taxonID	type	description	language	source
03B487E6FFF5FF92FC8FF9B7F8AEFABF.taxon	diagnosis	Emended diagnosis Mouth cone with 9 individual outer oral styles with broader base and thin and flexible anterior part; 14 trichoscalids each originating from a trichoscalid plate and accompanied anteriorly by two pointed filamentous appendages; 16 placids with broader midventral placid, neighboured by two narrower placids and alternatingly a broader and a narrower placid; all placids with knobby projections (= condyles) in one or two lateral rows in narrower placids and two or three lateral rows in broader placids; midventral placid with two or three apical condyles, zero or two in intermediate lateral row, and 3 ‾ 6 in basal lateral row; ventromedial to midventral free flap and primary pectinate fringe of segment 1 at least partly reduced; acicular spine middorsally on segments 1 ‾ 9, midterminally on segment 11 (= midterminal spine), lateroventrally on segments 2 ‾ 9, lateroventrally or in lateral accessory position on segment 1, laterodorsally on segment 10 in male and in stage- 1 female (in species with more than one adult life history stage), and lateral accessorily on segment 11 (= lateral terminal accessory spine); cuspidate spines present on two or more segments from 1 to 9; type- 3 sensory spot ventrolaterally next to lateral terminal accessory spine and subdorsally or laterodorsally on segment 11; type- 6 sensory spot elevated above trunk surface giving the impression to be half-drowned in the trunk cuticle; ventromedial appendage at least on two segments of segments 5 ‾ 8 in female.	en	Zotto, Matteo Dal, Neuhaus, Birger, Yamasaki, Hiroshi, Todaro, Antonio (2019): The genus Condyloderes (Kinorhyncha: Cyclorhagida) in the Mediterranean Sea, including the description of two new species with novel characters. Zoologischer Anzeiger 282: 206-231, DOI: 10.1016/j.jcz.2019.05.006
03B487E6FFF6FF9CFF88FA88FB7CFEF1.taxon	description	http: // zoobank. org / 8 D 20 EDE 4 - FD 6 E- 4251 - AF 4 F- 539 F 88 DC 1831. (Figs. 2 ‾ 7; Figs. S 1 and S 2; Tables 1 ‾ 3; Tables S 1, S 2).	en	Zotto, Matteo Dal, Neuhaus, Birger, Yamasaki, Hiroshi, Todaro, Antonio (2019): The genus Condyloderes (Kinorhyncha: Cyclorhagida) in the Mediterranean Sea, including the description of two new species with novel characters. Zoologischer Anzeiger 282: 206-231, DOI: 10.1016/j.jcz.2019.05.006
03B487E6FFF6FF9CFF88FA88FB7CFEF1.taxon	diagnosis	3.1.1. Diagnosis Condyloderes with two lateral rows of condyles in broader placids, two apical and three basal condyles in midventral placid, two basal and two apical condyles in remaining broader placids; longitudinal rows of cuticular hairs regularly arranged on trunk segments 1 ‾ 9; acicular spines in lateralaccessory position on segment 1; cuspidate spines in lateral accessory position on segment 2; subdorsal cuspidate spines on segment 3; ventrolateral slightly displaced ventromedially cuspidate spines on segment 5; pairs of paradorsal and sublateral cuspidate spines on segment 7; ventrolateral cuspidate spines on segment 8; lateral accessory cuspidate spines on segment 9; thick and short lateroventral acicular spines on segment 10 in females; ventromedial appendages on segments 6 ‾ 8 in females; laterodorsal acicular spines on segment 10 in males.	en	Zotto, Matteo Dal, Neuhaus, Birger, Yamasaki, Hiroshi, Todaro, Antonio (2019): The genus Condyloderes (Kinorhyncha: Cyclorhagida) in the Mediterranean Sea, including the description of two new species with novel characters. Zoologischer Anzeiger 282: 206-231, DOI: 10.1016/j.jcz.2019.05.006
03B487E6FFF6FF9CFF88FA88FB7CFEF1.taxon	materials_examined	3.1.2. Examined material Holotype: female, mounted as glycerol-paraffin slide on a Cobb aluminium frame, ZMB 11763. Paratypes: 14 females and 31 males, mounted as glycerol-paraffin slide on a Cobb aluminium frame. Additional non-type material consists of five females and 12 males, also mounted for light microscopy or on SEM stubs. Catalogue numbers for types and non-types: ZMB 11764 to 11825. One male non-type mounted for light microscopy, USNM 1484169. Information on localities of the type and non-type material is reported in Table 1. 3.1.3. Type locality Mediterranean Sea, southern Tyrrhenian Sea, Sicily, Gulf of Castellammare, off Castellamare del Golfo, 38 Ǫ 02 Į 59.80 ĮĮ N, 012 Ǫ 52 Į 44.91 ĮĮ E, 43 m depth, silt to very fine sand.	en	Zotto, Matteo Dal, Neuhaus, Birger, Yamasaki, Hiroshi, Todaro, Antonio (2019): The genus Condyloderes (Kinorhyncha: Cyclorhagida) in the Mediterranean Sea, including the description of two new species with novel characters. Zoologischer Anzeiger 282: 206-231, DOI: 10.1016/j.jcz.2019.05.006
03B487E6FFF6FF9CFF88FA88FB7CFEF1.taxon	etymology	3.1.4. Etymology The species is named after the first daughter of the first author, Agnese, and follows the Latinized wording Agnes and its genitive Agnetis.	en	Zotto, Matteo Dal, Neuhaus, Birger, Yamasaki, Hiroshi, Todaro, Antonio (2019): The genus Condyloderes (Kinorhyncha: Cyclorhagida) in the Mediterranean Sea, including the description of two new species with novel characters. Zoologischer Anzeiger 282: 206-231, DOI: 10.1016/j.jcz.2019.05.006
