identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03EB87BD8511FFEE8B69DAF7944A6E3B.text	03EB87BD8511FFEE8B69DAF7944A6E3B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Echinoderes remanei (Blake 1930)	<div><p>4.1. Comparison of E. remanei, E. tubilak and E. svetlanae</p> <p>E. remanei was originally described by Blake in 1930, and redescribed in 1964 by Higgins (Blake 1930; Higgins 1964). Although the redescription provided some more accurate information about the species, important taxonomic characters were still overlooked or interpreted incorrectly. This led to the species being characterized by having middorsal spines on segment 4 to 8 and lateral spines on segment 5 to 9, in addition to the spines on segment 11 (Higgins 1964). Therefore, when Higgins and Kristensen (1988) described E. tubilak several years later, the only morphological difference reported between E. remanei and a E. tubilak was the presence of ‘round to oval cuticular scars’ on segment 2 (Higgins &amp; Kristensen 1988). The authors highlighted the similarity between the two species, both regarding the presence of a prominent pectinate fringe on the ventral side of segment 1 and in the differences in the length of the lateral terminal spines between males and females. At present, we know that structures reported as “oval cuticular scars” in the earlier descriptions usually referred to glandular cell outlets type 2 - a diagnostic trait that was not known at that time (see discussion in Neuhaus &amp; Blasche 2006; Sørensen et al., 2012; Grzelak &amp; Sørensen 2018). The occurrence of glands in E. tubilak was also illustrated in the line art illustrations (see Figs 65–66, 68, 72 in Higgins &amp; Kristensen 1988), but only on the dorsal side of segments 2 and 4. The additional pairs in sublateral and ventrolateral positions on segment 2 were overlooked, as well as the pairs of gland openings on segment 5 and 8, which was subsequently reported by Grzelak &amp; Sørensen (2018). Presence of glandular cell outlets type 2 on segment 10 was not confirmed until this study, as there was previously no material available for SEM (visualising glandular cell outlets type 2 on segment 10 using LM only is very challenging). In the case of E. remanei, the presence of glands or cuticular scars has never been mentioned before, but the material deposited in the NMNH shows their presence, which is in agreement with the observation of the E. tubilak type material and additional specimens analysed in the present study. Echinoderes svetlanae that was described by Adrianov in 1999 represents a similar situation. According to the original description, the species was characterized by the presence of middorsal spines on segment 4 to 8, lateroventral spines on segment 6 to 9, and the absence of lateral tubes/spines on segment 5 (Adrianov &amp; Malakhov 1999). Again, thorough examination of the museum materials revealed a number of traits that were not reported previously, including the number and distribution pattern of glandular cell outlets type 2 matching that observed in E. tubilak, and even the presence of lateroventral tubes on segment 5. This character, however, due to the condition of E. svetlanae specimens was poorly visible, thus it is not really surprising that it was overlooked in previous studies (Adrianov &amp; Malakhov 1999; Grzelak &amp; Sørensen 2018).</p> <p>In addition to overlapping morphological characters in the three species, the specimens of E. remanei, E. tubilak, and E. svetlanae share distinct morphometric traits. When specimens of E. remanei from Maine were compared with specimens of E. tubilak from localities in Greenland, Alaska and Japan and with specimens of E. svetlanae from the White Sea, it became evident that most of the spine dimensions are highly comparable across the analysed populations (Fig. 8). The only exceptions were specimens from southern Alaska, of which middorsal spines on segment 8 were significantly shorter than those of specimens from Greenland; and lateroventral spines on segments 6 to 8 were slightly shorter than those from Greenland and Maine, likely due to the smaller body size of southern Alaskan specimens compared to Greenlandic individuals (Fig. 7). Nevertheless, a very clear pattern of the middorsal spine always being shorter on segment 8 in comparison to the middorsal spine on segment 7 was observed for all individuals (Fig. 8A). Differences in lateral terminal spines length between the sexes were also very prominent for all populations (Fig. 9), with no significant differences in spines length among males from different locations, and significant differences in spines length of females being detected only between specimens from southern Alaska and Maine.