An overview of the genus Plesionika Bate, 1888 (Decapoda, Caridea, Pandalidae) in Asian waters

Dr. Md. Yeamin Hossain

The pandalid genus Plesionika Bate, 1888 includes 93 species and has a widespread distribution including mainly subtropical and tropical waters and some temperate waters, but most species can be found in the Indo-West Pacific. The species of this genus are benthic or nektobenthic, feeding on pelagic and benthic resources. Up to the present, the occurrence of 45 species of the genus has been reported from Asian waters. However, studies on these species from this region are mainly concerning their taxonomic report and very less their biology. Herein a key for the 45 Asian Plesionika species, and brief notes on its taxonomy are provided.

Zootaxa 4221 (5): 575–593 http://www.mapress.com/j/zt/ ISSN 1175-5326 (print edition) Article Copyright © 2017 Magnolia Press ZOOTAXA ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4221.5.6 http://zoobank.org/urn:lsid:zoobank.org:pub:1CB61A37-23C5-4041-AB51-0EA76E62770F An overview of the genus Plesionika Bate, 1888 (Decapoda, Caridea, Pandalidae) in Asian waters FERDOUS AHAMED1,6, IRENE A. CARDOSO2, ZOARDER F. AHMED3, MD. Y. HOSSAIN4 & JUN OHTOMI5 1 Department of Fisheries Management, Patuakhali Science and Technology University, Patuakhali-8602, Bangladesh Setor de Carcinologia, Museu Nacional/UFRJ, Quinta da Boa Vista, São Cristóvão s/n, 20940-040, Rio de Janeiro, Brazil 3 Department of Fisheries Management, Bangladesh Agricultural University, Mymensingh-2202, Bangladesh 4 Department of Fisheries, Faculty of Agriculture, University of Rajshahi, Rajshahi-6205, Bangladesh 5 Faculty of Fisheries, Kagoshima University, 4-50-20 Shimoarata, Kagoshima 890-0056, Japan 6 Corresponding author. E-mail: ferdous@pstu.ac.bd 2 Abstract The pandalid genus Plesionika Bate, 1888 includes 93 species and has a widespread distribution including mainly subtropical and tropical waters and some temperate waters, but most species can be found in the Indo-West Pacific. The species of this genus are benthic or nektobenthic, feeding on pelagic and benthic resources. Up to the present, the occurrence of 45 species of the genus has been reported from Asian waters. However, studies on these species from this region are mainly concerning their taxonomic report and very less their biology. Herein a key for the 45 Asian Plesionika species, and brief notes on its taxonomy are provided. Key words: pandalid shrimp, Plesionika, taxonomy, biology, Asia Introduction Plesionika Bate, 1888 is the largest genus of the family Pandalidae, containing 93 species (Cardoso 2011; Li & Chan 2013). This genus has a wide spread distribution all over the world (except in high latitudinal areas) and consists mainly of deep water shrimps. The species of this genus mainly occur in subtropical and tropical waters and in some temperate waters (Chace 1985), but most species can be found in the Indo-West Pacific (Li 2006a). Plesionika species are small and intermediate sized shrimps ranging from 6–29 mm carapace length (Vafidis et al. 2005). Several studies reported that they have a clear segregation on size and depth and thus each species has a preferred depth range (King & Butler 1985; Company & Sardà 1997; Carbonell et al. 2003). The species of this genus are benthic or nektobenthic, distributed on the shelf and slope. This benthic fondness of Plesionika species often make their distribution more restricted than that of the pelagic species, which contributes to speciation within the genus (Vafidis et al. 2005). They play an important role in the marine ecosystem (Cartes 1998; Maynou & Cartes 1998; Cartes et al. 2002) as they constitute a major part of the diet of mesopelagic, suprabenthic, epibenthic and infaunal organisms, and are considered non-migratory macroplankton feeders (Cartes et al. 2002). Furthermore, they constitute a large part of the diet of demersal fish and cephalopods. The existence of Plesionika spp. have been studied extensively in Asian waters and several species of this genus have been reported mainly from Japan, Korea, China, Taiwan, Philippine, Indonesia, India and from few other countries. In the mixed demersal species fisheries of the Asian waters, most of the Plesionika spp. are bycatch of the trawl fishery, however; many of them are commercially important for small-scale bottom seine fishery. To the best of our knowledge, there is no information of the total number of Plesionika spp. recorded from Asian waters. Therefore, the current review paper aims to collect the existing published report to provide a list of Plesionika spp. inhabiting Asian waters along with their key taxonomic characters and present status of their biological aspects. Accepted by J. Goy: 16 Nov. 2016; published: 20 Jan. 2017 575 Occurrence of Plesionika spp. in Asian waters with their description Plesionika species are caridean shrimps and can be characterized by both sides of the pleura of the second somite of the abdomen extending over the first and third somites, respectively. They have dorsal rostral teeth on the base of the rostrum and along the length of the rostrum dorsally or on both sides. The carapace is smooth or covered with a thin layer of minute scales. The claws of the first pair of pereopods are either microscopic, or absent. The carpus of the second pair of pereopods is subdivided into many articles. The 3rd maxilliped is bearing an exopodite (Holthuis 1980). From Asian waters, 45 species have been reported so far (Table 1), which can be distinguished based on several meristic characters. Brief notes on the taxonomy of Plesionika spp. recorded from Asian waters along with their taxonomic position are as follows (species are ordered alphabetically). Also a key for the 45 Asian Plesionika species is provided. TABLE 1. List of species along with their synonyms belonging to the genus of Plesionika from Asian waters. Sl. No. Species name Synonyms 1. Plesionika acinacifer Chace, 1985 2. P. adensameri (Balss, 1914) Parapandalus adensameri Balss, 1914 3. P. alcocki (Anderson, 1896) Pandalus alcocki Anderson, 1896 Plesionika adenensis Timofeev, 1993 4. P. albocristata Chan & Chuang, 2002 5. P. assimilis De Man, 1917 6. P. bifurca Alcock & Anderson, 1894 7. P. binoculus (Bate, 1988) 8. P. brevirostris Bate, 1888 9. P. carsini Crosnier, 1986 10. P. crosnieri Chan & Yu, 1991 Plesionika dentirostris Tung, Wang & Li, 1988 11. P. edwardsii (Brandt, 1851) Pandalus (Parapandalus) longirostris Borradaile, 1900 Pandalus (Pontophilus) edwardsii Brandt, 1851 Pandalus guerinii Risso, 1844 Plesionika longirostris 12. P. ensis (Milne-Edwards, 1881) Acanthephyra ensis A. Milne-Edwards, 1881 Pandalus ensis (A. Milne-Edwards, 1881) 13. P. erythrocyclus Chan & Crosnier, 1997 14. P. fimbriata Chace, 1985 15. P. grandis Doflein, 1902 Plesionika spinipes var. grandis Doflein, 1902 16. P. heterocarpus (Costa, 1871) Pandalus heterocarpus A. Costa, 1871 Pandalus longicarpus A. Milne-Edwards, 1883 Pandalus sagittarius A. Milne-Edwards, 1883 17. P. hsuehyui Chan, 2004 18. P. hypanodon Doflein, 1902 19. P. indica De Man, 1917 20. P. intermedia Chace, 1985 21. P. izumiae Omori, 1971 22. P. kensleyi Chace, 1985 23. P. longicauda (Rathbun, 1901) 24. P. longidactylus Li & Komai, 2003 25. P. lophotes Chace, 1985 26. P. martia (Milne-Edwards, 1883) Nothocaris binoculus Bate, 1888 Plesionika longipes var. indica De Man, 1917 Pandalus longicauda Rathbun, 1901 Pandalus martius A. Milne-Edwards, 1883 Plesionika (Pandalus) sicherii Riggio, 1900 Plesionika martia martia (A. Milne-Edwards, 1883) ......continued on the next page 576 · Zootaxa 4221 (5) © 2017 Magnolia Press AHAMED ET AL. TABLE 1. (Continued) Sl. No. Species name Synonyms 27. P. narval (Fabricius, 1787) Astacus narual Fabricius, 1787 Nisea formosa Risso, 1844 Palaemon tarentinum O.G. Costa, 1844 Palemon pristis Risso, 1816 Pandalus escatilis Stimpson, 1860 Pandalus stylopus A. Milne-Edwards, 1883 Parapandalus narval (Fabricius, 1787) 28. P. orientalis Chace, 1985 Plesionika cottei Kotte, 1903 Plesionika martia orientalis Chace, 1985 29. P. ortmanni Doflein, 1902 30. P. parvimartia Chace, 1985 31. P. persica (Kemp, 1925) 32. P. philippinensis Chace, 1985 33. P. pumila Chace, 1985 34. P. quasigrandis Chace, 1985 35. P. reflexa Chace, 1985 36. P. rufomaculata Chan, 2004 37. P. semilaevis Bate 1888 38. P. simulatrix Chace, 1985 39. P. sindoi (Rathbun, 1906) 40. P. spinensis Chace, 1985 41. P. spinidorsalis (Rathbun, 1906) 42. P. taiwanica Chan & Yu, 2000 43. P. unidens Bate, 1888 Plesionika affinis Alcock & Anderson, 1899 44. P. williamsi Forest, 1963 Plesionika alaini Burukovsky, 1993 Plesionika crosnieri Burukovsky, 1992 45. P. yui Chan & Crosnier, 1991 Parapandalus filipes Calman, 1939 Parapandalus persicus Kemp, 1925 Parapandalus yugniroi Timofeev, 1993 Pandalus sindoi Rathbun, 1906 Pandalus spinidorsalis Rathbun, 1906 Taxonomic position Phylum: Arthropoda Sub-phylum: Crustacea Class: Malacostraca Order: Decapoda Infraorder: Caridea Family: Pandalidae Haworth, 1825 Genus: Plesionika Bate, 1888 Key to Asian species of Plesionika 1. 