03B487E6FFF6FF9CFF88FA88FB7CFEF1.taxon	description	3.1.5. Description The description refers to females. See chapter “ 3.1.6. Sexual dimorphism ” for male characters which are different from female ones. Figures and tables show both male and female characters (see figure and table legends). Adult specimens consist of head, neck, and 11 trunk segments (Figs. 2, 3 A — B, 4 A, 6 A, Band 7 A; Figs. S 1 A and B). See Table 2 for a summary of spine and sensory spot positions, and Table 3 for measurements and dimensions. Head. The head is made up of a retractable mouth cone and an introvert (Figs. 4 E, 5 and 6 C ‾ E; Figs. S 1 A and B). The pharyngeal crown consists of 10 lobular extensions at the anterior end of the pharynx and becomes visible in specimens with extremely protruded mouth cone (Fig. 6 D). The mouth cone is characterized by the presence of a mouth cone weir (Fig. 6 E). At least two rings with five inner oral styles in each were detected (rings — 02 and — 01; Fig. 6 D, E). Ring 00 is characterized by nine thin outer oral styles, consisting of a single element with a thin distal part (Fig. 6 E). The introvert has six rings of scalids and one ring of trichoscalids. Ring 01 bears 10 primary spinoscalids consisting of a sheath-like basis and an elongated distal part. The number of visible scalids in ring 06 is 14, no ring 06 scalids were observed in sectors 5 and 7 (Fig. 5). The posterior part of the introvert is characterized by 14 elongated and fringed trichoscalids. Apair of filamentous appendages is attached anteriorly to each trichoscalid. These structures are thinner than the scalids and do not show external sculpturing (Fig. 6 C). Neck. The neck consists of 16 placids, having irregular, knobby surfaces (condyles), varying in number between narrower and broader placids (Figs. 3 C, 4 F, 6 F and 7 B). Abroader midventral placid is neighboured by two narrower placids, and from thereon a broader and a narrower placid alternate. The midventral placid shows two condyles in the apical lateral row and three condyles in the basal lateral row. The remaining broader placids have two basal and two apical condyles. The narrower placids have one condyle in the basal lateral row, which splits apically into two condyles, whereas the paraventral narrower placids reveal only a single condyle (TableS 2). Trunk. The trunk is divided into 11 segments. It appears triangular in cross-section, with the tergosternal junctions located along the lateroventral angles. Segment 1 is formed by a closed cuticular ring and shows a deep broad and squared midventral indentation at the posterior edge, because the free flap is missing (Figs. 2 A and 7 B). This segment is characterized by the presence of a middorsal hirsute spine and two acicular spines placed in lateral accessory position (Figs. 4 B and 7 A; Fig. S 1 C). The middorsal spine from this segment through segment 10 is located posteriorly on its segment and originates from a sclerotized anterior keel on a trunk segment; the free flap is missing where a spine inserts (Fig. 3 D). Minute cuticular hairs are located along the anterior part of the segment. Paired sensory spots are located anteriorly in subdorsal, laterodorsal, midlateral, sublateral, and ventromedial position. These sensoryspots are type- 6, hence consisting of a structure emerging from the cuticle and giving the impression of being half-drowned in it (Fig. 6 H). In rare cases a cilium, ca. 2 M m long, juts outof the central pore of this sensory spot (Fig. 6 G). Similarly to the following segments, the posterior margin of the segment is straight, and shows a free flap overlapping the subsequent segment and terminating in a primary pectinate fringe (Fig. 4 D). The free flap is interrupted where the spines originate and ventrolaterally to midventrally. This condition is repeated through segments 2 to 11 except for the ventral lack of the free flap, even if the teeth of the primary pectinate fringe are shorter and smaller on segment 10 and, particularly, on segment 11 (Figs. 6 I and 7 B ‾ C, F — G). Elongated cuticular hairs (Figs. 6 J and 7 H) are arranged in longitudinal rows of one to three each originating from the middle part of the segment and continuing beyond the segment's posterior edge (Figs. 6 C and 7 B). Within each row the hairs are arranged one after the other from anterior to posterior. The rows are approximately 16 on the ventral side and 40 on the dorsal side, and are absent midventrally and paraventrally, and partially in a midlateral position. Other irregularly arranged cuticular hairs are present midventrally and ventrolaterally from anterior to posterior. Segment 2 consists of a tergal and two sternal plates (Figs. 3 D and 7 A). The same condition is repeated through segments 3 to 10. This segment bears a middorsal spine, two lateroventral acicular spines, and two cuspidate spines in lateral accessory position next to the edge of the sublateral position (Figs. 4 B and 7 A, B, C; Fig. S 1 C). Pairs of type- 6 sensory spots are located posteriorly in subdorsal, laterodorsal, midlateral, sublateral and ventromedial position (Figs. 3 D, 4 B and 7 B, C; Fig. S 1 C). Similarly to segment 1, long cuticular hairs arranged in longitudinal rows of three to five each originate from the middle part of the segment and continue beyond the segment's posterior edge. Approximately 9 ‾ 12 rows are present on each sternal plate and 38 ‾ 42 rows on the tergal plate. These rows are absent close to the midsternal junction and also in part of the midlateral area. The same condition is repeated through segments 3 to 9 (Figs. 3 E, 4 D, 6 A — B, Iand 7 A — F). Segments 10 and 11 do not bear the rows of cuticular hairs (Figs. 4 D, 6 I and 7 G). Segment 3 has a middorsal spine, two subdorsal cuspidate spines (Figs. 3 D, 6 A — B and 7 C — E) and two lateroventral acicular spines. Three pairs of type- 6 sensory spots are placed laterodorsally, sublaterally, and ventromedially (Figs. 4 B and 7 C; Fig. S 1 C). Segment 4 bears one middorsal spine and two lateroventral acicular spines. Paired type- 6 sensory spots are present paradorsally, a pair of type- 6 sensory spots is located laterodorsally and another pair ventromedially (Figs. 4 B and 7 C; Fig. S 1 C). Ovaries almost reaching the junction between this segment and segment 3 (Fig. 3 H). Segment 5 shows one middorsal spine, two lateroventral acicular spines, and two ventrolateral cuspidate spines slightly displaced ventromedially (Figs. 2 A and 7 A; Fig. S 1 D). The cuspidate spines are located next to the edge of the ventromedial position (Figs. 4 C, 6 B and 7 A, F). Pairs of type- 6 sensory spots are located laterodorsally and sublaterally. No