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Review of the Subfamily Cleonardopsinae Lowry, 2006 (Crustacea: Amphipoda: Amathillopsidae) with Description of a New Genus

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Review of the Subfamily Cleonardopsinae Lowry, 2006 (Crustacea: Amphipoda: Amathillopsidae) with Description of a New Genus Journal of the Marine Review of the subfamily Cleonardopsinae Biological Association of the United Kingdom Lowry, 2006 (Crustacea: Amphipoda: Amathillopsidae) with description of a new cambridge.org/mbi genus and species from Japan Masafumi Kodama* and Tomohiko Kawamura Original Article International Coastal Research Center, Atmosphere and Ocean Research Institute, The University of Tokyo, 1-19-8 *Present address: Faculty of Fisheries, Akahama, Otsuchi, Iwate 028-1102, Japan Kagoshima University, 4-50-20 Shimoarata, Kagoshima 890-0056, Japan Abstract Cite this article: Kodama M, Kawamura T The amathillopsid subfamily Cleonardopsinae Lowry, 2006 is reviewed. The only species of (2021). Review of the subfamily the subfamily, Cleonardopsis carinata K.H. Barnard, 1916, should be regarded as a species- Cleonardopsinae Lowry, 2006 (Crustacea: complex. A new genus and species of the subfamily, Carinocleonardopsis seisuiae gen. et Amphipoda: Amathillopsidae) with description of a new genus and species from Japan. sp. nov., is described from the Sea of Kumano, Japan as the second species of the subfamily Journal of the Marine Biological Association of Cleonardopsinae as well as the first record of the subfamily from the North Pacific. This new the United Kingdom 101,359–369. https:// genus can be easily distinguished from Cleonardopsis by the presence of distinct large eyes and doi.org/10.1017/S0025315421000357 the dorsal carination on head, pereonites and pleonites. Received: 5 September 2020 Revised: 1 April 2021 Accepted: 20 April 2021 Introduction First published online: 9 June 2021 The monotypic amphipod subfamily Cleonardopsinae Lowry, 2006 has a complex taxonomic Key words: Carinocleonardopsis seisuiae; Cleonardopsis; history. The only current genus of the subfamily, Cleonardopsis Barnard, 1916 was originally Sea of Kumano; species complex; taxonomy established in the family Eusiridae, however, it has been frequently placed in other families such as Amathillopsidae or Pleustidae by various authors (e.g. Pirlot, 1936; Barnard & Author for correspondence: Karaman, 1991a, 1991b; Coleman, 1998; Lowry, 2006). Masafumi Kodama, E-mail: [email protected] The only species of the genus, Cleonardopsis carinata Barnard, 1916,alsohasacomplextaxo- nomic history. This species was originally described from off South Africa (Barnard, 1916). Since then, several authors have reported the species from various localities (e.g. Schellenberg, 1926; Pirlot, 1934, 1936;Stephensen,1944). However, these reports sometimes lacked descriptions or illustrations, and moreover, sometimes showed different morphologies from each other. Lowry (2006) implied that the material of ‘Cl. carinata’ from different localities could be separate species. During a survey of the deep-sea benthic fauna in the Sea of Kumano and off Tanabe Bay by TRV ‘Seisui-maru’, Mie University (research cruise no. 1803; Kimura et al., 2019), an amphi- pod species attributed to the subfamily Cleonardopsinae was collected. The present study pro- vides a review of the subfamily Cleonardopsinae and compares our material with the description of Cleonardopsis carinata in previous studies. As a result, our specimens clearly revealed distinct characters from the genus Cleonardopsis, and therefore, we herein describe and illustrate the present species as Carinocleonardopsis seisuiae gen. et sp. nov. Materials and methods Fresh specimens of Carinocleonardopsis seisuiae gen. et sp. nov. were collected from sandy- muddy bottom in the Sea of Kumano, Japan (Figure 1). The body lengths of specimens exam- ined were measured by tracing individuals’ mid-trunk lengths (tip of the rostrum to end of © The Author(s), 2021. Published by telson); measuring this curved length and then converting this to actual animal body length Cambridge University Press on behalf of by correcting for magnification, following previous studies (e.g. Lörz et al., 2007, 2009). The Marine Biological Association of the United Kingdom. This is an Open Access article, specimens were dissected under a binocular stereomicroscope, and the appendages were distributed under the terms of the Creative mounted in Euparal on glass slides. Observations and line drawings were made using a Commons Attribution licence (http:// light microscope with the aid of a drawing tube (Y-IDT, Nikon, Tokyo, Japan). The specimens creativecommons.org/licenses/by/4.0/), which were deposited in the National Museum of Nature and Science, Tokyo (NSMT). permits unrestricted re-use, distribution, and reproduction in any medium, provided the original work is properly cited. Results Systematics Order AMPHIPODA Latreille, 1816 Family AMATHILLOPSIDAE Pirlot, 1934 [New Japanese name: Ryukotsu-yokoebi-ka] Subfamily Cleonardopsinae Lowry, 2006 Genus Cleonardopsis Barnard, 1916 Cleonardopsis Barnard, 1916: 175. —Schellenberg, 1926: 230. —Pirlot, 1936: 237. — Stephensen, 1944:7.