Anim. Syst. Evol. Divers. Vol. 32, No. 4: 272-280, October 2016 https://doi.org/10.5635/ASED.2016.32.4.037

Review article First Record of Two Cold-Water Jellyfishes Aurelia limbata and Parumbrosa polylobata (Scyphozoa: Semaeostomeae: Ulmaridae) in Korean Coastal Waters

Soo-Jung Chang1,*, Jung Nyun Kim1, Won-Duk Yoon2, Jang-Seu Ki3

1Fisheries Resources and Environment Division, West Sea Fisheries Research Institute, National Institute of Fisheries Science, Incheon 22383, Korea 2Southeast Sea Fisheries Research Institute, National Institute of Fisheries Science, Tongyeong 53085, Korea 3Department of Life Science, Sangmyung University, Seoul 03016, Korea

ABSTRACT

Most scyphomedusae jellyfishes recorded in Korean waters are temperate and subtropical species. In the present study, two cold-water jellyfishes from Korean waters are first described. Scyphomedusae were collected from the coasts of Gangneung and Ulsan of the East Sea, and Boryeong, and the eastern area of the Yellow Sea from June 2006 to May 2015. Scyphomedusae collected in Ulsan and Gangneung were identified morphologically as Aurelia limbata Brandt, 1835. Their umbrella was 250-500 mm in width and 30-50 mm in height; the exumbrella was white and the subumbrella was dark brown in color. The vascular system was complex with anastomosed branches. Individuals of this species lived at about 6.6-9.9°C and 30-50 m depth in Korean waters. Scyphomedusae collected from the center of the Yellow Sea and from Boryeong were identified as Parumbrosa polylobata Kishinouye, 1910. Their umbrella was disc-shaped, 80-200 mm in width and 20-40 mm in height. Its milky white gonads could be seen through the transparent exumbrella. The stomach cavity was round and flat with four interradial cavities connected by subgenital cavities, and the four oral arms were bifurcated and spear-head shaped. Individuals were found at about 6.4-10.0°C and 40-90 m depth in Korean waters. The identities of both species were confirmed by molecular analysis using nuclear ribosomal DNA sequences. Keywords: Aurelia limbata, Parumbrosa polylobata, Scyphozoa, jellyfish, cold-water species, Korean waters, Yellow Sea Cold Bottom Water (YSCBW), Korean Strait Bottom Cold Water (KSBCW)

INTRODUCTION water jellyfish species were found in Japanese waters (Mi­ yake et al., 2002, 2005), none have been recorded in Korean Ulmaridae is one of the three traditional jellyfish families waters yet. This might be due to most cold-water species included in the order Semaeostomeae, class Scyphozoa. Spe­ being distributed in deep-water, which is difficult to survey. cies within Ulmaridae differ from the other species within In the present study, cold-water jellyfishes were sampled the other two families (Pelagiidae and Cyaneidae) by their from Korean waters using set nets (fixed shore net, fish trap, branched radial canals and their circular canal (Mayer, 1910). and bottom trawl net). Based on examinations of their mor­ About one-third of the 40 species (14 genera) in Ulmaridae phology and DNA comparisons, individuals were identified belongs to the genus Aurelia, and the rest of them are most­ as Aurelia limbata Brandt, 1835 and Parumbrosa polylo- ly deep-water species (Daly et al., 2007). bata Kishinouye, 1910, both belonging to Ulmaridae (order Up to date, only six scyphomedusae species have been Semaeostomeae). reported from Korean waters (Park, 2000, 2002; Park and This study aimed to provide morphological descriptions of Chang, 2006; Ullah et al., 2015), most of which are temper­ two cold-water jellyfishes first reported from Korean waters. ate and subtropical species. Although some deep- and cold- Moreover, genetic information based on nuclear ribosomal

This is an Open Access article distributed under the terms of the Creative *To whom correspondence should be addressed Commons Attribution Non-Commercial License (http://creativecommons.org/ Tel: 82-32-745-0641, Fax: 82-32-745-0685 licenses/by-nc/3.0/) which permits unrestricted non-commercial use, distribution, E-mail: [email protected] and reproduction in any medium, provided the original work is properly cited. pISSN 2234-6953 eISSN 2234-8190 Copyright The Korean Society of Systematic Zoology First Record of Two Cold-Water Jellyfishes in Korean Waters

38

Gangneung

37 Yellow Sea East Sea

Boryeong (°N)

36 Ulsan Latitude

35

Parumbrosa polylobata Aurelia limbata 34 123 124 125 126 127 128 129 130 131 Longitude (°E)

Fig. 1. Aurelia limbata and Parumbrosa polylobata sampling sites in Korean waters.

