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1 Vol. 32(1):1-13 DOI: /OPR Ocean and Polar Research March 2010 Article ûwÿ «v jm p yá * Áx ¼Á t Á½x Áy w w ûw ( ) û 391 Distribution Characteristics and Community Structure of Phytoplankton in the Different Water Masses During Early Summer of Southern Sea of Korea Seung Ho Baek, Kyoungsoon Shin *, Bonggil Hyun, Pung-Guk Jang, Hyun-Su Kim, and Ok-Myung Hwang South Sea Research Institute, KORDI Geoje , Korea Abstract : To assess short-term variation of summer phytoplankton community structure in different water masses, phytoplankton and environmental factors were monitored from 31 stations on and off the southern coasts of Korea, from June 18 to June According to multidimensional scaling (MDS) and cluster analysis based on phytoplankton community data from each station, the southern sea was divided into two groups. The first group included stations in the south-eastern region of Jeju Island, which is strongly influenced by the Kuroshio warm current. The second group located along the coastal region of the southern sea, which was mainly comprised of Bacillariophyceae and Crytophyceae. Of these stations, St. 13 and 28 formed a temperature front caused by different hydrological conditions. In particular, nutrients and Chl.a concentrations in these two stations were significantly higher compared to those in the other stations. This indicates that phytoplankton population and subsequent microalgal growth under high nutrient concentrations vary in different water masses. Our results support the theory that phytoplankton community structure in the southern sea of Korea can be influenced on a short-term scale by different water masses and currents. Key words : phytoplankton community structure, canonical correspondence analysis (CCA), cluster analysis, multidimensional scaling (MDS), southern sea of Korea 1. w k v jm» w e w, w y y œ sp yw. v jm x sp q w w *Corresponding author. ksshin@kordi.re.kr wwš mw ( ) q w w. ù ûwÿ w 1990 ¾ w yw y k w w p w šƒ š(œ 1971; ½ 1982; 1988), 1990 z v jm sp w ww ƒ š ( 1990; 1990; 1993;

2 2 Baek, S. H. et al. 2003; ½ 2008). ûwÿ w i ù yw ù,, ûw«w w w w ƒ x, ƒ (front) x v jm w (Yang and Kim 1990; 1998; 1998). y w (ocean front) x jš, x e ƒ» w v jm w w sw y j w (Han et al. 1989; ½ 1998; 2003). ûw Ÿ «v jm ûw (Shim and Park 1984; ½ 2003), ûw w ( 2003; 2008), ûw ( 1998), wwx( œ 1985; Shon et al. 2008), ( 1990; ½ 2008) ù, ù w wwx sww ûw, Ÿ w Ÿ ww w k. w s w y p w w š m w» w p sƒw w. wwx sww ûw Ÿ «v jm s sp q wš ƒ x x k MDS (Multidimensional Scaling) Cluster ww. w v jm sp w e y y w» w CCA(Canonical Correspondence Analysis) ww v jm p wwš w. 2. w w ûw» (St. 1) w ü w (St. 2-3), wwx w ¾ (St. 4-8), ûw w (St. 9-14), w Ÿ (St ), Ÿ ƒ (St ) 31 ûwÿ «w, td w (Fig. 1).» w v jm s q w» w. ƒ, t sww, ü l. Fig. 1. Location of study sites in southern sea of Korea.

