Temporal Succession of Ecosystem Structure in the Kuroshio Extension Region: Are Gelatinous Zooplankton Species Indicators of Ecosystem Status?

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Temporal Succession of Ecosystem Structure in the Kuroshio Extension Region: Are Gelatinous Zooplankton Species Indicators of Ecosystem Status? H. Saito, K. Hidaka, M. Ichinomiya, Yuichiro Nishibe FRA K. Furuya, Yuta Nishibe, K. Takahashi The Univ. of Tokyo

Kuroshio Extension Region (KEX) KEX NO 3 (μm)

Nursery ground of small epipelagic fish Okunishi et al., in prep April spawned cohort in 2006 Chl-a (mg m -3 ) BL < 1 cm BL 1-3 cm BL 3-5 cm BL 5-7 cm BL 7-9 cm BL > 9 cm

SUPRFISH (Studies on Prediction and Application of Fish Species Alternation) Field Campaign 2008 WK0805 KEX

N Inventory (April 10 m, mmol m 3 ) PON 0.90+0.29 Phyto 0.62+0.19 Detritus 0.13+0.23 HNF 0.15+0.09 MicroZ 0.22+0.20 DON 3.28+0.09 Bacteria 0.19+0.05 N flow to fish NO 3 1.04+0.42 NH 4 +NO 2 0.21+0.13 DIN 1.25+0.46 Copepods 0.09+0.01 Other MZP 0.13+0.21 Daily PON Flux (0.02 mmol N d 1 ) TN 2.52 DIN:TN 0.49 POC:PON 7.3 Phy:Z 3.6

N Inventory (May 10 m, mmol m 3 ) PON 1.40+0.13 Detritus 0.94+0.21 HNF 0.07+0.04 DON 3.41+0.01 N flow to fish Phyto 0.27+0.15 MicroZ 0.14+0.10 NO 3 NH 4 +NO 2 0.01+0.01 Other MZ 0.15+0.11 0.04+0.02 Daily PON Flux (0.05 mmol N d 1 ) Bacteria 0.27+0.03 Copepods 0.10+0.05 DIN 0.16+0.12 TN 1.96 DIN:TN 0.07 POC:PON 8.3 Phy:Z 0.91

Cyanobacteria Dinoflagellates Cryptophytes Prasinophytes Chlorophytes Pelagophytes Haptophytes Diatoms WK0804 -KE1 WK0804 -KE2 SY0902 -S39 SY0902 -S5 WK0905 -T1 WK0905 -T3 WK1004 -SF3 Chlorophyll a (µg L -1 )

Primary production

KEX Copepods Community in spring Oithonidae Paracalanusparvuss.l. Clausocalanidae Oncaeidae Oithona similis Paracalanidae Oithona nana Calocalanus spp. Ctenocalanus vanus Mecynocera clausi Metridia sp. Neocalanus plumchrus Pseudocalanus others Paracalanidae Clanusocalanidae Oncaea After nutrient depletion Mecynocera clausi Calocalanus Clausocalanidae

Ecosystem succession in the KEX J F M A M J J Light Nutrient Phytoplankton Copepods Cryptophytes Diatoms Synechococcus Paracalanus Clanusocalanus Oncaea Nut+ Nano+microphyto. Hervivorous copepods Bottom up control of the ecosystem succession Clausocalanus Calocalanus Nut zero Picophyto. Detritivorous copepods Bactrial production

Gelatinous Mucus Feeder Doliolid Photo: K. Ide

Salp

Appendicularian

Prey size of Tunicates & Copepod Copepod size of food particle 0.2 2 10 (µm) Salp Bacteria HNF Ciliate Doliolid & Appendicularia Tunicates can grow in the absence of microphytoplankton (in which herbivorous copepods such as Paracalanus can not grow).

