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1 The Sea Journal of the Korean Society of Oceanography Vol. 12, No. 3, pp , August 2007 z to Œ n } qsp* x r t r~ ~d s 1 t 2 2 (v) nl Šn 1r m 2d p dn Interaction between Invertebrate Grazers and Seaweeds in the East Coast of Korea J. W. YOO*, H. J. KIM, H. J. LEE, C. G. LEE, C. S. KIM, J. S. HONG 1, J. P. HONG 2 AND D. S. KIM 2 Korea Institute of Coastal Ecology, Inc., Incheon , Korea 1 Department of Oceanography, Inha University, Incheon , Korea 2 East Fisheries Research Institute, NFRDI, Kangnung , Korea se- s qnp r o l 12sp sel l ~ p e (PCGR, per capita grazing rate, g seaweeds/individual/day)p, sp ep r p p l. sep ~ r PCGRp power curve(y=0.2310x )l r l p rp m. p sep PCGR p ~ pp, e pp ~ o l p v k p Žkl. s ~ r PCGRp r m, qp ~ sp prp eqp p r. e k ~ qn(grazing impact, mg/m 2 )p, sql q m p ~p (Aplysia kurodai, k 2,513 mg/m 2 )p p ˆ, pp (Strongylocentrotus nudus, k 1,500 mg/)m (S. intermedius, 733 mg/m 2 ) p p ˆ. n o r 4,000 ~ pp ˆmn, Elasmopus sp.m 2,000 ~ pp eovmnp, Jassa falcatap ep 3.435m mg/m 2 /dayp p ˆ. ll r se vp s s e p qnp p 5,045 mg/m 2 /dayp p ˆ. e l m nl rqp qnp p p p s sl v ppp r pl. p n 3 ~/m 2 p el ep } kelp s (k 5 ton/ha)p p ml. le p ek sl e v rqp ( s p k 16%)p p ml. sq l e sep ll p sr ppp k, lm se qnl pp (lp )p sql ƒ o p qqp v p m. We estimated the distribution of predator-prey interaction strengths for 12 species of herbivores (including amphipods, isopods, gastropods, and sea urchins) and made a regression model that may be applicable to other species. Laboratory experiments were used to determine per capita grazing rate (PCGR; g seaweeds/individual/ day). Relationship between the biomass of individual grazers and fourth-root transformed PCGR was fitted to power curve (y=0.2310x , r=0.8864). This finding supported that the grazing efficiency was not even as individual grazers increase in size (biomass). Therefore, the biomass-normalized PCGR was estimated and revealed that smaller size herbivores were more effective grazers. Grazing impact considering density of each taxon was calculated. The sea hare Aplysia kurodai had greatest grazing impact on the seaweed bed and the sea urchin Strongylocentrotus nudus and S. intermedius were ranked in descending order of the impact. The amount of seaweed grazed by the amphipod Elasmopus sp. (>4,000 ind./m 2 ) and Jassa falcata (>2,000 ind./m 2 ) were and mg/m 2 /day respectively. The combined grazing amount of herbivores was 5,045 mg/m 2 /day in the seaweed bed. Although sea hare and sea urchin had strong impacts on seaweeds, the effects of dense, smaller species could not be seen as negligible. Surprisingly, the calculated grazing potential of sea urchins with a mean density of 3 ind./m 2 exceeded the mean production of seaweed cultured in domestic coastal waters in Korea (ca., *Corresponding author: jwyoo@coastkorea.com 125