DNA (rDNA) was used to support identification of both spe­ tric pouch, manubrium, subgenital cavity, and canal system cies and establish their phylogenetic relationships with other of the sampled jellyfish were morphologically analyzed and medusae. photographed using a digital camera (Cannon G10; Cannon, Tokyo, Japan). The morphology of the individuals identified as A. limbata was compared to those previously published MATERIALS AND METHODS for this species (Brandt, 1835; Kishinouye, 1910; Uchida, 1954; Kramp, 1961; Yamaji, 1984; Larson, 1990; Miyake et Study sites and sample collection al., 2002) and for Aurelia labiata (Mayer, 1910; Gershwin, Thirty-four individuals of Scyphomedusae were collected 2001). Similarly, records obtained for the individuals iden­ from the coasts of Gangneung (37°47′44.06″N, 128°57′7.32″ tified as Parumbrosa polylobata were compared to those E), Ulsan (35°36′21.07″N, 129°28′37.73″E), and Boryeong previously published in Kishinouye (1910), Mayer (1910), (36°17′53.81″N, 126°28′33.94″E), and from the center and Uchida (1954), Kramp (1961), Yamaji (1984), and Miyake et the eastern Yellow Sea (34°15′00.00″N, 123°15′00.00″E- al. (2005). 37°19′50.00″N, 126°28′67.00″E) in Korean waters using fixed shore nets, fish traps, and bottom trawls, from June, Molecular analysis and species identification 2006 to May, 2015 (Fig. 1). The water temperature and depth Genomic DNA (gDNA) was extracted from the umbrella of at which scyphomedusae were sampled were recorded by a four A. limbata and two P. polylobata specimens. Before ex­ Sea-Bird conductivity-temperature-depth (CTD) instrument traction, alcohol-preserved tissues were washed in distilled (SBE-911; Sea-Bird Electronics Inc., Bellevue, WA, USA). water to remove all ethanol and to avoid contamination from Whole specimens (n=34) were preserved in 5% (v/v) for­ the zooplankton contained in the stomach of jellyfish. Total malin (Junsei, Tokyo, Japan) whereas umbrella tissue sam­ gDNA was extracted by using the method described by Asa­ ples (n=6) were preserved in 99.9% ethanol (Merck, Darm­ hida et al. (1996). Nuclear rDNA sequences spanning the in­ stadt, Germany) for their further use in nuclear DNA analy­ ternal transcribed spacer 1 (ITS1), partial sequence, 5.8S ri­ ses. bosomal RNA gene, complete sequence, internal transcribed spacer 2 (ITS2), complete sequence and 28S ribosomal RNA Morphological analysis and description gene, partial sequence were amplified using the forward The width and height of umbrellas were measured from each primer jfITS-5f (5′-GGTTTCCGTAGGTGAACCTGCGGA specimen. Using a stereomicroscope (Leica MZ 12; Leica AGGATC-3′) and the reverse primer 28S-2R (5′-GCTTTGG Microsystems, Wetzlar, Germany), the umbrella, marginal GCTGCAGTCCCAAGCAACCCACTC-3′) (Dawson, 2004). lobes, rhopalium, marginal tentacles, mouth, stomach, gas­ Amplified PCR fragments were purified and sequenced on

