Systematics of the genus Heterolaophonte (Crustacea, Copepoda, Harpacticoida), with redescription of H. uncinata and H. curvata Author Karaytuğ, Süphan text Zootaxa 2014 3780 3 503 533 journal article 46147 10.11646/zootaxa.3780.3.4 fc9d9984-f0a2-4043-baba-58ec9b5835ae 1175-5326 250197 BF90D095-4452-4222-84CB-232416BF7AE0 Heterolaophonte curvata ( Douwe, 1929 ) ( Figs. 10 –21) Synonymy . Laophonte curvata , Douwe (1929) , p. 286, figs. 4–9. Neotype designation. With the type materials lost, a neotype is designated here in order to clarify the taxonomic status of H. curvata . Neotype is a female, collected on 10.09.2002 from a beach in Yaroz Feneri Village (St. 11), Trabzon province, Turkey ( 41º05.677'N 39º23.718'E ), dissected on 8 Slides (deposited in the NHMUK reg. no. 2014.7). Material examined. St.11 ( 10.09.2002 ), one ♀ and one ♂, each dissected on eight slides (deposited in the NHMUK reg. no. 2014.8-9); St.2, one ♀ dissected on four slides, four ♀, one ♂, ( 01.05.2001 ); St.4, three ♀, two ♂ ( 13.09.2002 ); St.7, three ♀ ( 11.09.2002 ); St.8, six ♀ (1 ♀, dissected on 1 slide) and one ♂ ( 11.09.2002 ), St.12, one ♀ (dissected on 2 slides), three ♂ ( 10.09.2002 ); St. 13, two ♀ ( 09.09.2002 ); St. 14, three ♀, one ♂ ( 14.09.2002 ); St. 16, one ♂ ( 29.11.2007 ); St. 18, one ♀ ( 26.11.2007 ); St. 19, two ♀, one ♂ ( 08.04.2007 ), one ♀ ( 25.11.2007 ); St.20, two ♀ ( 24.11.2007 ); St. 21, one ♂ ( 24.11.2007 ); St. 22, one ♀ ( 24.11.2007 ). All other materials are deposited in the collection of Zoological Museum at the Biology Department of Mersin University. Undissected materials are whole mounted on slides. Redescription of female (based on neotype ) . Body ( Fig. 10 A, B). Total body length 686–706 µm (n = 20; mean = 694 µm). Shape, ornamentation and structure of body, rostrum and cephalothorax as in H. uncinata . Entire surface covered with tiny spinules (see insert on Fig. 10 A). Rostrum triangular ( Fig. 12 A), with 1 pair of welldeveloped sensilla near apex; midventral tube-pore in subapical position; with transverse incomplete surface ridge dorsally indicating original articulation; with ovoid patch of fine spinules located centrally. All prosomites without defined hyaline frills; posterior margins fringed with small spinules. Urosomites ( Fig. 11 A) with several rows of spinules on ventral surface extending laterally ( Fig. 10 B). Genital double-somite with transverse surface ridge dorsally and laterally ( Fig. 10 A, B), 2 pairs of pores located medially; completely fused ventrally ( Fig. 11 A). Copulatory pore located in proximal depression; gonophores fused medially forming single genital slit covered on either side by operculum derived from sixth leg; P6 with small protuberance bearing 1 bare seta ( Fig. 11 A). Anal somite ( Fig. 11 A, B); anal operculum flanked by 1 pair of sensilla; anal opening bordered by 1 frill bearing fine setular extensions. Caudal rami ( Fig. 11 A, B). Divergent, cylindrical, about twice as long as wide; each ramus with 7 bare setae: seta I subventral, bare and shortest; setae II and III bare; setae IV and V fused basally, and with fracture planes; seta VII tri-articulate at base with spinular row near insertion. Each ramus covered with hardly visible spinules on dorsal surface; additional spinular ornamentation present around ventral and dorsal distal margins; long tube-pore present near ventral distal margin. Antennule ( Fig. 12 B) 7-segmented; segmentation, setation and ornamentation as in H. uncinata except for 1 plumose seta on segment 2. Antenna ( Fig. 17 C); segmentation, setation and ornamentation as in H. uncinata except for allobasis with 1 transverse proximal spinular row near base of exopod. Mandible, maxillule, maxilla, maxilliped as in H. uncinata . Swimming legs P2–P4 (Fig. 