Bull. Kitakyushu Mus. Nat. Hist. Hum. Hist., Ser. A, 17: 49–56, March 31, 2019

Field surveys on the Indonesian , menadoensis using remotely been described. The aim of this study is to clear their habitat Allometric growth in the extant coelacanth lung during capture of a coelacanth off Madagascar. South African operated vehicles from 2005 to 2015 and distribution. ontogenetic development. Nature Communications, 6: Journal of Science 92: 150–151. 8222. DOI: 10.1038/ncomms9222 POUYAUD, L., WIRJOATMODJO, S., RACHMATIKA, I., TJAKRAWIDJAJA, DE VOS, L. and OYUGI, D. 2002. First capture of a coelacanth, A., HADIATY, R. and HADIE, W. 1999. A new species of Masamitsu IWATA 1, Yoshitaka YABUMOTO2, Toshiro SARUWATARI3,4, Shinya YAMAUCHI1, Kenichi FUJII1, METHODS Latimeria chalumnae SMITH, 1939 (Pisces, Latimeriidae), coelacanth. Genetic and morphologic proof. C. R. 1 1 5 5 6 7 off Kenya. South African Journal of Science, 98: 345–347. Academy of Science, 322: 261–267. Rintaro ISHII , Toshiaki MORI , Frensly D. HUKOM , DIRHAMSYAH , Teguh PERISTIWADY , Augy SYAHAILATUA , Two remotely operated vehicles (ROVs) (Kowa; ERDMANN, M. V., CALDWELL, R. L. and MOOSA, M. K. 1998. SMITH, J. L. B. 1939. The living coelacanth fish from South 8 8 8 1 Kawilarang W. A. MASENGI , Ixchel F. MANDAGI , Fransisco PANGALILA and Yoshitaka ABE VEGA300) were used for the surveys. The first ROV was Indonesian ‘king of the sea’ discovered. Nature, 395: 335. Africa. Nature, 143: 748–750. replaced with the second one in 2007. Our ROVs are able to overhang off Talise Island between 144 and 150 m deep. There again in October 2015 on a steep slope (Table 2: E25, E27). FRICKE, H., REINICKE, O., HOFER, H. and NACHTIGALL, W. SMITH, J. L. B. 1953. The second coelacanth. Nature, 171: 1Aquamarine Fukushima, Marine Science Museum, 50 Tatsumi-cho, Onahama, Iwaki, Fukushima, 971-8101, Japan dive up to 300 m depth. These had two vertical, two horizontal was almost no water current in this area. All individuals were The coelacanth ID 28 was found on the 1st November along 1987. Locomotion of the coelacanth Latimeria chalumnae 99–101. 2Kitakyushu Museum of Natural History and Human History, 2-4-1 Higashida, Yahata Higashi-ku, Kitakyushu, Fukuoka, and two right-left propellers and these were controlled from a close to each other under the same overhang and most of them one of the large rocks scattered on a gentle slope in a bay of in its natural environment. Nature, 329: 331–333. SMITH, C. L., RAND, C. S., SCHAEFFER, B. and ATZ, J. W. 1975. 805-0071, Japan boat on surface through a 400 m long tether cable. The stayed still with their head facing downward. Four days later, the Lolak Island by 125 m depth. FRICKE, H., HISSMANN, K., SCHAUER, J., REINICKE, O., KASANG, Latimeria, the living coelacanth, is ovoviviparous. Science, 3 Atmosphere and Ocean Research Institute, The University of Tokyo, 5-1-5 Kashimanohara, Kashiwa-shi, Chiba, 277-8564, Japan underwater operations were visualized on a screen with ID 9 was observed again, alone on a steep slope at 172.9 m The encountered Indonesian during these L. and PLANTE, R. 1991. Habitat and population size of 190: 1105–1106. 4Seikei Education and Research Center for Sustainable Development, Seikei Gakuen, 3-3-1 Kichijoji-Kitamachi, information including directions of ROVs, depth, water depth (Table 2: E12), which was located within several dozen surveys were in caves, alongside large rocks, under overhangs coelacanth Latimeria chalumnae at Grand Comoro. VENTER, P., TIMM, P., GUNN, G., LE ROUX, E., SERFONTEIN, E., Musashino-shi, Tokyo, 180-8633, Japan 5Research Center for Oceanography, Indonesian Institute of Science, Jl. Pasir Putih I, Ancol Timur, Jakarta, 14430, temperature, date and time. All data were directly recorded on meters far from Talise-2 and 30 m deeper. or on steep slopes (Table 2; Fig. 2). Environmental Biology of Fishes, 32: 287–300. SMITH, P., SMITH, E., BENSCH, M., HARDING, D. and 6Technical Implementation Unit Marine Biota Conservation Bitung, Indonesian Institute of Sciences, Jl. Colombo, Kec, Maesa, video-tapes on the boat. Since the 2007 survey, water Four new coelacanths were recorded from the 29th Most individuals were found under overhangs or FRICKE, H., HISSMANN, K., SCHAUER, J., ERDMANN, M., HEEMSTRA, P. 2000. Discovery of a viable population of Bitung Tengah, Maesa, Kota Bitung, Utara, 95511, Indonesia temperature and depth were independently recorded with more September to the 6th October 2009 in Talise and Banggka alongside large rocks. Some individuals were encountered not MOOSA, M. K. and PLANTE, R. 2000. Biogeography of the coelacanths (Latimeria chalumnae SMITH, 1939) at 7Research Center for Deep Sea, Indonesian Institute of Science, Jl. Y. Syaranamual, Guru-Guru, Poka, Ambon, 97233, Indonesia accuracy by external measuring memories (Alec Electronics, Islands and off Manado (Table 2: E13 and 14, ID 13, 14, 15). to hide in any shade and stayed just on rocky slopes. In Indonesian coelacanth. Nature, 403: 38. Sodwana Bay, South Africa. South African Journal of 8 Faculty of Fisheries and Marine Science, Sam Ratulangi University, Jl. Kampus UNSRAT Bahu, Manado, Sulawesi Utara, later named JFE Alec; MDS-MKV/T, MDS-MKV/D). In ID 15 is the only juvenile coelacanth filmed so far. It was in a overhangs and cracks, all individuals had their ventral side HEEMSTRA, P. C., FREEMAN, A. L., WONG, H. Y., HENSLEY, D. Science, 96: 567–568. 95115, Indonesia addition two laser beam irradiators, that provide line lasers of small crack between 164.6 and 170.9 m deep in a temperature alongside and close to a rock wall, but without touching it. A. and RABESANDRATANA, H. D. 1996. First authentic 20-cm distance, were attached to the second ROV and have between 14.5 and 15 °C. Estimated total lengths are shown in Table 3. Most (Received August 24, 2018; accepted November 9, 2018) been used to register the size of encountered . The The shallowest record of a coelacanth was at 115.6 m individuals are more than 1 m in total length. ID 15 is a surveys were conducted off northern parts of Sulawesi Island depth off the southern part of Manado (Table 2: E16, juvenile individual of 31.5 cm and ID 23 is 90 cm in total and along the Biak Island located in northwestern New Guinea Manado-3) on the 9th October in 2009. ID 16 (Table 2) was length. No individual beyond 140 cm has been recorded in ABSTRACT − Habitats of the Indonesian coelacanth, Latimeria menadoensis, were investigated by Remotely Island, Indonesia. The areas and time periods of each survey found at the end of the morning, alone alongside a large rock Indonesia so far. restricted as those of the African coelacanths, and so a broader and other staffs for our work on the research boat. Dr. BATUNA Operated Vehicles (ROVs) surveys in the northern coast of Sulawesi Island and southern coast of Biak Island by are shown in Table 1 and Fig. 1. on the edge of a shelf. The water temperature was about 20.0 distribution of the Indonesian coelacanth could be expected. and the Murex staffs supported our daily life during the collaboration of Aquamarine Fukushima (Japan), and Indonesian Institute of Sciences and Sam Ratulangi ºC. Occurrences of coelacanths along the eastern coast of the expeditions. We would like to express our sincere thanks to University (Indonesia) from 2005 to 2015. The Remotely Operated Vehicles operations were conducted 1173 From the 6th to the 16th November in 2010, the field of the propellers of the ROV. level changed, the water flow was reversed. Coelacanths African continent were recorded in South Africa (SMITH, colleagues at Aquamarine Fukushima, Japan, for their times and coelacanths were encountered 30 times. A total of 30 different individuals were observed at a depth range RESULTS survey was conducted around Biak Island (Table 1: R9). Adult DISCUSSION Temperature and depth data for each Indonesian always kept their head against the water flow, like other 1939), Comoros (SMITH, 1953), Madagascar (HEEMSTRA et al., understanding and supports. We would like to thank Dr. Gaël from 115.6 m to 218.9 m deep. The water temperature was between 12.4 to 21.5 ºC. Most of the individuals were coelacanths were found at two different sites (Table 2: coelacanth record are shown in Fig. 3. Though an average of observed actinopterygian fishes did. It means that as the 1996), Kenya (DE VOS and OYUGI, 2002), Tanzania (BENNO et CLÉMENT of Muséum national d’Histoire naturelle and Dr. found alone, however, schools of two, three and six individuals were also observed. The Indonesian coelacanth, L. The field survey of the Indonesian coelacanth, Latimeria E17–19). Two individuals (ID 17 and 18) were found under an During the field surveys by underwater ROV recording water temperature on the ocean surface was about 30 °C, the direction of the water flow changed, the swimming direction of al., 2006), and Mozambique (BRUTON et al., 1992). In Camila CUPELLO of Universidade do Estado do Rio de Janeiro menadoensis, was observed at a similar depth of the African coelacanth, Latimeria chalumnae, and the temperature menadoensis, was conducted 14 times from 2005 to 2015 with overhang between 212.5 and 218.9 m deep (Table 2: Biak-1) for Indonesian coelacanths, Latimeria menadoensis, from 2005 temperature often dropped into about 11 °C at 300 m water depth. each coelacanth individual changed accordingly. Indonesia, coelacanths have only been found off the northern as a referee for their critical reading of the manuscript and their range also seems to be very similar. However, Latimeria menadoensis was sometimes observed besides big rocks 1173 underwater operations around Sulawesi and Biak islands on the 11th November. Three other ones (ID 19, 20, 21) were to 2015, 30 different individuals were identified and three The temperatures changed widely even though the changes of On the 24th September 2009 (Table 2: E11) six coast of the Sulawesi Island and the southern coast of the Biak comments. or the steep wall. It seems to be less sensitive to daylight than L. chalumnae. Here we report also, for the first time in Indonesia (Fig. 1, Table 1). No coelacanth was encountered registered under an overhang between 193.2 and 195.9 m deep others were unidentified. Among those, six individuals were depth were small at several places. All Indonesian coelacanths, individuals were recognized under an overhang (Talise-2). The Island in northern New Guinea. Unfortunately one of our authors, Dr. Djoko Hadi KUNARSO in the world, a juvenile coelacanth was observed in a small crack at 165 to 171 m depth during these surveys. during the first surveys in 2005 although 452 underwater (Table 2: Biak-2) on the 13th November. Two days later, observed more than twice: ID 3 at Buol-2, 3, 5, 6 (Table 2: E3, L. menadoensis, were encountered during day time at a depth all had about the same size. There was almost no water flow, Field surveys of the Indonesian coelacanths should be has passed away during the process of this contribution. We operations were conducted around Manado-tua Island and spots pattern on its body, an ID number was allocated. The temperature changed up and down by five degrees during the between the maximum and the minimum was 6.7 °C, which Biak-2 was observed again and only ID 20 stayed there. E4, E6 and E7); ID 8 at Talise-1 (E9 and E10), ID 9 at Talise 2 between 115.6 and 218.9 m and a temperature between 12.4 and each individual was head downward. They were close to continued in all area of Indonesia and expanded to all pray for the response of his soul. KEY WORDS: Indonesian coelacanth, underwater survey, Latimeria menadoensis, Sulawesi, Biak, ROV other close Indonesian islands from the 17th to 30th of April individual which could not be distinguished was allocated an observation due to water flow reversals. When the water flow was the largest range of all surveys. Two new coelacanths were recorded off Manado in and 3 (E11 and E12), ID 20 at Biak-2 (E18 and E19), ID 24 at and 21.5 ºC. Indonesian coelacanths have exceptionally been each other but no direct contacts or interactions between Southeast Asia in order to increase our knowledge of the This study was supported in part by funding from survey. (Table 1, R1). The second field survey was held along the UN (unknown) + number. was reversed some times, all individuals always changed their On the next day, on the 15th December, the ID 3 was December 2010 (Table 1: R10; Table 2: E 20 and 21; ID 22 Manado-6 and 8 (E22 and E24), ID 26 at Lolak-1 and 3 (E25 recorded in an environment with a temperature exceeding 20 coelacanths were observed. No evidence of a social behavior distribution of this rare species and to define, in a close future, Interdisciplinary Collaborative Research program of Atmosphere northern coast of Sulawesi from the 6th to 19th May in 2006, In the end of the morning of the same day, a third directions against the water flow. observed again under an overhang (Table 2: E7, Buol-6) near and 23). and E27). ID 8 was observed on the 14th September 2009 and ºC, regardless of the depth. The African coelacanth, L. has been recognized in the Comorian populations of the most relevant conservation policies of the different known and Ocean Research Institute, the University of Tokyo. but once again despite of 107 underwater operations no individual (Table 2: E3, ID 3) was found in another cave During the field survey carried on December of the same other sites where it was previously found at Buol-2 , Buol-3 On the 4th May 2012, an individual (Table 2: E22, ID 24) was encountered again at the same place (Talise-1) the day chalumnae, was observed by submarine vehicles in steep coelacanths (FRICKE et al., 1991). populations. coelacanth was recorded. (Buol-2), larger and located 20 m deeper (Buol-2) than the first year, in the same Buol area (Table 1: R4) the sixth individual and Buol-5. Its posture was upside down with its ventral side was found on a steep slope at 169.4 m depth off Manado after. ID 24 was observed on the 4th May 2012 and then 13 slopes between 150 m and 253 m (FRICKE et al., 1991). Divers According to the observations of the African coelacanths, INTRODUCTION of the reproduction biology of the coelacanths is almost The first individual of Indonesian coelacanth (Table 2, ID cave (Buol-1). After 29 minutes of recording, this individual (Table 2: E5, ID 6) was recognized at 14:34 on the 12th facing the ceiling of the overhang. Such posture is also known (Table 2: Manado-6). It swam up and down 5.4 m between months later, on the 9th June 2013 at the same locations of recorded the presence of this species at 104 m depth at the species L. chalumnae is nocturnal and hides in cave in REFERENCES restricted to an ovoviviparity strategy since some caught large 1) was recorded during the third field survey, in the morning of hid inside the cave. Although the water temperature increased December alongside a large rock. This individual stayed in the in coelacanths from the Comorian Archipelago (FRICKE et al., 168.1 and 173.5 m deep for 17 minutes. Thirteen months later Manado area within several kilometers. Sodwana Bay, South Africa (VENTER et al., 2000). Latimeria daytime but comes out at night, supposedly in search of food ACKNOWLEDGMENTS The first living coelacanth, Latimeria chalumnae, was females carried unfertilized eggs or embryos in their oviduct the 30th May 2006 (Table 1, R3). The coelacanth was of more than five degrees, from 14.8 to 20.4 ºC, it stayed at the beginning of the observation and then moved ahead to deeper 1987). ID 3 was found totally four times in this area in two (the 9th June 2003), this individual (ID 24) was observed again All individuals observed more than twice were chalumnae seem to prefer a temperature range of 15–20 °C, (FRICKE et al., 1991). The living geomorphologic environment BENNO, B., VERHEIJI, E., STAPLEY, J., RUMISHA, C., NGATUNGA, discovered in South Africa in the in 1938 (SMITH et al., 1975; CUPELLO et al., 