ventral sensory spots were detected. Segment 6 bears one middorsal spine and two lateroventral acicular spines (Fig. 4 C). Pairs of type- 6 sensoryspots are located paradorsally, laterodorsally, sublaterally, and ventromedially (Fig. 7 F, I; Fig. S 1 D). Females bear ventromedial appendages on this segment (Figs. 4 C and 7 A, F, H). Segment 7 has one middorsal spine, two pairs of cuspidate spines located paradorsally and sublaterally (Figs. 3 E, 6 A and 7 D, E, I; Fig. S 1 G), and two lateroventral acicular spines. Apair of type- 6 sensory spots is located laterodorsally and a second pair ventromedially (Figs. 4 C, D, and 7 F, I; Fig. S 1 D). Females bear ventromedial appendages (Figs. 2 and 7 F, H). Segment 8 bears one middorsal spine, two lateroventral acicular, and two ventrolateral cuspidate spines (Figs. 3 E, F, 4 D, 6 A, and 7 A, D; Fig. S 1 A). Pairs of type- 6 sensory spots are placed paradorsally, laterodorsally, and ventromedially (Figs. 4 D and 7 F; Fig. S 1 D. Females bear ventromedial appendages (Figs. 2 A and 7 F). Segment 9 has one middorsal spine, two lateroventral acicular spines, and two cuspidate spines in lateral accessory position (Figs. 3 E ‾ G, 4 D, 6 A, I, and 7 A; Fig. S 1 E). Pairs of type- 6 sensoryspots are located paradorsally, laterodorsally, and ventromedially (Figs. 4 D, 6 I and 7 G; Fig. S 1 E). Astructure very likely representing the protonephridial opening, marked by minute cuticular papillae, is present in sublateral position (Fig. 6 I ‾ J). Ventromedial areas of micropapillae are absent. Segment 10 is characterized by the presence of a middorsal spine (Figs. 3 B, E, 6 B, and 7 G). Apair of paradorsal type- 6 sensory spot is present (Fig. 6 I). The teeth of the primary pectinate fringe are shorter and smaller than those on segments 1 ‾ 9. Females bear two thick and short lateroventral acicular spines (Figs. 3 A, G, 6 I, and 7 G; Fig. S 1 A). Short cuticular hairs are arranged irregularly especially on the sternal plates (Figs. 6 I and 7 G). Segment 11 is formed by one tergal and one sternal plate. The latter seems to show a midventral to paraventral cuticular thickening, appearing optically separated from the remaining plate because of a paraventral fold in the cuticle (Figs. 4 H and 7 G). This segment bears a relatively short midterminal spine and two elongated lateral terminal accessory spines each with two thin areas in the basal part (Figs. 3 A, 4 H and 6 A). Two pairs of type- 3 sensory spots, each composed of a conical base and terminal cuticular papillae, are placed laterodorsally and ventrolaterally (Fig. 7 G, J). The former are located more centrally on the segment, while the latter are placed posteriorly. Two ventromedial sensory spots are placed anteriorly on the segment, partially hidden by the free flap of segment 10. We could not assign these sensoryspots to any specific type of sensory spot, even though they resemble type- 6. Similar structures have been observed in Condyloderes shirleyi Neuhaus & Higgins, 2019 in Neuhausetal. (2019; fig. 42 D, F). The teeth of the primary pectinate fringe are much shorter and smaller than on segments 1 ‾ 9. Females show cuticularized gonopores at the anterior margin of the sternal plate, almost at the junction between segments 10 and 11 (Figs. 3 A, G, and 7 G). It was noted that a single female specimen had an indistinct mass of material which may be interpreted as a spermatophore attached to the terminal part of its trunk (Fig. S 1 A). This would be the second report of potential spermatophore in a cyclorhagid, after the finding in Centroderes drakei Neuhaus, Pardos, SØrensen & Higgins, 2014 (see Neuhaus et al. 2014). The spermatophore was present in both females and male in the species of this closely related genus. 3.1.6. Sexual dimorphism Males possess a pair of laterodorsal acicular spines on segment 10, extending beyond the posterior edge of segment 10 (Figs. 4 G, 6 A — B, and 7 J; Fig. S 1 F). The acicular spines on segment 10 are thinner than the acicular spines on other segments (Fig. 4 G, Fig. S 1 F). In addition, males lack gonopores on segment 11, lateroventral acicular spines on segment 10, and ventromedial appendages on segments 6, 7, and 8 (Fig. 4 C, D), all of which are present only in females (see chapter 3.1.5.). 3.1.7. Variation Out of 46 specimens mounted for light microscopy and in a condition to check variation of trunk characters, variation is observed in 22 specimens (= 48 %) (Fig. 2; Figs. S 2 A ‾ I; Table S 1). Ten specimens vary in one character, seven in two characters, three in three characters, and one each in five and seven characters. Most variation results from the absence of a cuspidate spine paradorsally on one or both sides of segment 7 and of a type- 6 sensory spot sublaterally on segment 5 (Fig. 7 E, I; Figs. S 2 A, D, E; Table 2; Table S 1). In two specimens, the paradorsal cuspidate spine is missing on one side of segment 7 and replaced by a sensory spot. One specimen reveals a very short middorsal and laterodorsal spine on segment 10 (Fig. S 2 H). Another specimen possesses a crippled lateral terminal accessory spine on one side (Fig. S 2 I). One specimen shows an acicular spine lateroventrally on the right side of segment 5 but a cuspidate spine on the left side of this segment; the ventrolateral cuspidate spine of the left side is missing (Fig. S 2 C). Three specimens lack a laterodorsal sensory spot on one side of segments 6 or 9 and possess an additional spot paradorsally on the same side of these segments (Figs. S 2 B and E; Table S 1). A type- 6 sensory spot may occur in an unusual position on one side only, e. g., lateroventrally on segment 1 (Fig. S 2 F). Little variation has been found in the arrangement of placids. One specimen shows two narrower placids instead of one broader placid next to the middorsal placid (Fig. S 2 G).	en	Zotto, Matteo Dal, Neuhaus, Birger, Yamasaki, Hiroshi, Todaro, Antonio (2019): The genus Condyloderes (Kinorhyncha: Cyclorhagida) in the Mediterranean Sea, including the description of two new species with novel characters. Zoologischer Anzeiger 282: 206-231, DOI: 10.1016/j.jcz.2019.05.006