3 Phytoplankton Assemblage Characteristics in Southern Sea of Korea 3 td ml GF/Fvl(47 mm Whatman glass fiber filters) w, vl chlorophyll-a(chl.a) d w 15 ml p þ w š, w 50 ml p š þ w w. v jm w 500 ml td 500 ml š š (Lugol) 1%ƒ š w. Chl.a 90% m š 24 þ w z xÿd»(turner Designs 10-AU Fluorometer) w (Parsons et al. 1984).»(Lachat Quick-chen 8000 auto-analyzer Lachat, Milwaukee, WI) w d w. w t (CSK standard solution, Wako Pure Chemical industries, Osaka), + (NO 3 +NO 2 ), (NH 4 ) (PO 3 4 ), ³ (SiO 3 ) ƒƒ w (Parsons et al. 1984). v jm x q w» w 500 ml v 20 ml k z µl w 200 w. p Ÿwx w š, 10 µm w r wù r š, r (Crytophyceae) wù r. v jm x x k Shannon-Wiener (H', species diversity index) ³ (J') PRIMER version 5 w w. w v jm» w ƒ xw v jm k Bray-Curtis w š, PRIMER version 5 w Cluster, MDS w (Clarke and Warwick 2001; 2008). w k v jm y w ƒ CANOCO 4.5 software w CCA(Canonical Correspondence Analysis) ww. 3. y y 6» ww 31 ûwÿ «17.2 o C(St. 28) š 23.0 o C(St.10) 5.8 o C š, s³ 21.01±1.4 o C ùkû PSU(St. 12) š PSU (St. 7) w š, s³ 33.49±0.36 PSU (Fig. 2). 1 wwx 5¾ ƒw ù, wwx û ww ùkû (St. 6-12). 13 w w p x y w. z Ÿ ww x ù. Ÿ d û ww 28 w w x ùkûš, z 29 l w w. wwx ù ûw 33 PSU z j. w w w p w. p w w» w T-S diagram Cluster ww (Fig. 3).» ûw Fig. 2. Horizontal distribution changes of temperature and salinity in southern sea of Korea, from June 18 to June Gray areas indicate night periods during the sampling cruise. St. 13 and 28, indicated by arrowheads, are strongly influenced by different water masses.

4 4 Baek, S. H. et al. Fig. 3. A T-S diagram (a) showing the relationship between temperature (T), salinity (S) and density. St. 13 and 28, indicated by arrows, are strongly influenced by different water masses. Gray areas are indicated based on cluster analyses. Joint plots of sites association analyses obtained by cluster analyses (b), using a data set of surface temperature and salinity during the study period in southern sea. Ÿ w p j 3w. š (>22 o C), š (>34 PSU) (7, 8, 9, 10). (<33.5 PSU)ƒ œ (>19, <22 o C) x œ. (>33.5, <34 PSU) (<19.5 o C) ƒ x œ ƒƒ. ƒ ³ ƒ š, + ùkû (Fig. 4a, 4b). ³ (St. 1) µm še» w š, 2 Ÿ (St. 24) 5µM z ù, z (St ) w (Fig. 4a). + še µm š, (St ) Ÿ w y ùkü (Fig. 4b). w µm še» w ù, 30 w ƒ¾ û ƒ (Fig. 4c).

5 Phytoplankton Assemblage Characteristics in Southern Sea of Korea 5 Fig. 4. Horizontal distribution changes of silicate (a), nitrate + nitrite (b), ammonia (c) and phosphate (d) in southern sea from June 18 to June Gray areas indicate night periods during the sampling cruise. St. 13 and 28, indicated by arrowheads, are strongly influenced by different water masses. ùkü µm še» w š, û (St. 27, 29). w, (St. 30, 31) ùkü. p ww x Ÿ µm z û. v jm s ƒ Chl.a s y Fig. 5 ùkþ. Chl.a (2.36 mg m ) 3 28 š, w w (Fig. 5a). w Ÿ š, ƒ ùkûš, 1. v jm x Chl.a y w w (Fig. 5b).» td xw v jm 111, ³ (Bacillariophyceae)ƒ , r (Dinophyceae)ƒ , ³ r (Chrysophyceae)ƒ 1 1 4, ù (Euglenophyceae) 1 2 2, v (Raphidophyceae) ùkû. r (Crytophyceae) wù crytomonads. v jm r, w 1, 28, 30, 31 ƒ x 13 ³ x v jm 50% ùk

