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A new species of echinoderid kinorhynchs, Echinoderes xiphophorus sp. nov. collected from oxidized brown silt at the deepest depression in the Sea of Japan, North-West Pacific, is described and illustrated using light and electron microscopy. This new representative of the most speciose kinorhynch genus is characterized by the unique set of spines and tubes and can easily be distinguished from most of its congeners. The second trunk segment bears three pairs of tubes in subdorsal, midlateral and ventrolateral position in both sexes; one pair of tubes on trunk segment 5 in lateroventral position and on trunk segment 8 in sublateral position; aciculate lateroventral spines on trunk segments 6–9; aciculate middorsal spines on trunk segments 4, 6, 8. This species is well recognized by very long tergal extensions of the posteriormost segment, some of the longest within the family Echinoderidae. Males of Echinoderes xiphophorus sp. nov. are well distinguished from all the congeners by extremely long sword-like appendages dorsally to three pairs of penile spines. The species constitutes the first deep-sea representative of the Echinoderidae in the Sea of Japan and the deepest representative of the Kinorhyncha in this sea.
Limited data are available for the kinorhynch fauna from the Southern Hemisphere, with little or no data from New Zealand. Here, we provide a first comprehensive overview of the diversity of mud dragons, with an emphasis on species of Echinoderes from the continental slope of New Zealand, from a variety of habitats such as slopes, canyons and seamounts located in the Hikurangi Margin region. The study revealed fifteen species of Echinoderes. Of these, ten are described as new to science: E. aragorni sp. nov., E. blazeji sp. nov., E. dalzottoi sp. nov., E. frodoi sp. nov., E. galadrielae sp. nov., E. gandalfi sp. nov., E. landersi sp. nov., E. leduci sp. nov., E. legolasi sp. nov. and E. samwisei sp. nov. Moreover, Echinoderes juliae Sørensen et al., 2018, Echinoderes sp. aff. E. balerioni, Echinoderes sp. aff. E. galadrielae/beringiensis, Echinoderes sp. aff. E. lupherorum and Echinoderes sp. aff. E. unispinosus are reported in the investigated region. The most abundant among all was E. gandalfi sp. nov., but it was found only in canyons. Interestingly, the second most common species was E. juliae that was found at several stations in canyons, seamount and on the slope. This species is one of the deep-sea species originally found on the abyssal plain off Oregon and along the continental rise off California, Northeast Pacific, recorded in polymetallic nodules in the tropical eastern Pacific, and recently found on the abyssal plains off Chile, east of the Atacama Trench. These findings, together with records of Echinoderes sp. aff. E. lupherorum and Echinoderes sp. aff. E. unispinosus indicate that, despite their low dispersal abilities, kinorhynchs, similar to other meiofaunal species, may exhibit a wider distribution pattern than previously assumed. The number of recorded species and numerous new species show that New Zealand sediments not only are inhabited by a diverse kinorhynch fauna, but Echinoderes, the most speciose genus, still holds much to discover.