03B487E6FFF6FF9CFF88FA88FB7CFEF1.taxon	biology_ecology	3.1.8. Ecology Beyond the type locality of C. agnetis sp. nov., the species was found also at four more sites off Castellammare del Golfo and off the town of Trappeto, a second location placed over 12 km to the east, within the Gulf of Castellammare, Sicily, southern Tyrrhenian Sea (see chapter 3.4.8. and Table 1). The species was collected from both the two areas (off Castellammare and off Trappeto) during four different surveys led from June 2006 to December 2007. The bottom depths of the sampling sites ranged from 32 to 50 m. Sediment was made up on average of silt to very fine sand off Castellammare del Golfo, and very fine sand to coarse silt off Trappeto. C. agnetis sp. nov. co-occurred with other kinorhynchs, i. e.: Paracentrophyes quadridentatus (Zelinka, 1928), Cristaphyes carinatus (Zelinka, 1928), Pycnophyes communis Zelinka, 1908, Pycnophyes giganteus (Zelinka, 1928), Pycnophyes robustus Zelinka, 1928, Semnoderes armiger Zelinka, 1928, Condyloderes multispinosus (off Trappeto only; see chapter 3.4.8), Echinoderes capitatus (Zelinka, 1928), Echinoderes ferrugineus Zelinka, 1928, and Echinoderes gerardi Higgins, 1978 (Dal Zotto & Todaro 2016). The other meiobenthic taxa associated with C. agnetis sp. nov. were mainly nematodes, harpacticoids, polychaetes, turbellarians, tanaidaceans, ostracods, amphipods, cumaceans, tardigrades, gastropods, bivalves, and cnidarians (Dal Zotto et al. 2016). The densities of C. agnetis sp. nov. ranged from 1 to 6 individuals / 10 cm 2. Comparedtothe otherassociatedkinorhynch species it was relatively scarce, but it was also rather common, because it was found at seven out of eight investigated sites off Castellammare del Golfo (Dal Zotto et al. 2016). Any clear correlations among the abundances of C. agnetis sp. nov. and other co-occurring kinorhynch species were found, with Pearson correlation coefficients varying from — 0.18 (C. agnetis sp. nov. ‾ S. armiger) to 0.44 (C. agnetis sp. nov. ‾ Pycnophyes spp. juveniles). One male of C. agnetis sp. nov. from a sample collected at 42 m depth off Rovinj (Croatia, northern Adriatic Sea) was found among the specimens of Condyloderes on loan from the National Museum of Natural History, Smithsonian Institution, Washington D. C. (see Table 1; Figs. S 1 B ‾ G).	en	Zotto, Matteo Dal, Neuhaus, Birger, Yamasaki, Hiroshi, Todaro, Antonio (2019): The genus Condyloderes (Kinorhyncha: Cyclorhagida) in the Mediterranean Sea, including the description of two new species with novel characters. Zoologischer Anzeiger 282: 206-231, DOI: 10.1016/j.jcz.2019.05.006
03B487E6FFF6FF9CFF88FA88FB7CFEF1.taxon	discussion	3.1.9. Note on epizoic protozoa Six epizoic protozoans attached on segments 1 and 7 ‾ 9 to a male specimen from off Rovinj, northern Adriatic Sea (Figs. S 1 B and G).	en	Zotto, Matteo Dal, Neuhaus, Birger, Yamasaki, Hiroshi, Todaro, Antonio (2019): The genus Condyloderes (Kinorhyncha: Cyclorhagida) in the Mediterranean Sea, including the description of two new species with novel characters. Zoologischer Anzeiger 282: 206-231, DOI: 10.1016/j.jcz.2019.05.006
03B487E6FFF8FF83FCF5FE5CFC7DFE15.taxon	description	http: // zoobank. org / A 8 E 22442 - 31 E 0 - 4 A 5 C-A 9 DD- CA 1 E 62880 B 56. (Figs. 5, 8 ‾ 12; Tables 1, 2, 4; TableS 2).	en	Zotto, Matteo Dal, Neuhaus, Birger, Yamasaki, Hiroshi, Todaro, Antonio (2019): The genus Condyloderes (Kinorhyncha: Cyclorhagida) in the Mediterranean Sea, including the description of two new species with novel characters. Zoologischer Anzeiger 282: 206-231, DOI: 10.1016/j.jcz.2019.05.006
03B487E6FFF8FF83FCF5FE5CFC7DFE15.taxon	diagnosis	3.2.1. Diagnosis Condyloderes with two lateral rows of condyles in broader placids, two apical and three basal condyles in midventral placid, two apical and two large basal condyles in remaining broader placids, and two basal condyles in narrower placids, apart from the two neighbouring the midventral placid with only one basal condyle; longitudinal rows of cuticular hairs regularly arranged on trunk segments 1 ‾ 10, fewer and with shorter hairs on segment 10; acicular spines lateral accessorily on segments 1 and 11, lateroventrally on segments 2 ‾ 9 (on segment 10 in females only), laterodorsally on segment 10 in males only; cuspidate spines ventrolaterally slightly displaced ventromedially on segment 5, ventrolaterally on segment 8, and lateral accessorily on segment 9; ventromedial appendages on segments 5 ‾ 7 in females only; type- 6 sensory spots paradorsally on segments 1 and 7, sublaterally on segments 3 and 6, and ventromedially on segment 8.	en	Zotto, Matteo Dal, Neuhaus, Birger, Yamasaki, Hiroshi, Todaro, Antonio (2019): The genus Condyloderes (Kinorhyncha: Cyclorhagida) in the Mediterranean Sea, including the description of two new species with novel characters. Zoologischer Anzeiger 282: 206-231, DOI: 10.1016/j.jcz.2019.05.006
03B487E6FFF8FF83FCF5FE5CFC7DFE15.taxon	materials_examined	3.2.2. Examined material Holotype: female, mounted as glycerol-paraffin slide on a Cobb aluminium frame, ZMB 11826. Paratypes: two males, mounted as glycerol-paraffin slide on a Cobb aluminium frame, ZMB 11827 and ZMB 11828 mounted as glycerol-paraffin slide on a Cobb aluminium frame. Non-type: one juvenile ZMB 11829, mounted as glycerol-paraffin slide on a Cobb aluminium frame. One non-type male ZMB 11830 mounted on aluminium stub for scanning electron microscopy. Information on localities of the type and non-type material is reported in Table 1. 3.2.3. Type locality Mediterranean Sea, Ligurian Sea, Tuscany, off Livorno (Leghorn), 43 Ǫ 37 Į 35 ĮĮ N, 009 Ǫ 59 Į 18 ĮĮ E, 112 m depth, sandy mud.	en	Zotto, Matteo Dal, Neuhaus, Birger, Yamasaki, Hiroshi, Todaro, Antonio (2019): The genus Condyloderes (Kinorhyncha: Cyclorhagida) in the Mediterranean Sea, including the description of two new species with novel characters. Zoologischer Anzeiger 282: 206-231, DOI: 10.1016/j.jcz.2019.05.006