14B; 16A, B) with wide intercoxal sclerites bearing hardly visible spinules and with well developed praecoxae. Praecoxae with spinules along outer margin. Exopods 3-segmented, endopods 2- segmented. P1 (Figs. 14A; 15A, B, C) with 2-segmented exopod. Ornamentation of intercoxal sclerite and praecoxae as in other swimming legs. Coxa large; with 3 spinular rows on anterior surface, inner and outer margins as figured. Basis with bipinnate seta near insertion of endopod; with spinular rows on anterior surface, along inner and outer margins; with bipinnate seta at outer margin. Exopodal segments with spinular rows as figured. Exp-2 (Figs. 14A; 15C) about twice as long as exp-1. Enp-1 about 5 times as long as width ( Fig. 15 A), and about 3 times as long as exopod, with spinular row located medially on anterior surface; enp-2 with one strong, minutely denticulate claw, and 1 small naked seta (arrowed in Fig. 15 B); several spinules along outer margin and around inner distal corner. P2–P4 (Figs 14B; 16A, B). Coxae and bases with spinular rows along outer margin and on anterior surface; basis with tube-pore on anterior surface; outer margin of basis with bipinnate seta; exopodal and endopodal segments with elaborate spinular/setular ornamentation along outer margins as figured; outer half of endopod segments of P2-P3 covered with fine spinules. P3–P4 enp-1 shorter than enp-2, P3 enp-2 about as long as enp-1, P2-P4 enp-1 without seta; P2 and P4 enp-2 with tube pore located terminally on anterior surface. Spine and setal formulae of swimming legs as follows:
Exopod Endopod
P2 0.1.0 23 0. 211
P3 0.1.0 23 0. 221
P4 0.1.0 22 [0.1.022 ♂] 0. 0 20 [0.120 ♂]
Fifth pair of legs ( Fig. 17 B). Similar to that of H. uncinata . Exopod and baseoendopod each with pattern of spinules on anterior surface as figured. Baseoendopod with 3 tubepores on anterior surface; endopodal lobe extending to middle of exopod, with 2 apical and 3 medial bipinnate setae, outermost minutely pinnate seta shortest, 2 proximal setae minutely pinnate. Exopod with 6 setae (one of which naked). Redescription of male (based on material from St.11) . Body ( Figs. 17 A; 18A) similar to H. uncinata . Body length 638–670 µm (n=10; mean 650 µm). Rostrum narrower than female ( Fig. 12 C) with 1 pore (arrowed in Fig. 18 B), surface with 1 group of spinules on anterior surface as figured. All urosomites with several spinular rows centrally, some rows extending laterally. Caudal rami as in female ( Figs. 11 C, D; 18A, D). Antennule ( Fig. 12 C; 13A, B, C) 8-segmented; subchirocer with geniculation between segments 5 and 6. Segment 1 with spinules along anterior margin and covered with tiny spinules dorsally as figured. Segment 5 swollen with 1 robust spinulose seta swollen at base ( Fig. 12 C; arrowed in Fig. 13 A) with 1 long aesthetasc (arrowed in Fig. 13 C) and 1 semispinulose seta. Segment 6 with 2 modified spinous element (arrowed in Fig. 13 B). Segmental homologies: 1-(I), 2-(II–VIII), 3-(IX–XII), 4-(XIII), 5-(XIV–XX), 6-(XXI–XXII), 7-(XXIII), 8- (XXIV–XXVIII). Armature formula: 1-[1], 2-[9], 3-[6], 4-[2], 5-[11 + 1 modified + (1 + ae)], 6-[2 modified spinous elements], 7-[1], 8-[7 + 1 modified + acrothek]. Apical acrothek consisting of 1 aesthetasc and 2 naked setae ( Fig. 12 C). Surface ornamentation of intercoxal sclerites and protopods of P1–P4 ( Figs 16 C; 19A, B) generally as in ♀. Many modifications on P1-P4. P2 ( Fig. 19 A). Exopod segments more robust than female. Exp-3 more sclerotised, outer spines stouter and naked, innermost spine remarkably shorter and stouter (homologous to terminal bipinnate spine of female). Enp-1 terminally elongated as 1 apophysis carrying 1 pore at tip and with 1 group of spinules along inner and outer margins. Enp-2 with longer spinules along outer