Thirteen species of Echinoderes with nearly identical spine/tube patterns, and apparently similar tergal extensions were re-examined and compared. Based on this, redescriptions and/or emended species diagnoses are provided for Echinoderes aureus, E. dujardinii, E. gerardi, E. imperforatus, E. pacificus, E. pilosus, E. sensibilis, E. sublicarum and E. worthingi, and new details about cuticular structures are added for E. kozloffi and E. gizoensis. The new information derived from the redescriptions, and the subsequent comparative studies revealed that: 1) the holotype of Echinoderes lanceolatus is identical with the types of Echinoderes aureus, and E. lanceolatus is thus a junior synonym of E. aureus; other potentially synonymous species that should be addressed further in the future include: E. dujardinii + E. gerardi; E. imperforatus + E. sensibilis, and E. pacificus + E. sublicarum; 2) the paratypes of E. lanceolatus represented a different yet undescribed species, here described as E. songae Sørensen & Chang sp. nov.; 3) a comparison with literature information about E. ehlersi showed that the species is so insufficiently described that a redescription of topotype material is required before the species should be considered for taxonomic comparison; 4) specimens from the Andaman Islands, India, that previously have been reported as Echinoderes cf. ehlersi represent two different undescribed species, of which one is described as E. chandrasekharai Sørensen & Chatterjee sp. nov. and the other is left undescribed due to the limited material available; 5) out of a total of fifteen addressed species, it is proposed that eleven represent a putatively monophyletic group that is named the Echinoderes dujardinii group. The group includes following species: E. dujardinii, E. ehlersi, E. gerardi, E. imperforatus, E. kozloffi, E. sensibilis, E. pacificus, E. sublicarum, E. songae Sørensen & Chang sp. nov., E. chandrasekharai Sørensen & Chatterjee sp. nov., and Echinoderes sp. from the Andaman Islands, and is supported by a similar spine/tube pattern (except for variation regarding the presence of lateral accessory tubes on segment 8); generally short middorsal spines, especially on segments 4 to 6; glandular cell outlets type 1 always present in middorsal positions on segments 1 to 3, and in subdorsal positions on segments 4 to 9; glandular cell outlets type 2 always present in laterodorsal or midlateral positions on segment 8, and sometimes in same positions on segment 9 but never at any other segments or positions; female papillae always present on sternal plates of segments 7 and 8, and occasionally also on segment 6; tergal extensions well-spaced, triangular, gradually tapered cones, and pectinate fringes of sternal extensions are differentiated into seta-like tufts. The comparisons furthermore showed potential taxonomic significance of two echinoderid character traits that previously have been slightly neglected as diagnostic traits, namely the presence and appearance of female papillae, and the dorsal pattern of glandular cell outlets type 1. Female papillae may occur on the sternal plates of segments 6 to 8, but the positions may differ from ventrolateral to ventromedial, and the morphology of the intracuticular substructure also differ at species level. Information about position and morphology of female papillae proved helpful for species recognition, but it might also provide information of phylogenetic importance. Analyses of glandular cell outlet type 1 patterns on the dorsal sides of segments 1 to 9 in species of Echinoderidae, revealed several apparently unique or rare patterns, but also three distinct patterns that applied to larger groups of species. One pattern is the one present in all species of the E. dujardinii group, whereas the other two common patterns included 1) middorsal outlets on segments 1 to 3, and paradorsal outlets on segments 4 to 9 (found in 27 species), and 2) middorsal outlets on segments 1 to 3, 5 and 7, and paradorsal outlets on segments 4, 6 and 8 to 9 (found in 27 species).
The bathyal kinorhynch fauna along the Northwest American continental rise is explored, with emphasis on species of Echinoderidae Zelinka, 1894. Seven species of Echinoderes Claparède, 1863 are described as new to science: E. anniae sp. nov., E. dubiosus sp. nov., E. hamiltonorum sp. nov., E. hviidarum sp. nov., E. juliae sp. nov., E. lupherorum sp. nov. and E. yamasakii sp. nov. Three known species, Echinoderes hakaiensis Herranz, Yangel & Leander, 2017, E. cf. unispinosus Yamasaki, Neuhaus & George, 2018 and Fissuroderes higginsi Neuhaus & Blasche, 2006, are reported. The numerous new species indicate that the deep-sea still holds a great, undiscovered diversity of kinorhynchs, and that Echinoderes, as is also the case in more shallow, coastal waters, represents an important component of the deep-sea kinorhynch fauna. The presence of E. hakaiensis in the deepsea sediments demonstrates that the species may occur at a great depth range, and suggests that depth may play a less important role for the distribution of some kinorhynch species. The finding of the Northeast Atlantic species E. cf. unispinosus and the Southwest Pacific species Fissuroderes higginsi could indicate that kinorhynch species in the deep-sea may cover considerably larger distributional ranges than is assumed for coastal species of Echinoderidae.