03B487E6FFF8FF83FCF5FE5CFC7DFE15.taxon	etymology	3.2.4. Etymology The species is named after the second daughter of the first author, Chiara, and follows the Latinized wording Clara and its genitive Clarae.	en	Zotto, Matteo Dal, Neuhaus, Birger, Yamasaki, Hiroshi, Todaro, Antonio (2019): The genus Condyloderes (Kinorhyncha: Cyclorhagida) in the Mediterranean Sea, including the description of two new species with novel characters. Zoologischer Anzeiger 282: 206-231, DOI: 10.1016/j.jcz.2019.05.006
03B487E6FFF8FF83FCF5FE5CFC7DFE15.taxon	description	3.2.5. Description The description refers to female. No variation was observed in the examined material except for the midventral placid (see below). See chapter “ 3.2.6. Sexual dimorphism ” for male characters that differ from female ones. Figures and tables show both male and female characters (see figure and table legends). The analysed adult specimen consists of head, neck, and 11 trunk segments (Figs. 8, 9 A, B, 10 A, and 11 A). See Table 2 for a summary of spine and sensory spot position, and Table 4 for measurements and dimensions. Head. The head is made up of a retractable mouth cone and an introvert (Fig. 10 B). The introvert has six rings of scalids and one ring of trichoscalids. Ring 01 bears 10 primary spinoscalids, consisting of a sheath-like basis and an elongated distal part. Rings 02 to 05 bear 5, 15, 15, and 15 scalids, respectively. The number of scalids in ring 06 is more than 12, even though the exact number could not be detected because those in sectors 3, 5, 7, and 9 could not be observed. Generally, the scalid arrangement appeared very similar and is most likely identical to C. agnetis sp. nov. (Fig. 5). The posterior part of the introvert is characterized by 14 elongated and fringed trichoscalids (Figs. 11 C and 12 F). Apair of filamentous appendages is attached anteriorly to each trichoscalid. These structures are thinner than the scalids and do not show external sculpturing (Fig. 10 B). Neck. The neck consists of 16 placids, having irregular, knobby surfaces (condyles), varying in number between narrower and broader placids (Figs. 9 C, 10 D, and 11 C). Abroader midventral placid is neighboured by two narrower placids, and from thereon a broader and a narrower placid alternate. The midventral placid shows 5 ‾ 6 condyles arranged in an apical lateral row with two condyles and a basal lateral row with three condyles in the female holotype and in the male allotype (Fig. 10 D), whereas the male mounted for SEM reveals four basal condyles (Fig. 11 C). The remaining broader placids have two apical and two basal condyles; the basal condyles are very large and seem to possess apically three smaller condyles (Fig. 11 C). The narrow placids have two basal condyles, apart from the two closest to the midventral placid, which bear only one basal condyle (Fig. 11 C; TableS 2). Trunk. The trunk is divided into 11 segments. It appears triangular in cross-section, with the tergosternal junctions located along two of the lateroventral angles. Segment 1 is formed by a closed cuticular ring and shows a broad and squared midventral indentation, because the free flap is missing (Figs. 8 A, 9 D, 10 D, and 11 C). This segment is characterized by a middorsal hirsute spine and two acicular spines placed in lateral accessory position (Fig. 9 D). The middorsal spine from this segment through segment 10 is located posteriorly on a segment and originates from a sclerotized anterior keel on a trunk segment; the free flap is missing where a spine inserts (Fig. 9 B). Minute cuticular hairs are located along the anterior part of the segment. Paired type- 6 sensory spots are located anteriorly in subdorsal, laterodorsal, midlateral, sublateral, and ventromedial position (Figs. 9 C, D, 10 D, 11 C, and 12 D). Similar to the following segments, the posterior margin of the segment is straight, and shows a free flap overlapping the subsequent segment and terminating in a primary pectinate fringe (Figs. 11 D and 12 C). The free flap is interrupted where the spines originate and ventrolaterally to midventrally. This condition is repeated through segments 2 to 11 except for the ventral lack of the free flap (Figs. 11 D and 12 C), even if the teeth of the primary pectinate fringe are shorter and smaller on segments 10 and 11. Very long cuticular hairs are arranged in longitudinal rows of three to five each originating from the anterior to the central part of the segment and continuing beyond the segment's posterior edge on the dorsal side (Fig. 9 F). There are approximately 18 ‾ 20 rows. In addition, many irregularly arranged cuticular hairs are present midventrally and ventrolaterally from anterior to posterior. Segment 2 consists of a tergal and two sternal plates. The same condition is repeated through segments 3 to 10. The segment bears a middorsal spine and two lateroventral acicular spines. Pairs of type- 6 sensoryspots are located in subdorsal, laterodorsal, midlateral, sublateral, and ventromedial position (Figs. 9 D, 11 C, Dand 12 D). The subdorsal and sublateral sensoryspots are placed posteriorly, whereas the laterodorsal, midlateral and ventromedial are located more medially on the segment (Fig. 11 D). Long cuticular hairs arranged in longitudinal rows of normally six each originate from the middle part of the segment and continue beyond the segment's posterior edge. There are approximately 6 ‾ 7 rows on each sternal plate and 24 ‾ 26 on the tergal plate. These rows are absent midventrally, paraventrally and midlaterally (Figs. 9 D ‾ F, 10 E, 11 A — B, D — E, and 12 A — B, D — E). Segment 3 bears a middorsal spine and two lateroventral acicular spines. Two pairs of type- 6 sensory