Doliolids and Salps (ind. m -2, 0-50m) April 2008 Doliolids Salps 40 38 1000 3000 10000 40 38 300 1000 3000 36 36 34 34 32 32 30 140 142 144 146 148 150 152 30 140 142 144 146 148 150 152 >4 order of magnitude in abundance (copepods<1 order)

Doliolids and Salps (ind. m -2, 0-50m) May 2008 Doliolids Salps >5000 1000-5000 500-1000 100-500 <100 (individ. m -2 )

Abundance of doliolids vs salps (ind m 2 ) 25000 20000 Doliolids 15000 10000 5000 0 0 2000 4000 6000 8000 10000 Salps Indicating the contribution of salps and doliolids on ecosystem dynamics is highly variable by stations. Doliolids are more adaptive at oligotrophic picoplankton dominated environments than salps

Clearance rate and daily ration Hidaka in prep. Length (mm) Weight (mg C ind 1 ) Clearance rate (l ind 1 d 1 ) Daily ration (%) Salps aggregate solitary 10.9 3.3 24.0 2.3 0.5 0.3 2.9 0.6 0.9 1.5 2.2 1.5 6.9 62.2 1.8 7.6 Doliolids oozoid 6.9 0.2 0.04 0.003 0.5 0.08 46.5 91.8 Length (mm) Weight (mg C ind 1 ) Clearance rate (l ind 1 d 1 ) Source Salps S. fusiformis 17.0 1.3 6.0 P. confederata 13 0.7 3.6 T. democratica 2.7 0.02 0.1 Doliolids D gegenbauri 6.5 35.0 0.6 3.4 10.5 0.3 Anderson (1985) Harbinson & Gilmer (1976) Deibel (1982) Gibson & Paffenhoffer (2000) Deibel (1982)

Grazing pressure (% d 1 ) Salps Doliolids Avg 1.1+2.6 3.6+5.2 Max 14.9 19.8

Ecological function of salps Producing strong feeding current Feeding on not only micro and nanophytoplankton but also protozoa microzooplankton, copepod nauplius, HNF (grazer of nano and picophytoplankton) Salps may accelerate phytoplankton succession from nanoplankton dominant assemblage to picophytoplankton dominant assemblage (driving trophic cascading). Egestion of fast sinking faecal pellet prevent regeneration of nitrogen and accelerate oligotrophication on the KEX The role of salps on transferring primary production to higher trophic levels is low, rather decreasing the ecological transfer efficiency of primary productoin.

Ecological function of doliolids Weak feeding current Fine mesh ( 0.2 μm) Grazer for pico and nanophytoplankton Asexual reproduction, high growth rate (g>1 d 1 ) Sapphirina spp. feed on doliolids Doliolids adapt to picoplankton dominated oligotrophic environment. Transferring picophytoplankton production to Sapphirina spp. and then juvenile fish. It is suggested the dominancy of doliolids in zooplankton assemblage occur after salp retreatment.

Horizontal distribution of appendicularians April May >10000 5000-10000 1000-5000 500-1000 <500 (individ. m -2 )

Grazing pressure of appendicularians (gut pigment method)

Oncaea spp are dependent on sinking particles, esp. for discarded appendicularian houses. Ecological function of appendicularian High grazing pressure on phytoplankton and production of sticky houses indicate the role of appendicularians is repackaging and gathering small non sinking particles Supporting the production of Oncaea spp., important prey for juvenile fish in the KEX. Appendicularians enhance the feeding ground for juvenile fish by transferring nanopicophytoplankton production to Oncaea. Increasing the ecological transfer efficiency in pico nanophytoplankton dominant environment.

Outburst of tunicates indicates the status of KEX ecosystem seasonal succession. Salp outburst: Driving the ecosystem from micro and nanophytoplankton dominated to picophytoplankton dominated status. Driving the ecosystem to more oligotrophic condition by activating biological pump (preventing regenerated production) Doliolid outburst: Indicating picophytoplankton dominated oligotrophic status of the ecosystem. Appendicularian outburst: Indicating ecosystem succession period from herbivorous copepod dominated condition to detritivore dominated condition. Driving the ecosystem to detritus PON dominated condition.

Seasonal ecosystem succession in the KEX Season Dominant zoopl. Herbivorous copepods Salps Detritivorous copepods Doliolids Appendicularians? N form NO3 NO3 Nanophy. Microphy. Nanophy. Detritus NH4 Detritus Bacteira Picopl. Bacteira Detritus Picopl. N dynamics N supply by ver. mix. Particulated by photosyn. Regenerated production Nitrate consumption by phytopl. Sinking by faecal pellets, mucus net

Remainign issue: Control factor(s) of the dominancy between appendicularians and salps/doliolids. Control factor(s) of the dominancy between copepods and tunicates This is due to limited understanding on the biology and ecology of tunicates. Most zooplanktologists (including myself) have been avoided to study gelatinous zooplankton..

Thank you!