2015). observed in a cave of 165 m depth (Table 2, Boul-1) for 10 same place, not moving away. Despite it disappeared from the slowly. Eventually it reached 12 m deeper during one hour and surveys during the seven months and seems to live in this area. near a cliff at 152.3 m depth (Table 2: E24, Manado-8), where encountered in the same area, at locations close from each where they tend to choose their daytime habitats (FRICKE et al., of L. menadoensis is similar to that of L. chalumnae, with We are grateful to Dr. Teruya UYENO for his valuable B., ABDALLAH, A. and KALOMBO, H. 2006. Coelacanth (SMITH, 1939). The first observation of the living coelacanth The Indonesian coelacanth, L. menadoensis, was first minutes (Table 2, E1). The adult individual was in stationary camera frame we kept setting the ROV there in front of the 42 minutes. During the 2007 survey (Table 1: R5) a coelacanth (Table located on the same slope of Manado-6. other within hundreds of meters. However, in the Comorian 1991). The suitable temperature for L. menadoensis seems to be rocky steep slopes with undermarine caves or overhangs. advice, guidance and encouragement throughout the present (Latimeria chalumnae SMITH, 1939) discoveries and habitats using submersibles was reported in Comoros (FRICKE discovered in Manado, Sulawesi Island, Indonesia, in 1997. position in a cave (Buol-1) slowly moving its pectoral, pelvic, cave until the next morning, however it did not appear again. On the 14th December, the individual ID 3 previously 2: ID 7) was recorded off Manado by 195 m depth, with a Another new coelacanth (Table 2: E23, ID 25) was Archipelago, same individuals of L. chalumnae were observed, almost the same as L. chalumnae. However, Latimeria menadoensis was also observed alongside study. Many thanks go to colleagues of the Faculty of Fisheries conservation in Tanzania. South African Journal of et al., 1987). Latimeria chalumnae hide in submarine caves The first specimen was captured in 1998 (ERDMANN et al., second dorsal and anal fins. The day after, another individual On the 4th June in 2006, three individuals, including ID 3 recorded at Buol-2 in a cave and at Buol-3 in a crack, was temperature of 12.4 °C, the coolest one registered during all recorded alone on a Manado site (Manado-7). within a two week period, in several caves distributed in a 8 Water temperature can increase or drop for more than 5 large rocks or along vertical wall in daytime. Some of these and Marine Sciences of the Sam Ratulangi University at Science, 102: 486–490. along steep slopes between 100 m and 200 m depth during 1998). According to a genetic study, the specimen differed (ID 2) and an unidentified one (UN1) were found in the same which was observed on the 31th May at Buol-2 and two new encountered again (Table 2: Buol-5) near a vertical wall at 145 surveys. Field surveys were conducted from the 19th to 30th May kilometers wide area (FRICKE et al., 1991). °C in a very short period of time (about 30 min according to our individuals were encountered above 200 m deep, in an Manado, Indonesia, for their generous supports to conduct BRUTON, M. N., CABRAL, A. J. P. and FRICKE, H. 1992. First daytime and come out and drift near the ocean floor for from L. chalumnae and a new species was erected (POUYAUD cave (Buol-1). The unidentified one immediately hid deeper in individuals, were observed in a vertical crack which was m depth (Table 2: E6). After a while, this individual (ID 3) The field survey in 2009 was carried out off Manado and 2015 off Lolak Island and from the 30th October to the 16th When individuals of L. menadoensis were encountered, recordings). The coelacanths however often stayed there environment that remains slightly bright. Latimeria expeditions. Also we express our thanks to Dr. SUHARSONO, Dr. capture of a coelacanth, Latimeria chalumnae (Pisces, hunting prey at night (FRICKE et al., 1991). However, since the et al., 1999). In 1999 two individuals were observed by the cave, where the ROV could not access, and only its lateral several hundred meters far from Buol-2 (Buol-3) for 2 hours moved upward about 18 m and stayed behind a large rock (Fig. around Talise and Bangka islands off the northern coast of November 2015 off Lolak and Bitung Islands (Table 1; R13 some of them stayed stationary at same place but some without moving away or significantly modifying their menadoensis seems to be less sensitive to the daylight than L. Zainal ARIFIN, Dr. DIRHAMSYAH and colleagues of the Indonesian Latimeriidae), off Mozambique. South African Journal of record of juvenile or small (below 80 cm long) individuals is submersible in a location 360 km away from Manado (FRICKE side of the body was recorded. The individual (ID 2) was and 14 minutes (Fig. 2C; Table 2: E4, ID 3, 4, 5). During this 3). The observation time was the longest of all surveys with Sulawesi Island from the 12th September to the 9th October and R14). Three new individuals (Table 2: ID 26–28) were individuals such as ID 6 on the 12th December 2006 (E5) and behavior. These water currents of different temperatures are chalumnae. Institute of Science for their advice and administrative work to Science, 88: 225–227. virtually absent, the growth, development and breeding et al., 2000). Today 8 individuals were officially caught in the observed for one minute before it hid inside the cave (Table 2: long period of time all individuals stayed near the rock wall four hours and 16 minutes of recording. The temperature (Table 1: R8). On the 24th September 2009, six individuals recognized at different places off Lolak Island (Lolak 1-4). All ID 24 on the 4th May 2012 (E22) swam away. It seemed to caused by the movement of the marine thermocline. During These first observations are potentially crucial since the obtain our research permit. Other thanks go to Mr. OPO, Mr. CUPELLO, C., BRITO, P. M., HERBIN, M., MEUNIER, F. J., biology of L. chalumnae remain poorly known. Our knowledge Indonesian seas. However, the details of their habitat have not E2). When a new individual could be identified by the white always keeping their head against the water flow. The water changed up and down several times (Fig. 4). Its difference (Table 2: E11, ID 9, 10, 11, 12, UN 2, 3) were found under an individuals were alone. ID 26 was observed in May and once avoid and escape from the brightness of the light or the sounds our surveys we observed in some area that as the thermocline habitats of the Indonesian coelacanths could be possibly not as REFRY and colleagues at Murex Dive Resort, and DAUD, AL, J ANVIER, P., DUTEL, H. and CLÉMENT, G. 2015. 50 IWATA et al. Field surveys on Latimeria menadoensis using ROV from 2005 to 2015 51

been described. The aim of this study is to clear their habitat Table 1. Periods and areas of the Latimeria menadoensis survey using ROV. Total of 14 Allometric growth in the extant coelacanth lung during capture of a coelacanth off Madagascar. South African and distribution. expeditions were conducted in the north of Sulawesi Island and Biak Island located in north west ontogenetic development. Nature Communications, 6: Journal of Science 92: 150–151. of New Guinea Island. The survey areas are shown in Fig. 1. 8222. DOI: 10.1038/ncomms9222 POUYAUD, L., WIRJOATMODJO, S., RACHMATIKA, I., TJAKRAWIDJAJA, DE VOS, L. and OYUGI, D. 2002. First capture of a coelacanth, A., HADIATY, R. and HADIE, W. 1999. A new species of No. Date Survey Area METHODS Latimeria chalumnae SMITH, 1939 (Pisces, Latimeriidae), coelacanth. Genetic and morphologic proof. C. R. R1 17 Apr. – 30 Apr. 2005 Manado-tua Isl. and other islands off Kenya. South African Journal of Science, 98: 345–347. Academy of Science, 322: 261–267. Two remotely operated vehicles (ROVs) (Kowa; R2 6 May – 19. May 2006 Northern coast of Sulawesi ERDMANN, M. V., CALDWELL, R. L. and MOOSA, M. K. 1998. SMITH, J. L. B. 1939. The living coelacanth fish from South VEGA300) were used for the surveys. The first ROV was R3 27 May – 5 Jun. 2006 Buol Indonesian ‘king of the sea’ discovered. Nature, 395: 335. Africa. Nature, 143: 748–750. replaced with the second one in 2007. Our ROVs are able to R4 11 Dec. – 20 Dec. 2006 Buol overhang off Talise Island between 144 and 150 m deep. There again in October 2015 on a steep slope (Table 2: E25, E27). FRICKE, H., REINICKE, O., HOFER, H. and NACHTIGALL, W. SMITH, J. L. B. 1953. The second coelacanth. Nature, 171: dive up to 300 m depth. These had two vertical, two horizontal was almost no water current in this area. All individuals were The coelacanth ID 28 was found on the 1st November along 1987. Locomotion of the coelacanth Latimeria chalumnae 99–101. R5 27 Jun. – 5 Jul. 2007 Manado and two right-left propellers and these were controlled from a close to each other under the same overhang and most of them one of the large rocks scattered on a gentle slope in a bay of in its natural environment. Nature, 329: 331–333. SMITH, C. L., RAND, C. S., SCHAEFFER, B. and ATZ, J. W. 1975. R6 9 Jul. – 12 Jul. 2007 Buol boat on surface through a 400 m long tether cable. The stayed still with their head facing downward. Four days later, the Lolak Island by 125 m depth. FRICKE, H., HISSMANN, K., SCHAUER, J., REINICKE, O., KASANG, Latimeria, the living coelacanth, is ovoviviparous. Science, underwater operations were visualized on a screen with R7 2 Dec. – 8 Dec. 2008 Talise and Bangka Isl. ID 9 was observed again, alone on a steep slope at 172.9 m The encountered Indonesian coelacanths during these L. and PLANTE, R. 1991. Habitat and population size of 190: 1105–1106. information including directions of ROVs, depth, water R8 12 Sep. – 9 Oct. 2009 Talise and Bangka Isl. Manado depth (Table 2: E12), which was located within several dozen surveys were in caves, alongside large rocks, under overhangs coelacanth Latimeria chalumnae at Grand Comoro. VENTER, P., TIMM, P., GUNN, G., LE ROUX, E., SERFONTEIN, E., temperature, date and time. All data were directly recorded on R9 6 Nov – 16 Nov. 2010 Biak Island meters far from Talise-2 and 30 m deeper. or on steep slopes (Table 2; Fig. 2). Environmental Biology of Fishes, 32: 287–300. SMITH, P., SMITH, E., BENSCH, M., HARDING, D. and video-tapes on the boat. Since the 2007 survey, water R10 5 Dec. – 18 Dec. 2010 Manado Four new coelacanths were recorded from the 29th Most individuals were found under overhangs or FRICKE, H., HISSMANN, K., SCHAUER, J., ERDMANN, M., HEEMSTRA, P. 2000. Discovery of a viable population of temperature and depth were independently recorded with more September to the 6th October 2009 in Talise and Banggka alongside large rocks. Some individuals were encountered not MOOSA, M. K. and PLANTE, R. 2000. Biogeography of the coelacanths (Latimeria chalumnae SMITH, 1939) at R11 2 May – 13 May 2012 Manado accuracy by external measuring memories (Alec Electronics, Islands and off Manado (Table 2: E13 and 14, ID 13, 14, 15). to hide in any shade and stayed just on rocky slopes. In Indonesian coelacanth. Nature, 403: 38. Sodwana Bay, South Africa. South African Journal of R12 3 Jun. – 10 Jun. 2013 Manado later named JFE Alec; MDS-MKV/T, MDS-MKV/D). In ID 15 is the only juvenile coelacanth filmed so far. It was in a overhangs and cracks, all individuals had their ventral side HEEMSTRA, P. C., FREEMAN, A. L., WONG, H. Y., HENSLEY, D. Science, 96: 567–568. addition two laser beam irradiators, that provide line lasers of R13 19 May – 30 May 2015 Bitung, Lolak small crack between 164.6 and 170.9 m deep in a temperature alongside and close to a rock wall, but without touching it. A. and RABESANDRATANA, H. D. 1996. First authentic 20-cm distance, were attached to the second ROV and have R14 30 Oct. – 16 Nov. 2015 Bitung, Lolak between 14.5 and 15 °C. Estimated total lengths are shown in Table 3. Most been used to register the size of encountered animals. The The shallowest record of a coelacanth was at 115.6 m individuals are more than 1 m in total length. ID 15 is a surveys were conducted off northern parts of Sulawesi Island depth off the southern part of Manado (Table 2: E16, juvenile individual of 31.5 cm and ID 23 is 90 cm in total and along the Biak Island located in northwestern New Guinea Table 2. Registers of Latimeria menadoensis. Time shows start, end and duration of each Manado-3) on the 9th October in 2009. ID 16 (Table 2) was length. No individual beyond 140 cm has been recorded in Island, Indonesia. The areas and time periods of each survey observation. When an individual could be identified by patterns of white spot on the body, an ID found at the end of the morning, alone alongside a large rock Indonesia so far. restricted as those of the African coelacanths, and so a broader and other staffs for our work on the research boat. Dr. BATUNA are shown in Table 1 and Fig. 1. number was allocated. Individuals that could not been distinguished were allocated UN (unknown) on the edge of a shelf. The water temperature was about 20.0 distribution of the Indonesian coelacanth could be expected. and the Murex staffs supported our daily life during the + number. Individuals observed several times were identified as numbers with underbar. Duration ºC. Occurrences of coelacanths along the eastern coast of the expeditions. We would like to express our sincere thanks to shows long of each observation time. Range indicates vertical movement of the individual during From the 6th to the 16th November in 2010, the field of the propellers of the ROV. level changed, the water flow was reversed. Coelacanths African continent were recorded in South Africa (SMITH, colleagues at Aquamarine Fukushima, Japan, for their survey was conducted around Biak Island (Table 1: R9). Adult MITH EEMSTRA RESULTS observation and not the distance traveled horizontally. Some individuals swam along the slope. DISCUSSION Temperature and depth data for each Indonesian always kept their head against the water flow, like other 1939), Comoros (S , 1953), Madagascar (H et al., understanding and supports. We would like to thank Dr. Gaël coelacanths were found at two different sites (Table 2: coelacanth record are shown in Fig. 3. Though an average of observed actinopterygian fishes did. It means that as the 1996), Kenya (DE VOS and OYUGI, 2002), Tanzania (BENNO et CLÉMENT of Muséum national d’Histoire naturelle and Dr. Time Depth (m) Temperature (ºC) The field survey of the Indonesian coelacanth, Latimeria Fig. 1. Localities of the Indonesian coelacanth survey using No. Date ID Configuration Place Site E17–19). Two individuals (ID 17 and 18) were found under an During the field surveys by underwater ROV recording water temperature on the ocean surface was about 30 °C, the direction of the water flow changed, the swimming direction of al., 2006), and Mozambique (BRUTON et al., 1992). In Camila CUPELLO of Universidade do Estado do Rio de Janeiro Start End Duration Max. Min. Range Max. Min. Ave. menadoensis, was conducted 14 times from 2005 to 2015 with ROV from 2005 to 2015. E1 30. May 06 8:30 8:40 10 165.0 165.0 0 17.1 16.4 16.8 1 cave Buol Buol-1 overhang between 212.5 and 218.9 m deep (Table 2: Biak-1) for Indonesian coelacanths, Latimeria menadoensis, from 2005 temperature often dropped into about 11 °C at 300 m water depth. each coelacanth individual changed accordingly. Indonesia, coelacanths have only been found off the northern as a referee for their critical reading of the manuscript and their 1173 underwater operations around Sulawesi and Biak islands E2 31. May 06 7:58 7:59 1 165.0 165.0 0 15.5 15.3 15.4 2,UN1 cave Buol Buol-1 on the 11th November. Three other ones (ID 19, 20, 21) were to 2015, 30 different individuals were identified and three The temperatures changed widely even though the changes of On the 24th September 2009 (Table 2: E11) six coast of the Sulawesi Island and the southern coast of the Biak comments. E3 31. May 06 11:15 11:44 29 183.0 183.0 0 20.4 14.8 17.5 3 cave Buol Buol-2 in Indonesia (Fig. 1, Table 1). No