has now been reached, with an attempt to approach the diverse composition of Echinoderes in these samples. Kinorhynchs from more than 70 samples taken in Korean or nearby waters have been examined, and the present paper represents the first contribution about Echinoderes new to Korea. ...
Article
Full-text available
A large collection of kinorhynch specimens from coastal and subtidal localities around the Korean Peninsula and in the East China Sea was examined, and the material included several species of undescribed or poorly known species of Echinoderes Claparède, 1863. The present paper is part of a series dealing with echinoderid species from this material, and inludes descriptions of four new species of Echinoderes, E. aspinosus sp. nov., E. cernunnos sp. nov., E. microaperturus sp. nov. and E. obtuspinosus sp. nov., and redescriprion of the poorly known Echinoderes tchefouensis Lou, 1934.