Anim. Syst. Evol. Divers. 32(4), 272-280 273 Soo-Jung Chang, Jung Nyun Kim, Won-Duk Yoon, Jang-Seu Ki an automated sequencer (ABI 3730Xl; Applied Biosystems, Material examined. Korea: 5 individuals, Gangwon-do: off Foster City, CA, USA). The four sequences obtained for A. Gangneung (37°47′44.06″N, 128°57′07.32″E), 31 Aug 2007, limbata and the two sequences obtained for P. polylobata Chang SJ; 7 individuals: off Ulsan (35°36′21.07″N, 129°28′ were compared with nuclear rDNA regions retrieved from 37.73″E), 9 Jun 2011, Chang SJ. the National Center for Biotechnology Information (NCBI) Morphological description. Umbrella hemispherical when database (http://www.ncbi.nlm.nih.gov). Sequences of A. contracted but flat when expanded, about 250-500 mm in limbata from Hokkaido, Japan (Genbank accession No. AY width and 30-50 mm in height. Exumbrella water-white 935215.1), A. limbata from Alaska, USA (AY935211.1), and subumbrella dark brown in color. Gelatinous substance A. aurita from Rhode Island, USA (AY935205.1), Aurelia (mesoglea) thick at center and thin toward margin. Umbrel­ sp. 1 from Miyazu Bay, Japan (AY935214.1), Aurelia sp. 3 la margin divided into 16 marginal lobes with 16 shallow (AY935209.1), Aurelia sp. 4 (AY935208.1), Aurelia sp. 5 notches at regular intervals, 8 rhopalia located between clefts (AY935210.1), Aurelia sp. 6 (AY935207.1), Aurelia sp. 7 of lappets. Numerous thin and long tentacles, dark yellow or (AY935212.1), Aurelia sp. 8 (AY935217.1), and A. labiata brown, rising from umbrella margin. from Alaska, USA (AY935202.1), all deposited by Dawson Single and simple mouth situated at center of subumbrel­ et al. (2005), Aurelia sp. 1 from Busan, Korea (EU332744.1) la. Manubrium cruciform from top view but triangular in (Ki et al., 2008) and Cyanea capillata (U65481.1) (Odorico lateral view. Four frilly oral arms extended from manubrium and Miller, 1997) were used as reference sequences for the to umbrella margin, nematocyst warts scattered on abaxial class Scyphozoa, Craspedacusta sowerbyi Lankester, 1880 side of oral arms. Stomach divided into 4 gastric pouches (FJ423631.1) was used as a reference sequence for the class with subgenital cavity and subgenital pore. Subgenital cavi­ Hydrozoa, and Parazoanthus axinellae (EU363364.2) (Rei­ ty enclosed with dark brown gonads. mer et al., 2008), a member of the class Anthozoa, was used Vascular system complex and branches anastomosed. Six­ as outgroup. teen radial canals extend to the circular canal around um­ The 22 sequences were aligned using Clustal X (Larkin brella margin. Four interradial canals begin from gonads, 4 et al., 2007) and their genetic distances were calculated in perradial canals begin from mouth and 8 adradial canals be­ MEGA 6 (Tamura et al., 2013). Analyses were conducted gin from central part of gonads. Each adradial canal reaches using the Kimura 2-parameter model. There were a total of circular canal without any confluence. Perradial canals and 730 positions in the final dataset. Maximum likelihood (ML) interradial canals branched around gonad part but anasto­ phylogenetic trees were reconstructed using a heuristic sear­ mosed at umbrella margin. Vascular system clear in 250 mm ch and the general time reversible model (GTR), as included width medusa whereas complex in 500 mm width medusa in MEGA 6. Branch support in the ML trees was obtained by due to canals’ confluence. bootstrap analysis using 1,000 replicates. Distribution. Pacific Ocean: Korea (East Sea), Japan (Hok­ kaido); Okhotsk Sea; Alaska; Bering Sea; Cold waters. Remarks. Umbrella of A. limbata (250-500 mm in width) SYSTEMATIC ACCOUNTS reported here is larger than that of A. aurita (40-110 mm in width) reported by Park (2000). Aurelia aurita have an eight- Class Scyphozoa Goette, 1887 scalloped umbrella margin whereas A. labiata and A. limbata Order Semaeostomeae L. Agassiz, 1862 have a 16-scalloped umbrella margin. Aurelia labiata and A. Family Ulmaridae Haeckel, 1880 limbata differ on the color of the umbrella (milky white vs. Genus Aurelia Lamarck, 1816 chocolate brown), on the shape of the manubrium (rounded vs. pyramidal) and on the secondary scalloping of the um­ 1*Aurelia limbata Brandt, 1835 (Fig. 2A-E) brella margin (present in A. labiata and absent in A. limbata) Aurelia limbata Brandt, 1835: 26; Haeckel, 1879: 558; (Gershwin, 2001). Kishinouye, 1910: 22; Uchida, 1954: 209-219; Kramp, Aurelia limbata were found between 6.6-9.9°C and 30- 1961: 341; Yamaji, 1984: 244-246, Pl. 78, fig. 7; Larson, 50 m depth in the coastal areas of Gangneung and Ulsan in 1990: 549-552, fig. 1-H; Miyake et al., 2002: 44-46, figs. the East Sea. Larval decapods are the basic prey of A. lim- 3-4, videograph. bata (Zavolokin et al., 2008). Aurelia (Diplocraspedon) limbata Brandt, 1838: 372; La­ In this study, DNA sequences of A. limbata of the region marck et al., 1840: 179; Agassiz, 1862: 160. spanning ITS1 (partial sequence), 5.8S, ITS2, and 28S (par­ Aurelia labiata Mayer, 1910: 628. tial sequence) rDNA were deposited in GenBank (accession