coelacanth was encountered registered under an overhang between 193.2 and 195.9 m deep others were unidentified. Among those, six individuals were depth were small at several places. All Indonesian coelacanths, individuals were recognized under an overhang (Talise-2). The Island in northern New Guinea. Unfortunately one of our authors, Dr. Djoko Hadi KUNARSO E4 4. Jun. 06 18:42 20:56 134 156.2 153.9 2.3 17.8 12.8 13.7 3,4,5 crack Buol Buol-3 during the first surveys in 2005 although 452 underwater E5 12. Dec. 06 14:34 16:16 102 163.8 151.0 12.8 16.6 14.2 15.7 6 wall Buol Buol-4 (Table 2: Biak-2) on the 13th November. Two days later, observed more than twice: ID 3 at Buol-2, 3, 5, 6 (Table 2: E3, L. menadoensis, were encountered during day time at a depth all had about the same size. There was almost no water flow, Field surveys of the Indonesian coelacanths should be has passed away during the process of this contribution. We operations were conducted around Manado-tua Island and spots pattern on its body, an ID number was allocated. The E6 1. Dec. 06 14:21 18:34 253 145.0 126.5 18.5 21.2 14.5 17.9 3 wall Buol Buol-5 temperature changed up and down by five degrees during the between the maximum and the minimum was 6.7 °C, which Biak-2 was observed again and only ID 20 stayed there. E4, E6 and E7); ID 8 at Talise-1 (E9 and E10), ID 9 at Talise 2 between 115.6 and 218.9 m and a temperature between 12.4 and each individual was head downward. They were close to continued in all area of Indonesia and expanded to all pray for the response of his soul. E7 15. Dec 06 14:02 14:24 22 154.2 152.3 1.9 15.6 15.2 15.4 3 over hang Buol Buol-6 other close Indonesian islands from the 17th to 30th of April individual which could not be distinguished was allocated an E8 27. Jun. 07 12:15 12:46 31 195.0 190.8 4.2 16.2 12.4 15.0 7 beside big rock Mando Mando-1 observation due to water flow reversals. When the water flow was the largest range of all surveys. Two new coelacanths were recorded off Manado in and 3 (E11 and E12), ID 20 at Biak-2 (E18 and E19), ID 24 at and 21.5 ºC. Indonesian coelacanths have exceptionally been each other but no direct contacts or interactions between Southeast Asia in order to increase our knowledge of the This study was supported in part by funding from (Table 1, R1). The second field survey was held along the UN (unknown) + number. E9 14. Sep. 09 10:33 11:00 27 172.2 159.8 12.4 15.8 14.3 15.5 8 over hang Talise Isl. Talise-1 was reversed some times, all individuals always changed their On the next day, on the 15th December, the ID 3 was December 2010 (Table 1: R10; Table 2: E 20 and 21; ID 22 Manado-6 and 8 (E22 and E24), ID 26 at Lolak-1 and 3 (E25 recorded in an environment with a temperature exceeding 20 coelacanths were observed. No evidence of a social behavior distribution of this rare species and to define, in a close future, Interdisciplinary Collaborative Research program of Atmosphere northern coast of Sulawesi from the 6th to 19th May in 2006, In the end of the morning of the same day, a third E10 15. Sep. 09 8:40 8:42 2 166.8 160.6 6.2 20.2 19.7 20.1 8 over hang Talise Isl. Talise-1 directions against the water flow. observed again under an overhang (Table 2: E7, Buol-6) near and 23). and E27). ID 8 was observed on the 14th September 2009 and ºC, regardless of the depth. The African coelacanth, L. has been recognized in the Comorian populations of the most relevant conservation policies of the different known and Ocean Research Institute, the University of Tokyo. 9,10,11,12 E11 24. Sep. 09 14:51 14:55 4 149.9 143.7 6.2 18.5 18.2 18.4 over hang Talise Isl. Talise-2 but once again despite of 107 underwater operations no individual (Table 2: E3, ID 3) was found in another cave UN2,UN3 During the field survey carried on December of the same other sites where it was previously found at Buol-2 , Buol-3 On the 4th May 2012, an individual (Table 2: E22, ID 24) was encountered again at the same place (Talise-1) the day chalumnae, was observed by submarine vehicles in steep coelacanths (FRICKE et al., 1991). populations. coelacanth was recorded. (Buol-2), larger and located 20 m deeper (Buol-2) than the first E12 28. Sep. 09 8:22 8:30 8 172.9 158.9 14 18.5 17.6 18.0 9 steep slope, beside big rock Talise Isl. Talise-3 year, in the same Buol area (Table 1: R4) the sixth individual and Buol-5. Its posture was upside down with its ventral side was found on a steep slope at 169.4 m depth off Manado after. ID 24 was observed on the 4th May 2012 and then 13 slopes between 150 m and 253 m (FRICKE et al., 1991). Divers According to the observations of the African coelacanths, of the reproduction biology of the coelacanths is almost E13 29. Sep. 09 9:22 9:26 4 158.6 156.5 2.1 17.1 16.7 16.9 13 over hang Talise Isl. Talise-4 INTRODUCTION The first individual of Indonesian coelacanth (Table 2, ID cave (Buol-1). After 29 minutes of recording, this individual E14 29. Sep. 09 14:26 14:33 7 212.6 211.5 1.1 13.0 12.5 12.7 14 over hang Banggka Isl. Bang-1 (Table 2: E5, ID 6) was recognized at 14:34 on the 12th facing the ceiling of the overhang. Such posture is also known (Table 2: Manado-6). It swam up and down 5.4 m between months later, on the 9th June 2013 at the same locations of recorded the presence of this species at 104 m depth at the species L. chalumnae is nocturnal and hides in cave in REFERENCES restricted to an ovoviviparity strategy since some caught large 1) was recorded during the third field survey, in the morning of hid inside the cave. Although the water temperature increased E15 6. Oct. 09 11:34 11:51 17 170.9 164.6 6.3 15.0 14.5 14.8 15 over hang Manado Manado-2 December alongside a large rock. This individual stayed in the in coelacanths from the Comorian Archipelago (FRICKE et al., 168.1 and 173.5 m deep for 17 minutes. Thirteen months later Manado area within several kilometers. Sodwana Bay, South Africa (VENTER et al., 2000). Latimeria daytime but comes out at night, supposedly in search of food ACKNOWLEDGMENTS The first living coelacanth, Latimeria chalumnae, was females carried unfertilized eggs or embryos in their oviduct the 30th May 2006 (Table 1, R3). The coelacanth was of more than five degrees, from 14.8 to 20.4 ºC, it stayed at the E16 9. Oct. 09 10:19 11:02 43 122.2 115.6 6.6 21.5 18.9 20.0 16 over hang Manado Manado-3 beginning of the observation and then moved ahead to deeper 1987). ID 3 was found totally four times in this area in two (the 9th June 2003), this individual (ID 24) was observed again All individuals observed more than twice were chalumnae seem to prefer a temperature range of 15–20 °C, (FRICKE et al., 1991). The living geomorphologic environment BENNO, B., VERHEIJI, E., STAPLEY, J., RUMISHA, C., NGATUNGA, E17 11. Nov. 10 14:22 14:46 24 218.9 212.5 6.4 17.6 17.2 17.4 17,18 over hang Biak Biak-1 discovered in South Africa in the Indian Ocean in 1938 (SMITH et al., 1975; CUPELLO et al., 2015). observed in a cave of 165 m depth (Table 2, Boul-1) for 10 same place, not moving away. Despite it disappeared from the E18 13. Nov. 10 16:48 17:05 17 195.9 193.2 2.7 17.1 16.8 16.9 19,20,21 over hang Biak Biak-2 slowly. Eventually it reached 12 m deeper during one hour and surveys during the seven months and seems to live in this area. near a cliff at 152.3 m depth (Table 2: E24, Manado-8), where encountered in the same area, at locations close from each where they tend to choose their daytime habitats (FRICKE et al., of L. menadoensis is similar to that of L. chalumnae, with We are grateful to Dr. Teruya UYENO for his valuable B., ABDALLAH, A. and KALOMBO, H. 2006. Coelacanth (SMITH, 1939). The first observation of the living coelacanth The Indonesian coelacanth, L. menadoensis, was first minutes (Table 2, E1). The adult individual was in stationary camera frame we kept setting the ROV there in front of the E19 15. Nov. 10 14:54 15:24 30 199.5 198.2 1.3 18.1 17.8 17.9 20 over hang Biak Biak-2 42 minutes. During the 2007 survey (Table 1: R5) a coelacanth (Table located on the same slope of Manado-6. other within hundreds of meters. However, in the Comorian 1991). The suitable temperature for L. menadoensis seems to be rocky steep slopes with undermarine caves or overhangs. advice, guidance and encouragement throughout the present (Latimeria chalumnae SMITH, 1939) discoveries and E20 5. Dec. 10 Manado Manado-4 habitats using submersibles was reported in Comoros (FRICKE discovered in Manado, Sulawesi Island, Indonesia, in 1997. position in a cave (Buol-1) slowly moving its pectoral, pelvic, cave until the next morning, however it did not appear again. 14:39 14:51 12 179.5 171.9 7.6 20.3 18.8 19.6 22 wall On the 14th December, the individual ID 3 previously 2: ID 7) was recorded off Manado by 195 m depth, with a Another new coelacanth (Table 2: E23, ID 25) was Archipelago, same individuals of L. chalumnae were observed, almost the same as L. chalumnae. However, Latimeria menadoensis was also observed alongside study. Many thanks go to colleagues of the Faculty of Fisheries conservation in Tanzania. South African Journal of E21 16. Dec. 10 14:38 14:59 21 199.6 187.4 12.2 17.8 17.6 17.7 23 crack Manado Manado-5 et al., 1987). Latimeria chalumnae hide in submarine caves The first specimen was captured in 1998 (ERDMANN et al., second dorsal and anal fins. The day after, another individual On the 4th June in 2006, three individuals, including ID 3 E22 4. May. 12 10:45 11:02 17 173.5 168.1 5.4 18.9 16.6 18.0 24 steep slope Manado Manado-6 recorded at Buol-2 in a cave and at Buol-3 in a crack, was temperature of 12.4 °C, the coolest one registered during all recorded alone on a Manado site (Manado-7). within a two week period, in several caves distributed in a 8 Water temperature can increase or drop for more than 5 large rocks or along vertical wall in daytime. Some of these and Marine Sciences of the Sam Ratulangi University at Science, 102: 486–490. along steep slopes between 100 m and 200 m depth during 1998). According to a genetic study, the specimen differed (ID 2) and an unidentified one (UN1) were found in the same which was observed on the 31th May at Buol-2 and two new E23 13. May, 12 14:32 14:39 7 149.1 148.7 0.4 18.7 18.7 18.7 25 slope Manado Manado-7 encountered again (Table 2: Buol-5) near a vertical wall at 145 surveys. Field surveys were conducted from the 19th to 30th May kilometers wide area (FRICKE et al., 1991). °C in a very short period of time (about 30 min according to our individuals were encountered above 200 m deep, in an Manado, Indonesia, for their generous supports to conduct BRUTON, M. N., CABRAL, A. J. P. and FRICKE, H. 1992. First E24 9. Jun. 13 13:31 13:48 19 154.0 147.1 6.9 20.8 16.5 18.6 24 wall Manado Manado-8 daytime and come out and drift near the ocean floor for from L. chalumnae and a new species was erected (POUYAUD cave (Buol-1). The unidentified one immediately hid deeper in individuals, were observed in a vertical crack which was E25 27. May. 15 14:48 14:55 7 121.2 116.6 4.6 19.8 19.2 19.4 26 beside big rock Lolak Lolak-1 m depth (Table 2: E6). After a while, this individual (ID 3) The field survey in 2009 was carried out off Manado and 2015 off Lolak Island and from the 30th October to the 16th When individuals of L. menadoensis were encountered, recordings). The coelacanths however often stayed there environment that remains slightly bright. Latimeria expeditions. Also we express our thanks to Dr. SUHARSONO, Dr. capture of a coelacanth, Latimeria chalumnae (Pisces, hunting prey at night (FRICKE et al., 1991). However, since the et al., 1999). In 1999 two individuals were observed by the cave, where the ROV could not access, and only its lateral several hundred meters far from Buol-2 (Buol-3) for 2 hours E26 28. May. 15 16:18 16:35 17 150.3 142.2 8.1 19.1 16.4 17.5 27 beside big rock Lolak Lolak-2 moved upward about 18 m and stayed behind a large rock (Fig. around Talise and Bangka islands off the northern coast of November 2015 off Lolak and Bitung Islands (Table 1; R13 some of them stayed stationary at same place but some without moving away or significantly modifying their menadoensis seems to be less sensitive to the daylight than L. Zainal ARIFIN, Dr. DIRHAMSYAH and colleagues of the Indonesian Latimeriidae), off Mozambique. South African Journal of record of juvenile or small (below 80 cm long) individuals is submersible in a location 360 km away from Manado (FRICKE side of the body was recorded. The individual (ID 2) was and 14 minutes (Fig. 2C; Table 2: E4, ID 3, 4, 5). During this E27 30. Oct. 15 15:32 15:40 8 131.0 122.7 8.3 21.2 20.7 20.9 26 steep slope Lolak Lolak-3 3). The observation time was the longest of all surveys with Sulawesi Island from the 12th September to the 9th October and R14). Three new individuals (Table 2: ID 26–28) were individuals such as ID 6 on the 12th December 2006 (E5) and behavior. These water currents of different temperatures are chalumnae. Institute of Science for their advice and administrative work to Science, : 225–227. E28 1. Nov. 15 14:53 14:58 5 125.6 122.5 3.1 17.4 15.8 16.2 28 beside big rock Lolak Lolak-4 88 virtually absent, the growth, development and breeding et al., 2000). Today 8 individuals were officially caught in the observed for one minute before it hid inside the cave (Table 2: long period of time all individuals stayed near the rock wall E29 16. Nov. 15 13:39 13:41 2 149.1 147.3 1.8 19.1 19.1 19.1 29 steep slope Bitung Bitung-1 four hours and 16 minutes of recording. The temperature (Table 1: R8). On the 24th September 2009, six individuals recognized at different places off Lolak Island (Lolak 1-4). All ID 24 on the 4th May 2012 (E22) swam away. It seemed to caused by the movement of the marine thermocline. During These first observations are potentially crucial since the obtain our research permit. Other thanks go to Mr. OPO, Mr. CUPELLO, C., BRITO, P. M., HERBIN, M., MEUNIER, F. J., biology of L. chalumnae remain poorly known. Our knowledge Indonesian seas. However, the details of their habitat have not E2). When a new individual could be identified by the white always keeping their head against the water flow. The water E30 16. Nov. 15 14:36 14:39 3 156.1 151.8 4.3 20.3 20.2 20.2 30 steep slope Bitung Bitung-2 changed up and down several times (Fig. 4). Its difference (Table 2: E11, ID 9, 10, 11, 12, UN 2, 3) were found under an individuals were alone. ID 26 was observed in May and once avoid and escape from the brightness of the light or the sounds our surveys we observed in some area that as the thermocline habitats of the Indonesian coelacanths could be possibly not as REFRY and colleagues at Murex Dive Resort, and DAUD, AL, J ANVIER, P., DUTEL, H. and CLÉMENT, G. 2015. 50 IWATA et al. Field surveys on Latimeria menadoensis using ROV from 2005 to 2015 51 been described. The aim of this study is to clear their habitat Table 1. Periods and areas of the Latimeria menadoensis survey using ROV. Total of 14 Allometric growth in the extant coelacanth lung during capture of a coelacanth off Madagascar. South African and distribution. expeditions were conducted in the north of Sulawesi Island and Biak Island located in north west ontogenetic development. Nature Communications, 6: Journal of Science 92: 150–151. of New Guinea Island. The survey areas are shown in Fig. 1. 