spots are placed posteriorly in subdorsal and sublateral position (Fig. 12 D, E). No ventral sensory spot was detected (Fig. 11 D). Segment 4 is characterized by one middorsal spine and two lateroventral acicular spines. Pairs of type- 6 sensory spots are placed paradorsally, laterodorsally, and ventromedially (Figs. 10 E, 11 B, D, and 12 E). Segment 5 shows one middorsal spine, two lateroventral acicular spines, and two ventrolateral cuspidate spines slightly displaced ventromedially (Figs. 8 A, 9 D, 10 E and 11 B). The cuspidate spines are located next to the edge of the ventromedial position (Figs. 9 A, Dand 10 E). Asingle pair of type- 6 sensory spots is located laterodorsally (Fig. 12 E). No ventral sensory spot was detected. The female bears ventromedial appendages on this segment (Fig. 9 D). Segment 6 bears one middorsal and two lateroventral acicular spines. Pairs of type- 6 sensory spots are located paradorsally, midlaterally, and ventromedially (Figs. 8 A, 11 B, and 12 A). The female bears ventromedial appendages on this segment (Fig. 9 D). Segment 7 has one middorsal spine and two lateroventral acicular spines. Apair of type- 6 sensory spots is located laterodorsally and a second pair of this same type of sensory spots is placed ventromedially (Figs. 11 B and 12 A). The female bears ventromedial appendages on this segment (Fig. 9 D). Segment 8 bears one middorsal spine, two lateroventral acicular, and two ventrolateral cuspidate spines (Fig. 12 A). These latter spines are noticeably longer than other cuspidate spines (Figs. 9 B, 10 C, and 11 A — B; Table 5). Pairs of type- 6 sensory spots are located in paradorsal, laterodorsal, and ventromedial position (Figs. 11 B and 12 A). Segment 9 has one middorsal spine, two lateroventral acicular spines, and two lateral accessory cuspidate spines (Figs. 9 B, 10 C and 11 A, E, 12 B, C). Pairs of type- 6 sensory spots are placed paradorsally, laterodorsally, and ventromedially (Figs. 11 E and 12 B, C). Astructure with cuticular papillae is present in sublateral position. We assume that this character represent the protonephridial opening (Fig. 12 B ‾ C). Ventromedial areas of micropapillae are absent. Segment 10 is characterized by a rather long middorsal spine (Figs. 9 B and 12 B). Apair of paradorsal type- 6 sensoryspots is present. The teeth of the primary pectinate fringe are shorter and smaller than on segments 1 to 9 (Fig. 11 E). Cuticular hairs shorter, and arranged in fewer rows. The female bears two lateroventral acicular spines on this segment (Fig. 9 B). Segment 11 is formed by one tergal and one sternal plate. The latter shows a midventral to paraventral cuticular thickening, appearing optically separated from the remaining plate because of a paraventral fold in the cuticle (Figs. 9 E and 11 E). This segment bears a midterminal spine and two elongated lateral terminal accessory spines each with two thin areas in the basal parts (Figs. 9 A, B, 10 A, 11 A, E, and 12 B). Two type- 3 sensoryspots, composed of a conical base and terminal cuticular papillae, are placed centrally on the segment in laterodorsal position. Another pair of type- 3 sensory spots is placed posteriorly in ventrolateral position (Figs. 10 C, G, 11 E, and 12 B). Two ventromedial sensory spots are placed anteriorly on the segment, partially hidden by the free flap of segment 10. We could not assign these structures to any specific type of sensory spot, even though they resemble type- 6. On this segment, the teeth of the primary pectinate fringe are shorter and smaller than on segments 1 to 10, and cuticular hairs are absent. The female shows cuticularized gonopores at the anterior margin of the sternal plate, almost at the junction between segments 10 and 11 (Fig. 9 E). 3.2.6. Sexual dimorphism The two males bear a pair of laterodorsal acicular spines on segment 10, ending after the posterior edge of the segment (Figs. 10 F and 12 B), and a ventromedial gland cell outlet on segment 10. In addition, the males lack gonopores on segment 11, lateroventral acicular spines on segment 10, and ventromedial appendages on segments 5, 6, and 7 (Fig. 10 C, E), all of which are present only in the female (see chapter 3.2.5.). 3.2.7. Ecology C. clarae sp. nov. was found at different sites off Livorno (Ligurian Sea, Tuscany, Central Italy). The bottom depth of the sampling sites ranged from 111 to 113 m. The sediment was made up on average of sandy mud or loamy sand (see Table 1). C. clarae sp. nov. co-occurred with other kinorhynchs, i. e.: P. quadridentatus, Fujuriphyes cf. rugosus (Zelinka 1928), P. communis, P. giganteus, S. armiger, Echinoderes cf. capitatus, Echinoderes sp. 2, and Echinoderes sp. 3 (see Dal Zotto & Todaro, 2016). The other meiobenthic taxa associated with C. clarae sp. nov. were mainly nematodes, polychaetes, harpacticoids, tanaidaceans, ostracods, amphipods, cumaceans, gastropods, bivalves, priapulids, and loriciferans. The densities of C. clarae sp. nov. were of 1 ‾ 2 individuals / 10 cm 2. Compared to theother associated kinorhynchspecies it was rare, found at four out of twelve investigated sites, and scarce.	en	Zotto, Matteo Dal, Neuhaus, Birger, Yamasaki, Hiroshi, Todaro, Antonio (2019): The genus Condyloderes (Kinorhyncha: Cyclorhagida) in the Mediterranean Sea, including the description of two new species with novel characters. Zoologischer Anzeiger 282: 206-231, DOI: 10.1016/j.jcz.2019.05.006