Korean name: 1*두빛보름달해파리

274 Anim. Syst. Evol. Divers. 32(4), 272-280 First Record of Two Cold-Water Jellyfishes in Korean Waters

A B

g oa

m

mt

C D oa o

g

sc pc

ac ic

E

mt oa

m g

o

rh

sc

pc ic

Fig. 2. Aurelia limbata Brandt, 1835. A, Whole body in side view; B, Aboral view; C, Radial canal system; D, Expanded oral arms; E, Diagram showing all structures observed in oral view. ac, adradial canal; g, gonad; ic, interradial canal; m, umbrella margin; mt, marginal tentacles; o, oral cavity; oa, oral arm; pc, perradial canal; rh, rhopalium; sc, stomach cavity. Scale bars: A=5 cm, B-E=3 cm.

Anim. Syst. Evol. Divers. 32(4), 272-280 275 Soo-Jung Chang, Jung Nyun Kim, Won-Duk Yoon, Jang-Seu Ki

Nos. KX943298 to KX943301). DISCUSSION

Genus Parumbrosa Kishinouye, 1910 Mayer (1910) defined three species (A. aurita, A. labiata, and A. solida Browne, 1905) and 13 varieties within the genus 1*Parumbrosa polylobata Kishinouye, 1910 (Fig. 3A-C) Aurelia. However, most scientists used Kramp’s (1961) clas­ Parumbrosa polylobata Kishinouye, 1910: 19-21, Pl. 4, sification concerning two species (the circum global A. aurita figs. 20-23; Mayer, 1910: 728, fig. 428; Uchida, 1954: and the arctic A. limbata) until the end of the 1990s (Larson, 209-219; Kramp, 1961: 346; Yamaji, 1984: 244-245, Pl. 1990; Arai, 1997). With the development of electronic jour­ 78, fig. 4; Miyake et al., 2005: 61, fig. 2, videograph. nals and photo sharing, three morphological species of A. aurita, A. limbata and A. labiata were distinguished (Ger­ Material examined. Korea: 5 individuals, Chungcheongnam- shwin, 2001) Moreover, with the development of molecular do: off Boryeong (36°17′53.81″N, 126°28′33.94″E), 25 Apr tools, at least 13 Aurelia species (Aurelia sp. 1-10) includ­ 2006, Chang SJ; 4 individuals, Korea-China Provisional ing A. aurita, A. labiata, and A. limbata were distinguished zone in the Yellow Sea (34°15′00″N, 123°15′00″E; 34°45′ based on nuclear rDNA and mitochondrial cytochrome oxi­ 00″N, 123°15′00″E; 36°15′00″N, 123°45′00″E), 28-30 Jul dase I (Dawson and Jacobs, 2001; Dawson et al., 2005) and 2007, Chang SJ; 13 individuals, the Eastern part of the Yel­ on ITS1 (Schroth et al., 2002; Dawson, 2003) sequence data. low Sea (34°15′00.00″N, 123°15′00.00″E-37°19′50.00″N, Based on morphological data, Park (2000) reported A. aurita 126°28′67.00″E), 1-6 May 2015, Kim JN, Kim JE. in Korean waters and these specimens were molecularly as­ Morphological description. Umbrella flat and disc-shaped, signed as Aurelia sp. 1 by Ki et al. (2008). 80-200 mm in width and 20-40 mm in height. Gelatinous The present study, the analysis of pairwise genetic dis­ umbrella colorless, transparent; small granulated spots scat­ tances from ITS1 to 28S rDNA among genus Aurelia (Table tered over exumbrella. Very weak muscle system in subum­ 1), revealed that sequences of the A. limbata specimens brella. Umbrella margin divided into octants, each compris­ collected in Gangneung were 99.5% identical to that of A. ing 1 divergent ocular lobe, 3 tentacles, and 8 velar lobes limbata collected in Hokkaido (AY935215.1) and 92.9% (64 in total). Cores of tentacles and lobes opaque white with similar to that