8222. DOI: 10.1038/ncomms9222 POUYAUD, L., WIRJOATMODJO, S., RACHMATIKA, I., TJAKRAWIDJAJA, DE VOS, L. and OYUGI, D. 2002. First capture of a coelacanth, A., HADIATY, R. and HADIE, W. 1999. A new species of No. Date Survey Area METHODS Latimeria chalumnae SMITH, 1939 (Pisces, Latimeriidae), coelacanth. Genetic and morphologic proof. C. R. R1 17 Apr. – 30 Apr. 2005 Manado-tua Isl. and other islands off Kenya. South African Journal of Science, 98: 345–347. Academy of Science, 322: 261–267. Two remotely operated vehicles (ROVs) (Kowa; R2 6 May – 19. May 2006 Northern coast of Sulawesi ERDMANN, M. V., CALDWELL, R. L. and MOOSA, M. K. 1998. SMITH, J. L. B. 1939. The living coelacanth fish from South VEGA300) were used for the surveys. The first ROV was R3 27 May – 5 Jun. 2006 Buol Indonesian ‘king of the sea’ discovered. Nature, 395: 335. Africa. Nature, 143: 748–750. replaced with the second one in 2007. Our ROVs are able to R4 11 Dec. – 20 Dec. 2006 Buol overhang off Talise Island between 144 and 150 m deep. There again in October 2015 on a steep slope (Table 2: E25, E27). FRICKE, H., REINICKE, O., HOFER, H. and NACHTIGALL, W. SMITH, J. L. B. 1953. The second coelacanth. Nature, 171: dive up to 300 m depth. These had two vertical, two horizontal was almost no water current in this area. All individuals were The coelacanth ID 28 was found on the 1st November along 1987. Locomotion of the coelacanth Latimeria chalumnae 99–101. R5 27 Jun. – 5 Jul. 2007 Manado and two right-left propellers and these were controlled from a close to each other under the same overhang and most of them one of the large rocks scattered on a gentle slope in a bay of in its natural environment. Nature, 329: 331–333. SMITH, C. L., RAND, C. S., SCHAEFFER, B. and ATZ, J. W. 1975. R6 9 Jul. – 12 Jul. 2007 Buol boat on surface through a 400 m long tether cable. The stayed still with their head facing downward. Four days later, the Lolak Island by 125 m depth. FRICKE, H., HISSMANN, K., SCHAUER, J., REINICKE, O., KASANG, Latimeria, the living coelacanth, is ovoviviparous. Science, underwater operations were visualized on a screen with R7 2 Dec. – 8 Dec. 2008 Talise and Bangka Isl. ID 9 was observed again, alone on a steep slope at 172.9 m The encountered Indonesian coelacanths during these L. and PLANTE, R. 1991. Habitat and population size of 190: 1105–1106. information including directions of ROVs, depth, water R8 12 Sep. – 9 Oct. 2009 Talise and Bangka Isl. Manado depth (Table 2: E12), which was located within several dozen surveys were in caves, alongside large rocks, under overhangs coelacanth Latimeria chalumnae at Grand Comoro. VENTER, P., TIMM, P., GUNN, G., LE ROUX, E., SERFONTEIN, E., temperature, date and time. All data were directly recorded on R9 6 Nov – 16 Nov. 2010 Biak Island meters far from Talise-2 and 30 m deeper. or on steep slopes (Table 2; Fig. 2). Environmental Biology of Fishes, 32: 287–300. SMITH, P., SMITH, E., BENSCH, M., HARDING, D. and video-tapes on the boat. Since the 2007 survey, water R10 5 Dec. – 18 Dec. 2010 Manado Four new coelacanths were recorded from the 29th Most individuals were found under overhangs or FRICKE, H., HISSMANN, K., SCHAUER, J., ERDMANN, M., HEEMSTRA, P. 2000. Discovery of a viable population of temperature and depth were independently recorded with more September to the 6th October 2009 in Talise and Banggka alongside large rocks. Some individuals were encountered not MOOSA, M. K. and PLANTE, R. 2000. Biogeography of the coelacanths (Latimeria chalumnae SMITH, 1939) at R11 2 May – 13 May 2012 Manado accuracy by external measuring memories (Alec Electronics, Islands and off Manado (Table 2: E13 and 14, ID 13, 14, 15). to hide in any shade and stayed just on rocky slopes. In Indonesian coelacanth. Nature, 403: 38. Sodwana Bay, South Africa. South African Journal of R12 3 Jun. – 10 Jun. 2013 Manado later named JFE Alec; MDS-MKV/T, MDS-MKV/D). In ID 15 is the only juvenile coelacanth filmed so far. It was in a overhangs and cracks, all individuals had their ventral side HEEMSTRA, P. C., FREEMAN, A. L., WONG, H. Y., HENSLEY, D. Science, 96: 567–568. addition two laser beam irradiators, that provide line lasers of R13 19 May – 30 May 2015 Bitung, Lolak small crack between 164.6 and 170.9 m deep in a temperature alongside and close to a rock wall, but without touching it. A. and RABESANDRATANA, H. D. 1996. First authentic 20-cm distance, were attached to the second ROV and have R14 30 Oct. – 16 Nov. 2015 Bitung, Lolak between 14.5 and 15 °C. Estimated total lengths are shown in Table 3. Most been used to register the size of encountered animals. The The shallowest record of a coelacanth was at 115.6 m individuals are more than 1 m in total length. ID 15 is a surveys were conducted off northern parts of Sulawesi Island depth off the southern part of Manado (Table 2: E16, juvenile individual of 31.5 cm and ID 23 is 90 cm in total and along the Biak Island located in northwestern New Guinea Table 2. Registers of Latimeria menadoensis. Time shows start, end and duration of each Manado-3) on the 9th October in 2009. ID 16 (Table 2) was length. No individual beyond 140 cm has been recorded in Island, Indonesia. The areas and time periods of each survey observation. When an individual could be identified by patterns of white spot on the body, an ID found at the end of the morning, alone alongside a large rock Indonesia so far. restricted as those of the African coelacanths, and so a broader and other staffs for our work on the research boat. Dr. BATUNA are shown in Table 1 and Fig. 1. number was allocated. Individuals that could not been distinguished were allocated UN (unknown) on the edge of a shelf. The water temperature was about 20.0 distribution of the Indonesian coelacanth could be expected. and the Murex staffs supported our daily life during the + number. Individuals observed several times were identified as numbers with underbar. Duration ºC. Occurrences of coelacanths along the eastern coast of the expeditions. We would like to express our sincere thanks to shows long of each observation time. Range indicates vertical movement of the individual during From the 6th to the 16th November in 2010, the field of the propellers of the ROV. level changed, the water flow was reversed. Coelacanths African continent were recorded in South Africa (SMITH, colleagues at Aquamarine Fukushima, Japan, for their survey was conducted around Biak Island (Table 1: R9). Adult MITH EEMSTRA RESULTS observation and not the distance traveled horizontally. Some individuals swam along the slope. DISCUSSION Temperature and depth data for each Indonesian always kept their head against the water flow, like other 1939), Comoros (S , 1953), Madagascar (H et al., understanding and supports. We would like to thank Dr. Gaël coelacanths were found at two different sites (Table 2: coelacanth record are shown in Fig. 3. Though an average of observed actinopterygian fishes did. It means that as the 1996), Kenya (DE VOS and OYUGI, 2002), Tanzania (BENNO et CLÉMENT of Muséum national d’Histoire naturelle and Dr. Time Depth (m) Temperature (ºC) The field survey of the Indonesian coelacanth, Latimeria Fig. 1. Localities of the Indonesian coelacanth survey using No. Date ID Configuration Place Site E17–19). Two individuals (ID 17 and 18) were found under an During the field surveys by underwater ROV recording water temperature on the ocean surface was about 30 °C, the direction of the water flow changed, the swimming direction of al., 2006), and Mozambique (BRUTON et al., 1992). In Camila CUPELLO of Universidade do Estado do Rio de Janeiro Start End Duration Max. Min. Range Max. Min. Ave. menadoensis, was conducted 14 times from 2005 to 2015 with ROV from 2005 to 2015. E1 30. May 06 8:30 8:40 10 165.0 165.0 0 17.1 16.4 16.8 1 cave Buol Buol-1 overhang between 212.5 and 218.9 m deep (Table 2: Biak-1) for Indonesian coelacanths, Latimeria menadoensis, from 2005 temperature often dropped into about 11 °C at 300 m water depth. each coelacanth individual changed accordingly. Indonesia, coelacanths have only been found off the northern as a referee for their critical reading of the manuscript and their 1173 underwater operations around Sulawesi and Biak islands E2 31. May 06 7:58 7:59 1 165.0 165.0 0 15.5 15.3 15.4 2,UN1 cave Buol Buol-1 on the 11th November. Three other ones (ID 19, 20, 21) were to 2015, 30 different individuals were identified and three The temperatures changed widely even though the changes of On the 24th September 2009 (Table 2: E11) six coast of the Sulawesi Island and the southern coast of the Biak comments. E3 31. May 06 11:15 11:44 29 183.0 183.0 0 20.4 14.8 17.5 3 cave Buol Buol-2 in Indonesia (Fig. 1, Table 1). No coelacanth was encountered registered under an overhang between 193.2 and 195.9 m deep others were unidentified. Among those, six individuals were depth were small at several places. All Indonesian coelacanths, individuals were recognized under an overhang (Talise-2). The Island in northern New Guinea. Unfortunately one of our authors, Dr. Djoko Hadi KUNARSO E4 4. Jun. 06 18:42 20:56 134 156.2 153.9 2.3 17.8 12.8 13.7 3,4,5 crack Buol Buol-3 during the first surveys in 2005 although 452 underwater E5 12. Dec. 06 14:34 16:16 102 163.8 151.0 12.8 16.6 14.2 15.7 6 wall Buol Buol-4 (Table 2: Biak-2) on the 13th November. Two days later, observed more than twice: ID 3 at Buol-2, 3, 5, 6 (Table 2: E3, L. menadoensis, were encountered during day time at a depth all had about the same size. There was almost no water flow, Field surveys of the Indonesian coelacanths should be has passed away during the process of this contribution. We operations were conducted around Manado-tua Island and spots pattern on its body, an ID number was allocated. The E6 1. Dec. 06 14:21 18:34 253 145.0 126.5 18.5 21.2 14.5 17.9 3 wall Buol Buol-5 temperature changed up and down by five degrees during the between the maximum and the minimum was 6.7 °C, which Biak-2 was observed again and only ID 20 stayed there. E4, E6 and E7); ID 8 at Talise-1 (E9 and E10), ID 9 at Talise 2 between 115.6 and 218.9 m and a temperature between 12.4 and each individual was head downward. They were close to continued in all area of Indonesia and expanded to all pray for the response of his soul. E7 15. Dec 06 14:02 14:24 22 154.2 152.3 1.9 15.6 15.2 15.4 3 over hang Buol Buol-6 other close Indonesian islands from the 17th to 30th of April individual which could not be distinguished was allocated an E8 27. Jun. 07 12:15 12:46 31 195.0 190.8 4.2 16.2 12.4 15.0 7 beside big rock Mando Mando-1 observation due to water flow reversals. When the water flow was the largest range of all surveys. Two new coelacanths were recorded off Manado in and 3 (E11 and E12), ID 20 at Biak-2 (E18 and E19), ID 24 at and 21.5 ºC. Indonesian coelacanths have exceptionally been each other but no direct contacts or interactions between Southeast Asia in order to increase our knowledge of the This study was supported in part by funding from (Table 1, R1). The second field survey was held along the UN (unknown) + number. E9 14. Sep. 09 10:33 11:00 27 172.2 159.8 12.4 15.8 14.3 15.5 8 over hang Talise Isl. Talise-1 was reversed some times, all individuals always changed their On the next day, on the 15th December, the ID 3 was December 2010 (Table 1: R10; Table 2: E 20 and 21; ID 22 Manado-6 and 8 (E22 and E24), ID 26 at Lolak-1 and 3 (E25 recorded in an environment with a temperature exceeding 20 coelacanths were observed. No evidence of a social behavior distribution of this rare species and to define, in a close future, Interdisciplinary Collaborative Research program of Atmosphere northern coast of Sulawesi from the 6th to 19th May in 2006, In the end of the morning of the same day, a third E10 15. Sep. 09 8:40 8:42 2 166.8 160.6 6.2 20.2 19.7 20.1 8 over hang Talise Isl. Talise-1 directions against the water flow. observed again under an overhang (Table 2: E7, Buol-6) near and 23). and E27). ID 8 was observed on the 14th September 2009 and ºC, regardless of the depth. The African coelacanth, L. has been recognized in the Comorian populations of the most relevant conservation policies of the different known and Ocean Research Institute, the University of Tokyo. 9,10,11,12 E11 24. Sep. 09 14:51 14:55 4 149.9 143.7 6.2 18.5 18.2 18.4 over hang Talise Isl. Talise-2 but once again despite of 107 underwater operations no individual (Table 2: E3, ID 3) was found in another cave UN2,UN3 During the field survey carried on December of the same other sites where it was previously found at Buol-2 , Buol-3 On the 4th May 2012, an individual (Table 2: E22, ID 24) was encountered again at the same place (Talise-1) the day chalumnae, was observed by submarine vehicles in steep coelacanths (FRICKE et al., 1991). populations. coelacanth was recorded. (Buol-2), larger and located 20 m deeper (Buol-2) than the first E12 28. Sep. 09 8:22 8:30 8 172.9 158.9 14 18.5 17.6 18.0 9 steep slope, beside big rock Talise Isl. Talise-3 year, in the same Buol area (Table 1: R4) the sixth individual and Buol-5. Its posture was upside down with its ventral side was found on a steep slope at 169.4 m depth off Manado after. ID 24 was observed on the 4th May 2012 and then 13 slopes between 150 m and 253 m (FRICKE et al., 1991). Divers According to the observations of the African coelacanths, of the reproduction biology of the coelacanths is almost E13 29. Sep. 09 9:22 9:26 4 158.6 156.5 2.1 17.1 16.7 16.9 13 over hang Talise Isl. Talise-4 INTRODUCTION The first individual of Indonesian coelacanth (Table 2, ID cave (Buol-1). After 29 minutes of recording, this individual E14 29. Sep. 09 14:26 14:33 7 212.6 211.5 1.1 13.0 12.5 12.7 14 over hang Banggka Isl. Bang-1 (Table 2: E5, ID 6) was recognized at 14:34 on the 12th facing the ceiling of the overhang. Such posture is also known (Table 2: Manado-6). It swam up and down 5.4 m between months later, on the 9th June 2013 at the same locations of recorded the presence of this species at 104 m depth at the species L. chalumnae is nocturnal and hides in cave in REFERENCES restricted to an ovoviviparity strategy since some caught large 1) was recorded during the third field survey, in the morning of hid inside the cave. Although the water temperature increased E15 6. Oct. 09 11:34 11:51 17 170.9 164.6 6.3 15.0 14.5 14.8 15 over hang Manado Manado-2 December alongside a large rock. This individual stayed in the in coelacanths from the Comorian Archipelago (FRICKE et al., 168.1 and 173.5 m deep for 17 minutes. Thirteen months later Manado area within several kilometers. Sodwana Bay, South Africa (VENTER et al., 2000). Latimeria daytime but comes out at night, supposedly in search of food ACKNOWLEDGMENTS The first living coelacanth, Latimeria chalumnae, was females carried unfertilized eggs or embryos in their oviduct the 30th May 2006 (Table 1, R3). The coelacanth was of more than five degrees, from 14.8 to 20.4 ºC, it stayed at the E16 9. Oct. 09 10:19 11:02 43 122.2 115.6 6.6 21.5 18.9 20.0 16 over hang Manado Manado-3 beginning of the observation and then moved ahead to deeper 1987). ID 3 was found totally four times in this area in two (the 9th June 2003), this individual (ID 24) was observed again All individuals observed more than twice were chalumnae seem to prefer a temperature range of 15–20 °C, (FRICKE et al., 1991). The living geomorphologic environment BENNO, B., VERHEIJI, E., STAPLEY, J., RUMISHA, C., NGATUNGA, E17 11. Nov. 10 14:22 14:46 24 218.9 212.5 6.4 17.6 17.2 17.4 17,18 over hang Biak Biak-1 discovered in South Africa in the Indian Ocean in 1938 (SMITH et al., 1975; CUPELLO et al., 2015). observed in a cave of 165 m depth (Table 2, Boul-1) for 10 same place, not moving away. Despite it disappeared from the E18 13. Nov. 10 16:48 17:05 17 195.9 193.2 2.7 17.1 16.8 16.9 19,20,21 over hang Biak Biak-2 slowly. Eventually it reached 12 m deeper during one hour and surveys during the seven months and seems to live in this area. near a cliff at 152.3 m depth (Table 2: E24, Manado-8), where encountered in the same area, at locations close from each where they tend to choose their daytime habitats (FRICKE et al., of L. menadoensis is similar to that of L. chalumnae, with We are grateful to Dr. Teruya UYENO for his valuable B., ABDALLAH, A. and KALOMBO, H. 2006. Coelacanth (SMITH, 1939). The first observation of the living coelacanth The Indonesian coelacanth, L. menadoensis, was first minutes (Table 2, E1). The adult individual was in stationary camera frame we kept setting the ROV there in front of the E19 15. Nov. 10 14:54 15:24 30 199.5 198.2 1.3 18.1 17.8 17.9 20 over hang Biak Biak-2 42 minutes. During the 2007 survey (Table 1: R5) a coelacanth (Table located on the same slope of Manado-6. other within hundreds of meters. However, in the Comorian 1991). The suitable temperature for L. menadoensis seems to be rocky steep slopes with undermarine caves or overhangs. advice, guidance and encouragement throughout the present (Latimeria chalumnae SMITH, 1939) discoveries and E20 5. Dec. 10 Manado Manado-4 habitats using submersibles was reported in Comoros (FRICKE discovered in Manado, Sulawesi Island, Indonesia, in 1997. position in a cave (Buol-1) slowly moving its pectoral, pelvic, cave until the next morning, however it did not appear again. 