03B487E6FFF8FF83FCF5FE5CFC7DFE15.taxon	diagnosis	3.3. Differential diagnosis Six species of Condyloderes have been described to date, i. e., C. multispinosus, C. paradoxus Higgins, 1969, Condyloderes setoensis Adrianov, Murakami & Shirayama, 2002, C. storchi Higgins, 2004 in Martorelli & Higgins, 2004, Condyloderes kurilensis Adrianov & Maiorova, 2016, and C. shirleyi. Two additional species are described from off California and the Gulf of Mexico by SØrensen etal. (2019). C. agnetis sp. nov. and C. clarae sp. nov. can be distinguished from their congeners bythe presence of (1) longitudinal rows of cuticular hairs on segments 1 ‾ 9 and (2) a lateral accessory acicular spine on segment 1 (Table S 2; Neuhaus et al. 2019; this paper). C. agnetis sp. nov. can be differentiated from all its congeners by (1) a subdorsal cuspidate spine on segment 3, (2) a paradorsal cuspidate spine on segment 7, (3) a sublateral cuspidate spine on segment 7, (4) the lack of a paradorsal type- 6 sensory spot on segment 5, and (5) a very short midterminal spine in comparison to the lateral terminal accessory spine (Table S 2; Neuhaus et al. 2019; this paper). C. clarae sp. nov. can be identified from all its congeners by (1) the lack of a paradorsal type- 6 sensory spot on segments 1 and 4, (2) the lack of a sublateral type- 6 sensory spot on segments 3 and 6, (3) the lack of a ventromedial type- 6 sensory spot on segment 8, (4) the existence of a ventromedial female-specific appendage on segment 5, and (5) the lack of a ventromedial female-specific appendage on segment 8 (Table S 2; Neuhaus et al. 2019; this paper). C. agnetis sp. nov. can be distinguished from C. clarae sp. nov. by (1) a larger trunk length (302 ‾ 327 M m versus 215 ‾ 229 M m length), (2) a shorter midterminal spine (18 ‾ 30 M m versus ca. 55 M m length; see Tables 4 and 5 for average values, MTS / TL, and MTS / LTAS of both species), (3) the existence of a lateral accessory cuspidate spine on segment 2 versus its lack, (4) the existence of a subdorsal cuspidate spine on segment 3 versus its lack, (5) the existence of paradorsal and sublateral cuspidate spines on segment 7 (but spines may be missing naturally) in C. agnetis sp. nov. versus their lack in C. clarae sp. nov., (6) the existence of a paradorsal type- 6 sensory spot on segments 1 and 4 versus its lack on these segments, (7) the existence of a ventromedial type- 6 sensory spot on segments 3, 8, and 11 versus its lack on these segments, (8) the existence of a sublateral type- 6 sensory spot on segments 3 and 6 versus its lack on these segments, (9) the existence of a laterodorsal type- 6 sensory spot on segments 3 (but may be missing naturally), 6, and 11 versus its lack on these segments, and (10) the existence of a ventromedial female-specific appendage on segments 6 ‾ 8 versus its existence on segments 5 ‾ 7 (Table S 2). Additional, though weaker, traits distinguishing C. agnetis sp. nov. from C. clarae sp. nov. are (1) shorter and less robust cuticular hairs as well as fewer hairs in each longitudinal row (3 ‾ 5 versus 6 hairs per row) and more rows of hairs both on the sternal plates (9 versus 6 ‾ 7 rows) and tergal plates (40 ‾ 44 versus 24 ‾ 26 rows), (2) the lack of rows of cuticular hairs on segment 10 versus their presence in C. clarae sp. nov.	en	Zotto, Matteo Dal, Neuhaus, Birger, Yamasaki, Hiroshi, Todaro, Antonio (2019): The genus Condyloderes (Kinorhyncha: Cyclorhagida) in the Mediterranean Sea, including the description of two new species with novel characters. Zoologischer Anzeiger 282: 206-231, DOI: 10.1016/j.jcz.2019.05.006
03B487E6FFE7FF86FCEFFE60FA55F9A6.taxon	diagnosis	3.4.1. Emended diagnosis Neck placids with condyles in one lateral row in narrower placids and three lateral rows in broader placids; each narrower placid with one basal condyle; each broader placid with two condyles both in apical, intermediate, and basal lateral row; midventral placid with three apical and three basal condyles and two in intermediate lateral row; acicular spine middorsally on segments 1 ‾ 9 (on segment 10 in male only), midterminally on segment 11, laterodorsally on segment 10 in male only, lateroventrally on segments 1 ‾ 9, and laterodorsally on segment 10; cuspidate spine ventrolaterally on segments 2, 5, and 9 and lateral accessorily on segment 8, sometimes also middorsally on segments 5 and / or 7; type- 6 sensory spot ventromedially on segments 1 (slightly more paraventrally), 2 ‾ 4 and 6 ‾ 9, lateroventrally on segment 10, sublaterally on segments 1 ‾ 3 and 5 ‾ 9, midlaterally on segments 1 and 2, laterodorsally on segments 1 ‾ 9, subdorsally on segment 10, and paradorsally on segments 1 (almost subdorsally), 2, and 4 ‾ 9; gland cell outlet ventromedially on segment 10; ventromedial appendage on segments 7 and 8 and area of micropapillae ventromedially on segment 9 in female only.	en	Zotto, Matteo Dal, Neuhaus, Birger, Yamasaki, Hiroshi, Todaro, Antonio (2019): The genus Condyloderes (Kinorhyncha: Cyclorhagida) in the Mediterranean Sea, including the description of two new species with novel characters. Zoologischer Anzeiger 282: 206-231, DOI: 10.1016/j.jcz.2019.05.006
03B487E6FFE7FF86FCEFFE60FA55F9A6.taxon	materials_examined	3.4.2. Examined material One male (USNM 1562541), two females (USNM 1562542, USNM 1562543), and one juvenile (USNM 1562544), from off Banyuls-sur-Mer, Gulf of Lion. One female from off Rovinj, northern Adriatic Sea (USNM 1562540). All specimens mounted for light microscopy (Figs. 13 and 14; Fig. S 3; Table 5). One male from off Trappeto, Gulf of Castellammare, Sicily, southern Tyrrhenian Sea, lost during observation; thus only few photographs were available for this specimen.	en	Zotto, Matteo Dal, Neuhaus, Birger, Yamasaki, Hiroshi, Todaro, Antonio (2019): The genus Condyloderes (Kinorhyncha: Cyclorhagida) in the Mediterranean Sea, including the description of two new species with novel characters. Zoologischer Anzeiger 282: 206-231, DOI: 10.1016/j.jcz.2019.05.006