of A. limbata from Alaska (AY935211.1). In their transparent margins. addition, the rDNA region examined differed about 11.1% One single and simple mouth at center of subumbrella. to 11.6% between Korean A. limbata and Korean Aurelia Stomach cavity round and flat with four interradial cavities sp. 1 (AY935244.1), about 16.8% to 17.0% between Korean connected by subgenital cavities. Four oral arms bifurcated, A. limbata and A. aurita (AY935205.1), and about 17.2% to spear-head shape, and nearly as long as umbrella diameter. 17.5% between Korean A. limbata and A. labiata (AY93502. Gonads opaque, white-pink, and lotus shaped. Gonads 1). Thus, these results seem to contradict the hypothesis that enclosed around stomach cavity and reflected through trans­ A. limbata is a color morph of A. labiata (Gershwin, 2001). parent exumbrella. In interradial cavity, gonads triangularly In this study, A. limbata, A. aurita, A. labiata, and Aurelia layered in transvers section and mixed with numerous long sp. 1 are monophyletic according to the ML tree reconstruct­ gastric cirri. ed based on their ITS1 to 28S DNA sequences. ML phylo­ Simple vascular system extended to circular canal around genetic tree constructed with Semaeostomeae members, in­ umbrella margin. Four interradial canals, 8 adradial canals, cluding A. limbata, Aurelia sp. 1, A. aurita, A. labiata, and and 4 perradial canals. Adradial canals located between in­ P. polylobata showed that identical species were clustered terradial and perradial canals, reaching circular canal in a together, but different species were clearly seperated, which straight line. Interradial canal split from gonads to the um­ were supported by high bootstrap values (Fig. 4). brella margin. Aurelia limbata is distributed in the East Sea (this study), Distribution. Pacific Ocean: Korea (Yellow Sea), Japan Alaskan waters (Larson, 1990), Tohoku region to Hokkaido, (Japanese coast of East Sea, Suruga Bay); cold waters. the Okhotsk Sea, the Bering Sea (Kishinouye, 1910; Uchi­ Remarks. Parumbrosa polylobata were found at 6.4-10.0°C da, 1954; Kramp, 1961; Larson, 1990; Wrobel and Mills, and 40-90 m depth in Korean waters. Euphausia pacifica 1998; Zavolokin et al., 2008; Zavolokin, 2010, 2011), and were found in the stomach cavity of P. polylobata. DNA se­ the northern part of the Japanese waters (Pogodin, 1998; quences of P. polylobata of the region spanning ITS1 (partial Miyake et al., 2002; Zavolokin, 2010). Individuals were sequence), 5.8S, ITS2, and 28S (partial sequence) rDNA found in the epipelagic and mesopelagic zone of the sea of were deposited in GenBank (accession No. KX943302). Okhotsk and Bering Sea (Zavolokin et al., 2008; Zavolokin,

Korean name: 1*유리햇살해파리

276 Anim. Syst. Evol. Divers. 32(4), 272-280 First Record of Two Cold-Water Jellyfishes in Korean Waters

A B

ic rh

mt

pc oa

g sc o

C

rh

mt

oa

o

g

sc

pc

ic

Fig. 3. Parumbrosa polylobata Kishinouye, 1910. A, Whole body; B, Margin of umbrella; C, Diagram showing all structures observed in oral view. g, gonad; ic, interradial canal; mt, marginal tentacles; o, oral cavity; oa, oral arm; pc, perradial canal; rh, rhopalium; sc, stomach cavity. Scale bars: A–C=2 cm.

