Observations of Large Muraenid Leptocephali in Coastal Indonesia: Locations of Sightings and Behaviour of the Larvae Michael J

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Observations of Large Muraenid Leptocephali in Coastal Indonesia: Locations of Sightings and Behaviour of the Larvae Michael J Marine Biodiversity Records, page 1 of 8. # Marine Biological Association of the United Kingdom, 2013 doi:10.1017/S1755267213000079; Vol. 6; e82; 2013 Published online Observations of large muraenid leptocephali in coastal Indonesia: locations of sightings and behaviour of the larvae michael j. miller1, rob rutgers2, barry haythorne3, tunc‚ yavuzdog'an4, seishoku obata5, tony wu6, henny rutgers2, jerry powell7 and katsumi tsukamoto1 1Atmosphere and Ocean Research Institute, The University of Tokyo, 5-1-5 Kashiwanoha, Kashiwa, Chiba, 277-8564, Japan, 2Hoensbroek, The Netherlands, 3Rotherham, South Yorkshire, United Kingdom, 4Ayıs¸ıg˘ı Diving Center, Bag˘dat Cad, Istanbul, Turkey, 5Omiya-shi, Saitama-ken, Japan, 6Yokohama-shi, Kohoku-ku, Kanagawa-ken, Japan, 7Winterthur Museum & Country Estate, Winterthur, Delaware, USA A large often greenish-appearing type of leptocephalus larva has been observed and photographed or filmed by divers at 4 different locations within the Indonesian Archipelago from 2008 to 2011, and this paper documents these sightings and evalu- ates what can be learned about this type of unusual eel larvae. These leptocephali (300–400 mm long) appear to be excep- tionally large metamorphosing larvae of some type of moray eel of the family Muraenidae. Because of the morphological similarities between these larvae and ribbon eels, Rhinomuraena quaesita, they are hypothesized to be their larvae, which have never been distinguished among leptocephali of the Muraenidae. These leptocephali were observed at Sangeang Island, in Lembeh Strait adjacent to north-eastern Sulawesi Island, in two different years at Ambon Island, and on two con- secutive days at Bali, between December and June of 4 different years. All observations of the large leptocephali were at depths shallower than about 23 m in areas with mixed coral rubble or coarse sand and silt substrates during daylight hours. The larvae used various styles of anguilliform swimming and swam near the bottom or in the lower water column while being observed. One larva tried unsuccessfully several times to enter small crevices. They were all at the metamorphosing stage and may have been entering coastal habitats to find a hiding place where they could transform into juvenile eels. Further studies are needed to clearly identify these large leptocephali and to learn about their life history. Keywords: Anguilliformes, leptocephali, moray eels, ribbon eels, metamorphosis, recruitment, swimming behaviour, Indonesia Submitted 10 April 2012; accepted 18 January 2013 INTRODUCTION leptocephalus species in the Atlantic Ocean (Blache, 1977; Smith, 1979;Bo¨hlke, 1989; Leiby, 1989), but even in the Leptocephali are the larvae of the true eels of the order Atlantic the leptocephali of moray eels of the family Anguilliformes and their close relatives (Smith, 1989a; Muraenidae are especially hard to identify, because of a lack Miller & Tsukamoto, 2004; Miller, 2009). They are remarkably of morphological features that can be linked to the adults transparent because they have greatly reduced body muscula- (Smith, 1989b). ture and organs, and their highly laterally compressed bodies Leptocephali live in the upper few hundred metres of the open are filled with transparent jelly-like materials (Smith, 1989a; ocean or over the continental shelf, with some species showing Pfeiler, 1999; Miller, 2009). They grow larger than most fish diel vertical migrations (Castonguay & McCleave, 1987; Miller, larvae, with maximum sizes ranging from about 50 mm to 2009).Theycanbeabundantalongtheedgeofthecontinental more than 300 mm, but probably ,120 mm for most shelf (Miller et al., 2002; Miller & McCleave, 2007) or in offshore species (Bo¨hlke, 1989). The leptocephali of some species of areas where anguillid and mesopelagic eels spawn (Tsukamoto, congrids can occasionally reach about 400 mm (Mochioka 1992; Miller & McCleave, 1994; Miller et al., 2006). The behav- et al., 1982; Strehlow et al., 1998) and some notacanthiform ioural ecology of eel larvae is very poorly known though. larvae have been collected at sizes up to 1.8 m (Bertin, 1954; Leptocephali enter shallow tropical habitats for recruitment Castle 1959, 1967; Tabeta, 1970). using tidal currents (e.g. Thorrold et al., 1994a, b;Dufouret al., Because their morphology is so different from the juveniles 1996;Harndenet al., 1999;McIlwain,2003;Nolan& and adults, they are difficult to match with the adults of their Danilowicz, 2008), but there have been few reported observations species, so few types of leptocephali have been identified to the of live leptocephali. Divers have recently filmed a few leptocephali species level in the Indo-Pacific (Miller & Tsukamoto, 2004, at night over deep water, providing some new insights into their 2006). Much greater progress has been achieved in identifying behaviour and morphology (Miller et al., 2010, 2013). One of the few reported