03B487E6FFE7FF86FCEFFE60FA55F9A6.taxon	description	3.4.3. Brief description The morphology of the specimens from the Mediterranean Sea agrees widely with that of the recently re-described specimens of C. multispinosus from the North Sea, so only differences are mentioned for the Mediterranean specimens here. For illustrations, data and morphological measurements see Figs. 13, 14, Fig. S 3, Tables 2 and 5, and Table S 2. Our findings report for the first time a female adult stage for C. multispinosus. The female exhibits a ventromedial appendage on each sternal plate of segments 7 and 8, which gives the impression of an extremely elongated sensory spot covered by numerous cuticular micropapillae (Fig. 13 E). At its base, each structure shows a sclerotized cuticular duct penetrating the trunk cuticle. The duct seems to be surrounded by a cavity. The structures occur ventromedially on segment 7 and slightly more paraventrally on segment 8 (Fig. 13 E; Tables 2 and 5; Table S 2). Each sternal plate of segment 9 possesses a more or less rectangular area with rounded edges and prominent cuticular micropapillae (Fig. 13 E; Tables 2 and 5; Table S 2). Pores and ducts penetrating the trunk cuticle do not seem to exist in this area. Alarge circular gonopore with a more sclerotized edge at its anterior margin and a less sclerotized edge at the posterior margin occurs on each sternal plate of segment 11 (Fig. 13 G). 3.4.4. Sexual dimorphism Males can be distinguished from females by the lack of the ventromedial appendage on each sternal plate of segments 7 and 8 versus its existence in females (Fig. 13 E; Tables 2 and 5; Table S 2), the lack of a more or less rectangular area with prominent cuticular micropapillae on each sternal plate of segment 9 versus its existence in females (Fig. 13 E; Tables 2 and 5; Table S 2), the existence of a ventromedial gland cell outlet with a short sclerotized cuticular duct and an intra- and subcuticular cuticle-lined cavity on each sternal plate of segment 10 versus the lack of this outlet in females, the possession of a middorsal and a laterodorsal acicular spine on segment 10 thinner than on the remaining segments (Fig. 14 G) versus the lack of these spines in females (Fig. 14 F; Tables 2 and 5; Table S 2), and by the lack of a large circular gonopore with a sclerotized edge at its anterior and posterior margin on each sternal plate of segment 11 versus its existence in females (Fig. 13 G). 3.4.5. Variation Out of the three specimens from the Gulf of Lion, one male (USNM 1562541) shows a middorsal cuspidate spine on segment 5 (Fig. 14 D), whereas two females (USNM 1562542, USNM 1562543) reveal a middorsal cuspidate spine on segments 5 and 7 (Fig. 14 E). The female from the northern Adriatic Sea (USNM 1562540) lacks any middorsal cuspidate spine. 3.4.6. Juvenile stage One small specimen (trunk length = 149 M m) possesses a very thin cuticle, has segments 10 and 11 still fused, lacks a free flap but shows a series of postmarginal spicula with a stronger base, and reveals papilla-like sensory spots elevated above the surface of the cuticle (Fig. S 3 A — D). Based on these observations, this specimen is regarded as a juvenile stage. Placids with condyles exist but are difficult to separate from each other. The trunk is composed of segments 1 — 9 separated from each other and from segments 10 + 11 which are still fused (Fig. S 3 B). Segment 1 reveals a single cuticular plate, whereas segments 2 ‾ 11 probably possess one tergal and one sternal plate (Fig. S 3 A). Short cuticular hairs with a stronger base cover all segments except ventromedially and midlaterally, where the dorsoventral muscles originate (Figs. S 3 A and B). A type- 3 sensory spot occurs ventrolaterally and subdorsally on segment 11 (Fig. S 3 B). Type- 6 sensory spots appear at least ventromedially on segments 2, 5, and 9, sublaterally on segments 1, 3, 6, 7, and 9, and paradorsally on segments 1 (almost subdorsally), 4, and 6 ‾ 9 (Fig. S 3 A — D). Acicular spines are found lateroventrally and middorsally on segments 1 ‾ 10 as well as in a lateral accessory position and midterminally on segment 11 (Fig. S 3 A — D). Cuspidate spines are located ventrolaterally on segments 2, 5, and 9, in a lateralaccessory position on segments 4 and 8, and middorsally on segment 5 (Fig. S 3 A — D). 3.4.7. Comparison with previous descriptions The species was originally described as Centroderes multispinosus by McIntyre (1962) and later placed in a new genus and combined as Condyloderes multispinosus (McIntyre, 1962) by Higgins (1969). Most recently, Neuhaus et al. (2019) re-described the male holotype and three additional males from the North Sea. The specimens from the Mediterranean Sea differ from the specimens from the North Sea in the possession of a type- 6 sensory spot ventromedially on segments 5 and 11 versus the lack of these spots in the specimens from the North Sea and in the existence of a middorsal cuspidate spine on segments 5 and / or 7 at least in some specimens versus their lack in the northern population. The juvenile specimen from the Mediterranean Sea possesses a cuspidate spine in a lateral accessory position on segment 4, which is missing in any adult specimen from the North Sea and from the Mediterranean Sea. We regard the above mentioned differences as variation within one and the same species. It should be kept in mind that few specimens of this species are known both from the North Sea and from the Mediterranean Sea (Table S 2). In any case, this study expands the distribution record of C. multispinosus considerably from the North Sea to the Mediterranean Sea suggesting that the species should also occur in the northeastern Atlantic Ocean. 3.4.8. Co-occurrence of C. multispinosus and C. agnetis sp. nov. Asingle male of C. multispinosus was collected together with C. agnetis sp. nov. at a station off Trappeto, Gulf of Castellammare, Sicily, southern Tyrrhenian Sea (Table 1). The two species also occurred in almost the same location off Rovinj, northern Adriatic Sea, but were collected in different years (Table 1).	en	Zotto, Matteo Dal, Neuhaus, Birger, Yamasaki, Hiroshi, Todaro, Antonio (2019): The genus Condyloderes (Kinorhyncha: Cyclorhagida) in the Mediterranean Sea, including the description of two new species with novel characters. Zoologischer Anzeiger 282: 206-231, DOI: 10.1016/j.jcz.2019.05.006