margin, terminal seta longer and bipinnate, proximal inner seta transformed into 1 naked spine (arrowed in Fig. 19 A; indicated with upper right arrows in Fig 20 A). P3 ( Fig. 19 B). Exp-3 slightly curved and sclerotised, outer spines naked and robust (posterior fine spinular ornamentations only visible with SEM, arrowed in Fig. 20 D), middle outer spine modified into 1 longer naked spine, outer terminal spine modified into 1 shorter naked spine. Enp-1 with long setules along inner margin. Enp-2 with long spinules along inner margin, outermost seta transformed into 1 apophysis (arrowed in Fig. 19 B; indicated with lower left arrow in Fig. 20 A), anterior surface with 2 stout spinules. P4 ( Fig. 16 C). Inner seta of exp-2 minutely pinnate and smaller than that of female. Terminal inner spine of female reduced to 1 small naked seta. Enp-1 with 2 small inner spinules. Enp-2 ornamented with long spinules along outer margin and 1 patch of anterior minute spinules near base of tube pore; with small pinnate inner seta; terminal seta semispinulose about twice longer than outer terminal seta. Fifth pair of legs ( Fig. 11 C). Baseoendopods fused medially ( Figs. 11 C; 18C), with setophore bearing outer naked basal seta; endopodal and exopodal lobes vestigial bearing 2 and 4 small naked setae respectively; with spinules and 1 tube-pore near base of setophore (indicated with upper arrow in Fig. 20 B). Sixth pair of legs ( Fig. 11 C) symmetrical; represented by 1 plate fused to ventral wall of supporting somite ( Figs. 11 C; 18C); outer distal corner produced into small process bearing several spinules at base and 2 naked setae (indicated with lower arrow in Fig. 20 B). FIGURE 10 . Heterolaophonte curvata , ♀. A, habitus, dorsal (insert indicates ornamentation on the body surface); B, habitus, lateral. FIGURE 11 . Heterolaophonte curvata . A, ♀ urosome, ventral; B, ♀ anal somite and caudal rami, dorsal; C, ♂ urosome, P5 and P6, ventral; D, ♂ anal somite and caudal rami, dorsal. FIGURE 12 . Heterolaophonte curvata . A, ♀ rostrum, dorsal; B, ♀ antennule, ventral; C, ♂ antennule and rostrum, dorsal. FIGURE 13 . Heterolaophonte curvata , ♂. SEM micrographs of antennule. A, dorsal (arrow indicates the modified seta); B, distal portion (arrow indicates the modified elements on segment 5); C, swollen segment 5 (arrow indicates the aesthetasc). Variability. No significant variation was observed among the examined specimens. Distribution. Based on the examined materials and confirmed records, it can be assumed that H. curvata has a wide distribution both in the Black Sea and the Mediterranean Sea. Remarks. Heterolaophote curvata was originally described from Bay of Cavaliere and Cette along the Mediterranean French coast by Douwe (1929) . The present specimens differ from the original description in the presence of four setae (instead of one) on antennary exopod and in having one small inner terminal seta on the third exopodal segment of the male P3. But, it is highly possible that these setae have been overlooked by Douwe (1929) . Further comparisons about the spinular ornamentation on the appendages cannot be made since the original FIGURE 14 . Heterolaophonte curvata , ♀. A, P1, anterior; B, P3, anterior. FIGURE 15 . Heterolaophonte curvata , ♀. SEM micrographs of P1. A, exopod and endopod; B, terminal endopod segment and the distal claw (arrow indicates the small seta); C, exopod. FIGURE 16 . Heterolaophonte curvata . A, ♀ P2, anterior; B, ♀ P4, anterior; C, ♂ P4, anterior FIGURE 17 . Heterolaophonte curvata . A, habitus, lateral ♂; B, ♀ P5, anterior; C, ♀ antenna. FIGURE 18 . Heterolaophonte curvata , ♂. SEM micrographs. A, habitus, ventral; B, rostrum, dorsal (arrow indicates the pore); C, plate-like structure of P6; D, caudal rami, ventral. FIGURE 