14:39 14:51 12 179.5 171.9 7.6 20.3 18.8 19.6 22 wall On the 14th December, the individual ID 3 previously 2: ID 7) was recorded off Manado by 195 m depth, with a Another new coelacanth (Table 2: E23, ID 25) was Archipelago, same individuals of L. chalumnae were observed, almost the same as L. chalumnae. However, Latimeria menadoensis was also observed alongside study. Many thanks go to colleagues of the Faculty of Fisheries conservation in Tanzania. South African Journal of E21 16. Dec. 10 14:38 14:59 21 199.6 187.4 12.2 17.8 17.6 17.7 23 crack Manado Manado-5 et al., 1987). Latimeria chalumnae hide in submarine caves The first specimen was captured in 1998 (ERDMANN et al., second dorsal and anal fins. The day after, another individual On the 4th June in 2006, three individuals, including ID 3 E22 4. May. 12 10:45 11:02 17 173.5 168.1 5.4 18.9 16.6 18.0 24 steep slope Manado Manado-6 recorded at Buol-2 in a cave and at Buol-3 in a crack, was temperature of 12.4 °C, the coolest one registered during all recorded alone on a Manado site (Manado-7). within a two week period, in several caves distributed in a 8 Water temperature can increase or drop for more than 5 large rocks or along vertical wall in daytime. Some of these and Marine Sciences of the Sam Ratulangi University at Science, 102: 486–490. along steep slopes between 100 m and 200 m depth during 1998). According to a genetic study, the specimen differed (ID 2) and an unidentified one (UN1) were found in the same which was observed on the 31th May at Buol-2 and two new E23 13. May, 12 14:32 14:39 7 149.1 148.7 0.4 18.7 18.7 18.7 25 slope Manado Manado-7 encountered again (Table 2: Buol-5) near a vertical wall at 145 surveys. Field surveys were conducted from the 19th to 30th May kilometers wide area (FRICKE et al., 1991). °C in a very short period of time (about 30 min according to our individuals were encountered above 200 m deep, in an Manado, Indonesia, for their generous supports to conduct BRUTON, M. N., CABRAL, A. J. P. and FRICKE, H. 1992. First E24 9. Jun. 13 13:31 13:48 19 154.0 147.1 6.9 20.8 16.5 18.6 24 wall Manado Manado-8 daytime and come out and drift near the ocean floor for from L. chalumnae and a new species was erected (POUYAUD cave (Buol-1). The unidentified one immediately hid deeper in individuals, were observed in a vertical crack which was E25 27. May. 15 14:48 14:55 7 121.2 116.6 4.6 19.8 19.2 19.4 26 beside big rock Lolak Lolak-1 m depth (Table 2: E6). After a while, this individual (ID 3) The field survey in 2009 was carried out off Manado and 2015 off Lolak Island and from the 30th October to the 16th When individuals of L. menadoensis were encountered, recordings). The coelacanths however often stayed there environment that remains slightly bright. Latimeria expeditions. Also we express our thanks to Dr. SUHARSONO, Dr. capture of a coelacanth, Latimeria chalumnae (Pisces, hunting prey at night (FRICKE et al., 1991). However, since the et al., 1999). In 1999 two individuals were observed by the cave, where the ROV could not access, and only its lateral several hundred meters far from Buol-2 (Buol-3) for 2 hours E26 28. May. 15 16:18 16:35 17 150.3 142.2 8.1 19.1 16.4 17.5 27 beside big rock Lolak Lolak-2 moved upward about 18 m and stayed behind a large rock (Fig. around Talise and Bangka islands off the northern coast of November 2015 off Lolak and Bitung Islands (Table 1; R13 some of them stayed stationary at same place but some without moving away or significantly modifying their menadoensis seems to be less sensitive to the daylight than L. Zainal ARIFIN, Dr. DIRHAMSYAH and colleagues of the Indonesian Latimeriidae), off Mozambique. South African Journal of record of juvenile or small (below 80 cm long) individuals is submersible in a location 360 km away from Manado (FRICKE side of the body was recorded. The individual (ID 2) was and 14 minutes (Fig. 2C; Table 2: E4, ID 3, 4, 5). During this E27 30. Oct. 15 15:32 15:40 8 131.0 122.7 8.3 21.2 20.7 20.9 26 steep slope Lolak Lolak-3 3). The observation time was the longest of all surveys with Sulawesi Island from the 12th September to the 9th October and R14). Three new individuals (Table 2: ID 26–28) were individuals such as ID 6 on the 12th December 2006 (E5) and behavior. These water currents of different temperatures are chalumnae. Institute of Science for their advice and administrative work to Science, : 225–227. E28 1. Nov. 15 14:53 14:58 5 125.6 122.5 3.1 17.4 15.8 16.2 28 beside big rock Lolak Lolak-4 88 virtually absent, the growth, development and breeding et al., 2000). Today 8 individuals were officially caught in the observed for one minute before it hid inside the cave (Table 2: long period of time all individuals stayed near the rock wall E29 16. Nov. 15 13:39 13:41 2 149.1 147.3 1.8 19.1 19.1 19.1 29 steep slope Bitung Bitung-1 four hours and 16 minutes of recording. The temperature (Table 1: R8). On the 24th September 2009, six individuals recognized at different places off Lolak Island (Lolak 1-4). All ID 24 on the 4th May 2012 (E22) swam away. It seemed to caused by the movement of the marine thermocline. During These first observations are potentially crucial since the obtain our research permit. Other thanks go to Mr. OPO, Mr. CUPELLO, C., BRITO, P. M., HERBIN, M., MEUNIER, F. J., biology of L. chalumnae remain poorly known. Our knowledge Indonesian seas. However, the details of their habitat have not E2). When a new individual could be identified by the white always keeping their head against the water flow. The water E30 16. Nov. 15 14:36 14:39 3 156.1 151.8 4.3 20.3 20.2 20.2 30 steep slope Bitung Bitung-2 changed up and down several times (Fig. 4). Its difference (Table 2: E11, ID 9, 10, 11, 12, UN 2, 3) were found under an individuals were alone. ID 26 was observed in May and once avoid and escape from the brightness of the light or the sounds our surveys we observed in some area that as the thermocline habitats of the Indonesian coelacanths could be possibly not as REFRY and colleagues at Murex Dive Resort, and DAUD, AL, J ANVIER, P., DUTEL, H. and CLÉMENT, G. 2015. 52 IWATA et al. Field surveys on Latimeria menadoensis using ROV from 2005 to 2015 53 been described. The aim of this study is to clear their habitat Table 3. Total length of each coelacanth estimated with 20 cm distance parallel laser lines applied Allometric growth in the extant coelacanth lung during capture of a coelacanth off Madagascar. South African and distribution. from the ROV. ontogenetic development. Nature Communications, 6: Journal of Science 92: 150–151. 8222. DOI: 10.1038/ncomms9222 POUYAUD, L., WIRJOATMODJO, S., RACHMATIKA, I., TJAKRAWIDJAJA, ID number 7 8 14 15 16 17 19 20 22 23 24 DE VOS, L. and OYUGI, D. 2002. First capture of a coelacanth, A., HADIATY, R. and HADIE, W. 1999. A new species of METHODS Estimated TL (cm) 123 115 110 31.5 113 125 114 113 106 90 137 Latimeria chalumnae SMITH, 1939 (Pisces, Latimeriidae), coelacanth. Genetic and morphologic proof. C. R. off Kenya. South African Journal of Science, 98: 345–347. Academy of Science, 322: 261–267. Two remotely operated vehicles (ROVs) (Kowa; ERDMANN, M. V., CALDWELL, R. L. and MOOSA, M. K. 1998. SMITH, J. L. B. 1939. The living coelacanth fish from South VEGA300) were used for the surveys. The first ROV was Indonesian ‘king of the sea’ discovered. Nature, 395: 335. Africa. Nature, 143: 748–750. replaced with the second one in 2007. Our ROVs are able to overhang off Talise Island between 144 and 150 m deep. There again in October 2015 on a steep slope (Table 2: E25, E27). FRICKE, H., REINICKE, O., HOFER, H. and NACHTIGALL, W. SMITH, J. L. B. 1953. The second coelacanth. Nature, 171: dive up to 300 m depth. These had two vertical, two horizontal was almost no water current in this area. All individuals were The coelacanth ID 28 was found on the 1st November along 1987. Locomotion of the coelacanth Latimeria chalumnae 99–101. and two right-left propellers and these were controlled from a close to each other under the same overhang and most of them one of the large rocks scattered on a gentle slope in a bay of in its natural environment. Nature, 329: 331–333. SMITH, C. L., RAND, C. S., SCHAEFFER, B. and ATZ, J. W. 1975. boat on surface through a 400 m long tether cable. The stayed still with their head facing downward. Four days later, the Lolak Island by 125 m depth. FRICKE, H., HISSMANN, K., SCHAUER, J., REINICKE, O., KASANG, Latimeria, the living coelacanth, is ovoviviparous. Science, underwater operations were visualized on a screen with ID 9 was observed again, alone on a steep slope at 172.9 m The encountered Indonesian coelacanths during these L. and PLANTE, R. 1991. Habitat and population size of 190: 1105–1106. information including directions of ROVs, depth, water depth (Table 2: E12), which was located within several dozen surveys were in caves, alongside large rocks, under overhangs coelacanth Latimeria chalumnae at Grand Comoro. VENTER, P., TIMM, P., GUNN, G., LE ROUX, E., SERFONTEIN, E., temperature, date and time. All data were directly recorded on meters far from Talise-2 and 30 m deeper. or on steep slopes (Table 2; Fig. 2). Environmental Biology of Fishes, 32: 287–300. SMITH, P., SMITH, E., BENSCH, M., HARDING, D. and video-tapes on the boat. Since the 2007 survey, water Four new coelacanths were recorded from the 29th Most individuals were found under overhangs or FRICKE, H., HISSMANN, K., SCHAUER, J., ERDMANN, M., HEEMSTRA, P. 2000. Discovery of a viable population of temperature and depth were independently recorded with more September to the 6th October 2009 in Talise and Banggka alongside large rocks. Some individuals were encountered not MOOSA, M. K. and PLANTE, R. 2000. Biogeography of the coelacanths (Latimeria chalumnae SMITH, 1939) at accuracy by external measuring memories (Alec Electronics, Islands and off Manado (Table 2: E13 and 14, ID 13, 14, 15). to hide in any shade and stayed just on rocky slopes. In Indonesian coelacanth. Nature, 403: 38. Sodwana Bay, South Africa. South African Journal of later named JFE Alec; MDS-MKV/T, MDS-MKV/D). In ID 15 is the only juvenile coelacanth filmed so far. It was in a overhangs and cracks, all individuals had their ventral side HEEMSTRA, P. C., FREEMAN, A. L., WONG, H. Y., HENSLEY, D. Science, 96: 567–568. addition two laser beam irradiators, that provide line lasers of small crack between 164.6 and 170.9 m deep in a temperature alongside and close to a rock wall, but without touching it. A. and RABESANDRATANA, H. D. 1996. First authentic 20-cm distance, were attached to the second ROV and have between 14.5 and 15 °C. Estimated total lengths are shown in Table 3. Most been used to register the size of encountered animals. The The shallowest record of a coelacanth was at 115.6 m individuals are more than 1 m in total length. ID 15 is a surveys were conducted off northern parts of Sulawesi Island depth off the southern part of Manado (Table 2: E16, juvenile individual of 31.5 cm and ID 23 is 90 cm in total and along the Biak Island located in northwestern New Guinea Manado-3) on the 9th October in 2009. ID 16 (Table 2) was length. No individual beyond 140 cm has been recorded in Island, Indonesia. The areas and time periods of each survey found at the end of the morning, alone alongside a large rock Indonesia so far. restricted as those of the African coelacanths, and so a broader and other staffs for our work on the research boat. Dr. BATUNA are shown in Table 1 and Fig. 1. on the edge of a shelf. The water temperature was about 20.0 distribution of the Indonesian coelacanth could be expected. and the Murex staffs supported our daily life during the ºC. Occurrences of coelacanths along the eastern coast of the expeditions. We would like to express our sincere thanks to From the 6th to the 16th November in 2010, the field of the propellers of the ROV. level changed, the water flow was reversed. Coelacanths African continent were recorded in South Africa (SMITH, colleagues at Aquamarine Fukushima, Japan, for their RESULTS survey was conducted around Biak Island (Table 1: R9). Adult DISCUSSION Temperature and depth data for each Indonesian always kept their head against the water flow, like other 1939), Comoros (SMITH, 1953), Madagascar (HEEMSTRA et al., understanding and supports. We would like to thank Dr. Gaël coelacanths were found at two different sites (Table 2: coelacanth record are shown in Fig. 3. Though an average of observed actinopterygian fishes did. It means that as the 1996), Kenya (DE VOS and OYUGI, 2002), Tanzania (BENNO et CLÉMENT of Muséum national d’Histoire naturelle and Dr. The field survey of the Indonesian coelacanth, Latimeria Fig. 2. Configuration of encounter point. A, over hang (E9); B, over hang (E17); C, crack (E4); D, crack (E21); E, beside big rock E17–19). Two individuals (ID 17 and 18) were found under an During the field surveys by underwater ROV recording water temperature on the ocean surface was about 30 °C, the direction of the water flow changed, the swimming direction of al., 2006), and Mozambique (BRUTON et al., 1992). In Camila CUPELLO of Universidade do Estado do Rio de Janeiro menadoensis, was conducted 14 times from 2005 to 2015 with (E28); F, beside big rock (E25); G, wall (E20); H, steep slope (E12); I, steep slope, beside big rock (E13). Red triangles overhang between 212.5 and 218.9 m deep (Table 2: Biak-1) for Indonesian coelacanths, Latimeria menadoensis, from 2005 temperature often dropped into about 11 °C at 300 m water depth. each coelacanth individual changed accordingly. Indonesia, coelacanths have only been found off the northern as a referee for their critical reading of the manuscript and their 1173 underwater operations around Sulawesi and Biak islands indicate coelacanths. on the 11th November. Three other ones (ID 19, 20, 21) were to 2015, 30 different individuals were identified and three The temperatures changed widely even though the changes of On the 24th September 2009 (Table 2: E11) six coast of the Sulawesi Island and the southern coast of the Biak comments. in Indonesia (Fig. 1, Table 1). No coelacanth was encountered registered under an overhang between 193.2 and 195.9 m deep others were unidentified. Among those, six individuals were depth were small at several places. All Indonesian coelacanths, individuals were recognized under an overhang (Talise-2). The Island in northern New Guinea. Unfortunately one of our authors, Dr. Djoko Hadi KUNARSO during the first surveys in 2005 although 452 underwater (Table 2: Biak-2) on the 13th November. Two days later, observed more than twice: ID 3 at Buol-2, 3, 5, 6 (Table 2: E3, L. menadoensis, were encountered during day time at a depth all had about the same size. There was almost no water flow, Field surveys of the Indonesian coelacanths should be has passed away during the process of this contribution. We operations were conducted around Manado-tua Island and spots pattern on its body, an ID number was allocated. The temperature changed up and down by five degrees during the between the maximum and the minimum was 6.7 °C, which Biak-2 was observed again and only ID 20 stayed there. E4, E6 and E7); ID 8 