Recruitment processes of jack mackerel (Trachurus

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Recruitment processes of jack mackerel (Trachurus japonicus) ) in the East China Sea (ECS) in relation to environmental conditions Chiyuki Sassa 1, Youichi Tsukamoto 1, Yoshinobu Konishi 1* Songguang Xie 2, Yoshiro Watanabe 2 and Hideaki Nakata 3 1 Seikai National Fisheries Research Institute, Fisheries Research Agency 2 Ocean Research Institute, The University of Tokyo 3 Nagasaki University * Present address: Southeast Asian Fisheries Development Center (SEAFDEC) 1*,

Introduction (1) The shelf-break regions of the ECS have been considered to be an important spawning ground for various commercially valuable pelagic fishes. jack mackerel chub mackerel spotted chub mackerel yellowtail common squid A large proportion of their eggs, larvae and juveniles are transported to the northern ECS, Pacific and Japan Sea coasts by currents. However, no comprehensive investigation has been made on their recruitment processes in the ECS.

Introduction (2) Taiwan The jack mackerel is one of the most exploited fishery resources in southern Japan The annual catch during 1990 2004 ranged 180 310 thousand metric tons in Japanese waters Annual catch in recent years Korea = several ten thousand t Taiwan = several thousand t

Contents To understand the mechanisms of year-to-year variation in jack mackerel recruitment 1. The main spawning ground and transport processes of larvae and juveniles 2. Key developmental stages and key area determining their year-class strength 3. The relationship between the yearly fluctuation of survival rate and environmental conditions

Sampling area We conducted sampling in the shelf-break region of the East China Sea during February March and April from 2001 to 2004. The total number of sampling stations during each month ranged from 89 to 284. 2001 2002 N = 240 N = 284 February March N = 89 N = 107 Yoko-Maru April

How to estimate the location of the main spawning ground At present, the identification of jack mackerel eggs by morphological characters is very difficult.? Therefore, based on the catch of the newly-hatched larvae < 3 mm notochord length, we estimated their main spawning ground. This size of larvae is reported to be less than 10 d old after the fertilization. Larval sampling Bongo net mouth diameter, 60-cm; mesh size, 0.33-mm Double-oblique tows (0 150 m layer)

The larvae < 3 mm were concentrated in the southern East China Sea south of 28 N during both periods. Horizontal distribution of the newly-hatched larvae < 3 mm during February March and April, 2002.

122 E124 E126 E 128 E130 E 132 E134 E 136 E138 E 140 E142 E 37 N 36 N Japan 35 N 34 N 33 N 32 N 31 N 30 N 29 N 28 N 27 N 26 N 25 N 100 10 250 50 1 Horizontal distribution of the jack mackerel larvae < 3 mm during February March, 2003. Data in the Pacific was after Uehara & Mori (2004).

The egg production of jack mackerel is most abundant in the southern East China Sea during February to March, although they also spawn on a small scale in the coastal region off southern Japan. Comparison of the larval abundance between the southern East China Sea and the coastal region off Kyushu Island.

The following current systems in the East China Sea (1) Kuroshio (2) Kuroshio Branch Current north of Taiwan (3) Kuroshio Branch Current west of Kyushu are active in the continental shelf-break region. Amami Island Large spawning ground

Northeastward larval transport from the main spawning ground (February March, 2002) < 11 d old 11-16 d old 17-22 d old Horizontal distribution of larvae are shown for three body length categories, < 3 mm, 3 5 mm, and 5 10 mm, in relation to the current features.

Juveniles occurred in the downstream area of the main currents of the southern ECS where a large spawning ground was not observed. Juveniles (10 30 mm SL) Main spawning ground 22-44 d old Horizontal distribution of the juveniles (10 30 mm SL) in April. Neuston net mouth size, 1.3 0.75-m mesh size, 1.0-mm February March

The drifter with drogue at 15 m depth Juveniles (30 90 mm NL) In May June Kyushu Juveniles (20 50 mm SL) In April Drifter A Pacific coast of Japan Drifter B During February to March Komatsu & Kawasaki 2003 coastal region off Kyushu Japan Sea A small difference in the distribution of fish larvae A remarkable difference in the transport route in the ECS

Conclusion (1) Taiwan Schema showing the main spawning ground, larval transport, and recruitment into the fishing grounds, in relation to the current features.

Contents To understand the mechanisms of year-to-year variation in jack mackerel recruitment 1. The main spawning ground and transport processes of larvae and juveniles 2. Key developmental stages and key area determining their year-class strength 3. The relationship between the yearly fluctuation of survival rate and environmental conditions

Juvenile abundance vs. Age-0 recruitment

(a) ECS vs. Pacific coast of Japan Age-0 (b) ECS vs. Japan Sea Age-0 Pacific coast of Japan Japan Sea ECS ECS Juvenile Springtime juvenile abundance in the ECS is important for age-0 recruitment into the southern Japan. Survival processes in the ECS are key to understanding the fluctuations in their recruitment.

Jack mackerel s survival during the early life stages in the ECS We compared the abundance of newly-hatched larvae (< 3 mm) in the main spawning ground and juveniles (10 30 mm) in the nursery grounds.