</p> <p>All of our results provide strong arguments for considering the studied specimens of E. remanei, E. tubilak and E. svetlanae as conspecific, and they should now be regarded as E. remanei.</p> </div>	https://treatment.plazi.org/id/03EB87BD8511FFEE8B69DAF7944A6E3B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Grzelak, Katarzyna;Yamasaki, Hiroshi;Mincks, Sarah;Phillips, Anna J.;Sørensen, Martin V.	Grzelak, Katarzyna, Yamasaki, Hiroshi, Mincks, Sarah, Phillips, Anna J., Sørensen, Martin V. (2023): Revision of an Arctic kinorhynch species: Echinoderes svetlanae and E. tubilak are junior synonyms of E. remanei. Zoologischer Anzeiger 302: 75-89, DOI: 10.1016/j.jcz.2022.11.001, URL: http://dx.doi.org/10.1016/j.jcz.2022.11.001
03EB87BD8510FFE88B69D84197F46EAF.text	03EB87BD8510FFE88B69D84197F46EAF.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Echinoderes remanei (Blake 1930)	<div><p>4.2. Diagnostic features of E. remanei</p> <p>Echinoderes remanei belongs to the most common group of Echinoderes regarding its spine pattern. Five middorsal spines and lateroventral tubes/spines on segments 5 to 9 are shared with 46 congeners (Yamasaki et al., 2020). However, when we combine these characters with the presence of glandular cell outlets type 2 in subdorsal, laterodorsal, midlateral/sublateral and ventrolateral positions on segment 2, we shorten the possibilities to 10 species, i.e., Echinoderes angustus Higgins &amp; Kristensen, 1988, Echinoderes aquilonius Higgins &amp; Kristensen, 1988, Echinoderes beringiensis Adrianov &amp; Maiorova, 2022, Echinoderes cernunnos Sørensen et al., 2012, Echinoderes galadrielae Grzelak and Sørensen, 2022, Echinoderes juliae Sørensen et al., 2018, Echinoderes obtuspinosus Sørensen et al., 2012, Echinoderes pennaki Higgins, 1960, Echinoderes romanoi Landers &amp; Sørensen, 2016, and Echinoderes xalkutaat Cepeda et al., 2019 (Higgins 1960; Sørensen et al., 2012; Landers &amp; Sørensen 2016; Grzelak &amp; Sørensen 2018; Herranz et al., 2018; Sørensen et al., 2018; Cepeda et al., 2019; Adrianov &amp; Maiorova 2022; Grzelak &amp; Sørensen 2022). Nevertheless, E. beringiensis, E. cernunnos, E. galadrielae, E. romanoi and E. xalkutaat can immediately be distinguished from E. remanei by the absence of glandular cell outlets type 2 on segment 4. Instead, E. cernunnos shows a pair of glands on segment 7 and very characteristic long and spiniform tergal extensions, while E. romanoi and E. xalkutaat are (in addition to their lack of gco2 on segment 4) much smaller and show different shapes of tergal extensions (Sørensen et al., 2012; Landers &amp; Sørensen 2016; Cepeda et al., 2019). E. juliae and E. obtuspinosus, although possessing a subdorsal pair of glandular cell outlets type 2 on segment 4, have an extra pair of glands on segment 3 (E. juliae) or can be easily differentiated from E. remanei by the presence of very short and stout lateral terminal spines (E. obtuspinosus) (Sørensen et al., 2012; 2018).</p> <p>Of the abovementioned species, E. remanei shows a much closer general resemblance with E. aquilonius, E. angustus and E. pennaki. Interestingly, the first two have been described from Disko Island, western Greenland, along with E. tubilak (Higgins &amp; Kristensen 1988). The latter one, on the other hand, has been described from the San Juan Archipelago in northwest Washington, USA, and belongs to one of the first Echinoderes species described by Higgins (1960). Several years later, the species was redescribed by Higgins (1977) and recently by Herranz et al. (2018). Echinoderes remanei share identical spine patterns, distribution of glandular cell outlets type 2, and even midventral partial articulation of segment 2 (Grzelak &amp; Sørensen 2018; Herranz et al., 2018) with E. aquilonius, E. angustus and E. pennaki; the only exception is the absence of laterodorsal glandular cell outlets type 2 on segment 10 in E. pennaki; the lack of this trait has been confirmed using SEM by Herranz et al. (2018). Therefore, the easiest and most reliable way to Abbreviation s: LA lateral accessory; LD laterodorsal; LV lateroventral; MD middorsal; ML midlateral; PD paradorsal; PV paraventral; SD subdorsal; SL sublateral; VL ventrolateral; VM ventromedial; ac acicular spine; gco1/2 glandular cell outlet type 1/2; ltas lateral terminal accessory spine; lts lateral terminal spine; pe penile spine; pr protuberance; si, sieve plate; ss sensory spot; tu tube; ♀ female condition of sexually dimorphic characters; 6 male condition of sexually dimorphic characters.