Research Article |
Corresponding author: Charles Oliver Coleman ( oliver.coleman@mfn.berlin ) Academic editor: Carsten Lüter
© 2014 Charles Oliver Coleman, Michael H. Thurston.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Coleman C, Thurston M (2014) A redescription of the type species of Oedicerina Stephensen, 1931 (Crustacea, Amphipoda, Oedicerotidae) and the description of two new species. Zoosystematics and Evolution 90(2): 225-247. https://doi.org/10.3897/zse.90.8559
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The poorly known species Oedicerina ingolfi Stephensen, 1931 (Crustacea: Amphipoda: Oedicerotidae) is redescribed, based on new material from the Norwegian Sea. Oedicerina vaderi sp. n. from the northeast Atlantic Ocean and Oedicerina loerzae sp. n. from New Zealand waters are described raising the number of species in the genus to five. The three species treated here together with Oedicerina megalopoda Ledoyer, 1986 and Oedicerina denticulata Hendrycks & Conlan, 2003 are separated by characters of the rostrum, maxilliped, gnathopods, epimera, and by the dorsal armature of pleonites and urosomites. The genus is recorded from the Atlantic, Indian and Pacific Oceans, mainly at bathyal depths.
Taxonomy, Amphipoda , Oedicerotidae , Oedicerina , new species, Norwegian Sea, North Atlantic Ocean, New Zealand, Pacific Ocean
Participants in a workshop organised by Professor Wim Vader at the University of Tromsø field station at Skibotn in 2009 sorted and identified amphipods from the Natural History Collections of the University Museum of Bergen collected from the Norwegian Sea. Among the extensive material in these collections were specimens of the poorly known oedicerotid genus Oedicerina Stephensen, 1931. The genus was based on a single individual collected from the Norwegian Sea by The Danish-Ingolf Expedition (
The new material from the Bergen Museum has made possible a re-description of O. ingolfi. Specimens belonging to Oedicerina from the northeast Atlantic Ocean and from New Zealand waters demonstrate further diversity within the genus and require the recognition of two new species which are described below.
This paper is the third to utilize material sorted at Skibotn in 2009, following
Norwegian Sea material assigned to O. ingolfi from the Natural History Collection of the University Museum of Bergen (ZMBN) was collected with an RP sledge (
Specimens from the Discovery Collections at the National Oceanography Centre, Southampton were obtained in the East Iceland Basin on an RRS Discovery cruise that contributed to the Institute of Oceanographic Sciences investigations of mid-water and benthic faunas in the eastern North Atlantic Ocean (1965–1977). The material, from an epibenthid sledge, was fixed in 4% formaldehyde and later transferred to 70% Industrial Methylated Spirits. These specimens have been deposited in the Amphipoda collections at The Natural History Museum, London.
The New Zealand material was collected during the Ocean Survey 2020 expeditions with RV Tangaroa to the Chatham Rise and the Challenger Plateau (
For habitus drawings the specimens were transferred into glycerol on a cavity slide. Specimens were then dissected under a stereomicroscope (Leica M205 or Wild M5) using dissecting needles. Mouthparts and appendages were mounted temporarily in glycerol on slides for microscopic examination and drawing. Appendages were later mounted as permanent slides with glycerol jelly, or transferred into small glass microvials. Microvials were stoppered with a cotton ball wrapped in Japan paper to avoid the appendages being entangled in the cotton fibres.
After dissection, mouthparts and appendages of Discovery Collection material were made directly into permanent mounts using Polyvinyl-lactophenol stained with lignin-pink. Drawings of habitus and appendages were made using a camera lucida attached to a compound microscope (Leica DMLB or Wild M20). Pencil drawings were scanned, inked digitally and arranged to figs using the methods described in
Body lengths were measured along the dorsal outline from the tip of the rostrum to the end of the telson. Lengths of individual articles of gnathopods and pereopods measured along anterior or posterior margins can vary depending on the degree of flexure of the appendage. All articulations except those between coxae and tergites and between merus and carpus of gnathopods are bicondylar. Measurements made between condyles gives a length that is not affected by limb flexure. Length ratios herein have been derived using this principle.