Anim. Syst. Evol. Divers. 32(4), 272-280 277 Soo-Jung Chang, Jung Nyun Kim, Won-Duk Yoon, Jang-Seu Ki

Table 1. Pairwise genetic distances calculated for the region spanning ITS1 (partial sequence), 5.8S rDNA, ITS2, and 28S (partial sequence), among species of genus Aurelia 1 2 3 4 5 6 7 8 9 10 11 12 13 14 1. A_limbata_JH_AY935215 2. A_limbata_Gn 0.006 3. A_limbata_Al_AY935211 0.073 0.072 4. A_aurita_Rd_AY935205 0.171 0.170 0.159 5. Aurelia_aurita_AY935205 0.171 0.170 0.159 0.000 6. Aurelia_sp1_Bs_EU332744 0.118 0.111 0.129 0.169 0.169 7. Aurelia_sp1_JM_AY935214 0.102 0.096 0.112 0.147 0.147 0.002 8. Aurelia_sp5_AY935210 0.192 0.187 0.188 0.212 0.212 0.189 0.166 9. A_labiata_Al_AY935202 0.174 0.172 0.161 0.194 0.194 0.148 0.129 0.183 10. Aurelia_sp7_AY935212 0.120 0.117 0.113 0.133 0.133 0.118 0.100 0.168 0.162 11. Aurelia_sp6_AY935207 0.173 0.170 0.178 0.188 0.188 0.158 0.138 0.180 0.150 0.167 12. Aurelia_sp8_AY935217 0.163 0.165 0.168 0.195 0.195 0.153 0.139 0.207 0.106 0.147 0.176 13. Aurelia_sp4_AY935208 0.158 0.158 0.158 0.171 0.171 0.161 0.142 0.162 0.156 0.144 0.152 0.179 14. Aurelia_sp3_AY935209 0.156 0.153 0.156 0.177 0.177 0.146 0.130 0.192 0.154 0.173 0.116 0.164 0.135

A_limbata, Aurelia limbata; A_aurita, Aurelia aurita; A_labiata, Aurelia labiata; Gn, Gangneung, Korea; JH, Japan, Hokkaido; Al, Alaska, USA; Rd, Rhode Island, USA; Bs, Busan, Korea; JM, Japan, Miyazu Bay.

Fig. 4. Phylogenetic maximum likelihood (ML) tree constructed for Aurelia limbata, Parumbrosa polylobata, and the other species (class Scyphozoa, class Hydrozoa, and class Anthozoa), based on ITS1 to 28S nuclear DNA sequences, using a general time revers- ible model of evolution. Numerals above the branches refer to the percentage of the 1,000 bootstrap replications supporting each node. Bootstrap indices under ML are shown at each node. A aurita, Aurelia aurita; A labiata, Aurelia labiata; A limbata, Aurelia limbata; Cr sowerbyi, Craspedacusta sowerbyi; Cy capillata, Cyanea capillata; Pa axinellae, Parazoanthus axinellae; P polylobata, Parumbrosa polylobata; Gn, Gangneung, Korea; JH, Japan, Hokkaido; Al, Alaska, USA; Bs, Busan, Korea; JM, Japan, Miyazu Bay; Rd, Rhode Island, USA; YS, Yellow Sea.

2010, 2011) and at about 2.0°C and 33.4 psu from 200 to These coastal areas of the East Sea are influenced by the 250 m depth in Hokkaido, northern Japan (Miyake et al., Korean Strait Bottom Cold Water (KSBCW), which is a 2002). Moreover, A. limbata were found at about 6.6-9.9°C less saline water mass below 10°C (Na et al., 2010) known and 30-50 m depth in Gangneung and Ulsan in this study. to originate from the North Korean Cold Water (NKCW),