observations of leptocephali in shallow water was of a large greenish-appearing metamor- Corresponding author: M.J. Miller phosing leptocephalus that was recently photographed adja- Email: [email protected] cent to Sangeang Island in the Flores Sea (Figure 1)of 1 2 michael j. miller et al. 2011 in the Indonesian Archipelago that appear to be the same species of large metamorphosing moray eel larvae. The distribution of these sightings is documented and the video recordings and photographs are evaluated to help advance understanding of these unusual eel larvae that are present in Indonesia and probably elsewhere in the Indo-Pacific. MATERIALS AND METHODS Recording imagery of larvae Observations of the large leptocephali were made by SCUBA divers equipped with underwater cameras or video cameras at 4 general locations (Table 1; Figure 1). The larvae were encountered during dives in shallow water during daytime, and each larva was photographed or filmed for several minutes as it swam along the bottom or in the water Fig. 1. Map showing the locations where large leptocephali were observed column. The larva photographed at Sangeang Island in June along the north-eastern side of Sangeang Island in June 2008, in Lembeh 2008 was not seen at an established dive site, so the exact Strait adjacent to north-eastern Sulawesi Island in late December 2009, on location at the island could not be precisely determined the north shore of Ambon Bay in April 2010 and March 2011, and at Bali (Table 1). Larvae were observed on 2 different dates at dive on two consecutive days in April 2010 (Table 1). sites along the north shore of Ambon Bay of Ambon Island called Rhino City (April 2010) and Laha 3 (March 2011). southern Indonesia (Miller et al., 2009). The leptocephalus The photographs of the larva in the second sighting are not appeared to be about 400 mm in length and was moving included in this study. On the north-eastern shore of Bali at along the bottom (Figure 2). It differed in morphology from a dive site called Seraya between Amed and Tulamben, a lep- other leptocephali described previously (Blache, 1977; Smith, tocephalus was seen on two consecutive days. 1989a; Miller & Tsukamoto, 2004), because it appeared to be a remarkably large leptocephalus of the family Muraenidae. Muraenid leptocephali are rarely collected at Analyses of observations sizes greater than 90 mm (Smith, 1989b; Miller et al., 2006), Video footage was obtained from 4 of the observations so that leptocephalus appeared to be a completely new type (Sulawesi, Ambon and 2 sightings at Bali) and converted of moray eel larva. into video clips. Video from the sightings were converted The present study examines 6 observations of this type of from three different formats and resolutions into a single large leptocephalus that were made by divers from 2008 to master video file. This video clip is permanently available Fig. 2. Photographs showing the body (A–C) and head region (D–F) of the large leptocephalus that was observed along the coast of Sangeang Island, Indonesia on 26 June 2008. External nostrils are seen in (D). large muraenid leptocephali in coastal indonesia 3 online on the website of the Australian Museum (see link in complex structure of the bottom that included coarse sand Results section). and coral rubble. The leptocephalus filmed at Ambon Island Still photographs were taken of the Sangeang, Sulawesi and on 6 April 2010 was seen in the afternoon at a depth of Bali leptocephali. As shown by Miller et al. (2009) some 23 m along the north shore of Ambon Bay in a similar photographs were lightened through a filtering process to complex rubble substrate area. The habitat of the other remove haziness and increase clarity of the images. This larva, observed at an adjacent dive site in Ambon Bay makes it easier to see the morphology of the leptocephali as almost one year later on 27 March 2011, was likely similar seen in Figure 3. based on the location and the background of the photograph (not shown). Less than two weeks after the first sighting at Ambon, single leptocephali were seen at Bali (Figure 1) on the two con- RESULTS secutive days of 18–19 April 2010 (Table 1). It is not certain if these two sightings were of the same leptocephalus, because Locations and habitats of sightings the larvae were apparently about the same size and in the same general area, but in different types of habitats. Large leptocephali with the same basic appearance were However, the leptocephalus on the second day had distinct observed 6 times from 2008 to 2011 at 4 different general scratches on the side of its body, and a twisted region of its locations within the Indonesian Archipelago (Table 1). All notochord, so it was probably a different individual unless of the sightings were made from late morning to mid- the first larva was damaged between sightings. The first sight- afternoon, at a range of depths from about 5 m to 23 m. ing was made in areas with rubble and other types of structure The water temperatures in all of these coastal areas ranged on the bottom (Figure 5C), but the second sighting was made from 27–308C during the sightings from December to June.
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