2. 3. 4. Carapace armed with dorsal teeth in two thirds of its length . . . . . . . . . . . . . . . . . . . . . . . . . . . P. spinidorsalis (Rathbun, 1906) Carapace without dorsal teeth in two thirds of its length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Rostrum with high basal crest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Rostrum with low basal crest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Rostrum unarmed dorsally (excluding basal crest and subapical teeth) or armed with one or two teeth . . . . . . . . . . . . . . . . . 4 Rostrum armed dorsally (excluding basal crest and subapical teeth) with more than two teeth . . . . . . .P. lophotes Chace, 1985 Rostrum unarmed dorsally (excluding basal crest and subapical teeth) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 THE GENUS PLESIONIKA BATE, 1888 IN ASIAN WATERS Zootaxa 4221 (5) © 2017 Magnolia Press · 577 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. Rostrum armed dorsally (excluding basal crest and subapical teeth) with one or two teeth . . . . . . P. rufomaculata Chan, 2004 Rostrum with two subapical teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. hsuehyui Chan, 2004 Rostrum without subapical teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .P. albocristata Chan & Chuang, 2002 Rostrum short, not reaching or slightly overreaching level of distal end of antennal scale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Rostrum long, far overreaching level of distal end of antennal scale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Rostrum unarmed dorsally (excluding that teeth before orbital margin and the subapical teeth) . . . . . . . . . . . . . . . . . . . . . . . . 8 Rostrum armed dorsally (excluding that teeth before orbital margin and the subapical teeth) . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Rostrum ventral margin with 7–9 evenly distributed teeth between level of eye and tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. erythrocyclus Chan & Crosnier, 1997 Rostrum ventral margin with about 10 obscure serrations near apex . . . . . . . . . . . . . . . . . . . . . . . . . P. brevirostris Bate, 1888 Rostrum ventral margin unarmed or with appressed serrations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Rostrum ventral margin armed with teeth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Rostrum ventral marginunarmed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. hypanodon Doflein, 1902 Rostrum ventral margin with 7–13 appressed serrations becoming obscure anteriorly . . . . . . . . . . . P. acinacifer Chace, 1985 Rostrum ventral margin armed with 3–5 teeth in anterior half of rostrum . . . . . . . . . . . . . . . . . . . . . . . P. kensleyi Chace, 1985 Rostrum ventral margin armed with 9–10 small teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. persica (Kemp, 1925) Rostrum dorsally unarmed after antennular peduncle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Rostrum dorsally armed with teeth after antennular peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 Fourth pair of pereopods without epipods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Fourth pair of pereopods with epipods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 With epipods on pereopods 1–3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .P. intermedia Chace, 1985 Without epipods on pereopods 1–3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. simulatrix Chace, 1985 Teeth on rostral crest with barbed tips. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. pumila Chace, 1985 Teeth on rostral crest with sharp point . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 Abdominal somite 3 tergum with right angle or acute posterodorsal tooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Abdominal somite 3 tergum posterodorsally rounded and unarmed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Rostrum relatively short, about 1.7 carapace length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. williamsi Forest, 1963 Rostrum relatively long, more than twice carapace length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 Third abdominal somite not recurved dorsally; pereopod 3 dactyl usually less than 0.25 as long as propod . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .P. ensis (Milne-Edwards, 1881) Third abdominal somite usually with posterodorsal tooth recurved dorsally; pereopod 3 dactyl usually less than 0.25 as long as propod . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. reflexa Chace, 1985 Rostrum ventrally armed with few teeth (8–23) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. alcocki (Anderson, 1896) Rostrum ventrally armed with many teeth (19–53) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 Eye kidney shaped. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. semilaevis Bate, 1888 Eye ovoid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .21 Pereopod 5 propod thread like, extremely long, at least three times as long as carapace . . . . . . P. taiwanica Chan& Yu, 2000 Pereopod 5 propod not thread like, long but less than three times as long as carapace . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 Telson subequal to abdominal somite 6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. orientalis Chace, 1985 Telson longer than abdominal somite 6. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. martia (Milne-Edwards, 1883) Rostrum armed with numerous closely set teeth almost the entire length of both borders . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 Rostrum not armed with numerous closely set teeth almost the entire length of both borders . . . . . . . . . . . . . . . . . . . . . . . . . 29 Abdominal somite 5 pleura with posteroventral denticle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 Abdominal somite 5 pleura without posteroventral denticle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 Pereopods 1–4 with epipods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. indica De Man, 1917 Pereopods 1–4 without epipods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 Posterior 10 ventral rostral teeth corresponding to 8 or fewer dorsal teeth, penultimate segment of maxilliped 3 usually less than 1.5 times as long as terminal segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. quasigrandis Chace, 1985 Posterior 10 ventral rostral teeth corresponding to more than 8 dorsal teeth, penultimate segment of maxilliped 3 more than 1.5 times as long as terminal segment. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. grandis Doflein, 1902 Epipod well developed at maxilliped 3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .P. narval (Fabricius, 1787) Epipod absent or rudimentar at maxilliped 3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 Maxilliped 3 without epipod, rostrum with posterior 10 ventral teeth corresponding to more than 7 dorsal teeth, pereopod 3 dactylus about 0.10 as long as propod . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. longicauda (Rathbun, 1901) Maxilliped 3 with rudimentary epipod, rostrum with posterior 10 ventral teeth corresponding to less than 8 dorsal teeth, pereopod 3 dactylus more than 0.20 as long as propod . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. yui Chan & Crosnier, 1991 Rostrum armed dorsally with widely set teeth at basal region and closely set teeth at distal region . . . . . . . . . . . . . . . . . . . . . 30 Rostrum armed dorsally with widely set teeth along its entire length or with very few teeth (1–5) after antennular peduncle level . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 Pereopods 3–4 without epipods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .P. adensameri (Balss, 1914) Pereopods 3–4 with epipods reduced or well developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 Pereopods 3–4 with well-developed epipods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. spinensis Chace, 1985 Pereopods 3–4 with reduced epipods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 Rostrum dorsally armed with one single fixed tooth before post-orbital margin leveland 16–29 teeth posterior to post-orbital 578 · Zootaxa 4221 (5) © 2017 Magnolia Press AHAMED ET AL. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42. 43. 44. - margin level, ventrally armed with 30–43 teeth on ventral border . . . . . . . . . . . . . . . . . . . . . . . . . P. crosnieri Chan & Yu, 1991 Rostrum dorsally armed with 2–4 movable teeth before post-orbital margin level and 27–36 teeth posterior to post-orbital margin level, ventrally armed with 37–52 teeth on ventral border . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. edwardsii (Brandt, 1851) Rostrum armed ventrally with less than 10 teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 Rostrum armed ventrally with more than 10 teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 Rostrum armed dorsally with 5 teeth, anterior 3 with barbed tips . . . . . . . . . . . . . . . . . . . . . P. longidactylus Li & Komai, 2003 Rostrum dorsal teeth not barbed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 Rostrum armed dorsally with 5–10 teeth (including that before post-orbital margin level) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. bifurca Alcock & Anderson, 1894 Rostrum armed dorsally with more than 10 teeth (including that before post-orbital margin level) . . . . . . . . . . . . . . . . . . . . 36 Pereopod 3 with very short dactyl 0.08 to 0.16 times as long as propod . . . . . . . . . . . . . . . . . . . . . . . P. sindoi (Rathbun, 1906) Pereopod 3 with long dactyl 0.25 to 0.33 times as long as propod . . . . . . . . . . . . . . . . . . . . . . . . . . . P. fimbriata Chace, 1985 Rostrum armed ventrally with 10–15 teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 Rostrum armed ventrally with more than 15 teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 Pereopods 2 subequal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 Peropods 2 unequal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 Rostrum dorsally armed throughout entire length with 17–18 teeth . . . . . . . . . . . . . . . . . . . . . . . . . . P. ortmanni Doflein, 1902 Rostrum dorsally armed only until the end of scaphocerite level with 10–12 teeth . . . . . . . . . . . . . . . P. carsini Crosnier, 1986 Rostrum armed dorsally with less than ten teeth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. assimilis De Man, 1917 Rostrum armed dorsally with more than ten teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 Pereopod 2 left carpus with about 110 articles and right carpus with 27 or more articles . . . . . . . . . . .P. binoculus (Bate, 1988) Pereopod 2 left carpus with 50–94 articles and right carpus with 18–24 articles . . . . . . . . . . . . . . . . . . P. izumiae Omori, 1971 Pereopods 2 subequal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. parvimartia Chace, 1985 Peropods 2 unequal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 Rostrum armed dorsally with 6–7 teeth before antennal peduncle margin level and one isolated tooth in anterior ¼ of rostrum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. unidens Bate, 1888 Rostrum armed dorsally throughout entire length with 11 –13 teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 Rostrum armed dorsally with 11–13 teeth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. philippinensis Chace, 1985 Rostrum armed dorsally with 15–23 teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. heterocarpus (Costa, 1871) Comments on the key: Herein we include some comments on aspects that should be considered carefully when using the key above. The genus Plesionika is highly speciose and includes many species groups such as P. narval; P. martia; P. rostricrescentis and P. ocellus groups. Species included on those groups are very similar morphologically. Many Plesionika species also shows great variation in rostrum size and number of teeth according with the size of the specimen. The high basal crest, when present, is more visible in large specimens and in females, in juveniles (ontogenetic variation) and in males (sexual dimorphism in some species) it can be low. We consider that the rostrum dorsal basal crest extends until the end of orbital margin level. When we mention rostrum dorsally armed without any further comment we are considering all dorsal teeth (including that on basal crest and that on subapical teeth, when present). Plesionika ensis and P. reflexa are very similar morphologically and sometimes very hard to identify, for a complete discussion on the distinction between these species see Chan & Crosnier (1997). The same for P. edwardsii and P. crosnieri, see at Chan & Yu (1991) a table with distinctive characters between these two species. For a distinction between P. williamsi and its close related species and also for a complete discussion on the close related species P. sindoi and P. ocellus see Chan & Crosnier (1997). The identification of members of P. martia species group is very complex (here included P. semilaevis, P. orientalis, P. martia and P. taiwanica) for more details on this group see Chace (1985) Crosnier & Forest (1973); Chan & Crosnier (1997); Chan& Yu (2000) and Kim et al.(2012). For the part of the key treating the P. narval species group we used the key of Chan & Crosnier (1991). P. edwardsii and P. crosnieri are extremely similar in morphology (Chan & Yu, 1991); for a more complete discussion see Chan & Yu (1991). Plesionika acinacifer Chace, 1985 Description. Rostrum typically falling short of or slightly overreaching level of distal end of antennal scale, armed dorsally with 5 teeth on basal crest, including 3 movable spines on carapace at or posterior to level of orbital margin, dorsal teeth absent from more than anterior half of rostrum and none of teeth with barbed tips, armed ventrally with 7–13 closely appressed serrations becoming obscure anteriorly; abdomen without posteromesial tooth or median dorsal carina on 3rd somite; eye subpyriform, maximum diameter slightly more than one-fifth of THE GENUS PLESIONIKA BATE, 1888 IN ASIAN WATERS Zootaxa 4221 (5) © 2017 Magnolia Press · 579 carapace length, without true ocellus; 3rd maxilliped with epipod, penultimate segment subequal to terminal segment; 2nd pereopods subequal with epipod, bearing 27–30 carpal articles (Chace 1985). Distribution. Philippine and Malaysia; at depths of 518–790 m (Chace 1985). P. adensameri (Balss, 1914) Description. Rostrum double curved with dorsal teeth distributed over entire length; abdomen without posteromesial tooth or median dorsal carina on 3rd somite; eye broadly subpyriform and ocellus longitudinally oval; 3rd maxilliped long and slender, reaching beyond scaphocerite; 2nd pereopods reaching distal margin of 1st pereiopod (Rajool Shanis et al. 2012). Distribution. Red Sea, Gulf of Aden, Maldives and India; at depths of 200–1308 m (Rajool Shanis et al. 2012). P. alcocki (Anderson, 1896) Description. The rostrum is nearly 1.5 to 2.0 times of the carapace length, it is armed dorsally at its basal end with usually 5 teeth, three or four of which are movable and very small and stand close together on the gastric crest, with two large isolated ones just in front of them, ventrally it is armed throughout, beyond the antennular peduncle, with a series of distant teeth not concealed by setae to the number of from 4 to 8; the 6th abdominal tergum is not quite twice as long as the 5th; there is no distinct ocellus on the dorsal margin of the eye; 1st perepods are not shorter than the external maxillipeds, 2nd perepods in the male alone are very slightly unequal in length: one of them reaches almost to the other very slightly beyond the tips of the external maxillipeds (Alcock 1901). Distribution. Arabian Sea, off the south and south-west coast of India, Bay of Bengal, Andaman Sea, Indonesia, Japan and Philippines; at depths of 496–1092 m (Alcock 1901; Holthuis 1980; De Grave & Fransen 2011). P. albocristata Chan & Chuang, 2002 Description. Rostrum overreaching scaphocerite