19 . Heterolaophonte curvata , ♂. A, P2, anterior (arrow indicates modified seta); B, P3, anterior (arrow indicates apophysis). FIGURE 20 . Heterolaophonte curvata , ♂. SEM micrographs. A, modified element on P2 endopod (indicated with 2 upper arrows) and straight, spiniform apophysis on P3 endopod surrounded at base by row of coarse spinules (indicated with a lower arrow); B, setal elements on P5 and P6 (indicated with arrows); C, P3 exopod, anterior; D, exopodal spines of P3 exp-3, posterior (arrow indicates the minute pinnules). figures lack sufficient detail. On the other hand our specimens match well with the previous descriptions ( Lang 1948 ; Apostolov 1990 ; Apostolov & Marinov 1988 ). But, the setation on the figures and the setal formulae given in the text by Apostolov & Marinov (1988) contain several discrepancies which might have resulted from typing errors. Therefore the population reported by Apostolov & Marinov (1988) from the Black Sea is accepted as conspecific with the present species presented herein. The subspecies H. curvata micrarthros has an interesting history of description. This subspecies was originally created as a new variety of H. curvata from Yalta (Crimea) by Marcus & Por (1960) . But, they determined that a previous report of H. curvata by Por (1960) from Romanian Black Sea coast (Eforie) was conspecific with the specimens from Yalta. Although Por (1960) noted some differences between Romanian specimens and the typical H. curvata he decided at that time that creating a new variety was not justified since he had only male specimens from a single locality. Later Marcus & Por (1960) found female specimens in Yalta (Crimea) and created H. curvata var. micrarthros and concluded that their specimens differed from typical H. curvata by the following five features: i ) Exopod of the antenna more developed than as that described by Douwe (1929) and bearing 4-5 setae instead of one, ii ) terminal exopod segment of P3 has an extra seta and the P3 enp-2 a very weak thumb-shaped spine, iii ) anal operculum with spinules on free border, iv ) inner modified spine of P2 enp-2 of the male is less chitinized and weaker, v ) the P4 endopod of the male is 2-segmented instead of one. But, these diagnostic features of H. curvata micrarthros are also found in the present redescription of H. curvata . On the other hand, several new differences can now be defined between H. curvata micrarthros and presently redescribed H. curvata ; i ) caudal rami shorter in female, ii ) terminal exopodal segment of P4 with 5 setae/spines in male, iii ) P4 enp-2 of male with one long seta, iv ) structure and the ornamentation of setal elements of female P5 are different. On the other hand, it should be pointed out that the setal formula of P4 differs between female and male of H. curvata micrarthros , viz, terminal exopodal segment of P4 with 5 setae/spines in female but with 4 setae/spines in male and P4 enp-2 of male with one long seta in female but with 2 setae in male. This might mean that H. curvata micrarthros was described on the basis of male and female specimens belonging to different species. As mentioned above, the female specimens of H. curvata micrarthros were collected from Eforie ( Romania ) but the male specimens were obtained from Yalta. The description of the male possibly belongs to a taxon closely related to H. curvata (Por, 1960) but the female specimens described from Yalta ( Marcus & Por 1960 ) may not even belong to the genus Heterolaophonte but another genus such as Paralaophonte . Therefore, the position of H. curvata micrarthros within the genus should provisionally be considered doubtful. The reports of H. curvata micrarthros given by Marinov (1971) and Apostolov & Marinov (1988) should also be confirmed.