High dispersal of moray eel larvae to the open ocean : Early life history estimated from ocean-wide distribution patterns Atsushi Tawa 1, Taku Yoshimura 2 and Noritaka Mochioka 1 1 Laboratory of Fisheries Science, Graduate school of Bioresources and Bioenvironmental Science, Kyushu University, 6-10-1, Hakozaki, Higashi-ku, Fukuoka, 812-8581, Japan. E-mail: [email protected]; 2 Seikai National Fisheries Research Institute, Fisheries Research Agency, 1551 –8, Taira, Nagasaki 851 –2213, Japan.

80 Japan Introduction Results and Discussion n=267 60 A1 A1 A2 C 40 Larval fish ecology and early life history are very important All leptocephali (914 ind.) are identified 2 East 57 ind. 48 ind. 20 ・ China 22 sp. 17 sp. 15 information for resource management and the conservation of subfamily, 9 genus , 33 species ・・43 types 0 Sea C n=48 10 30 50 70 90 110 10 marine biodiversity. In addition, larval recruitment of demersal by morphology and DNA barcording. 15 West Madagascar 5 n=57 fishes determines the adult distribution patterns. Moray eels are Species A2 Mariana 10 0 Echidna nebulosa Gymnothorax pseudothyrsoideus 10 30 50 70 90 110 important elements in tropical to temperate coastal ecosystems Guam Western Echidna polyzona Gymnothorax richardsoni 5 339 ind. Indian due to their role as a higher-level predator. Some species are Enchelycore kamara Gymnothorax rueppellii 0 Taiwan Ocean Mauritius Evenchelys macrurus Gymnothorax sagmacephalus 10 30 50 70 90 110 reported to spawn in coastal areas, however, their larval ecology 46 sp. Reunion 80 Gymnomuraena zebra Gymnothorax sp.1 n=210

Number of leptocephali of Number B1 and early life histories are mostly unknown. Therefore, we first Gymnothorax albimarginatus Gymnothorax sp.2 60 Selebes Sea examined the distribution patterns of the leptocephali of Gymnothorax berndti Gymnothorax thyrsoideus 40 B1 Gymnothorax buroensis Gymnothorax ypsilon 20 25 ind. different species or types in the Indo-Pacific to estimate the Gymnothorax chilospilus Gymnothorax zonipectis 210 ind. D 10 0 13 sp. Sumatera D n=25 Gymnothorax chlorostigma Strophidon ui 10 30 50 70 90 110 14 sp. general early life history of the moray eels. Jawa 5 Gymnothorax fimbriatus Scaticaria tigrina 60 n=207 Gymnothorax flavimarginatus B2 Sulawesi 40 B2 Madagascar Gymnothorax isingteena Tyeps 0 Materials and Methods Tomini bay 10 30 50 70 90 110 Gymnothorax kidako Anarchias sp.1-3 20 211 ind. Eastern Indian Ocean Gymnothorax margaritopholus Channomuraena sp. 0 1 Sampling sites (open ocean area) 10 30 50 70 90 110 24 sp. Gymnothorax melanospilos Muraeninae sp.1-24 A1 Around Japan of the A1 Gymnothorax meleagris Uropterygiinae sp.1-15 Total length (mm) western North Pacific (84stations) Gymnothorax mieroszewskii Distribution patterns and lengh-frequency distributions of muraenid leptocphali (Muraenidae spp.) on each Gymnothorax minor sampling site. A2 West Mariana of the Gymnothorax monochlous 76 species in leptocephali ・ western North Pacific (238stns.) B1 A2 Gymnothorax nudivomer The individuals of 20-70 mm TL are often collected on each sampling sites. Gymnothorax intesi 150 Indo-Pacific morays ・ B1 Celebes Sea in Indonesia (27stns.) B2 Most muraenid leptocephali use open ocean in early life history. D B2 Gulf of Tomini in Indonesia (21stns.) C G. minor C Eastern Indian Ocean (12stns.) (Adult informations) Larval distribution G. pseudothyrsoideus >64 46.9 G. buroensis 16 n=99 29.2 D Western Indian Ocean (40stns.) Distribution 65 31.2 ・ South Japan to East China Sea ・ Indo-west Pacific n=8 ・ Indo-Pacific n=23 50~62.1 Open circle show station Larval sampling are Habitat and Taiwan 60~80mmTL RMT IKMT MOHT ORI MOCNESS 11~67 conducted by 5 larval nets Larval net ・ Sandy or muddy bottom ・ Coral reef to inner bay ・ Coral reef 2 19~80 on each station (Table). Size(m ) 1.0/8.0 8.7 5.1 7.1 10.0 Depth (muddy bottom) 60.7 Some specimens collected Mesh(mm) 0.33/4.5 0.5 1.6 0.5 3.0 ・ 50-150 m (moderately deep) ・ 1-50 m (shallow to middle) ・ 10-25 m (shallow) during a seine netting Site A1 B1, B2 A1 A2 A1 Maximam size 40.2~74.6 40.3~61.8 operation for anchovy ・ 60 cm (small to middle) ・ 80 cm (middle to large) 35 ・ 25 cm (small) 25.6 larvae in coastal areas, Leptocephalus Japan(A1) Evidence of larval migration to 18 A20 further research Metamorphosis Na/Ca General early life history and 16 A7 5’ GGCCGCG ～ the open ocean ? A19 14 A16 Migrational route × A17 2 DNA barcoding by mtDNA 16S ○○○ 12 A18 Adult life zone × The value of 3 element showed the Egg ・ Distribution patterns by more sample rRNA partial region (530bp) Adult clearly different between position 1 10 ・ Age and growth study by larval otolith (leptocephalus stage) to 2 (juvenile 8 Return Sr/Ca ・ Otolith trace element analysis Adults 46 species vs. Leptocephli stage). 4 Dispersal and return mechanism 5’ GCGCGAA ～ 5’ AACCGCG ～ 5’ GGCCGCG ～ 3 ・ This result may show that morays Behavioral experiments of olfactory 3 Otolith trace element analysis by using the LA-ICP-MS 2 inhabit the different sea area in Leptocephalus life zone and visual sense Sample: 6 adult specimens ( Gymnothorax kidako ) mmol/mol 1 ・ larva and juvenile stage. However, Ba/Ca Numerical simulation by physical and Otolith We compared the trace element it is possible that this is 1.2 -3 biological parameters composition (3 elements) of four physiological difference between Dispersal Timing of metamorphosis 1 2 3 4 0.8 -3 positions (right figure) for proving larva and juvenile. Furthermore, an ・ physiological study the larval migration from coastal additional experiment is necessary 0.4 -3 area to open ocean. 1 Leptocephalus stagestagestage to clarify the detail migration of 0 We estimated that small leptocephali born in the coastal areas were dispersing widely across the 2juvenile 3young3young3young 4adult4adult4adult 1 2 3 4 moray’s larvae. open ocean. Their leptocephali returned to the coastal area after growing. stagestagestagestage stagestagestage Position