and 1.2–1.4 times as long as carapace, anterodorsal carapace and basal part of rostrum above orbit strongly elevated and crest-like, basal rostral crest beainng 2 large lamellate fixed teeth and rather horizontal row of 7 or 8 densely packed small movable teeth, all movable teeth situated posterior to orbital margin, other part of dorsal border of rostrum generally devoid of tooth, ventral border of rostrum evenly distributed with 9–13 teeth, those near tip sometimes small and indistinct; eye subglobular and bearing distinct ocellus; 3rd maxilliped bearing well developed epipod and long strip-like exopod, penultimate segment 0.8–0.9 times as long as distal segment, overreaching scaphocerite by half to nearly entire length of distal segment; 2nd pereiopods subequal with 19–24 carpal articles, exceeding scaphocerite by 0.2 to very short length of carpus and entire chela (Chan & Chuang 2002). Distribution. Known only from eastern Taiwan at depths of 350–423 m (Chan & Chuang 2002). Plesionika assimilis De Man, 1917 Description. Rostrum overreaching antennal scale, it is armed dorsally with 5–7 teeth on basal crest, including 4 or 5 on carapace posterior to level of orbital margin, and 3 or 4 subapical teeth, 4 or 5 posterior most teeth with distinct basal sutures but none with barbed tips, armed ventrally with 10–12 teeth; abdomen without posteromesial tooth or median dorsal carina on 3rd somite; eye broadly subpyriform, maximum diameter about l/4 carapace length, ocellus longitudinally oval, constricted at juncture with cornea; 3rd maxilliped with penultimate segment about 7/10 as long as terminal one; 2nd pereopods unequal with epipod and extremely slender or thread-like (Chace 1985) Distribution. Known only from Indonesia at depths of 54–90 m (Chace 1985) 580 · Zootaxa 4221 (5) © 2017 Magnolia Press AHAMED ET AL. P. bifurca Alcock & Anderson, 1894 Description. Rostrum from half to two-thirds the length of the rest of the carapace, it is armed almost all its extent both dorsally with 8 or 9 teeth and ventrally with 3 to 5 teeth, all being fixed; the 3rd abdominal tergum has the posterior border convex, but not acutely produced; the ocellus is quite indistinct; the external maxillipeds are stouter and very much longer than the first pair of pereopods, the 1st pair of pereopods are the shortest of all, and the left pereopod of 2nd pair is the longest of all, reaching far beyond the end of the external maxillipeds and being as long as the carapace and rostrum and first 4 abdominal somites combined (Alcock 1901). Distribution. Arabian Sea, Bay of Bengal, Andaman Sea, Indonesia, Philippines, South and East China seas and Japan; at depths of 220–1412 m (Li & Komai 2003). P. binoculus (Bate, 1988) Description. Rostrum somewhat curved dorsally, distinctly overreaching antennal scale, armed dorsally with 12 irregularly spaced teeth, including four on carapace posterior to level of orbital margin, four posteriormost teeth basally articulated but none with barbed tips, ventral margin of rostrum armed with 13 teeth; 3rd maxilliped bearing hooked epipods with penultimate segment 0.60 times as long as ultimate segment; 2nd pereopods greatly unequal, shorter (right) one with 30 carpal articles, longer (left) one with about 110 carpal articles (Li & Komai 2003). istribution. Arafura Sea and South China Sea; at depths of 90–140 m (Chace 1985; Li & Komai 2003). P. brevirostris Bate, 1888 Description. Rostrum reaching level of distal end of antennal scale, armed dorsally with 6 teeth on dorsal crest, including 2 on carapace posterior to or in line with orbital margin, and obscure serrations near tip of rostrum, none of teeth movable or with barbed tips, armed ventrally with about 10 obscure serrations near apex; abdomen without posteromesial tooth or median dorsal carina on 3rd somite; eye apparently quite broad, about one-fourth carapace length, ocellus not expanded beyond juncture with cornea; 3rd maxilliped with penultimate segment nearly 9/10 as long as terminal one; 2nd pereopods subequal and not extremely slender or thread-like (Chace 1985). Distribution. Western central Pacific: between Philippines and Borneo at depths of 457 m (Chace 1985). P. carsini Crosnier, 1986 Description. Rostrum 1.28–1.61 times length of carapace, only slightly recurved, overreaching the scaphocerite by half or 2/5 of its length, armed with 3–5 dorsal and 11–14 ventral fixed teeth; abdomen without posteromesial tooth or median dorsal carina on 3rd somite; eye sub-pyriform, maximum diameter about 1/5 carapace length, ocellus very clear; 3rd maxilliped bearing epipod, with penultimate segment as long as distal one; 2nd pereiopods subequal with 19–21 carpal articles (Chan & Crosnier 1997). Distribution. Known only from Taiwan at depths of 300 m (Chan & Crosnier 1997). P. crosnieri Chan & Yu, 1991 Description. Rostrum curved downwards basally, then strongly curved upwards beyond antennular peduncle, and occasionally bending slightly downwards again near apex, far overreaching scaphocerite, 1.8–2.5 times as long as carapace, dorsal border armed with 16–29 teeth over entire length, anterior teeth closely set but posterior ones larger and spaced farther apart, ventral border bearing 30–43 closely packed teeth on upturned section; dorsal surface of abdominal 3rd somite slightly arched but not sharply angular; eye large and subspherical, with distinct ocellus; 3rd maxilliped with distal two segments subequal, bearing rudimentary epipod and extending to about distal margin of scaphocerite; 2nd pereiopods subequal with 19–27 carpal articles (Chan & Yu 1991). THE GENUS PLESIONIKA BATE, 1888 IN ASIAN WATERS Zootaxa 4221 (5) © 2017 Magnolia Press · 581 Distribution. Indo-west-Pacific and southern Ocean, Crozet Islands, Indonesia, Philippines, Taiwan and Japan; at depths of 80–350 m (Chan & Yu 1991). P. edwardsii (Brandt, 1851) Description. Rostrum curved downward in basal region, then strongly curved upwards beyond antennular peduncle, slightly more than 2 times as long as carapace, armed dorsally with 28–36 teeth over entire length, anterior teeth closely spaced, posterior teeth larger and well spaced, ventral border bearing 37–52 closely spaced teeth; dorsal surface of abdominal 3rd somite slightly arched but not sharply angular; eye large and subspherical, with distinct ocellus; 3rd maxilliped with distal two segments subequal, bearing rudimentary epipod; 2nd Pereiopods subequal with 18–28 carpal articles (Chace 1985). Distribution. Indo-Pacific, Indonesia, Philippines and Taiwan; at depths of 50–680 m (Chace 1985). P. ensis (Milne-Edwards, 1881) Description. Rostrum nearly 1.6 to 2 times of the carapace length, armed dorsally with 5 or 6 teeth, three or four close together, two isolated anteriorly, ventrally it is closely and evenly serrated, the teeth are fixed; the posterior border of the 3rd abdominal tergum is acutely produced in the middle line to form a spine; the ocellus is distinct but not independent; the external maxillipeds with well-developed exopodite, little longer and stouter than the 1st pair of pereopods, reach a short way beyond the tip of the antennal scale (Alcock 1901). Distribution. Arabian Sea, Bay of Bengal and Andaman Sea; at depths of 250–400 m (Alcock 1901). P. erythrocyclus Chan & Crosnier, 1997 Description. Rostrum more or less as long as carapace and just overreaching scaphocerite, curving downward at base but abruptly upturned after passing the eye, dorsal border with only 2 small apical teeth and 1–2 fixed basal teeth above eye, ventral border with 7–9 evenly distributed teeth between level of eye and tip; eye subspherical and bearing distinct ocellus; 3rd maxilliped bearing well developed epipod and a long exopod, with penultimate segment 0.6–0.7 times as long as distal segment, overreaching scaphocerite by 1/3–1/2 of distal segment; 2nd pereiopods bearing 20–22 carpal articles on the right side and 76–83 carpal articles on the left side (Chan & Crosnier 1997). Distribution. Known only from Taiwan at depths of 267–360 m (Chan 2004). P. fimbriata Chace, 1985 Description. Rostrum far overreaching antennal scale, armed dorsally throughout length with 11–14 teeth, including 3 or 4 on carapace posterior to level of orbital margin, 2–4 posteriormost teeth with distinct basal sutures but none with barbed tips, armed ventrally with 5–8 teeth; abdomen without posteromesial tooth or median dorsal carina on 3rd somite; eye broadly subpyriform, maximum diameter about one-fourth carapace length, ocellus large, subcircular, in broad contact with cornea; 3rd maxilliped with epipod, penultimate segment 1.2–1.5 times as long as terminal segment; 2nd peropods subequal with epipod, bearing 18–30 carpal articles (Chace 1985). Distribution. Known only from the Philippines at depths of 95–250 m (Chace 1985). P. grandis Doflein, 1902 Description. Rostrum moderately curving dorsally, 1.4–1.7 times as long as carapace; dorsal margin with 37–44 teeth, including 6 on carapace, 2 posteriormost teeth with distinct basal sutures, ventral margin with 21–24 teeth; abdomen smooth and rounded dorsally; eye subpyriform, maximum diameter more or less than 0.2 carapace length, cornea broader than eyestalk; 3rd maxilliped slender, overreaching distal margin of antennal scale by entire ultimate segment and anterior 0.1 penultimate segment, without epipod; 2nd pereopods subequal without epipod, bearing 22–23 carpal articles (Kim et al. 2012). 