Larval abundance (< 3 mm) vs. juvenile abundance February Survival during larval to Abundance juvenile stages of larvae is < important 3 mm March for determining their year-class = an indicator strength. of the egg production Survival index = Juvenile abundance Larval abundance 10000 April Year-to-year fluctuation of survival during postlarval and juvenile stages

Relationship between the larval growth rate and survival in the ECS using otolith analysis Comparison of the growth trajectory based on the otolith increment widths between larvae and juveniles 3 7 d 8 12 d 18 22 d 13 17 d 3 7 d 8 12 d 18 22 d 13 17 d (Sagittal otolith) Larvae (as Original population) Juvenile (as Survivors )

Larvae (as Original population) Juvenile (as Survivors ) 26% 0% 33% 0 1 2 3 4 5 6 0% 0 1 2 3 4 5 6 26% The faster-growing larvae survived better to the juvenile stage than the slower-growing larvae. 0% 25% 0 2 4 6 8 10 3 7 d 8 12 d Frequency 0% 40% 0% 25% 0 2 4 6 8 10 0 4 8 12 16 20 13 17 d 0% 0 4 8 12 16 20 24% Frequency distributions of increment width were compared between larvae and juveniles. 0% 18% 0 4 8 12 16 20 24 0% 0 4 8 12 16 20 24 Increment width (µm) 18 22 d

Conclusion (2) 1. Survival during postlarval and juvenile stages in the ECS was important for determining their year-class strength. 2. Survival rate was very high in 2001 during our study period. 3. The early growth is one of the important factors determining recruitment success of jack mackerel.

Contents To understand the mechanisms of year-to-year variation in jack mackerel recruitment 1. The main spawning ground and transport processes of larvae and juveniles 2. Key developmental stages and key area determining their year-class strength 3. The relationship between the yearly fluctuation of survival rate and environmental conditions

Why was the survival of jack mackerel so different between 2001 and the other three years Possibilities Year-to-year variation of (1) Habitat temperature (2) Food availability (3) Larval transport condition (4) Medusa abundance Year-to-year fluctuation of survival during postlarval and juvenile stages

Possibilities (1) Year-to-year variation of habitat temperature FEB-MAR APR Spawning ground Nursery ground Survival index No clear relationship was recognized between survival index and habitat temperature in this study. Nursery ground Spawning ground

Possibilities (2) Possibilities (2) Year-to-year variation of food availability What do the jack mackerel larvae eat? Percent of the total number of major prey taxa in relation to size of jack mackerel larvae.

Paracalanus spp. We estimated the egg production rate of Paracalanus spp. which are dominant copepods in the southern ECS (Hsieh & Chiu 2002)

Model of the copepod egg production (Prestidge et al. 1995) Egg Production Rate (EP) = MaxEP e (T-24)/θ (eggs/female/day) Chl. a Chl. a + Chl. a h Parameter values for Paracalanus 1. MaxEP: 60 d -1 at 24 C (Uye & Shibuno 1992) 2. θ : temperature coefficient 6 C (Prestidge et al. 1995) 2 3. T: Monthly average sea surface temperature (after AVHRR [NOAA] ) 4. Chl. a: Monthly average chlorophyll a concentration (after SeaWiFS [NASA] ) Chlorophyll a (μg/l) 1.5 1 0.5 0.4 0.5 0.6 0.7 0.3 0.2 0.1 0 10 15 20 25 0.8 0.9 Temperature ( C ) Characteristics of this model 1

50 40 30 20 10 0 Year-to-year variation of the copepod egg production rate March 2001 2002 2003 2004 30 30 30 30 25 25 25 25 125 130 125 130 125 130 125 130 eggs/female/day Larval distribution The copepod egg production rate in 2001 was much higher than the rates in 2002 2004. Year-to-year fluctuation of survival during postlarval and juvenile stages The higher survival in 2001 might be related to the high food availability.

Possibilities (3) Year-to-year variation of larval transport condition 2001 2002 along the Kuroshio front off northeast Taiwan February March Comparison of larval distribution pattern between 2001 and 2002. Transport Good Poor condition

In 2002 larvae were transported to area outside of their nursery ground, i.e. suboptimal condition? Low survival? Release point 2001 2002 The particle-tracking model by Komatsu Larval transport conditions in the southern ECS February March

Horizontal distribution of the juveniles in April 2001 & 2003 vs. 2004 Different pattern Concentrated in the south A large spawning Juveniles (10 30 mm SL) Larval transport condition from the spawning ground to nursery ground varied largely year-to-year.

Possibilities (4) Year-to-year variation of medusa (Pelagia noctiluca) abundance Late stage larvae and juveniles of jack mackerel associate with medusa. Positive relationships 2001 2001 Kingsford 1993 Why do jack Trachurus mackerel spp. juveniles associate off New with Zealand medusae? The lack of gelatinous material in the stomachs Relationship between densities of jack of jack mackerel mackerel juveniles and the medusae in the ECS collected by the Neuston net. Jack mackerel juvenile associate to gain shelter from predation?

Year-to-year variation of the survival index vs. medusae abundance In 2001 the medusae abundance in the neuston net samples was very high compared to 2002 2004. This might be related to the higher survival in 2001. Survival index Medusae If these jack mackerel juveniles substantially decrease their vulnerability to predation by seeking shelter within medusae tentacles, this commensal behavior with medusae may have important implications relevant to jack mackerel recruitment dynamics in the East China Sea.

Conclusion (3) 1. The copepod egg production rate in 2001 was higher than in 2002 2004, corresponding well with the jack mackerel survival. 2. Yearly fluctuations of larval transport condition were observed, which would affect the survival and recruitment processes. 3. Yearly fluctuations of medusae abundance and jack mackerel survival rate corresponded well in the ECS.

Summary 1. We have found the location of the jack mackerel s main spawning ground and described the larval transport processes. 2. Survival during postlarval to juvenile stages in the ECS was important for determining their year-class strength. 3. We discussed the relationship between the yearly fluctuation of survival during early life stages and the environmental conditions.

Thank you very much Acknowledgment: This work was partially supported by grants from the FRECS2 project of the Japanese Ministry of Agriculture, Forestry and Fisheries.