</p> <p>distinguish E. remanei from its three congeners is by using morphometric differences in combination with a comparison of the pectinate fringe on segments 1 and 2. Unlike E. remanei, none of these species is characterized by the middorsal spine on segment 8 being significantly shorter than the one on segment 7 (Table 2; Fig. 8) (Higgins &amp; Kristensen 1988; Herranz et al., 2018). This feature, which is unusual among Echinoderes, is consistently found in all individuals in E. remanei, irrespective of sampling location. Moreover, E. angustus and E. pennaki, in contrast to E. remanei, do not show significant differences in lateral terminal spine lengths between males and females (Higgins &amp; Kristensen 1988; Herranz et al., 2018). Even if only the length of spines for males would be compared, both species have longer spines in comparison to those of E. remanei (avg. LTSm:&gt; 170 μm vs 149 μm, respectively). Echinoderes aquilonius is, however, similar to E. remanei in sexual dimorphism expressed in longer lateral terminal spines in males than in females (Higgins &amp; Kristensen, 1988; own data), but still both species can be distinguished from one another since the lateral terminal spines in E. aquilonius are shorter than those in E. remanei (avg. LTS ♀: 70 μm vs 90 μm, respectively; avg. LTS 6: 135 μm vs. 149 μm, respectively). In addition, all three species differ from E. remanei by being larger (avg.TL: 405–460 μm vs. 369 μm), and having markedly less conspicuous and more uniform midventral pectinate fringe on segment 1 and 2.</p> <p>It seems that the morphometric pattern for middorsal and terminal spines itself represents a rather unique feature for E. remanei. In species of Echinoderes segment 8 usually has the longest middorsal spine, or is at least similar in length to the spine on segment 7. Therefore, a significantly shorter spine on segment 8 certainly stands out and distinguishes E. remanei among other Echinoderes species. Also, the sexual dimorphism expressed in lateral terminal spine lengths is not a very common trait. According to our knowledge, this trait has only been observed for five other species so far, i.e., E. aquilonius, Echinoderes bengalensis (Timm, 1958), Echinoderes blazeji Grzelak &amp; Sørensen, 2022, Echinoderes coulli Higgins, 1977, and Echinoderes lusitanicus Neves et al., 2016 (Higgins 1977; Higgins &amp; Kristensen 1988; Neves et al., 2016; Grzelak &amp; Sørensen 2022; Sørensen, unpubl. obs). The difference in length due to sexual dimorphism observed in E. remanei is very conspicuous (Table 2; Figs. 2 and 9) and the size ranges between the sexes never overlapped (Fig. 9); males were characterized by considerably longer lateral terminal spines than those in females, and this pattern was very consistent for all localities. Consequently, when all morphometric data were taken into account, regardless of sampling location males were plotted closer together and form a distinct group (Fig. 10, left side of the plot) in comparison to females (Fig. 10, right side of the plot). This result only emphasizes that the specimens, despite the large geographic distance between their locations, are indeed morphologically highly similar, and that differences between the sexes are consistent and driven by the length of lateral terminal spines.</p> </div>	https://treatment.plazi.org/id/03EB87BD8510FFE88B69D84197F46EAF	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Grzelak, Katarzyna;Yamasaki, Hiroshi;Mincks, Sarah;Phillips, Anna J.;Sørensen, Martin V.	Grzelak, Katarzyna, Yamasaki, Hiroshi, Mincks, Sarah, Phillips, Anna J., Sørensen, Martin V. (2023): Revision of an Arctic kinorhynch species: Echinoderes svetlanae and E. tubilak are junior synonyms of E. remanei. Zoologischer Anzeiger 302: 75-89, DOI: 10.1016/j.jcz.2022.11.001, URL: http://dx.doi.org/10.1016/j.jcz.2022.11.001