2 126 oqoë vëp rëp}ë}ë që r Ë 5 ton/ha). Small crustaceans were also expected to consume up to 16% of the seaweed production if their densities were rising under weak predation conditions. Considering that the population density of herbivores are strongly controlled by fish, human interference like overfishing may have strong negative effects on persistence of seaweeds communities. Keywords: Macroalgal Bed, Herbivore, Seaweed-Grazer Interaction, Grazer Impact, Per Capita Grazing Rate sq(macroalgal bed)p lklp p kp ov l l, k rl vp, om r l oq koqp r, pl p 2 p lklp pp v l t n l p (Steneck et al., 2002). qvp ll l klp sep sql ƒ m p v, sq (p p)p tn np n v p(estes et al., 1998; Dean et al., 2000; Duffy and Hay, 2000). r n p(m l, ENSO event rr m )l p sq lrp, v p v kp l, m p se ekl p p n krrp, m v p vp(steneck et al., 2002). lklp pp p vm k s l e k rp. lkl p tp n lkl pp ql p p p. p r p, pr kp re o sel p ekl ~rp k r n, q l p r r l(, 2001; o, 2004)p sq. Duffy and Hay(2000)l p nl k sp, q p ~ mesograzer mesoherbivore( copepod size 2.5 cm, p t se; Brawley, 1992)l p ek le e tp k p r. k ~ ekl r o l tnp. sl v se p ekp ˆ o e- e plv p(-)p qn(negative interaction)p p. Sala and Graham(2002)p s p r p q~ p e r l l se s 1 ~ qnp (per capita interaction strength) r m. pp r qnp sp sl r (%, se m se sq/q e q~p p ) sep ~ r (v, Michaelis-Menten function) p p ˆ. lvp k sep ~p r ~ qnp r p, rr qpp e p rr p ~ e(per capita grazing rate)p m le p m. Sala and Graham(2002)p p s qnp r m p p (1) v l t r l p sp mkl Žk (2) edšp kr n l r p p m t p l n tn pp p l p. k l rr p ~ ekp m p v, lm l l lql p ek l m vep r, lklp sqp s l pl tn r r pp p. l p se ekl r q kvv kp se p edš l l p k s p sep rr e m ~p r edš qnp ˆ r p Žk l np t pp p. l (1) t ˆ e (mesocosm experiment)p l 12sp ~ 1p e(per capita grazing rate, g seaweeds/individual/day, p PCGR)p r p ˆp s PCGRp m p r, (2) r p r l e np se qnp rl n, (3) sq edš se s qnp m p Žk p trp. m m e sep l~ sp 8s, p 1s, 2s p 1s 12spl. e sp, sp (Aplysia kurodai), (Barleeia sp.), l(cantharidus callichroa), (Chlorostoma argyrostoma turbinata), Žk(Homalopoma sangarense), r(nordotis discus), (Omphalius rusticus), (Turbo sp.)m p e q(dynoides dentisinus) p mn(amphithoe spp.), (Caprella spp.) p (Strongylocentrotus nudus) pl. se p e e r 7p pl o } l q }v p, }v v q d pn l lv nl. lp s el e vrv e p, s l ˆ v k m. se p ~p l v t, v t se (,, s ) se(, s ) l. pl p p. t sep n p el sp e(laminaria japonica) m, o 9ol sp

3 k se } s qn 127 (Pachymeniopsis elliptica) sp (Ecklonia stolonifera) pn l e m. sep n, op sm s nl l(undaria pinnatifida)p q(sargassum fulvellum) p sm sp n(gelidium amansii) p m. se p s lp, t se tl sp 500 ml, se tl sp 20 lp o nl sm l e p m. p o n 1 mm p }p p s m 24e k l e p m. ms ep rr qnp rp o l p s pn l t ˆ e s( sp t l p eq ˆ sp p r ) l e m. e p o(m e ), 5o, 6o, 7o, 9o 11o p 6 l ~ e 24e k p e qpp e l. e k lp s l r m p - o op 11.2 C 7op o 18.7 o C m, o 11ov r rp r ˆl. m e e rp plvv kkp, l p r op k sqp lk rr 4()m tv lk rr 5(l nq )l r m ql p - o p 1.5 pmp e l por m p r k p Ž m. o n l se ~ sm l 24 e s dt p r mp, r ˆ e (blank, s l o n)p pm, r m. sp dtp p el p ˆ l e m p tnpp p p Žl. e rl m e p l 4 sp dt p 30 p r pp el rp mp, dtp 0.05±0.03%(n=52) p ˆ. e el 30 pl r dtp r, p 3 l r p ˆe l r m. o n p sep ~ o 1~14~m. ~ ol e s, Barleeia sp.p pl. l ~p o nl p po pp s ep p ml p. p sp o qll }v e(l. japonica) gl 7~ p (e 2 g k 1 ~ e, 1 ~ ~ g )lp, k 6 gp e p e p s 1 g k 2 ~m. ql p m ~ ~ tp pr p o n l s qp o r v k k. n p ep ~ l 24e kp 1 ~ ep ˆl. e e p ~ 2~3p o n n l ep r m o 11ovp e p k sep p PCGRp r p pn l s ~ dt PCGR r m. p PCGRp p Ž p l r (4th root transformation, )p m. r p p t p, p qr, v, p Ž p p rp se t, p p p p o l ˆt r p ov. dtp, PCGRp s ql (y=a+bx) p Michaelis-Menten function(saturation growthrate model, y=ax/(b+x)) power function model(y=ax b )p rn mp, r Š l ˆ p r m. l e p v kkp, r p qlp l PCGRp ~, ~ m pp p p p psp r pp p Ž m. m r, l, pp p l p r e q~ dtp m/rr v kp pp rr p, lqp p rp ˆ l r p p p rp p Ž m. e, rr e el n p r p ln p m mp, p / p r dtp p p j n p v, n~ ep le e p p n pp t p v kpp ve p km. se e p pp s o l ~ p r PCGR(biomass-normalized PCGR, v, se 1g 1p s e, p ~ r PCGR)p r m. p p ~ slp s el p sq o p. ~ r PCGRp s ~p op sq v Ž k o l Pearson n m. l k sqp o r se ~ ~ ~ q( 29s) n l ~ p vp 1p e(grazing impact, mg/m 2 /day) p r m. sep rm ~ ~p v k~(1999)m l(2002, 2005)p q s m. r r l o l em ~p pnlk p p r p ql e el p p Œplk. r r s om e p n lp r,, l r p lp p p p p. o z PCGR Table 1p sep r PCGR(g seaweeds/individual/

4 128 oqoë vëp rëp}ë}ë që r Ë Table 1. Comparison of linear/non-linear fitting results on the relationship between grazer biomassg (wet weight) and 4th root transformed PCGRG(per capita grazing rate, g seaweeds/individual/day) Model type Equation Standard error Correlation Linear Fit y= x Saturation Growth-Rate Model (Michaelis-Menten Model) y=1.453x/( x) Power Fit y=0.231x Fig. 1. Relationship between the individual biomass(g) and 4th root-transformed PCGR(g) data fitted by non-linear model(power curve fit). day) ~(dt, g WWt/individual) ql rn v p p v ppp lt p. n p q p (r=0.92)p ˆ. v p yrp p p r l ~p 0p k 0.006gp e p ml, p 0p lk se ~ PCGR rr p Žl. p ˆ p v v n r n p power curve (0.89 vs. 0.83)pl. Fig. 1p power curvel p r ˆ p, ~ r (y=0.2310x )p t ep vrp p ˆ. z o PCGR op r sep PCGRp ~ r l pp,, ~ o l, e pp p v kp p Žkl. Fig. 2 ~ r PCGRp r, s s nv m l p. m e p rn o 11ovp l 2 p e p s(12s n 5s) p n m. sp (Aplysia kurodai), r(nordotis discus), Žk (Homalopoma sangarense), (S. nudus), mn (Amphithoe spp.) pl. Fig. 2 sp (A. kurodai) (S. nudus)v ~ r PCGRp ±0.007 g p p p ˆ. Fig. 2. Comparions of biomass-normalized PCGR among species. Cha, Ch. argyrostoma turbinata; Ak, A. kurodai; Or, O. rusticus; Nd, N. discus; Hs, H. sangarense; Cc, C. callichroa; Sn, S. nudus; Dd, D. dentisinus, As, Amphithoe spp.; Bs, Barleeia sp.; Cs, Caprella spp.; Ts, Turbo sp. (S. nudus)p n 5o, 7o, 9o 11ol e p lp gp o( 0.015±0.007 g) p ˆ. mn, Amphithoe spp.p n 5o, 6 o 11ol e p lp gp o (, 0.253±0.081 g) p ˆ. ~ r PCGRp s ~ l op sq p Ž pl(n=36, Pearson s r = , p-value = 0.068). o z sm mk }o r PCGR s p p ~ e n l ~ qn(grazing impact, mg/m )p 2 m(table 2). sql q m p ~p (A. kurodai, k 2,513 mg/m )p p 2 ˆ, pp (S. nudus, k 1,500 mg/m )m 2 (S. intermedius, 733 mg/m 2 ) p p ˆ. (S. nudus)p e 2 ~/m p p 2 rlp, p Sala and Graham(2002) p Strongylocentrotus purpuratusp e p o tp. ~ g p p ~ dtp p ˆ. p n o r 4,000 ~ pp ˆmn(Elasmopus sp.)m 2,000~ pp eovmnp(jassa falcata)p ep 3.435m mg/m /dayp p 2 ˆ. ~ ep p 5,045 mg/m /dayp p 2 rl.