582 · Zootaxa 4221 (5) © 2017 Magnolia Press AHAMED ET AL. Distribution. Indo-West Pacific, Korea, Japan, East and South China seas, Philippines and Indonesia; at depths of 92–375 m (Kim et al. 2012). P. heterocarpus (Costa, 1871) Description. Rostrum long, about two times as long as scaphopcerite, slender and slightly curved upward, dorsal margin with 15–23 teeth of which the proximal 4–6 are movable and situated behind the orbit, ventral margin bears 16–23 teeth; posterior part of the 3rd abdominal somite is slightly concave; eye with maximum diameter about l/5 of carapace length, ocellus longitudinally oval, constricted at juncture with cornea; 3rd maxilliped with well developed strap-like epipod, penultimate segment sub-equal to terminal segment; 2nd pair of pereiopod distinctly unequal, left one much longer than the right one, overreaching scaphocerite with chela, carpus and distal half of merus, with 107–215 carpal segments, 80–111 meral segments and about 25 ischial segments (Holthuis 1980). Distribution. Cyprus, Israel, Syria and Turkey; at depths of 10–850 m (Holthuis 1980). P. hsuehyui Chan, 2004 Description. Rostrum 1.3–2.0 times as long as carapace, bending downwards near base but recurved upwards after passing eyes, dorsal rostrum proper unarmed except for 2 small subapical teeth, ventral rostrum generally bearing 9 or 10 teeth; abdominal 3rd somite slightly arched dorsally; 3rd maxilliped overreaching scaphocerite by up to 2/3 length of distal segment, distal segment 1.4–1.8 times as long as penultimate segment; 2nd pereopods bearing 15– 22 carpal articles on right side and 53–70 carpal articles on left side, shorter one exceeding scaphocerite by about half chela (Chan 2004). Distribution. Taiwan and Philippines; at depths of 115–300 m (Chan 2004). P. hypanodon Doflein, 1902 Description. Rostrum is only half as long as the carapace and quite straight, except for the tip, which is bent upwards, dorsal margin with 7 teeth, the anterior three are behind the eye basis, smaller and stand narrower to each other than the other 4, which are arranged in similar distances towards the tip, ventral margin is smooth and without teeth; abdomen is smooth, compressed, and the hindmargin of the third segment is slightly recessed; eyes are spherical-shaped, semi-large with distinct ocellus; 2nd pereiopods equally long and slender with 15 carpal articles (Doflein 1902). Distribution. Known only from Sagami Bay, Japan (Doflein 1902). P. indica De Man, 1917 Description. Rostrum 1.8 times as long as carapace, far overreaching antennal scale, armed dorsally throughout length with 27–33 teeth, including 4–6 on carapace posterior to level of orbital margin, 2–4 posteriormost teeth with distinct basal suture but none with barbed tips, armed ventrally with 22–27 teeth; abdomen without posteromesial tooth or median dorsal carina on 3rd somite; eye subpyriform, maximum diameter fully 1/6 carapace length, ocellus prominent, somewhat constricted at juncture with cornea; 3rd maxilliped with epipod, penultimate segment 1.2–1.4 times as long as terminal segment; 2nd pereopods subequal with prominent epipods, bearing 31– 34 carpal articles (De Man 1920; Chace 1985). Distribution. Indonesia, South China Sea, Philippines and Japan; at depths of 238–472 m. (Li & Komai 2003). THE GENUS PLESIONIKA BATE, 1888 IN ASIAN WATERS Zootaxa 4221 (5) © 2017 Magnolia Press · 583 P. intermedia Chace, 1985 Description. Rostrum far overreaching antennal scale, armed dorsally on basal crest with 6 or 7 teeth, including 3 on carapace posterior to level of orbital margin, 1 or more posteriormost teeth with faint, incomplete basal sutures but none movable or with barbed tips, armed ventrally with 34–43 teeth; abdomen without posteromesial tooth or median dorsal carina on 3rd somite; eye considerably broader than long, maximum diameter fully 1/5 carapace length, ocellus represented by tapering, subtruncate lobe, broadening rather than constricted at base; 3rd maxilliped with epipod, penultimate segment nearly 1.2 times as long as terminal segment; 2nd pereopods subequal with 19 or 20 carpal articles (Chace 1985). Distribution. Known only from the Philippines at depths of 472–574 m (Chace 1985). P. izumiae Omori, 1971 Description. Rostrum noticeably curving dorsally, 1.5–1.7 times as long as carapace, dorsal margin with 11–13 teeth, including 4 on carapace, 5–7 posteriormost teeth movable, ventral margin with 13–14 teeth; abdomen rounded dorsally, pleura of fourth and fifth somites with posteroventral tooth, sixth somite 1.7 times as long as maximum height; eye moderately large, maximum diameter about 0.2 carapace length; 3rd maxilliped overreaching antennal scale by half of ultimate segment, penultimate segment 0.7 as long as ultimate segment, with epipod; 2nd pereopods very unequal with prominent epipod and not extremely slender or threadlike (Omori 1971). Distribution. Japan, Korea, South and East China seas and Philippines; at depths of 17–300m (Kim et al. 2012). P. kensleyi Chace, 1985 Description. Rostrum not overreaching antennal scale, armed dorsally throughout length with 9–13 teeth, including 2–4 on carapace posterior to level of orbital margin, 2 or 3 posteriormost teeth with basal sutures but none with barbed tips, armed ventrally with 3–5 teeth in anterior half of rostrum; abdomen without posteromesial tooth or median dorsal carina on 3rd somite; eye subpyriform, maximum diameter about one-fourth carapace length, ocellus prominent, only slightly constricted at juncture with cornea; 3rd maxilliped with epipod, penultimate segment 1.4–1.5 times as long as terminal segment; 2nd pereopods subequal with well-developed epipods, bearing 18–23 carpal articles (Chace 1985). Distribution. South China Sea and Philippines; at depths of 118–333 m (Li 2006a). P. longicauda (Rathbun, 1901) Description. Rostrum 1.7–2.1 times as long as carapace, with basal region horizontal or slightly upturned but lacking ventral notch, curved slightly and directed dorsad, armed with 36–46 dorsal and 25–33 ventral teeth, rostral teeth somewhat well-spaced and with posterior 10 ventral teeth corresponding to 8–10.5 dorsal teeth; abdominal 6th somite with longitudinal dorsal groove more or less pronounced ; 3rd maxilliped without epipod, penultimate segment 1.3–1.55 times longer than terminal segment, two segments combined 1.1–1.35 times as long as carapace; 2nd pereiopods subequal without epipod, bearing 18–26 carpal articles (Chan & Crosnier 1991). Distribution. Known only from Bay of Bengal at depths of 10–50 m (Rajool Shanis et al. 2012). P. longidactylus Li & Komai, 2003 Description. Rostrum far overreaching antennal scale, 1.1–1.2 times as long as carapace, armed dorsally with 5 moderately large teeth, each with obscurely barbed tip and with 2 tiny subapical teeth, ventral margin with 10–13 teeth; 3rd abdominal somite not compressed dorsally, unarmed on posterodorsal margin; 3rd maxilliped with ultimate segment about 1.4 times as long as penultimate segment; 2nd pereopods greatly unequal with well 584 · Zootaxa 4221 (5) © 2017 Magnolia Press AHAMED ET AL. developed strap-like epipod each bearing terminal hook, left with 25–30 carpal articles and right with 8 carpal articles (Li & Komai 2003). Distribution. Known only from South China Sea at depths of 55–144 m (Li 2006a). P. lophotes Chace, 1985 Description. Rostrum remarkably curving