6 6 Baek, S. H. et al. Fig. 5. Horizontal distribution changes of Chl.a (a) and standing crop (b) in the southern sea from June 18 to June Gray areas indicate night periods during the sampling cruise. St. 13 and 28, indicated by arrowheads, are strongly influenced by different water masses. Fig. 6. Horizontal distribution changes of phytoplankton abundance (upper), and relative contribution (lower) in the southern sea from June 18 to June

7 Phytoplankton Assemblage Characteristics in Southern Sea of Korea 7 Fig. 7. Horizontal distribution changes of number of observed species (a), Shannon-Wiener diversity indices (H') (b) and evenness (J') (c) in southern sea of Korea from June 18 to June Gray areas indicate night periods during the sampling cruise. St. 13 and 28, indicated by arrowheads, are strongly influenced by different water masses. ü (Fig. 6). wwx w û v jm š, r ƒ w. r Ÿ w Ÿ û w ù kû. ³ r, ù, v x ûš û. x Shannon-Wiener v jm x ùkû ù, ³. x ƒ wš p ùkù ³ ƒ û w (Fig. 7).» ³ Chaetoceros curvisetus, Chaetoceros socialis, Letocylindrus danicus, Pseudo-nitzschia delicatissima, Skeletonema costatum š, r Ceratium fusus, Gyrodinium spirale, Heterocapsa niei, Heterocapsa tirquetra xw. š 5-10 µm j» ƒ r Ÿ w.» xw v jm k Cluster MSD w Fig. 8 ù kü. v jm x x yw ù, j 2 w. 1 wwx w š, 2 Ÿ ƒƒ. 4. š ù ûw w td š š p ƒ ù (Tsushima Warm Current Water) ƒ ûw ƒ ù ƒ¾ w w x, w v jm sw ( 1990; Kang et al. 2002; 2003). Seung (1992) w wx wwx ù

8 8 Baek, S. H. et al. ù ûw w e. s(t-s diagram) p j 3 w. 1 ù w w wx w (St. 6-8) û w (St , 17, 18). 2 ûw w w p. 1 2 p ƒ (St. 13 St. 28 z ) 3 ù. ûw 9 12 ww ù, w w w š(fig. 1, 2), w w û. w, ƒ ƒ¾ 13 w w, w (St. 14, 15, 16) û ùkûš, w w 33.5 PSU z w.» (St. 12) 2 o C x w w w yw ƒ x w ( ) w š w ( 2003; 2003).» p 28, š ( +, )ƒ.» ƒ œ 2ƒ dw., w y w ü yw ƒ. w y w x w, 28 ƒ y w œ š q w». p kg m 3» ƒ š, ( ) w p (Fig. 3, 4). 28 w yw ƒ š q. ù w w y w x yw q w» w z w d w ƒ v w. v jm w p x, j» ùkù. w v jm wš(riley 1942; Cushing 1989), w d w d t w Ÿd œ v jm x û (Joint et al. 1986; Baek et al. 2009). w ùw x v jm(micro and nano size) x (Smayda and Reynolds 2001), v jm(pico size) w (Johnson and Sieburth 1979; Furuya et al. 2003; 2005). p v jm sp. p 2 w w x j» ³ ƒ x, 1 w s ƒ û. wr, wwx û ww x w š, r x. Furuya CHEMTAX» w w (Furuya et al. 2003), û w d x 6 td v jm û. Prochlorophytes, Chrysophytes, Prymnesiophytes, Chlorophytes xw š, Chl.a 24-63% Prochlorophytesƒ w. ù w w wwx û w Chl.a ƒ mg m 3 û š, w Furuya et al. (2003) û w w» w. (2005) û» l w w, µm w Chl.a ƒ v jm 32-78% wš, Picocyanobacteria Synechococcusƒ w. wwx û 8 3 µm w j» Chl.a 45% w ( œ ). ³ w w ƒ¾ ùkü v jm w š, û ƒ w v jm w š. s 2 r ƒ r ³ r w k w (Hill 1991). s p ù w 2 w (St ) r w w. Ÿ w mw Ÿ s w yƒ mw w m w w w w v jm x œ ( 1984; 2003). Weng et al. (2007) w, Pearl River Estuary r Cryptomonas