5 k se } s qn 129 Table 2. Average biomass, density, PCGR and impact of grazers on seaweeds Taxa Scientific name Biomass (g/ind.) Density (ind./m 2 ) PCGR (mg/ind./day) Impact (mg/m 2 /day) MPo Acanthochiton spp CAm Aorcho sp CAm Aorides sp CAm Amphithoe spp MGs Aplysia kurodai MGs Barleeia sp MGs Cantharidus callichroa CAm Caprella spp MGs Chlorostoma argyrostoma turbinata CAm Ceradocus sp <0.001 CAm Corophium spp CIs Dynoides dentisinus CAm Elasmopus sp CAm Ericthonius brasiliensis CAm Gammaropsis spp MGs Homalopoma sangarense CAm Hyale spp MPo Ischinochiton comptus CAm Jassa falcata MPo Lepidozona spp MGs Littoraria strirata CAm Maera serratipalma MGs Nordotis discus MGs Omphalius rusticus CAm Podocerus inconspicuus CAm Stenothoe sp EEc Strongylocentrotus nudus EEc Strongylocentrotus intermedius MGs Turbo sp <0.001 CAm, Amphipoda; CIs, Isopoda; MGs, Gastropoda; MPo, Polyplacophora; EEc, Echinoidea }om m l PCGRp 1~ 1p s e p t mp, (1) ~ l s ql p l, (2) eq l ekp m p r, (3) e e pp qp pp l s pp rr mp p 1p e(maximum per capita grazing rate) p ˆ (Sala and Graham, 2002). se ~-PCGR p rl pl r (y=0.2310x )p tep vrp p ˆ. k e(v, e ep k m s)p p lpl lvp ˆ p p p lvp el rr ~ pl p l v k rp ˆ ppp ˆ p p. p sl se p qnp ~ power curvep p k plp, m k pl. y l ~ ekp r l p k kl e (S. nudus)p n pl r p pl. ql e}l nˆ e (enclosure experiment)p o (2004)p ol 0.160, 10ol 0.025, g seaweeds/g sea urchin/dayp r mp, el e p (2001)p n 3sp (Hemicentrotus pulcherrimus, S. nudus, S. intermedius)p o 9o e ep , g seaweeds/g sea urchin/dayp p rl. o (2004)p 8o 10o e r p p rrp Ž p sp pl p p k. l p p r el lp g seaweeds/g sea urchin/dayp p ˆ. l p p g seaweeds/g sea urchin/day 4 p Fig. 3p rrp m p o(mg-g)m o tl p plp, ˆp p Ž pl. k l m p, Sala and Graham(2002)p e l se q~ qnp p Michaelis-Menten

130 oqoë vëp rëp}ë}ë që r Ë Fig. 3. Frequency distribution of grazing impact of 29 populations. l r p ˆ. p p rp v s p Ž p p, p r p q~ p p ~ se p pr ep p l p sekp p (size refuge) p l p o p r v p l m.