dorsally, 1.5 times as long as carapace, dorsal margin with 14 teeth including 5–6 on carapace, all teeth on carapace with distinct basal sutures and movable, ventral margin with 13– 18 teeth; abdomen smooth and rounded dorsally, pleura of fourth and fifth somites with small marginal tooth posteriorly sixth somite 1.5 times as long as maximum height; eye broadly subpyriform, maximum diameter about 0.2 carapace length; 3rd maxilliped overreaching distal margin of antennal scale by half of ultimate segment, with epipod, penultimate segment about 0.7 as long as ultimate segment; 2nd pereopods unequal with epipods, not extremely slender or thread-like, left overreaching antennal scale by distal 3 segment and anterior 0.3 of merus, with 147 carpal articles, right overreaching antennal scale by entire chela and anterior 0.7 of carpus, with 40 carpal articles (Kim et al. 2012). Distribution. Indo-West Pacific: Korea, Japan, Philippines, Vietnam and southern Arabia; at depths of 105– 329 m (Kim et al. 2012). P. martia (Milne-Edwards, 1883) Description. Rostrum nearly 1.6 to 2.0 times as long as carapace, basal portion armed dorsally with 5 to 8 teeth, of which 5 or 6 form a series that gradually increase in size from behind forwards while the anterior two are usually somewhat isolated, ventrally it is very closely finely and evenly serrated, the teeth being a good deal concealed in a short fringe of cilia, all the teeth are fixed; the posterior border of the 3rd abdominal tergum though convex, is not acutely produced; the ocellus is distinct but not independent; the external maxillipeds with well-developed exopodite, which are a little longer and stouter than the 1st pair of pereopods, reach a short way beyond the tip of the antennal scale (Alcock 1901). Distribution. Andaman Sea, Bay of Bengal, Indian Ocean and Arabian Sea; at depths of 180–400 m (Alcock 1901). P. narval (Fabricius, 1787) Description. Rostrum slightly curving dorsally, elongate, 2.2–2.3 times as long as carapace, dorsal margin with 58–60 serrated teeth, including 5 on carapace, all teeth on carapace with distinct basal sutures and movable, ventral margin with 41–44 serrated teeth; abdomen smooth and rounded dorsally, pleuron of fourth somite rounded that of fifth somites with small marginal tooth posteriorly, sixth somite 1.8 times as long as maximum height; eye broadly subpyriform, maximum diameter about 0.2 carapace length; 3rd maxilliped slender, overreaching distal margin of antennal scale by ultimate segment and anterior 0.2 of penultimate segment, with epipod, penultimate segment about 1.7 as long as ultimate segment; 2nd pereopods subequal without epipod, not extremely slender or threadlike, bearing 27–29 carpal articles (Kim et al. 2012). Distribution. Indo-West Pacific, Indian Ocean, Japan, Korea and Red Sea; at depths of 35–910 m (Kim et al. 2012). P. orientalis Chace, 1985 Description. Rostrum moderately curving dorsally, elongate, 2.2 times as long as carapace, dorsal margin with 8 teeth including 3 on carapace, all teeth on partial basal sutures, ventral margin with 42 serrated teeth; abdomen smooth and rounded dorsally, pleura of first to fifth somites without distinct marginal tooth or denticle, sixth somite THE GENUS PLESIONIKA BATE, 1888 IN ASIAN WATERS Zootaxa 4221 (5) © 2017 Magnolia Press · 585 about 2.2 times as long as maximum height; eye very broadly subpyriform, maximum diameter aobut 0.25 carapace length; 3rd maxilliped overreaching distal margin of antennal scale by anterior 0.3 of ultimate segment, with epipod, well developed exopod present, penultimate segment 1.3 times as long as ultimate segment; 2nd pereopods subequal with epipod, not extremely slender or thread-like, bearing 18–21 carpal articles (Kim et al. 2012). Distribution. Korea, Japan, East and South China seas, Philippines and Indonesia; at depths of 66–686 m (Kim et al. 2012). P. ortmanni Doflein, 1902 Description. Rostrum rather curving dorsally, 1.5–1.6 times as long as carapace, dorsal margin with 17–18 teeth including 3–4 on carapace, all teeth with partial or complete basal sutures, ventral margin with 6–9 teeth; abdomen without carina or projection dorsally, pleuron of fifth somite with posteroventral tooth, sixth somite 1.5 times as long as maximum height; eye subpyriform, maximum diameter barely 0.2 carapace length; 3rd maxilliped overreaching distal margin of antennal scale by entire ultimate segment and anterior 0.2 of penultimate segment, with epipod, penultimate segment 1.4 as long as ultimate segment; 2nd pereopods subequal with prominent epipod, not extremely slender or thread-like, bearing 28–33 carpal articles (Kim et al. 2012). Distribution. Korea, Japan, East and South China Seas, Philippines and Indonesia; at depths of 29–400 m (Kim et al. 2012). P. parvimartia Chace, 1985 Description. Rostrum far overreaching antennal scale, armed dorsally, on posterior section only, with 6–9 teeth, including 2 or 3 on carapace posterior to level of orbital margin, none with distinct basal sutures or barbed tips, armed ventrally with 25–50 teeth; abdomen without posteromesial tooth or median dorsal carina on 3rd somite; eye somewhat kidney-shaped, maximum diameter from more than l/4 to 1/3 carapace length, ocellus represented by ungulate lobe, slightly broadening rather than constricted at junctiure with cornea; 3rd maxilliped with epipod, penultimate segment from 9/8 to more than 4/3 times as long as terminal segment; 2nd pereopods subequal with epipod, bearing 18–25 carpal articles (Chace 1985). Distribution. Known only from the Philippine at depths of 176–366 m (Chace 1985). P. persica (Kemp, 1925) Description. Rostrum is slightly shorter than the carapace and reaching just to the end of the antennal scale, downwards in its proximal half but the distal half is straight or slightly inclined upwards, dorsal margin with 7–8 fixed teeth, 3 of which are placed on the carapace behind the orbit, ventral margin bears 9–10 small teeth; abdomen without posteromesial tooth or median dorsal cariana on 3rd somite; eye broadly subpyriform, maximum diameter fifth of carapace length; 3rd maxilliped with slender exopod, penultimate segment 4/3 times as long as the ultimate segment; 2nd pereopods are equal and reached almost to the end of the scale, the merus is composed of 2 segments of which the distal is about 3 times as long as the proximal (Kemp 1910; Fransen 2006). Distribution. Gulf of Oman, Red Sea, South Arabian Coast and Gulf of Aden; at depths of 25–955 m (Fransen 2006; De Grave & Fransen 2011). P. philippinensis Chace, 1985 Description. Rostrum distinctly overreaching antennal scale, armed dorsally throughout length with 11–13 teeth including 4 to 5 on carapace posterior to orbital margin, the fifth one usually just above the margin, 4–6 posteriormost teeth with distinct basal suture, and ventrally with 10–16 teeth; fourth and fifth abdominal somites 586 · Zootaxa 4221 (5) © 2017 Magnolia Press AHAMED ET AL. each with posteroventral tooth on pleuron; ocellus skewed somewhat laterad, in rather broad contact with cornea but distinctly constricted at juncture with cornea; 3rd maxilliped with penultimate segment 0.75–0.80 times as long as terminal segment, with hooked epipods; 2nd pereopods strongly unequal, right (shorter) one with 18 carpal articles, left (longer) one with 92 carpal articles (Li & Komai 2003). Distribution. Philippines and South China Sea; at depths of 103–135 (Li & Komai 2003). P. pumila Chace, 1985 Description. Rostrum armed dorsally with 4 teeth in basal part, including one on the carapace posterior to orbital margin, all with bluntly barbed tips; fourth and fifth abdominal somites each with posteroventral tooth on pleuron; 3rd maxilliped with epipod, penultimate segment 0.60 as long as terminal segment; 2nd pereopods with hooked epipods, left 2nd pereopods with carpus consisting of about 37 articles (Li & Komai 2003). Distribution. Philippines and South China Sea; at depths of 20–79 m (Li & Komai 2003). P. quasigrandis Chace, 1985 Description. Rostrum 1.1–1.5 times as long as carapace, densely armed with abutting fixed teeth along almost entire dorsal and ventral margins, bearing 38–53 dorsal (including 4–7 teeth on carapace) and 31–44 ventral teeth; posterior 10 ventral teeth corresponding to 5.5–8 dorsal teeth; abdomen with 3rd somite posteriorly unarmed, without median dorsal carina; eye with cornea wider than long