at Talise-1 (E9 and E10), ID 9 at Talise 2 between 115.6 and 218.9 m and a temperature between 12.4 and each individual was head downward. They were close to continued in all area of Indonesia and expanded to all pray for the response of his soul. other close Indonesian islands from the 17th to 30th of April individual which could not be distinguished was allocated an observation due to water flow reversals. When the water flow was the largest range of all surveys. Two new coelacanths were recorded off Manado in and 3 (E11 and E12), ID 20 at Biak-2 (E18 and E19), ID 24 at and 21.5 ºC. Indonesian coelacanths have exceptionally been each other but no direct contacts or interactions between Southeast Asia in order to increase our knowledge of the This study was supported in part by funding from (Table 1, R1). The second field survey was held along the UN (unknown) + number. was reversed some times, all individuals always changed their On the next day, on the 15th December, the ID 3 was December 2010 (Table 1: R10; Table 2: E 20 and 21; ID 22 Manado-6 and 8 (E22 and E24), ID 26 at Lolak-1 and 3 (E25 recorded in an environment with a temperature exceeding 20 coelacanths were observed. No evidence of a social behavior distribution of this rare species and to define, in a close future, Interdisciplinary Collaborative Research program of Atmosphere northern coast of Sulawesi from the 6th to 19th May in 2006, In the end of the morning of the same day, a third directions against the water flow. observed again under an overhang (Table 2: E7, Buol-6) near and 23). and E27). ID 8 was observed on the 14th September 2009 and ºC, regardless of the depth. The African coelacanth, L. has been recognized in the Comorian populations of the most relevant conservation policies of the different known and Ocean Research Institute, the University of Tokyo. but once again despite of 107 underwater operations no individual (Table 2: E3, ID 3) was found in another cave During the field survey carried on December of the same other sites where it was previously found at Buol-2 , Buol-3 On the 4th May 2012, an individual (Table 2: E22, ID 24) was encountered again at the same place (Talise-1) the day chalumnae, was observed by submarine vehicles in steep coelacanths (FRICKE et al., 1991). populations. coelacanth was recorded. (Buol-2), larger and located 20 m deeper (Buol-2) than the first year, in the same Buol area (Table 1: R4) the sixth individual and Buol-5. Its posture was upside down with its ventral side was found on a steep slope at 169.4 m depth off Manado after. ID 24 was observed on the 4th May 2012 and then 13 slopes between 150 m and 253 m (FRICKE et al., 1991). Divers According to the observations of the African coelacanths, INTRODUCTION of the reproduction biology of the coelacanths is almost The first individual of Indonesian coelacanth (Table 2, ID cave (Buol-1). After 29 minutes of recording, this individual (Table 2: E5, ID 6) was recognized at 14:34 on the 12th facing the ceiling of the overhang. Such posture is also known (Table 2: Manado-6). It swam up and down 5.4 m between months later, on the 9th June 2013 at the same locations of recorded the presence of this species at 104 m depth at the species L. chalumnae is nocturnal and hides in cave in REFERENCES restricted to an ovoviviparity strategy since some caught large 1) was recorded during the third field survey, in the morning of hid inside the cave. Although the water temperature increased December alongside a large rock. This individual stayed in the in coelacanths from the Comorian Archipelago (FRICKE et al., 168.1 and 173.5 m deep for 17 minutes. Thirteen months later Manado area within several kilometers. Sodwana Bay, South Africa (VENTER et al., 2000). Latimeria daytime but comes out at night, supposedly in search of food ACKNOWLEDGMENTS The first living coelacanth, Latimeria chalumnae, was females carried unfertilized eggs or embryos in their oviduct the 30th May 2006 (Table 1, R3). The coelacanth was of more than five degrees, from 14.8 to 20.4 ºC, it stayed at the beginning of the observation and then moved ahead to deeper 1987). ID 3 was found totally four times in this area in two (the 9th June 2003), this individual (ID 24) was observed again All individuals observed more than twice were chalumnae seem to prefer a temperature range of 15–20 °C, (FRICKE et al., 1991). The living geomorphologic environment BENNO, B., VERHEIJI, E., STAPLEY, J., RUMISHA, C., NGATUNGA, discovered in South Africa in the Indian Ocean in 1938 (SMITH et al., 1975; CUPELLO et al., 2015). observed in a cave of 165 m depth (Table 2, Boul-1) for 10 same place, not moving away. Despite it disappeared from the slowly. Eventually it reached 12 m deeper during one hour and surveys during the seven months and seems to live in this area. near a cliff at 152.3 m depth (Table 2: E24, Manado-8), where encountered in the same area, at locations close from each where they tend to choose their daytime habitats (FRICKE et al., of L. menadoensis is similar to that of L. chalumnae, with We are grateful to Dr. Teruya UYENO for his valuable B., ABDALLAH, A. and KALOMBO, H. 2006. Coelacanth (SMITH, 1939). The first observation of the living coelacanth The Indonesian coelacanth, L. menadoensis, was first minutes (Table 2, E1). The adult individual was in stationary camera frame we kept setting the ROV there in front of the 42 minutes. During the 2007 survey (Table 1: R5) a coelacanth (Table located on the same slope of Manado-6. other within hundreds of meters. However, in the Comorian 1991). The suitable temperature for L. menadoensis seems to be rocky steep slopes with undermarine caves or overhangs. advice, guidance and encouragement throughout the present (Latimeria chalumnae SMITH, 1939) discoveries and habitats using submersibles was reported in Comoros (FRICKE discovered in Manado, Sulawesi Island, Indonesia, in 1997. position in a cave (Buol-1) slowly moving its pectoral, pelvic, cave until the next morning, however it did not appear again. On the 14th December, the individual ID 3 previously 2: ID 7) was recorded off Manado by 195 m depth, with a Another new coelacanth (Table 2: E23, ID 25) was Archipelago, same individuals of L. chalumnae were observed, almost the same as L. chalumnae. However, Latimeria menadoensis was also observed alongside study. Many thanks go to colleagues of the Faculty of Fisheries conservation in Tanzania. South African Journal of et al., 1987). Latimeria chalumnae hide in submarine caves The first specimen was captured in 1998 (ERDMANN et al., second dorsal and anal fins. The day after, another individual On the 4th June in 2006, three individuals, including ID 3 recorded at Buol-2 in a cave and at Buol-3 in a crack, was temperature of 12.4 °C, the coolest one registered during all recorded alone on a Manado site (Manado-7). within a two week period, in several caves distributed in a 8 Water temperature can increase or drop for more than 5 large rocks or along vertical wall in daytime. Some of these and Marine Sciences of the Sam Ratulangi University at Science, 102: 486–490. along steep slopes between 100 m and 200 m depth during 1998). According to a genetic study, the specimen differed (ID 2) and an unidentified one (UN1) were found in the same which was observed on the 31th May at Buol-2 and two new encountered again (Table 2: Buol-5) near a vertical wall at 145 surveys. Field surveys were conducted from the 19th to 30th May kilometers wide area (FRICKE et al., 1991). °C in a very short period of time (about 30 min according to our individuals were encountered above 200 m deep, in an Manado, Indonesia, for their generous supports to conduct BRUTON, M. N., CABRAL, A. J. P. and FRICKE, H. 1992. First daytime and come out and drift near the ocean floor for from L. chalumnae and a new species was erected (POUYAUD cave (Buol-1). The unidentified one immediately hid deeper in individuals, were observed in a vertical crack which was m depth (Table 2: E6). After a while, this individual (ID 3) The field survey in 2009 was carried out off Manado and 2015 off Lolak Island and from the 30th October to the 16th When individuals of L. menadoensis were encountered, recordings). The coelacanths however often stayed there environment that remains slightly bright. Latimeria expeditions. Also we express our thanks to Dr. SUHARSONO, Dr. capture of a coelacanth, Latimeria chalumnae (Pisces, hunting prey at night (FRICKE et al., 1991). However, since the et al., 1999). In 1999 two individuals were observed by the cave, where the ROV could not access, and only its lateral several hundred meters far from Buol-2 (Buol-3) for 2 hours moved upward about 18 m and stayed behind a large rock (Fig. around Talise and Bangka islands off the northern coast of November 2015 off Lolak and Bitung Islands (Table 1; R13 some of them stayed stationary at same place but some without moving away or significantly modifying their menadoensis seems to be less sensitive to the daylight than L. Zainal ARIFIN, Dr. DIRHAMSYAH and colleagues of the Indonesian Latimeriidae), off Mozambique. South African Journal of record of juvenile or small (below 80 cm long) individuals is submersible in a location 360 km away from Manado (FRICKE side of the body was recorded. The individual (ID 2) was and 14 minutes (Fig. 2C; Table 2: E4, ID 3, 4, 5). During this 3). The observation time was the longest of all surveys with Sulawesi Island from the 12th September to the 9th October and R14). Three new individuals (Table 2: ID 26–28) were individuals such as ID 6 on the 12th December 2006 (E5) and behavior. These water currents of different temperatures are chalumnae. Institute of Science for their advice and administrative work to Science, 88: 225–227. virtually absent, the growth, development and breeding et al., 2000). Today 8 individuals were officially caught in the observed for one minute before it hid inside the cave (Table 2: long period of time all individuals stayed near the rock wall four hours and 16 minutes of recording. The temperature (Table 1: R8). On the 24th September 2009, six individuals recognized at different places off Lolak Island (Lolak 1-4). All ID 24 on the 4th May 2012 (E22) swam away. It seemed to caused by the movement of the marine thermocline. During These first observations are potentially crucial since the obtain our research permit. Other thanks go to Mr. OPO, Mr. CUPELLO, C., BRITO, P. M., HERBIN, M., MEUNIER, F. J., biology of L. chalumnae remain poorly known. Our knowledge Indonesian seas. However, the details of their habitat have not E2). When a new individual could be identified by the white always keeping their head against the water flow. The water changed up and down several times (Fig. 4). Its difference (Table 2: E11, ID 9, 10, 11, 12, UN 2, 3) were found under an individuals were alone. ID 26 was observed in May and once avoid and escape from the brightness of the light or the sounds our surveys we observed in some area that as the thermocline habitats of the Indonesian coelacanths could be possibly not as REFRY and colleagues at Murex Dive Resort, and DAUD, AL, J ANVIER, P., DUTEL, H. and CLÉMENT, G. 2015. 52 IWATA et al. Field surveys on Latimeria menadoensis using ROV from 2005 to 2015 53 been described. The aim of this study is to clear their habitat Table 3. Total length of each coelacanth estimated with 20 cm distance parallel laser lines applied Allometric growth in the extant coelacanth lung during capture of a coelacanth off Madagascar. South African and distribution. from the ROV. ontogenetic development. Nature Communications, 6: Journal of Science 92: 150–151. 8222. DOI: 10.1038/ncomms9222 POUYAUD, L., WIRJOATMODJO, S., RACHMATIKA, I., TJAKRAWIDJAJA, ID number 7 8 14 15 16 17 19 20 22 23 24 DE VOS, L. and OYUGI, D. 2002. First capture of a coelacanth, A., HADIATY, R. and HADIE, W. 1999. A new species of METHODS Estimated TL (cm) 123 115 110 31.5 113 125 114 113 106 90 137 Latimeria chalumnae SMITH, 1939 (Pisces, Latimeriidae), coelacanth. Genetic and morphologic proof. C. R. off Kenya. South African Journal of Science, 98: 345–347. Academy of Science, 322: 261–267. Two remotely operated vehicles (ROVs) (Kowa; ERDMANN, M. V., CALDWELL, R. L. and MOOSA, M. K. 1998. SMITH, J. L. B. 1939. The living coelacanth fish from South VEGA300) were used for the surveys. The first ROV was Indonesian ‘king of the sea’ discovered. Nature, 395: 335. Africa. Nature, 143: 748–750. replaced with the second one in 2007. Our ROVs are able to overhang off Talise Island between 144 and 150 m deep. There again in October 2015 on a steep slope (Table 2: E25, E27). FRICKE, H., REINICKE, O., HOFER, H. and NACHTIGALL, W. SMITH, J. L. B. 1953. The second coelacanth. Nature, 171: dive up to 300 m depth. These had two vertical, two horizontal was almost no water current in this area. All individuals were The coelacanth ID 28 was found on the 1st November along 1987. Locomotion of the coelacanth Latimeria chalumnae 99–101. and two right-left propellers and these were controlled from a close to each other under the same overhang and most of them one of the large rocks scattered on a gentle slope in a bay of in its natural environment. Nature, 329: 331–333. SMITH, C. L., RAND, C. S., SCHAEFFER, B. and ATZ, J. W. 1975. boat on surface through a 400 m long tether cable. The stayed still with their head facing downward. Four days later, the Lolak Island by 125 m depth. FRICKE, H., HISSMANN, K., SCHAUER, J., REINICKE, O., KASANG, Latimeria, the living coelacanth, is ovoviviparous. Science, underwater operations were visualized on a screen with ID 9 was observed again, alone on a steep slope at 172.9 m The encountered Indonesian coelacanths during these L. and PLANTE, R. 1991. Habitat and population size of 190: 1105–1106. information including directions of ROVs, depth, water depth (Table 2: E12), which was located within several dozen surveys were in caves, alongside large rocks, under overhangs coelacanth Latimeria chalumnae at Grand Comoro. VENTER, P., TIMM, P., GUNN, G., LE ROUX, E., SERFONTEIN, E., temperature, date and time. All data were directly recorded on meters far from Talise-2 and 30 m deeper. or on steep slopes (Table 2; Fig. 2). Environmental Biology of Fishes, 32: 287–300. SMITH, P., SMITH, E., BENSCH, M., HARDING, D. and video-tapes on the boat. Since the 2007 survey, water Four new coelacanths were recorded from the 29th Most individuals were found under overhangs or FRICKE, H., HISSMANN, K., SCHAUER, J., ERDMANN, M., HEEMSTRA, P. 2000. Discovery of a viable population of temperature and depth were independently recorded with more September to the 6th October 2009 in Talise and Banggka alongside large rocks. Some individuals were encountered not MOOSA, M. K. and PLANTE, R. 2000. Biogeography of the coelacanths (Latimeria chalumnae SMITH, 1939) at accuracy by external measuring memories (Alec Electronics, Islands and off Manado (Table 2: E13 and 14, ID 13, 14, 15). to hide in any shade and stayed just on rocky slopes. In Indonesian coelacanth. Nature, 403: 38. Sodwana Bay, South Africa. South African Journal of later named JFE Alec; MDS-MKV/T, MDS-MKV/D). In ID 15 is the only juvenile coelacanth filmed so far. It was in a overhangs and cracks, all individuals had their ventral side HEEMSTRA, P. C., FREEMAN, A. L., WONG, H. Y., HENSLEY, D. Science, 96: 567–568. addition two laser beam irradiators, that provide line lasers of small crack between 164.6 and 170.9 m deep in a temperature alongside and close to a rock wall, but without touching it. A. and RABESANDRATANA, H. D. 1996. First authentic 20-cm distance, were attached to the second ROV and have between 14.5 and 15 °C. Estimated total lengths are shown in Table 3. Most been used to register the size of encountered animals. The The shallowest record of a coelacanth was at 115.6 m individuals are more than 1 m in total length. ID 15 is a surveys were