Oedicerina Stephensen, 1931: 250. —
Rostrum well-developed, moderately to strongly deflexed. Antennae sexually dimorphic or not, length medium. Antenna 1 about as long as head and pereonites 1–4 combined, peduncle article 1 longer than articles 2 and 3. Antenna 2 subequal to or weakly longer than antenna 1; peduncle article 4 longer than article 5. Lower lip, inner lobes prominent, separate. Mandible, molar triturative; incisor 5-dentate. Maxilla 1, outer fig 9-dentate; palp slender, article 2 subequal to or longer than article 1. Maxilla 2, figs short, inner broader than outer. Maxilliped, palp article 2 sub-triangular, breadth greatest at half-length, inner margin strongly convex; article 3 produced mediodistally; article 4 longer than article 3.
Coxal figs 1–4 deep, as long or longer than height of corresponding pereonite. Gnathopod 1, coxa expanded distally; carpus and propodus subequal in length, strongly expanded posterodistally. Gnathopod 2, carpus longer than propodus, both strongly expanded posterodistally. Pereopods 3 and 4 fossorial (setose); coxa 4 deeply excavate posteriorly, posterodistal lobe strong, subrectangular. Pereopod 5, coxa bilobate, posterior lobe as long as coxa 4. Pereopod 6, coxa bilobate, posterior lobe strong. Pereopod 7, basis expanded.
Pleonites, some or all carinate or toothed. Epimera 1–3, 1 and 3 rounded, 2 obtusely rounded, posterior margin convex or sinuous. Uropods 1–2, outer ramus subequal to or shorter than inner ramus. Uropod 3, peduncle short; rami subequal, not extending as far as apices of uropods 1–2. Telson notched 30–40%, apices acute.
Oedicerina ingolfi Stephensen, 1931
Oedicerina denticulata Hendrycks & Conlan, 2003; Oedicerina ingolfi Stephensen, 1931; Oedicerina loerzae sp. n.; Oedicerina megalopoda Ledoyer, 1986; Oedicerina vaderi sp. n.
1 | Rostrum massive, spatulate, longer than peduncle article 1 of antenna 1 | O. megalopoda Ledoyer, 1986 |
– | Rostrum at most as long as article 1 of antenna 1 | 2 |
2 | Pleonites 1–2 carinate, pleonite 3 with small tooth | O. ingolfi Stephensen, 1931 |
– | Some pleonites smooth dorsally | 3 |
3 | Pleonite 1 smooth | 4 |
– | Pleonite 1–2 carinate, pleonite 3 smooth | O. loerzae sp. n. |
4 | Pleonite 1–2 smooth | O. vaderi sp. n. |
– | Pleonite 2 carinate, pleonite 3 with short process | O. denticulata Hendrycks & Conlan, 2003 |
Oedicerina ingolfi Stephensen, 1931, p. 250, fig. 72.
Oedicerotidae gen. et sp. n.
3 ovig. females, 6 females, 3 males, 2 unknown sex, 4 juveniles, ZMBN 95143, St. 81.08.14.5, 64°16.9‘N, 00°11.7‘W, 2630 m, F/F Håkon Mosby, RP-sledge, T. Brattegard, 14 August 1981.
1 female, 1 male, 1 juvenile, ZMBN 95144, St. 81.08.14.1, 65°19.7‘N 01°02.7‘E, 2908 m, F/F Håkon Mosby, RP-sledge, T. Brattegard, 14 August 1981.
1 ovig. female, 4 females, 7 males, 3 unknown sex, 12 juveniles, ZMBN 95145, St. 82.11.24.1 64°48.2‘N 01°33.0‘W, 3000 m, F/F Håkon Mosby, RP-sledge, T. Brattegard, 24 November 1982.
1 male, 1 juvenile, ZMBN 95146, St. 81.06.03.5, 67°47.0‘N 07°43.9‘E, 2025 m, F/F Håkon Mosby, RP-sledge, T. Brattegard, 3 June 1981.
1 female, ZMBN 95147, St. 86.07.26.1, 69°36.4‘N 09°54.6‘W; 2212 m, F/F Håkon Mosby, RP-sledge, T. Brattegard, 26 July 1986.
2 adult females, 1 female, 2 juveniles, NORBI St. 2, DS05, 65°22.9‘N 00°02.1‘E–65°22.4‘N 00°02.2‘E; 2970 m, N.O. Jean Charcot, 21 July 1975.