and with distinct ocellus; 3rd maxilliped without epipod, overreaching scaphocerite by terminal segment and little penultimate segment, penultimate segment 1.2– 1.4 times as long as terminal segment, 2 segments combined more or less as long as carapace; 2nd pereiopods subequal without epipods, bearing 21–22 carpal articles (Chakraborty et al. 2015). Distribution. Indo-West Pacific from Japan to the Philippines, Indonesia, India and Gulf of Aden; at depths of 164–501 m (Chakraborty et al. 2015). P. reflexa Chace, 1985 Description. Rostrum far overreaching antennal scale, usually armed dorsally with 6–8 teeth, including 2 or 3 on carapace posterior to level of orbital margin and 1 subapical tooth, otherwise without dorsal teeth anterior to level of distal end of antennular peduncle, posteriormost tooth usually with indistinct basal suture but none with barbed tips, armed ventrally with 26–48 teeth; abdomen with strong, frequently recurved posteromesial tooth but without median dorsal carina on 3rd somite; eye very broadly subpyriform, maximum diameter nearly 1/3 carapace length, ocellus Ungulate, not constricted at juncture with cornea; 3rd maxilliped with epipod, penultimate segment about 9/10 as long as terminal segment; 2nd pereopods subequal with well-developed epipod, bearing 15–26 carpal articles (Chace 1985). Distribution. Indo west Pacific, South China Sea, Eastern China Sea, Indonesia and Philippine; at depths of 315–494 m (Chace 1985). P. rufomaculata Chan, 2004 Description. Rostrum curving downwards near base but recurved upwards after passing eyes, 1.1–1.6 times as long as carapace, basal rostral crest high in females and slightly lower in males, bearing 2 or 3 fixed and 6 movable teeth, posterior 5 or 6 of them situated posterior to orbital margin, dorsal rostrum proper bearing 1 or 2 teeth, usually restricted to proximal half of rostrum, and 2 (rarely 1) small apical teeth, 11–13 teeth generally distributed on ventral rostrum; abdominal 3rd somite slightly arched dorsally; 3rd maxilliped with distal segment 1.4–1.6 times as long as penultimate segment, overreaching scaphocerite by 1/2–2/3 length of distal segment; 2nd pereiopods bearing 21–30 carpal articles on right side and 106–121 carpal articles on left side, shorter one exceeding scaphocerite by 1/2–1/4 length ol carpus and entire chela (Chan 2004). THE GENUS PLESIONIKA BATE, 1888 IN ASIAN WATERS Zootaxa 4221 (5) © 2017 Magnolia Press · 587 Distribution. Japan and Taiwan; at depths of 180–300 m (Chan 2004). P. semilaevis Bate 1888 Description. Rostrum far overreaching antennal scale, armed dorsally, on basal crest only, with 5–9 teeth, including 2 or 3 on carapace posterior to level of orbital margin, none with distinct basal suture or barbed tip, armed ventrally with 19–52 teeth; abdomen without posteromesial tooth or median dorsal carina on 3rd somite; eye kidney-shaped, maximum diameter usually nearly 1/3 carapace length, ocellus represented by tapering, subtruncate lobe, broadening rather than constricted at juncture with cornea; 3rd maxilliped with epipod, penultimate segment from subequal to 7/5 as long as terminal segment; 2nd pereopods subequal with epipods, bearing 19–36 carpal articles (Chace 1985). Distribution. Indonesia, Philippines, South and East China Seas and Japan; at depths of 176–700 m (Li 2006a). P. simulatrix Chace, 1985 Description. Rostrum far overreaching antennal scale, armed dorsally, on basal crest only, with 6–9 teeth, including 2–4 on carapace posterior to level of orbital margin, 1 or more posteriormost teeth sometimes with faint, incomplete basal suture, none with barbed tips, armed ventrally with 29–49 teeth; abdomen without posteromesial tooth or median dorsal carina on 3rd somite; eye broadly subpyriform, maximum diameter about 1/4 carapace length, ocellus represented by tapering, subtruncate lobe, broadening slightly rather than constricted at juncture with cornea; 3rd maxilliped with epipod, penultimate segment from 11/10 to more than 4/3 times as long as terminal segment; 2nd pereopods subequal without epipod, bearing 14–21 carpal articles (Chace 1985). Distribution. Known only from the Philippine at depths of 216–472 m (Chace 1985). P. sindoi (Rathbun, 1906) Description. Rostrum 1.5 times as long as carapace, from the orbital margin the rostrum runs horizontally forward as far as the distal extremity of the antennular peduncle and from here it is a little and obliquely turned upward, 5 movable teeth on the carapace, lower margin is armed with 6 teeth; abdomen nearly 4-times as long as the carapace; the eyes are little more than one-fourth the length of the carapace and this diameter is slightly longer than the axial; external maxillipeds reach by their terminal joint and one-fourth of the penultimate beyond the antennal scale, exopodite small, not yet reaching to the middle of the antepenultimate joint; 1st pereopods reach by 4/5 their terminal joint (or propodus ) beyond the external maxillepeds the penultimate joint or carpus is almost twice as long as the terminal, these joints being respectively 9.1 mm. and 5 mm. long in the adult female (Chace1985). Distribution. Japan, South China Sea, Philippines and Indonesia; at depths of 122–800 m (Li & Komai 2003). P. spinensis Chace, 1985 Description. Rostrum far overreaching antennal scale, armed dorsally throughout length with 13–24 teeth, including 2 or 3 on carapace posterior to level of orbital margin, posteriormost tooth with faint, incomplete basal suture, none with barbed tips, armed ventrally with 2539 teeth; abdomen typically with posteromesial tooth but without median dorsal carina on 3rd somite; eye very broadly subpyriform, maximum diameter nearly or quite 1/3 carapace length, ocellus subcircular, somewhat constricted at juncture with cornea; 3rd maxilliped with epipod, penultimate segment slightly shorter or longer than terminal segment; 2nd pereopods subequal with welldeveloped epipod, bearing 14–23 carpal articles (Chace 1985). Distribution. South China Sea and Philippines; at depths of 199–472 m (Li 2006a) 588 · Zootaxa 4221 (5) © 2017 Magnolia Press AHAMED ET AL. P. spinidorsalis (Rathbun, 1906) Description. Rostrum overreaching antennal scale, armed dorsally throughout length with 12–15 teeth, including 7–10 on carapace posterior to level of orbital margin, none with basal suture or barbed tip, armed ventrally with 4– 8 teeth; orbital margin with subquadrate lobe, thence trending posterodorsally in nearly straight line and, finally, bending sharply anterodorsally at extreme dorsal limit; abdomen without posteromesial tooth or median dorsal carina on 3rd somite; eye pyriform, maximum diameter about 1/8 carapace length, without ocellus; 3rd maxilliped with epipod, penultimate segment slightly more than 1/2 to 2/3 as long as terminal segment; 2nd pereopods unequal and dissimilar with epipod, right one shorter, more robust, and with 5 or 6 carpal articles, left one with 13– 16 carpal articles (Chace 1985). Distribution. South China Sea, Philippines and Indonesia; at depths of 100–1250 m (Li 2006a). P. taiwanica Chan & Yu, 2000 Description. Rostrum 1.3–2.1 times as long as carapace, slightly curving downward at basal part but gently recurved upwards and straight after passing antennular peduncle, dorsal border generally armed with 6–8 teeth, with posterior 4–6 teeth restricted at basal part and forming a low crest while the anterior 1–3 teeth located anterior to level of antennular peduncle, lower border densely serrated with 22–38 small teeth; abdominal 3rd somite feebly arched dorsally and with posterior margin convex; eye subspherical and probably bearing indistinct ocellus; 3rd maxilliped bearing well developed epipod and long strip-like exopod, penultimate segment 1.2–1.5 times as long as distal segment, overreaching scaphocerite by 1/3 to almost entire distal segment; 2nd pereiopods subequal with well developed epipod, bearing 16–25 carpal articles (Chan & Yu 2000). Distribution. Known only from Taiwan at depths of 150–300 m (Chan & Yu 2000). P. unidens Bate, 1888 Description. Rostrum far overreaching antennal scale, armed dorsally with 7 or 8 teeth, including 3 on carapace posterior to level of orbital margin and 1 isolated in anterior 1/4 of rostrum, 3 posteriormost teeth with distinct basal sutures but none with barbed tips, armed ventrally with 10–20 teeth; abdomen with 3rd somite lacking posteromesial tooth