278 Anim. Syst. Evol. Divers. 32(4), 272-280 First Record of Two Cold-Water Jellyfishes in Korean Waters a coastal mode of the East Sea Intermediate Water (ESIW) ambient temperature using buffers containing high concen­ (Cho and Kim, 1998). Therefore, the cold-water masses tration of urea. Fisheries Science, 62:727-730. https://doi. from ESIW to KSBCW might restrict with the distribution org/10.2331/fishsci.62.727 of A. limbata in the East Sea. Brandt JF, 1835. Prodomus descriptionis animalium ab H. Mer­ In addition, Parumbrosa polylobata were recorded from tensio in orbis terrarum circumnavigatione observatorum. Fasc. I. Rec. Act. Acad. Imp. Sci. St. Petersburg, 1834:1-75. Toyama Bay deep waters (>130 m depth) in 1907 (Kishino­ Brandt JF, 1838. Ausführliche Beschreibung der von C. H. uye, 1910). Individuals observed at 267-509 m depth off the Mertens auf seiner Weltumsegelung beobachteten Schirm­ Pacific coast of Japan (Miyake et al., 2005) were found at quallen nebst allgemeinen Bemerkungen über die Schirm­ - - 6.6 9.2°C and 34.2 34.4 psu. In the present study, P. poly- quallen überhaupt. Mémoires de l’Académie impériale des - - lobata were found at about 6.4 10.0°C and 40 90 m depth Sciences de St.-Pétersbourg. Sér. 6. Sciences Naturelles, 2: in the center of the Yellow Sea. This species fed on several 237-411. gelatinous zooplankton such as Salpa fusiformis, Siphono­ Cho YK, Kim K, 1998. Structure of the Korea Strait bottom phora, and Solmissus sp. (Miyake et al., 2005), and on Eu- cold water and its seasonal variation in 1991. Continental phausia sp. in this study. During winter, P. polylobata is Shelf Research, 18:791-804. https://doi.org/10.1016/S0278- distributed from the western coast of Korean waters to the 4343(98)00013-2 center of the Yellow Sea whereas during summer it is distri­ Daly M, Brugler MR, Cartwright P, Collins AG, Dawson MN, buted in the center of the Yellow Sea, where the Yellow Sea Fautin DG, France SC, Mcfadden CS, Opresko DM, Rodri­ Cold Bottom Water (YSCBW) remains. Thus, the YSCBW guez E, Romano SL, Stake JL, 2007. The phylum Cnidaria: a review of phylogenetic patterns and diversity 300 years seems to affect the distribution of P. polylobata. after Linnaeus. Zootaxa, 1668:127-182. This study is the first step to report cold-water jellyfishes Dawson MN, 2003. Macro-morphological variation among from Korean waters. Moreover, records of genetic informa­ cryptic species of the moon jellyfish, Aurelia (Cnidaria: tion of nuclear rDNA from two cold-water jellyfishes will Scyphozoa). Marine Biology, 143:369-379. https://doi.org/ be a useful identification key to discriminate species. In 10.1007/s00227-003-1070-3 future, monitoring these two cold-water species might be Dawson MN, 2004. Some implications of molecular phyloge­ important to evaluate climate changes in the East Sea and in netics for understanding biodiversity in jellyfishes, with the Yellow Sea. emphasis on Scyphozoa. Hydrobiologia, 530/531:249-260. https://doi.org/10.1007/s10750-004-2659-3 Dawson MN, Gupta AS, England MH, 2005. Coupled biophys­ ACKNOWLEDGMENTS ical global ocean model and molecular genetic analyses identify multiple introductions of cryptogenic species. Pro­ ceedings of the National Academy of Sciences of the Unit­ We thank Hye-Eun Lee and Ji-Eun Kim for their collecting ed States of America, 102:11968-11973. https://doi.org/ of specimens and comments on our manuscript. We also 10.1073/pnas.0503811102 heartily thank that Im-Sung Hwang and Jong-Hyun Lee Dawson MN, Jacobs DK, 2001. Molecular evidence for cryptic participated in Digital eco photo club (DEPC) for their species of Aurelia aurita (Cnidaria, Scyphozoa). The Biol­ photographs (Fig. 2A and 2B, respectively). This work was ogical Bulletin, 200:92-96. https://doi.org/10.2307/1543089 funded by the National Institute of Fisheries Science, Korea Haeckel E, 1879. Das System der Medusen. pt. 2, System der (R2016032) and ‘Management of marine organisms causing Acraspeden. In: Monographie der Medusen. Vol. 1. G. ecological disturbance and harmful effects’ by Korea Insti­ Fischer, Jena, pp. 1-672. Gershwin LA, 2001. Systematics and Biogeography of the jel­ tute of Marine Science and Technology Promotion/Ministry of Oceans and Fisheries, Korea. lyfish Aurelia labiata (Cnidaria: Scyphozoa). Biological Bulletin, 201:104-119. https://doi.org/10.2307/1543531 Ki JS, Hwang DS, Shin K, Yoon WD, Lim D, Kang YS, Lee Y, Lee JS, 2008. 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