6 130 oqoë vëp rëp}ë}ë që r Ë Fig. 3. Frequency distribution of grazing impact of 29 populations. l r p ˆ. p p rp v s p Ž p p, p r p q~ p p ~ se p pr ep p l p sekp p (size refuge) p l p o p r v p l m. sp m~ n l l p sekp p (size refuge) ll ll power curve ˆp r p r p pp o (q~ m~) rp, e se ~l e pl r p p ppp ve p p Žl. Sala and Graham(2002)p e rl r q~l v p(-)p qnp r p rn o s m~ l rn lv sp m~ p lp t k ˆ r p l ~ m~l se p impact r l pn pp p. s ~ r PCGRp Fig. 2 sp 7 ~p n p tplp, np e q(dynoides dentisinus), mn(amphithoe spp.), (Caprella spp.) s Barleeia sp., Turbo sp. l o p p l. pp sl sl ~p qp s p. ~ r PCGRp s ~ s qp np sp s prp eqp p r plv p. s mkm j qk l re ~ qnp p 5,045 mg/m /dayp p 2 ˆ. p se vp s p p o r k 5gp s l ep p p. pm p r p p rp er qp m p pp p m p. ~w, ll r PCGRp, k l m p, eq ps qqp p l p l s p l n p Ž pl. w, ~p 1m l 2 Table 2p p s e pp p n. p ~(m l ˆ mn(elasmopus sp.), spmn(hyale spp.) p n e} p tep sl n~ v s p e}l p n p p rl. w, dp p sp m (epiphytes) s e qep, lp r pp sp e nl p. p rp ql e}l se ~p qn r p p 5g p p Ž p. vp ~ qnp vp s m ˆ p n tn l. sql e Table 2p 29 se ~p p s PCGR ~p qn r rp k qnp sp n mp pp qnp p ˆ(Fig. 3). p e r l Žk edš ~ qnp l l m o p. m l Paine(1992)p Berlow(1999) m p lqp ˆ p ~ (per capita effect) k qnq ol p qnqp sq Ž p p r k m. p pr l(m l, May, 1973)p p edšp qn n edšl p psl krp p p m p. Berlow(1999)p ll p k qnqp p kr l noise-dampening rolep p ˆp, p po pp p ql ˆp vl ƒ o p p m m. qv sq l kl m p qnql rp lr plp, sm p k qnql l r p p. pm, nl l lql p pp qqr p pv srnpl l p (Duffy and Hay, 2000; Christie and Kraufvelin, 2003). k qnq l sq edšp krp l p p o e r l n p. m m omi i sq ˆl e k sep er p n p, kvv pl p s p. lp s rp p m rp sqp sep sl qqr p p m pl. e p sql v p(-)p qn n m ~ p qnql p p p v p ˆp p nl sl p p s pp p l p rqp qnp v pp, prp ep p r pl. e Sala and Graham(2002)p e p o prp, sqp l rrp m p p l q~p pl rp qp ~ qnqp e l p p

7 k se } s qn 131 m pl. kl o (2004)p e l p q q l sqp (S. nudus)p em ~p 3 ~/m 2, g/~, 1 ~/m 2, g/~m. l p p rn l r p q ql sqp qnp m mg/m 2 pl. p ha l ep, ton/ha/yearpl p } kelll p sp p rp k 5 ton/ha(, 621,156 ton, } kell r, 124,668 ha; k, k 3 ~/m 2 p(k 210 g)p e tp s p pl v p p p p v. p (S. nudus) 1m g p e n s e p m p p ˆ ž(2001)p e m o tp p. qp ~ qnq n p ep sql e sep ek ol r p t l p ˆ. p ~p e ~ qnp r l tn m p pp p ˆ p p. Christie and Kraufvelin(2003)p t ˆ e l porp s p ek s (p Carcinus maenasm p Asterias rubens, m l )l m 2l t sep e l 500,000~2,000,000 ~/8 m p ˆ. k l m p, erp p lp m e r (k 10,000 ~/m ) n ek r g/m 2 /daym. k p ek s l pp 25 r v pl p ekp k 0.21 g/m 2 / day, 0.8 ton/ha/yearl p p } kelll p sp (5 ton/ha) p e l tp. ql sq edšp ll p ep } p e sr l tn l p (Christie and Kraufvelin, 2003). ol l se n l ekp pp e l m p t, rp sqp o ppp p l lqp q kr p (o, 2004; McClanahan and Muthiga, 1989; Sala and Zabala, 1996; Estes et al., 1998). qv kv pp eq, s,, l(u,,,, l ) p p(o, 1996; o, 2004; McClanahan and Muthiga, 1989; Sala and Zabala, 1996). le ll p vtrp pn p tn pop p p o n l q k r pp( l, 2002, 2005;, 2003), p ~ p l eql p prp sr p p(fjφsne and Gjφsæter, 1996; McCurdy et al., 2005). o (2004)p eq-se- sq e sr(topdown control) p r mp, Estes et al.