9 Phytoplankton Assemblage Characteristics in Southern Sea of Korea 9 sp. n l (phosphorus) (iron) w w. s mw ƒ w w ƒ j. (2000) š w, yy r w š, Chroomonas spp., Cryptomonas spp., Rhodomonas spp. ùkû. p ql ³ew ù, jš ù z xw. Ÿ w (St ) Shannon-Wiener (H': 0.3 w) ³ (J': 0.3 w)ƒ û. r ƒ xw q, (2000) š ew. w w r ³ew w ƒƒ yw z w, ³ ( r sy ƒ j) r ( r sy ƒ û ƒ û y ƒ w) v jm wš w (Smayda 1997; Baek et al. 2008b). ûwÿ «v jm s s» w Cluster MDS ww. w ww s s p ƒ mw. ww x Yd w d ³ w (Fig. 9 ). p v jm sp Cluster (Group 1). 1 w š Áš w i» ù w w v jm x û. v jm û w š, û ƒ w k xw. j»ƒ r ƒ xw š d w ƒ w e. v jm sp j 2 ù, p Fig. 8. Joint plots of sites association analyses obtained by cluster (a) and non-metric multidimensional scaling (MDS) (b) ordination analyses, using a data set of the surface species abundances during the study period in southern sea of Korea.

10 10 Baek, S. H. et al. Fig. 9. Score plot by principal component analysis (PCA) based on phytoplankton communities of all stations during the study period. Ÿ ù. w Ÿ X d, X d ƒƒ ew (Fig. 8). X d Ÿ r Cryptomonas sp.ƒ xw œ. w Cryptomonas sp. Ÿ s mw. X d ew ³ ƒ x w. ƒ, r ³ ƒ w w ³ ƒ w ƒ w (Smayda 1997). ³ w s s Fig. 10. Relationship between nutrient, temperature, salinity and phytoplankton community composition by canonical correspondence analysis (CCA). Effect of population development on each group species (a: dominant diatom group, b: Crytophyceae, Euglenophyceae and Chrysophyceae, c: dominant dinoflagellate group) was assessed.

11 Phytoplankton Assemblage Characteristics in Southern Sea of Korea 11 w ù(turner 1984; Welschmeyer and Lohrenzen 1985), œ ( yw ƒ ) w ³ s s w ƒ w» w ƒ ü s d x v w. w w 13 ƒ x œ v jm (St. 12, 14) x e. yw ƒ w (St. 28) w sw, p x ƒ yw w d ƒ td y yw» v jm x w s. 1 w û i w x š x w v jm x û., 2 w Ÿ w ³ r ƒ z w xw š, w ƒ¾ w w( N, P). v jm 111 xw ù, û xw š, p CCA w. ƒ x w ƒ ey k ƒ d y ƒ x w (diagram) y w y r m w. ³ w CCA r (Fig. 10A), 27, 28, 29 xw Pseudo-nitzschia spp., Chaetoceros spp. w ewš š,, (+) ƒ x ( ) ƒ x xw Skeletonema spp.( Skeletonema costatum) Leptocylindrus spp.( Leptocylindris danicus) w w wwš, ( ). Skeletonema costatum Ÿ Ÿ» w sw ù w xwš, w w (Han et al. 1992, 1994). Leptocylindrus danicus ûw w x š( 1976; Kang et al. 1999), m w w ( 2008). S. costatumƒ x w ƒ x š, w ew. ³ r, ù, r w CCA r (Fig. 10b), r w xw š, p ù y w ùkù. ³ r ³ ƒ š, ù + û (+) w. r xw w CCA Prorocentrum spp. Heterocapsa spp.ƒ w ew š, ³ x w (Fig. 10c). ³ ƒ 2 w w, r vƒ ³. Ceratium spp. Scrippsiella spp. (+) ùkü. Ceratium Ceratium fususƒ Ÿ üd(st. 23) (St. 1) w. Ceratium fusus w w w ù (Baek et al. 2008a), w sy ƒ r w û, w w w y w bloom x w (Baek et al. 2008b). r û» ƒ w. sy ƒ û»» w» ƒ. 5. ûw Ÿ w w v jm s q w» w l x ww, v jm 111 xw. s v jm k m w, w ûw Ÿ w j 2 ù. Group 1 w i w x w v jm x û., Group 2 w w ³ r ƒ xw. ew 13 ew 28 w Á š w š,