conducted off northern parts of Sulawesi Island depth off the southern part of Manado (Table 2: E16, juvenile individual of 31.5 cm and ID 23 is 90 cm in total and along the Biak Island located in northwestern New Guinea Manado-3) on the 9th October in 2009. ID 16 (Table 2) was length. No individual beyond 140 cm has been recorded in Island, Indonesia. The areas and time periods of each survey found at the end of the morning, alone alongside a large rock Indonesia so far. restricted as those of the African coelacanths, and so a broader and other staffs for our work on the research boat. Dr. BATUNA are shown in Table 1 and Fig. 1. on the edge of a shelf. The water temperature was about 20.0 distribution of the Indonesian coelacanth could be expected. and the Murex staffs supported our daily life during the ºC. Occurrences of coelacanths along the eastern coast of the expeditions. We would like to express our sincere thanks to From the 6th to the 16th November in 2010, the field of the propellers of the ROV. level changed, the water flow was reversed. Coelacanths African continent were recorded in South Africa (SMITH, colleagues at Aquamarine Fukushima, Japan, for their RESULTS survey was conducted around Biak Island (Table 1: R9). Adult DISCUSSION Temperature and depth data for each Indonesian always kept their head against the water flow, like other 1939), Comoros (SMITH, 1953), Madagascar (HEEMSTRA et al., understanding and supports. We would like to thank Dr. Gaël coelacanths were found at two different sites (Table 2: coelacanth record are shown in Fig. 3. Though an average of observed actinopterygian fishes did. It means that as the 1996), Kenya (DE VOS and OYUGI, 2002), Tanzania (BENNO et CLÉMENT of Muséum national d’Histoire naturelle and Dr. The field survey of the Indonesian coelacanth, Latimeria Fig. 2. Configuration of encounter point. A, over hang (E9); B, over hang (E17); C, crack (E4); D, crack (E21); E, beside big rock E17–19). Two individuals (ID 17 and 18) were found under an During the field surveys by underwater ROV recording water temperature on the ocean surface was about 30 °C, the direction of the water flow changed, the swimming direction of al., 2006), and Mozambique (BRUTON et al., 1992). In Camila CUPELLO of Universidade do Estado do Rio de Janeiro menadoensis, was conducted 14 times from 2005 to 2015 with (E28); F, beside big rock (E25); G, wall (E20); H, steep slope (E12); I, steep slope, beside big rock (E13). Red triangles overhang between 212.5 and 218.9 m deep (Table 2: Biak-1) for Indonesian coelacanths, Latimeria menadoensis, from 2005 temperature often dropped into about 11 °C at 300 m water depth. each coelacanth individual changed accordingly. Indonesia, coelacanths have only been found off the northern as a referee for their critical reading of the manuscript and their 1173 underwater operations around Sulawesi and Biak islands indicate coelacanths. on the 11th November. Three other ones (ID 19, 20, 21) were to 2015, 30 different individuals were identified and three The temperatures changed widely even though the changes of On the 24th September 2009 (Table 2: E11) six coast of the Sulawesi Island and the southern coast of the Biak comments. in Indonesia (Fig. 1, Table 1). No coelacanth was encountered registered under an overhang between 193.2 and 195.9 m deep others were unidentified. Among those, six individuals were depth were small at several places. All Indonesian coelacanths, individuals were recognized under an overhang (Talise-2). The Island in northern New Guinea. Unfortunately one of our authors, Dr. Djoko Hadi KUNARSO during the first surveys in 2005 although 452 underwater (Table 2: Biak-2) on the 13th November. Two days later, observed more than twice: ID 3 at Buol-2, 3, 5, 6 (Table 2: E3, L. menadoensis, were encountered during day time at a depth all had about the same size. There was almost no water flow, Field surveys of the Indonesian coelacanths should be has passed away during the process of this contribution. We operations were conducted around Manado-tua Island and spots pattern on its body, an ID number was allocated. The temperature changed up and down by five degrees during the between the maximum and the minimum was 6.7 °C, which Biak-2 was observed again and only ID 20 stayed there. E4, E6 and E7); ID 8 at Talise-1 (E9 and E10), ID 9 at Talise 2 between 115.6 and 218.9 m and a temperature between 12.4 and each individual was head downward. They were close to continued in all area of Indonesia and expanded to all pray for the response of his soul. other close Indonesian islands from the 17th to 30th of April individual which could not be distinguished was allocated an observation due to water flow reversals. When the water flow was the largest range of all surveys. Two new coelacanths were recorded off Manado in and 3 (E11 and E12), ID 20 at Biak-2 (E18 and E19), ID 24 at and 21.5 ºC. Indonesian coelacanths have exceptionally been each other but no direct contacts or interactions between Southeast Asia in order to increase our knowledge of the This study was supported in part by funding from (Table 1, R1). The second field survey was held along the UN (unknown) + number. was reversed some times, all individuals always changed their On the next day, on the 15th December, the ID 3 was December 2010 (Table 1: R10; Table 2: E 20 and 21; ID 22 Manado-6 and 8 (E22 and E24), ID 26 at Lolak-1 and 3 (E25 recorded in an environment with a temperature exceeding 20 coelacanths were observed. No evidence of a social behavior distribution of this rare species and to define, in a close future, Interdisciplinary Collaborative Research program of Atmosphere northern coast of Sulawesi from the 6th to 19th May in 2006, In the end of the morning of the same day, a third directions against the water flow. observed again under an overhang (Table 2: E7, Buol-6) near and 23). and E27). ID 8 was observed on the 14th September 2009 and ºC, regardless of the depth. The African coelacanth, L. has been recognized in the Comorian populations of the most relevant conservation policies of the different known and Ocean Research Institute, the University of Tokyo. but once again despite of 107 underwater operations no individual (Table 2: E3, ID 3) was found in another cave During the field survey carried on December of the same other sites where it was previously found at Buol-2 , Buol-3 On the 4th May 2012, an individual (Table 2: E22, ID 24) was encountered again at the same place (Talise-1) the day chalumnae, was observed by submarine vehicles in steep coelacanths (FRICKE et al., 1991). populations. coelacanth was recorded. (Buol-2), larger and located 20 m deeper (Buol-2) than the first year, in the same Buol area (Table 1: R4) the sixth individual and Buol-5. Its posture was upside down with its ventral side was found on a steep slope at 169.4 m depth off Manado after. ID 24 was observed on the 4th May 2012 and then 13 slopes between 150 m and 253 m (FRICKE et al., 1991). Divers According to the observations of the African coelacanths, INTRODUCTION of the reproduction biology of the coelacanths is almost The first individual of Indonesian coelacanth (Table 2, ID cave (Buol-1). After 29 minutes of recording, this individual (Table 2: E5, ID 6) was recognized at 14:34 on the 12th facing the ceiling of the overhang. Such posture is also known (Table 2: Manado-6). It swam up and down 5.4 m between months later, on the 9th June 2013 at the same locations of recorded the presence of this species at 104 m depth at the species L. chalumnae is nocturnal and hides in cave in REFERENCES restricted to an ovoviviparity strategy since some caught large 1) was recorded during the third field survey, in the morning of hid inside the cave. Although the water temperature increased December alongside a large rock. This individual stayed in the in coelacanths from the Comorian Archipelago (FRICKE et al., 168.1 and 173.5 m deep for 17 minutes. Thirteen months later Manado area within several kilometers. Sodwana Bay, South Africa (VENTER et al., 2000). Latimeria daytime but comes out at night, supposedly in search of food ACKNOWLEDGMENTS The first living coelacanth, Latimeria chalumnae, was females carried unfertilized eggs or embryos in their oviduct the 30th May 2006 (Table 1, R3). The coelacanth was of more than five degrees, from 14.8 to 20.4 ºC, it stayed at the beginning of the observation and then moved ahead to deeper 1987). ID 3 was found totally four times in this area in two (the 9th June 2003), this individual (ID 24) was observed again All individuals observed more than twice were chalumnae seem to prefer a temperature range of 15–20 °C, (FRICKE et al., 1991). The living geomorphologic environment BENNO, B., VERHEIJI, E., STAPLEY, J., RUMISHA, C., NGATUNGA, discovered in South Africa in the Indian Ocean in 1938 (SMITH et al., 1975; CUPELLO et al., 2015). observed in a cave of 165 m depth (Table 2, Boul-1) for 10 same place, not moving away. Despite it disappeared from the slowly. Eventually it reached 12 m deeper during one hour and surveys during the seven months and seems to live in this area. near a cliff at 152.3 m depth (Table 2: E24, Manado-8), where encountered in the same area, at locations close from each where they tend to choose their daytime habitats (FRICKE et al., of L. menadoensis is similar to that of L. chalumnae, with We are grateful to Dr. Teruya UYENO for his valuable B., ABDALLAH, A. and KALOMBO, H. 2006. Coelacanth (SMITH, 1939). The first observation of the living coelacanth The Indonesian coelacanth, L. menadoensis, was first minutes (Table 2, E1). The adult individual was in stationary camera frame we kept setting the ROV there in front of the 42 minutes. During the 2007 survey (Table 1: R5) a coelacanth (Table located on the same slope of Manado-6. other within hundreds of meters. However, in the Comorian 1991). The suitable temperature for L. menadoensis seems to be rocky steep slopes with undermarine caves or overhangs. advice, guidance and encouragement throughout the present (Latimeria chalumnae SMITH, 1939) discoveries and habitats using submersibles was reported in Comoros (FRICKE discovered in Manado, Sulawesi Island, Indonesia, in 1997. position in a cave (Buol-1) slowly moving its pectoral, pelvic, cave until the next morning, however it did not appear again. On the 14th December, the individual ID 3 previously 2: ID 7) was recorded off Manado by 195 m depth, with a Another new coelacanth (Table 2: E23, ID 25) was Archipelago, same individuals of L. chalumnae were observed, almost the same as L. chalumnae. However, Latimeria menadoensis was also observed alongside study. Many thanks go to colleagues of the Faculty of Fisheries conservation in Tanzania. South African Journal of et al., 1987). Latimeria chalumnae hide in submarine caves The first specimen was captured in 1998 (ERDMANN et al., second dorsal and anal fins. The day after, another individual On the 4th June in 2006, three individuals, including ID 3 recorded at Buol-2 in a cave and at Buol-3 in a crack, was temperature of 12.4 °C, the coolest one registered during all recorded alone on a Manado site (Manado-7). within a two week period, in several caves distributed in a 8 Water temperature can increase or drop for more than 5 large rocks or along vertical wall in daytime. Some of these and Marine Sciences of the Sam Ratulangi University at Science, 102: 486–490. along steep slopes between 100 m and 200 m depth during 1998). According to a genetic study, the specimen differed (ID 2) and an unidentified one (UN1) were found in the same which was observed on the 31th May at Buol-2 and two new encountered again (Table 2: Buol-5) near a vertical wall at 145 surveys. Field surveys were conducted from the 19th to 30th May kilometers wide area (FRICKE et al., 1991). °C in a very short period of time (about 30 min according to our individuals were encountered above 200 m deep, in an Manado, Indonesia, for their generous supports to conduct BRUTON, M. N., CABRAL, A. J. P. and FRICKE, H. 1992. First daytime and come out and drift near the ocean floor for from L. chalumnae and a new species was erected (POUYAUD cave (Buol-1). The unidentified one immediately hid deeper in individuals, were observed in a vertical crack which was m depth (Table 2: E6). After a while, this individual (ID 3) The field survey in 2009 was carried out off Manado and 2015 off Lolak Island and from the 30th October to the 16th When individuals of L. menadoensis were encountered, recordings). The coelacanths however often stayed there environment that remains slightly bright. Latimeria expeditions. Also we express our thanks to Dr. SUHARSONO, Dr. capture of a coelacanth, Latimeria chalumnae (Pisces, hunting prey at night (FRICKE et al., 1991). However, since the et al., 1999). In 1999 two individuals were observed by the cave, where the ROV could not access, and only its lateral several hundred meters far from Buol-2 (Buol-3) for 2 hours moved upward about 18 m and stayed behind a large rock (Fig. around Talise and Bangka islands off the northern coast of November 2015 off Lolak and Bitung Islands (Table 1; R13 some of them stayed stationary at same place but some without moving away or significantly modifying their menadoensis seems to be less sensitive to the daylight than L. Zainal ARIFIN, Dr. DIRHAMSYAH and colleagues of the Indonesian Latimeriidae), off Mozambique. South African Journal of record of juvenile or small (below 80 cm long) individuals is submersible in a location 360 km away from Manado (FRICKE side of the body was recorded. The individual (ID 2) was and 14 minutes (Fig. 2C; Table 2: E4, ID 3, 4, 5). During this 3). The observation time was the longest of all surveys with Sulawesi Island from the 12th September to the 9th October and R14). Three new individuals (Table 2: ID 26–28) were individuals such as ID 6 on the 12th December 2006 (E5) and behavior. These water currents of different temperatures are chalumnae. Institute of Science for their advice and administrative work to Science, 88: 225–227. virtually absent, the growth, development and breeding et al., 2000). Today 8 individuals were officially caught in the observed for one minute before it hid inside the cave (Table 2: long period of time all individuals stayed near the rock wall four hours and 16 minutes of recording. The temperature (Table 1: R8). On the 24th September 2009, six individuals recognized at different places off Lolak Island (Lolak 1-4). All ID 24 on the 4th May 2012 (E22) swam away. It seemed to caused by the movement of the marine thermocline. During These first observations are potentially crucial since the obtain our research permit. Other thanks go to Mr. OPO, Mr. CUPELLO, C., BRITO, P. M., HERBIN, M., MEUNIER, F. J., biology of L. chalumnae remain poorly known. Our knowledge Indonesian seas. However, the details of their habitat have not E2). When a new individual could be identified by the white always keeping their head against the water flow. The water changed up and down several times (Fig. 4). Its difference (Table 2: E11, ID 9, 10, 11, 12, UN 2, 3) were found under an individuals were alone. ID 26 was observed in May and once avoid and escape from the brightness of the light or the sounds our surveys we observed in some area that as the thermocline habitats of the Indonesian coelacanths could be possibly not as REFRY and colleagues at Murex Dive Resort, and DAUD, AL, J ANVIER, P., DUTEL, H. and CLÉMENT, G. 2015. 54 IWATA et al. Field surveys on Latimeria menadoensis using ROV from 2005 to 2015 55 been described. The aim of this study is to clear their habitat Allometric growth in the extant coelacanth lung during capture of a coelacanth off Madagascar. South African and distribution. ontogenetic development. Nature Communications, 6: Journal of Science 92: 150–151. 