(1998)p k l p p p. Christie and Kraufvelin(2003)p e l p tp mkm ~l se ~p n pp mp p l e e sr(bottom-up control)p p p p r p p. p ekp k npl p o eq s p p t l l p m p p p lpp k p pp v ol p p. k m p e- e p m e srp p rr p qo p sel v ekp p, pp e p v eˆ p. l p p sql ƒ oeˆ p qqp v p mp rk l p }p p. p r p s k s l rp plv p, n lk p sqp l on n p e p, r r l p v p. m l 2005 op sq s r se ˆ l p p p ld. p Š l np te t) lk ˆlp rdo lo op o. p e e l vrp te } p eoo. y o, e-r36 ( ). o, 780 pp. l, sq se l( s). o l, 163 pp. l, sq s r se ˆ l. o l, 115 pp. p, p} rp vsm p. p p o, 273 pp. n, k sq s ep. k p o sq s o, v o l, pp oqo, n, p}, r, }, }, m,, q, k } l e, Strongylocentrotus nudusp m e Ž. k v,, 9: v k~, Žr q m s. v k~, 176pp. ÓÔÇ Ô -³Â ÄÔ -. Éª. žé³, Berlow, E.L., Strong effects of weak interactions in ecological communities. Nature, 398: Brawley, S.H., Mesoherbivores. In: Price, J.H., D.M. John, S.J. Hawkins (eds.), Plant-animal interactions in the marine benthos. Systematics Association Special Volume 46. Clarendon Press, Oxford,

8 132 oqoë vëp rëp}ë}ë që r Ë Christie, H. and P. Kraufvelin, Mechanisms regulating amphipod population density within macroalgal communities with low predator impact. Scientia Marina, 67: Dean, T.A., J.L. Bodkin, S.C. Jewett, D.H. Monson and D. Jung, Changes in sea urchin and kelp following a reduction in sea otter density as a result of the Exxon Valdez oil spill. Mar. Ecol. Prog. Ser., 199: Duffy, J.E. and M.E. Hay, Strong impacts of grazing amphipods on the organization of a benthic community. Ecological Monograph, 70: Estes, J.A., M.T. Tinker, T.M. Williams and D.F. Doak, Killer whale predation on sea otters linking oceanic and nearshore ecosystems. Science, 282: Fjφsne K. and Gjφsæter J., Dietary composition and the potential of food competition between 0-group cod (Gadus morhual.) and some other fish species in the littorial zone. ICES Journal of Marine Science, Volume 53, Issue 5, pp May, R.T., Stability and complexity in model ecosystems. Princeton Univ. Press, Princeton, NJ. McClanahan, T.R. and N.A., Muthiga, Patterns of predation on a sea urchin, Echinometra mathaei (de Blainville), on Kenyan coral reefs. J. Exp. Mar. Biol. Ecol., 126: McCurdy, D.G., M.R. Forbes, S.P. Logan, D. Lancaster, S.I. Mautner, Foraging and impacts by benthic fish on the intertidal amphipod Corophium volutator. Journal of Crustacean Biology, 25(4): Paine, R.T., Food-web analysis through field measurement of per capita interaction strength. Nature, 355: Sala, E. and M.H. Graham, Community-wide distribution of predator-prey interaction strength in kelp forests. PNAS, 99: Sala, E. and M. Zabala, Fish predation and the structure of the sea urchin Paracentrotus lividus populations in the NW Mediterranean. Mar. Ecol. Prog. Ser., 140: Steneck, R.S., M.H. Grahan, B.J. Bourque, D. Corbett, J.M. Erlandson, J.A. Estes and M.J. Tegner, Kelp forest ecosystems: biodiversity, stability, resilience and future. Environmental Conservation, 29: o 15p or o 15p r }ˆ voo: e

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<30372E31362D323028BDC5C7F6C5C32DB9CCB1B970626D292E666D> k m v r1 r1 16~20 (2009) J. Korean Academy of Managed Care Pharmacy Vol. 1, No. 1 Pharmacy Benefit Management iç 1# x sƒ, 1 w w w Overview on Pharmacy Benefit Management in USA Kyung Eob ChoiGand Hyun