12 12 Baek, S. H. et al. Chl.a ƒ w ùkû. ûw Ÿ w v jm sp p w w q. w w ks w w( wwx) e w (PE98445)»z yƒ ûw«w k e w» sƒ» (PM55400) w. Ì w. š x œ (1971) w ûw w. w w wz 6:25-36 ½» (1982) w ûw. w w wz 17:74-84 ½x,, (2008) w t Ornithocercus r x w w ùy. w w wz Õ Ö 13: z,,, ½x, w (2005) x, Flow Cytometer, HPLC k w d» v jm. Ocean and Polar Res 27: ³, x, (1998) w w ûw v j m». Algae 13: , (1990) w ûw v jm s p. w w 23: y,, w, jie m g (2008) w ƒ v jm e wsƒ. Algae 23: », ½, (2000) yy» v jm k. w wz 33: x,, y (1984) Ÿ v jm w. w w wz 19: , ½ x, y¼ (1998) w û, w w w xw. w wz 31: Ÿ, y (1976) y w 2. v jm y. w w wz 11:34-38 y (1993) w v jm». w. w š 17:33-56 y, ½ (2003) û w v jm œ sp. y wz 21:8-17 y,, y, y (2003) w v j m œ s r. y wz 21: , š, x (1990) w v jm k. w wz 5: ,, ½,, ½w³, (2003) ûw v jm» y. Algae 18:49-58, «, y (2008) m w v jm x. Algae 23:53-61 ³, ³ (1988) ù x y. w wz 21: s, œ (1985) wwx v jm». w wz 18:74-83 z, ½ (1998) w ûw ù e ù, e w. w wz 31: Baek SH, Shimode S, Han MS, Kikuchi T (2008a) Population development of the dinoflagellates Ceratium furca and Ceratium fusus in Iwa Harbor, Sagami Bay, Japan. Ocean Sci J 43:49-59 Baek SH, Shimode S, Han MS, Kikuchi T (2008b) Growth of dinoflagellates, Certium furca and Ceratium fusus in Sagami Bay, Japan: the role of nutrients. Harmful Algae 7: Baek SH, Shimode S, Kim HC, Han MS, Kikuchi T (2009) Strong bottom-up effects on phytoplankton community caused by a rainfall during spring and summer in Sagami Bay, Japan. J Mar Syst 75: Clarke KR, Warwick RM (2001) Change in Marine Communities: An approach to statistical analysis and interpretation, 2nd ed. PRIMER-E, Plymouth, pp 179 Cushing DH (1989) A difference in structure between ecosystems in strongly stratified waters and in those that are only weakly stratified. J Plankton Res 11:1-13 Furuya K, Hayashi M, Yabushita Y, Ishikawa A (2003) Phytoplankton dynamics in the East China Sea in spring and summer as revealed by HPLC-derived pigment signatures. Deep-Sea Res II 50: Han MS, Furuya K, Nemoto T (1989) Phytoplankton distribution in frontal region of Tokyo Bay, in November J Oceanogr Soc Japan 45: Han MS, Furuya K, Nemoto T (1992) Species-specific productivity of Skeletonema costatum in the inner part of Tokyo Bay. Mar Ecol Prog Ser 79: Han MS, Kim OK, Jeon JK (1994) Changes in phytoplankton communities in the two different hydrological condition of semi-enclosed Chinhae Bay Korea. Bull Plankton Soc Japan 41:43-55

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