but with median dorsal carina typically forming obtuse tooth anterior to posterior margin of somite; eye kidney-shaped, maximum diameter about 1/3 carapace length, ocellus obliquely oval, slightly constricted at junction with cornea; 3rd maxilliped with epipod, penultimate segment about 4/5 as long as terminal segment; 2nd pereopods very unequal with epipods, left one longer with more than 200 carpal articles, right with 33–36 (Chace 1985). Distribution. Bay of Bengal, South and East China seas, Japan, Philippines and Indonesia; at depths of 184– 400m (Li 2006b). P. williamsi Forest, 1963 Description. Rostrum long, extending far beyond scaphocerite, about 1.5 times as long as carapace, curved downwards upto middle of antennular peduncle and thereafter gradually ascending, armed with 11 dorsal teeth arranged closely on basal crest, proximal 3 or 4 placed behind orbit, first tooth microscopic and movable, next 2 or 3 of smooth, ventral margin serrated with 11–15 teeth spaced more widely than dorsal; all abdominal segements non-carinate; eyes large with distinct ocellus; 3rd maxillipeds with well developed exopods, surpassing scaphocerite by whole length of utlimate and 1/4 of penultimate segments, ultimate segment shorter than penultimate, exopod reaching about middle of basal article, epipod small and terminating in a sickle-shaped hook and laterally bent spine; 2nd pereopods equal, longer and slender, reaching tip of rostrum, carpus divided into 27– 30 joints, merus slightly more than half, the length of carpus (Suseelan 1990). Distribution. Indian Ocean, southwest coast of India, Japan and Taiwan; at depths of 300–455 m (Suseelan 1990; Komai et al. 2005). THE GENUS PLESIONIKA BATE, 1888 IN ASIAN WATERS Zootaxa 4221 (5) © 2017 Magnolia Press · 589 P. yui Chan & Crosnier, 1991 Description. Rostrum directed somewhat dorsad, 0.95–1.85 times as long as carapace and armed with 26–45 dorsal teeth and 18–40 ventral teeth, posterior 10 ventral teeth corresponding to 4–8 dorsal teeth; 3rd maxilliped with rudimentary epipod, penultimate segment 1.4–1.6 times longer than terminal segment, two segments combined slightly longer than carapace; 2nd pereiopods subequal with 21–38 carpal articles (Chan & Crosnier 1991). Distribution. Known only from Taiwan at depths of 130 m (Chan & Crosnier 1991). Biological aspects Numerous studies have been conducted on the biological aspects focused mainly in the Mediterranean Sea and northeastern Atlantic Ocean (Ahamed & Ohtomi, 2011) of Plesionika spp. recorded from European waters (Vafidis et al. 2005). However, a few studies (Table 2) have been conducted on the biological aspects (e.g., reproduction, growth etc.) from Asian waters though near about 50% species of this genus inhabit this region. TABLE 2. Biological characteristics of some Plesionika species reported from Asian waters. Species Location Sexual maturity (female) Fecundity P. ensis India EEZ 83b** 2625 P. izumiae Kagoshima Bay 8.5a 4405 P. martia India EEZ P. quasigrandis south-west coast of India P. semilaevis Kagoshima Bay P. spinipes India EEZ b** 80 b** 81 Longevity References (years) Rajool Shanis et al. 2012 1.5 2833 Ahamed & Ohtomi 2011, 2012, 2014 Rajool Shanis et al. 2012 17376 3.0 Chakraborty et al. 2014 8702 3.0 Ohtomi 1997 4657 Rajool Shanis et al. 2012 a sexual maturity based on the 50% method; b sexual maturity based on the smallest size of ovigerous female; * mm CL; ** mm TL. Usually Plesionika spp. have continuous reproduction throughout the year (Orton 1920; Bauer 1989; Ahamed & Ohtomi 2011). However, there is an increasing seasonality in the reproductive periods when going from the shallowest living species to the deepest species (Company & Sardà 1997). Increasing seasonality in the reproductive periods also occur for the species inhabiting subtropical and tropical regions than those inhabiting temperate waters (Bauer 1992), because of relatively stable and high temperature in the subtropical and tropical regions (Orton 1920). In comparison between Asian and Atlantic species, both have an extended reproductive period. Because the Asian region has subtropical and tropical climate, on the other hand Atlantic species are deeper water species compared to Asian species though the Atlantic climate is very cold with elongated winter. Company & Sardà (1997) reported that continuous reproduction throughout the year of Plesionika species is a general trend of life-history adaptation with depth. The longevity of Plesionika spp. varied from about one year and a half to four years (Company & Sardà 2000; Maiorano et al. 2002). The longevity varies depending on the body size and the depths of distribution. Larger-sized species inhabit deeper waters compared to smaller-size species (King & Butler 1985; Company & Sardà 1997) and deeper water species exhibit longer life cycles compared to shallow water species (King & Butler 1985; Ahamed & Ohtomi 2012). Therefore, the species inhabiting Atlantic are larger with longer lifespan compared to Asian species. Conclusion The exact identification of many species of Plesionika are still confusing due to large number of species or similar forms with wide geographical distribution (Chan & Crosnier 1991, 1997; Chan & Yu 2000). Morphological 590 · Zootaxa 4221 (5) © 2017 Magnolia Press AHAMED ET AL. variation is observed among specimens from different localities. Therefore, doubtful identification may occur sometimes. Several species of Plesionika have been misidentified earlier from Asian waters. For example, P. ocellus reported by Chace (1995) actually represents P. sindoi (Chan & Crosnier 1997). Similarly P. rostricrescentis and P. serratifrons reported by Chace (1985) are also doubtful identification because of their restricted distribution and are only found in the South-West Pacifie (Chan & Crosnier 1991; Chan 2004). In Indian waters, one of the most dominant Plesionika species was previously reported as “Parapandalus (or Plesionika) spinipes” (Suseelan & Mohamed 1968; Suseelan 1974; Rajan et al. 2001; Kurup et al. 2008; Rajool Shanis et al. 2012) which is now known to be a misidentification of P. quasigrandis (Chakraborty et al. 2015). Several studies (Chan & Crosnier 1991, 1997; Chan 2004) reported that coloration in life may also be useful in recognizing species. Therefore, accumulation of data on both morphology and coloration may eventually be able to clarify the taxonomy of this genus. Finally, the present paper will provide an important basis for the shrimp taxonomist/ biologist working in this region and other parts of the world as well. Though near about 50% species of Plesionika genus inhabit Asian waters, studies on species from this region are limited to taxonomic report and very few studies have been conducted on the biological aspects on a few species. Consequently the lack of information and adequate knowledge on the biology of these species remains an obstacle for the definition of proper management strategies for these shrimp fisheries. Therefore, studies on the biological aspects of these species are needed to manage these populations properly as well as to compare the results with the populations studied from other parts of the world. Acknowledgments The authors would like to express their gratitude to Dr. Verena Kutschera, Biosystematic Documentation, University of Ulm, Germany for helping with translation of articles written in German language to English. Thanks are also due to i) Prof. Dr. Tin-Yam Chan, National Taiwan Ocean University, Taiwan and ii) Dr. Charles Fransen, Senior Researcher, Department of Taxonomy & Systematics, Naturalis Biodiversity Center, Netherlands for sending us some valuable articles to complete this review work. TWAS research grant (Ref: 15-082 RG/BIO/ AS_I—FR3240287053) is also acknowledged. References Ahamed, F. & Ohtomi, J. (2011) Reproductive biology of the pandalid shrimp Plesionika izumiae (Decapoda: Caridea). Journal of Crustacean Biology, 31, 441–449. https://doi.org/10.1651/10–3429.1 Ahamed, F. & Ohtomi, J. (2012) Growth patterns and longevity of the pandalid shrimp Plesionika izumiae (Decapoda: Caridea). Journal of Crustacean Biology, 32, 733–740. https://doi.org/10.1163/193724012X645853 Ahamed, F. & Ohtomi, J. (2014) Relative growth and sexual maturity of the pandalid shrimp Plesionika izumiae (Decapoda, Caridea) in Kagoshima Bay, southern Japan. 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