1 ovig. female, 1 adult male, NORBI St. 6, DS12, 76°54.4‘N 01°44.6‘E–76°54.0‘N 01°46.3‘E; 3200 m, N.O. Jean Charcot, 2 August 1975.
Based on ovigerous female, 10.3 mm, St. 81.08.14.5.
Head (Fig.
Oedicerina ingolfi, ovig. female, 10.3 mm; Norwegian Sea, ZMBN 95143, St. 81.08.14.5. a) habitus (uropods rolled up, not illustrated); b) antenna 1; c) antenna 2; d) upper lip; e) maxilla 1; f) maxilla 1, setal teeth of outer fig; g) maxilla 1, distal setation of palp article 2; h) telson; i) mandibular palp. Scale bars: a; 2 mm: b, c, e, h; 200 µm: d, i; 100 µm.
Pereon.Pereonite 1 (Fig.
Pleon.Pleonites 1–2 (Fig.
Urosome.Urosomite 1 (Fig.
Male antenna 1 with shorter peduncle articles in the ratio 1:0.7:0.3 and more numerous flagellum articles compared to female. Article 1 of the flagellum is elongate, about as long as peduncle article 3. Subsequent proximal articles are shorter than wide. The 1-articulate slender accessory flagellum is about 1/3 as long as article 1 of the primary flagellum.
Between the Faroes and Jan Mayen (
Male holotype, 7.3 mm; NHMUK 2014. 398, Discovery Stn 7709#73.
North Atlantic, East Iceland Basin: 60°07.1‘N 19°30.3‘W – 60°06.1‘N 19°24.8‘W, 5 May 1971, BN 2.4, 2636–2646 m.
1 female, 6.3 mm; 2 specimens of unknown sex, 4.2 mm and 5 mm; NHMUK 2014. 399-401, Discovery Stn 7709#73, from the type locality.
The specific name vaderi recognises the important contributions to amphipod studies made by Professor Wim Vader.
Holotype male, 7.3 mm. Head (Fig.
a, c–g) Oedicerina vaderi sp. n., male holotype, 7.3 mm; northeast Atlantic Ocean, NHMUK 2014. 398, Discovery Stn 7709#73. a) habitus; c) head; d) upper lip; e) epimeral figs 1-3; f) antenna 1; g) antenna 2. Female paratype, 6.3 mm; same locality. b) antenna 1. Scale bars: a; 1 mm: b–g; 200 μm.
Pereon.Pereonite 1 (Fig.
a–d, f) Oedicerina vaderi sp. n., male holotype, 7.3 mm; northeast Atlantic Ocean, NHMUK 2014. 398, Discovery Stn 7709#73. a) pereopod 6; b) pereopod 7; c) uropod 3; d) uropod 2, peduncle; f) telson; g) uropod 1. Female paratype, 6.3 mm; same locality. e) pleonite 3 and urosome with telson, uropods not shown, Scale bars: a–d; 200 μm: e; 1 mm.
Pleon.Pleonites 1–3 (Fig.
Urosome.Urosomite 1 (Fig.
The paratypes bear a small posteriorly directed tooth on pleonite 3. It may be that this process has been present and is worn down in the holotype. Antenna 1 of the female (Fig.
North Atlantic, south of Iceland, 2636–2646 m.
Male, 8.5 mm; NIWA 84727, TAN 0705-41, Chatham Rise, 43°50‘10.8“S 176°42‘33.0“E, 478–479 m, 5 April 2007. Paratypes. Male?, 7.7 mm; NIWA 89970, TAN 0705-251, Chatham Rise, 42°59‘45.0“S 178°59‘44.4“E, 520–530 m, 24 April 2007; ovig. female, 7 mm; NIWA 84740, TAN0705-83, Chatham Rise, 43°50‘10.8“S 176°42‘33.0“E, 529–530 m; 9 April 2007.
The species is named for Dr. Anne-Nina Lörz to acknowledge her significant contributions to amphipod systematics.
Holotype male, 8.5 mm. Head (Fig.
Pereon.Pereonite 1 (Fig.
Pleon.Pleonites: 1–2 (Fig.
Urosome.Urosomite 1 (Fig.
Chatham Rise, east of New Zealand.
The female specimen has the same antenna 1 morphology as the male: short peduncle articles and numerous flagellum articles. The proximal articles of the flagellum are shorter than wide.