—Barnard, 1969: 223. —Griffiths, 1975: 118. —Ledoyer, 1986: 1052. Downloaded from https://www.cambridge.org/core. IP address: 170.106.33.42, on 02 Oct 2021 at 02:33:36, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0025315421000357 360 Masafumi Kodama and Tomohiko Kawamura Fig. 1. Geographic distribution of the subfamily Cleonardopsinae Lowry, 2006. —Barnard & Karaman, 1991a: 315. —Barnard & Karaman, follows: South Africa (Barnard, 1916; Schellenberg, 1926); 1991b: 646. —Elizalde et al., 1993: 252. —Dauvin & Sorbe, Indonesia (Pirlot, 1934); Greenland (Stephensen, 1944); Bay of 1995: 454. —Coleman, 1998: 31. —Lowry, 2006: 12. —Coleman, Biscay (Elizalde et al., 1993;Dauvin&Sorbe,1995;Frutos&Sorbe, 2007: 20. —Frutos & Sorbe, 2014: 299, 304. —Frutos & Sorbe, 2014, 2017); Iceland (Brix et al., 2018;Jażdżewska et al., 2018). 2017: 36. —Brix et al., 2018:7.—Jażdżewska et al., 2018: 62. Amathillopleustes Pirlot, 1934: 205 [synonymized with Remarks Cleonardopsis by Pirlot (1936)] Detailed taxonomic history of the genus Cleonardopsis is reviewed below (see Review and discussion of the subfamily Cleonardopsinae). Diagnosis after Lowry (2006) Head deeper than long; lateral cephalic lobe subquadrate, head truncated apically; anteroventral margin straight, anteroventral Cleonardopsis carinata Barnard, 1916 margin moderately recessed, anteroventral margin moderately (Figures 2–4) excavate; rostrum short or moderate length; eyes absent. Body Cleonardopsis carinata Barnard, 1916: 176. —Schellenberg, without setae; dorsally carinate. Antenna 1 subequal in length 1926: 230. —Pirlot, 1936: 237. —Stephensen, 1944:7.— or longer than antenna 2; peduncle with sparse slender setae; ped- Griffiths, 1975: 118. —Ledoyer, 1986: 1052. —Barnard & uncular article 1 longer than article 2; article 2 longer than article Karaman, 1991a: 316. —Barnard & Karaman, 1991b: 646. — 3; article 3 shorter than article 1; accessory flagellum minute, Elizalde et al., 1993: 252. —Dauvin & Sorbe, 1995: 454. — 1-articulate; calceoli present. Antenna 2 medium length; peduncle Coleman, 1998: 31. —Lowry, 2006: 12. —Frutos & Sorbe, 2014: with sparse slender setae or none; flagellum longer than peduncle. 299, 304. —Frutos & Sorbe, 2017: 36. Pereon. Coxae 1–4 longer than broad, overlapping, coxae not Amathillopleustes alticoxa Pirlot, 1934: 205. [Synonymized ventrally acute. Coxae 1–3 progressively larger. Gnathopod 1 sub- with Cleonardopsis carinata by Pirlot (1936)] chelate; carpus shorter than propodus. Gnathopod 2 subchelate; coxa smaller than but not hidden by coxa 3; carpus short, shorter Distribution than propodus. Pereopods not prehensile. Pereopod 4 coxa not South Africa (Barnard, 1916; Schellenberg, 1926), Indonesia ventrally acute, with small posteroventral lobe. Pereopod 5 coxa (Pirlot, 1934), Greenland (Stephensen, 1944), Bay of Biscay without lobes; basis slightly expanded. Pereopod 6 basis slightly (Elizalde et al., 1993; Dauvin & Sorbe, 1995; Frutos & Sorbe, expanded. Pereopod 7 basis expanded, subrectangular. 2014, 2017)(Figure 1). Pleon. Urosomites not carinate. Uropods 1–2 apices of rami without robust setae. Telson weakly cleft; dorsal or lateral robust Remarks setae absent; apical robust setae absent. Several authors reported the occurrence of this species, however, only Barnard (1916) and Pirlot (1934) provided morphological Species composition description and illustration (Figures 2–4). Detailed taxonomic Cleonardopsis carinata Barnard, 1916 (monotypic) history of this species is reviewed below (see Review and discus- sion of the subfamily Cleonardopsinae). Distribution Some previous studies mentioned the distribution of the genus as ‘cosmopolitan’ (e.g. Barnard, 1969;Lowry,2006). Detailed records Genus Carinocleonardopsis gen. nov. of collecting localities are summarized in Figure 1 and shown as [New Japanese name: Mino-yokoebi-zoku] Downloaded from https://www.cambridge.org/core. IP address: 170.106.33.42, on 02 Oct 2021 at 02:33:36, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0025315421000357 Journal of the Marine Biological Association of the United Kingdom 361 margin moderately excavate; rostrum short or moderate length; distinct eyes present. Body without setae; dorsal carination present on head, pereonites and pleonites. Antenna 1 subequal in length or longer than antenna 2; pedunclewith sparse slender setae; peduncular article 1 longer than article 2; article 2 longer than article 3; article 3 shorter than article1; accessory flagellum minute,1-articulate; calceoli
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