1 incomplete female; NHMUK 2014. 402, Discovery Stn 7845: north-eastern Atlantic, off the coast of Western Sahara: 23°50.5‘N 17°05.9‘W – 23°51.0‘N 17°05.4‘W, 24 March 1972, BN 2.4, 947–958 m.
Only the head and pereonites 1–2 are present. Coxae 1–2 bear long setae along the distal margins. The animal appears similar to O. ingolfi, but as a result of incompleteness it is impossible to attribute it to any species.
The three species described herein are morphologically very similar. Mouthparts and appendages show only minor and subtle differences and the species are best discriminated by habitus characters. Two of the species, O. ingolfi and O. loerzae sp. n. have mid-dorsal carinae on pleonites 1–2. Oedicerina ingolfi differs from O. loerzae sp. n. in having a small, slender, acute, upright tooth on pleonite 3 and a small pointed process on the posterior margin of urosomite 3 both of which are absent in O. loerzae sp. n. Oedicerina vaderi sp. n. has a small pointed process on the posterior margin of urosomite 3, as found in O. ingolfi, but pleonites 1–2 are evenly vaulted lacking any trace of a carina. Pleonite 3 of the holotype of O. vaderi sp. n. appears to be dorsally unarmed, but the paratypes have a small acute process (see female paratype, 6.3 mm in Fig.
Oedicerina sp. indet. was collected from the warm-temperate east Atlantic Ocean off Western Sahara. The unique specimen is incomplete, only the head and pereonites 1 and 2 are present, preventing a full identification.
Apart from the type species O. ingolfi, two other species had been described in the genus prior to this study: Oedicerina megalopoda Ledoyer, 1986, collected close to Mayotte, Mozambique Channel, western Indian Ocean (200–500 m) and Oedicerina denticulata Hendrycks & Conlan, 2003 from the northeast Pacific Ocean off California (4050 m). Knowledge of O. megalopoda is limited as the unique specimen is incomplete, but the massive rostrum of this species differs markedly from all other species of the genus. The palm of gnathopod 2 of O. megalopoda is straight, similar to that of O. loerzae sp. n., and thus different from the convex pattern seen in both North Atlantic species. In Oedicerina denticulata pleonite 1 is smooth and pleonites 2 and 3 have a posteromarginal process. The process on pleonite 3 is directed posteriorly and is reminiscent of that seen in paratype material of O. vaderi sp. n. thus contrasting with the upright condition found in O. ingolfi. The posterior margin of urosomite 3 bears a small process (as do all species except for O. loerzae sp. n.) and urosomite 1 has a small upright process in the male but not the female, a character unique within the genus. Coxa 5 appears to be longer than wide, as in O. ingolfi.
The mouthparts and appendages of all species of this genus are remarkably similar to each other. Examination of the extensive Norwegian Sea material which we attribute to O. ingolfi indicates that intraspecific variability is minimal, except for sexual dimorphism in antenna 1. In females of both Atlantic species peduncle articles of antenna 1 are longer and more slender than in males. Flagellum articles in females are uniformly much longer than wide and relatively few in number, whereas in males they are more numerous and proximally wider than long, forming an incipient callynophore. This sexual dimorphism is not apparent in O. loerzae sp. n. where the structure of antenna 1 is very similar in males and the one ovigerous female paratype.
Because of minimal differences among appendages and mouthparts in Oedicerina species, differentiation within the genus relies significantly on patterns of ornamentation of pleonites and urosomites. As the posterior segments of the type material of O. ingolfi are missing, the question remains as to which of the two Atlantic species represents the species that Stephensen described. We allocate the material from the museums in Bergen and Lund studied herein to O. ingolfi on geographical grounds in that it was collected much closer to the type locality of that species, and on the morphological grounds of the dense fringe of setae on the distal margins of coxae 1 and 2 and the shape of the rostrum that our material shares with Stephensen‘s original description.
We are very grateful to Wim Vader for inviting us to the highly successful identification workshop at Skibotn which provided the material that stimulated the present paper. We acknowledge with thanks Mr. Jon Anders Kongsrud (University of Bergen), Dr Lars Lunqvist (Museum of Zoology, Lund University) and Ms Sadie Mills (National Institute of Water and Atmosphere Research, Wellington, New Zealand) for the loan of material. We thank Ms Camilla Norrag, who drew the appendages of O. ingolfi and made the line drawings of O. vaderi sp. n.