8222. DOI: 10.1038/ncomms9222 POUYAUD, L., WIRJOATMODJO, S., RACHMATIKA, I., TJAKRAWIDJAJA, DE VOS, L. and OYUGI, D. 2002. First capture of a coelacanth, A., HADIATY, R. and HADIE, W. 1999. A new species of METHODS Latimeria chalumnae SMITH, 1939 (Pisces, Latimeriidae), coelacanth. Genetic and morphologic proof. C. R. off Kenya. South African Journal of Science, 98: 345–347. Academy of Science, 322: 261–267. Two remotely operated vehicles (ROVs) (Kowa; ERDMANN, M. V., CALDWELL, R. L. and MOOSA, M. K. 1998. SMITH, J. L. B. 1939. The living coelacanth fish from South VEGA300) were used for the surveys. The first ROV was Indonesian ‘king of the sea’ discovered. Nature, 395: 335. Africa. Nature, 143: 748–750. replaced with the second one in 2007. Our ROVs are able to overhang off Talise Island between 144 and 150 m deep. There again in October 2015 on a steep slope (Table 2: E25, E27). FRICKE, H., REINICKE, O., HOFER, H. and NACHTIGALL, W. SMITH, J. L. B. 1953. The second coelacanth. Nature, 171: dive up to 300 m depth. These had two vertical, two horizontal was almost no water current in this area. All individuals were The coelacanth ID 28 was found on the 1st November along 1987. Locomotion of the coelacanth Latimeria chalumnae 99–101. and two right-left propellers and these were controlled from a close to each other under the same overhang and most of them one of the large rocks scattered on a gentle slope in a bay of in its natural environment. Nature, 329: 331–333. SMITH, C. L., RAND, C. S., SCHAEFFER, B. and ATZ, J. W. 1975. boat on surface through a 400 m long tether cable. The stayed still with their head facing downward. Four days later, the Lolak Island by 125 m depth. FRICKE, H., HISSMANN, K., SCHAUER, J., REINICKE, O., KASANG, Latimeria, the living coelacanth, is ovoviviparous. Science, underwater operations were visualized on a screen with ID 9 was observed again, alone on a steep slope at 172.9 m The encountered Indonesian coelacanths during these L. and PLANTE, R. 1991. Habitat and population size of 190: 1105–1106. information including directions of ROVs, depth, water depth (Table 2: E12), which was located within several dozen surveys were in caves, alongside large rocks, under overhangs coelacanth Latimeria chalumnae at Grand Comoro. VENTER, P., TIMM, P., GUNN, G., LE ROUX, E., SERFONTEIN, E., temperature, date and time. All data were directly recorded on meters far from Talise-2 and 30 m deeper. or on steep slopes (Table 2; Fig. 2). Environmental Biology of Fishes, 32: 287–300. SMITH, P., SMITH, E., BENSCH, M., HARDING, D. and video-tapes on the boat. Since the 2007 survey, water Four new coelacanths were recorded from the 29th Most individuals were found under overhangs or FRICKE, H., HISSMANN, K., SCHAUER, J., ERDMANN, M., HEEMSTRA, P. 2000. Discovery of a viable population of temperature and depth were independently recorded with more September to the 6th October 2009 in Talise and Banggka alongside large rocks. Some individuals were encountered not MOOSA, M. K. and PLANTE, R. 2000. Biogeography of the coelacanths (Latimeria chalumnae SMITH, 1939) at accuracy by external measuring memories (Alec Electronics, Islands and off Manado (Table 2: E13 and 14, ID 13, 14, 15). to hide in any shade and stayed just on rocky slopes. In Indonesian coelacanth. Nature, 403: 38. Sodwana Bay, South Africa. South African Journal of later named JFE Alec; MDS-MKV/T, MDS-MKV/D). In ID 15 is the only juvenile coelacanth filmed so far. It was in a overhangs and cracks, all individuals had their ventral side HEEMSTRA, P. C., FREEMAN, A. L., WONG, H. Y., HENSLEY, D. Science, 96: 567–568. addition two laser beam irradiators, that provide line lasers of small crack between 164.6 and 170.9 m deep in a temperature alongside and close to a rock wall, but without touching it. A. and RABESANDRATANA, H. D. 1996. First authentic 20-cm distance, were attached to the second ROV and have between 14.5 and 15 °C. Estimated total lengths are shown in Table 3. Most been used to register the size of encountered animals. The The shallowest record of a coelacanth was at 115.6 m individuals are more than 1 m in total length. ID 15 is a Fig. 3. Water temperature and depth during observations of coelacanths in each locality. The orange box shows depth. The top and Fig. 4. Change of the water temperature at Buol-5 off Buol of Sulawesi Island. surveys were conducted off northern parts of Sulawesi Island depth off the southern part of Manado (Table 2: E16, juvenile individual of 31.5 cm and ID 23 is 90 cm in total bottom of the box mean a minimum depth and maximum depth of coelacanths stay. The blue bar shows a maximum and and along the Biak Island located in northwestern New Guinea Manado-3) on the 9th October in 2009. ID 16 (Table 2) was length. No individual beyond 140 cm has been recorded in minimum water temperatures. The open circle shows an average of temperature during observation. All individuals of L. Island, Indonesia. The areas and time periods of each survey found at the end of the morning, alone alongside a large rock Indonesia so far. menadoensis were observed in depth between 115.6 and 218.9 m and water temperature between 12.4 and 21.5 ºC. restricted as those of the African coelacanths, and so a broader and other staffs for our work on the research boat. Dr. BATUNA are shown in Table 1 and Fig. 1. on the edge of a shelf. The water temperature was about 20.0 distribution of the Indonesian coelacanth could be expected. and the Murex staffs supported our daily life during the ºC. Occurrences of coelacanths along the eastern coast of the expeditions. We would like to express our sincere thanks to From the 6th to the 16th November in 2010, the field of the propellers of the ROV. level changed, the water flow was reversed. Coelacanths African continent were recorded in South Africa (SMITH, colleagues at Aquamarine Fukushima, Japan, for their RESULTS survey was conducted around Biak Island (Table 1: R9). Adult DISCUSSION Temperature and depth data for each Indonesian always kept their head against the water flow, like other 1939), Comoros (SMITH, 1953), Madagascar (HEEMSTRA et al., understanding and supports. We would like to thank Dr. Gaël coelacanths were found at two different sites (Table 2: coelacanth record are shown in Fig. 3. Though an average of observed actinopterygian fishes did. It means that as the 1996), Kenya (DE VOS and OYUGI, 2002), Tanzania (BENNO et CLÉMENT of Muséum national d’Histoire naturelle and Dr. The field survey of the Indonesian coelacanth, Latimeria E17–19). Two individuals (ID 17 and 18) were found under an During the field surveys by underwater ROV recording water temperature on the ocean surface was about 30 °C, the direction of the water flow changed, the swimming direction of al., 2006), and Mozambique (BRUTON et al., 1992). In Camila CUPELLO of Universidade do Estado do Rio de Janeiro menadoensis, was conducted 14 times from 2005 to 2015 with overhang between 212.5 and 218.9 m deep (Table 2: Biak-1) for Indonesian coelacanths, Latimeria menadoensis, from 2005 temperature often dropped into about 11 °C at 300 m water depth. each coelacanth individual changed accordingly. Indonesia, coelacanths have only been found off the northern as a referee for their critical reading of the manuscript and their 1173 underwater operations around Sulawesi and Biak islands on the 11th November. Three other ones (ID 19, 20, 21) were to 2015, 30 different individuals were identified and three The temperatures changed widely even though the changes of On the 24th September 2009 (Table 2: E11) six coast of the Sulawesi Island and the southern coast of the Biak comments. in Indonesia (Fig. 1, Table 1). No coelacanth was encountered registered under an overhang between 193.2 and 195.9 m deep others were unidentified. Among those, six individuals were depth were small at several places. All Indonesian coelacanths, individuals were recognized under an overhang (Talise-2). The Island in northern New Guinea. Unfortunately one of our authors, Dr. Djoko Hadi KUNARSO during the first surveys in 2005 although 452 underwater (Table 2: Biak-2) on the 13th November. Two days later, observed more than twice: ID 3 at Buol-2, 3, 5, 6 (Table 2: E3, L. menadoensis, were encountered during day time at a depth all had about the same size. There was almost no water flow, Field surveys of the Indonesian coelacanths should be has passed away during the process of this contribution. We operations were conducted around Manado-tua Island and spots pattern on its body, an ID number was allocated. The temperature changed up and down by five degrees during the between the maximum and the minimum was 6.7 °C, which Biak-2 was observed again and only ID 20 stayed there. E4, E6 and E7); ID 8 at Talise-1 (E9 and E10), ID 9 at Talise 2 between 115.6 and 218.9 m and a temperature between 12.4 and each individual was head downward. They were close to continued in all area of Indonesia and expanded to all pray for the response of his soul. other close Indonesian islands from the 17th to 30th of April individual which could not be distinguished was allocated an observation due to water flow reversals. When the water flow was the largest range of all surveys. Two new coelacanths were recorded off Manado in and 3 (E11 and E12), ID 20 at Biak-2 (E18 and E19), ID 24 at and 21.5 ºC. Indonesian coelacanths have exceptionally been each other but no direct contacts or interactions between Southeast Asia in order to increase our knowledge of the This study was supported in part by funding from (Table 1, R1). The second field survey was held along the UN (unknown) + number. was reversed some times, all individuals always changed their On the next day, on the 15th December, the ID 3 was December 2010 (Table 1: R10; Table 2: E 20 and 21; ID 22 Manado-6 and 8 (E22 and E24), ID 26 at Lolak-1 and 3 (E25 recorded in an environment with a temperature exceeding 20 coelacanths were observed. No evidence of a social behavior distribution of this rare species and to define, in a close future, Interdisciplinary Collaborative Research program of Atmosphere northern coast of Sulawesi from the 6th to 19th May in 2006, In the end of the morning of the same day, a third directions against the water flow. observed again under an overhang (Table 2: E7, Buol-6) near and 23). and E27). ID 8 was observed on the 14th September 2009 and ºC, regardless of the depth. The African coelacanth, L. has been recognized in the Comorian populations of the most relevant conservation policies of the different known and Ocean Research Institute, the University of Tokyo. but once again despite of 107 underwater operations no individual (Table 2: E3, ID 3) was found in another cave During the field survey carried on December of the same other sites where it was previously found at Buol-2 , Buol-3 On the 4th May 2012, an individual (Table 2: E22, ID 24) was encountered again at the same place (Talise-1) the day chalumnae, was observed by submarine vehicles in steep coelacanths (FRICKE et al., 1991). populations. coelacanth was recorded. (Buol-2), larger and located 20 m deeper (Buol-2) than the first year, in the same Buol area (Table 1: R4) the sixth individual and Buol-5. Its posture was upside down with its ventral side was found on a steep slope at 169.4 m depth off Manado after. ID 24 was observed on the 4th May 2012 and then 13 slopes between 150 m and 253 m (FRICKE et al., 1991). Divers According to the observations of the African coelacanths, INTRODUCTION of the reproduction biology of the coelacanths is almost The first individual of Indonesian coelacanth (Table 2, ID cave (Buol-1). After 29 minutes of recording, this individual (Table 2: E5, ID 6) was recognized at 14:34 on the 12th facing the ceiling of the overhang. Such posture is also known (Table 2: Manado-6). It swam up and down 5.4 m between months later, on the 9th June 2013 at the same locations of recorded the presence of this species at 104 m depth at the species L. chalumnae is nocturnal and hides in cave in REFERENCES restricted to an ovoviviparity strategy since some caught large 1) was recorded during the third field survey, in the morning of hid inside the cave. Although the water temperature increased December alongside a large rock. This individual stayed in the in coelacanths from the Comorian Archipelago (FRICKE et al., 168.1 and 173.5 m deep for 17 minutes. Thirteen months later Manado area within several kilometers. Sodwana Bay, South Africa (VENTER et al., 2000). Latimeria daytime but comes out at night, supposedly in search of food ACKNOWLEDGMENTS The first living coelacanth, Latimeria chalumnae, was females carried unfertilized eggs or embryos in their oviduct the 30th May 2006 (Table 1, R3). The coelacanth was of more than five degrees, from 14.8 to 20.4 ºC, it stayed at the beginning of the observation and then moved ahead to deeper 1987). ID 3 was found totally four times in this area in two (the 9th June 2003), this individual (ID 24) was observed again All individuals observed more than twice were chalumnae seem to prefer a temperature range of 15–20 °C, (FRICKE et al., 1991). The living geomorphologic environment BENNO, B., VERHEIJI, E., STAPLEY, J., RUMISHA, C., NGATUNGA, discovered in South Africa in the Indian Ocean in 1938 (SMITH et al., 1975; CUPELLO et al., 2015). observed in a cave of 165 m depth (Table 2, Boul-1) for 10 same place, not moving away. Despite it disappeared from the slowly. Eventually it reached 12 m deeper during one hour and surveys during the seven months and seems to live in this area. near a cliff at 152.3 m depth (Table 2: E24, Manado-8), where encountered in the same area, at locations close from each where they tend to choose their daytime habitats (FRICKE et al., of L. menadoensis is similar to that of L. chalumnae, with We are grateful to Dr. Teruya UYENO for his valuable B., ABDALLAH, A. and KALOMBO, H. 2006. Coelacanth (SMITH, 1939). The first observation of the living coelacanth The Indonesian coelacanth, L. menadoensis, was first minutes (Table 2, E1). The adult individual was in stationary camera frame we kept setting the ROV there in front of the 42 minutes. During the 2007 survey (Table 1: R5) a coelacanth (Table located on the same slope of Manado-6. other within hundreds of meters. However, in the Comorian 1991). The suitable temperature for L. menadoensis seems to be rocky steep slopes with undermarine caves or overhangs. advice, guidance and encouragement throughout the present (Latimeria chalumnae SMITH, 1939) discoveries and habitats using submersibles was reported in Comoros (FRICKE discovered in Manado, Sulawesi Island, Indonesia, in 1997. position in a cave (Buol-1) slowly moving its pectoral, pelvic, cave until the next morning, however it did not appear again. On the 14th December, the individual ID 3 previously 2: ID 7) was recorded off Manado by 195 m depth, with a Another new coelacanth (Table 2: E23, ID 25) was Archipelago, same individuals of L. chalumnae were observed, almost the same as L. chalumnae. However, Latimeria menadoensis was also observed alongside study. Many thanks go to colleagues of the Faculty of Fisheries conservation in Tanzania. South African Journal of et al., 1987). Latimeria chalumnae hide in submarine caves The first specimen was captured in 1998 (ERDMANN et al., second dorsal and anal fins. The day after, another individual On the 4th June in 2006, three individuals, including ID 3 recorded at Buol-2 in a cave and at Buol-3 in a crack, was temperature of 12.4 °C, the coolest one registered during all recorded alone on a Manado site (Manado-7). within a two week period, in several caves distributed in a 8 Water temperature can increase or drop for more than 5 large rocks or along vertical wall in daytime. Some of these and Marine Sciences of the Sam Ratulangi University at Science, 102: 486–490. along steep slopes between 100 m and 200 m depth during 1998). According to a genetic study, the specimen differed (ID 2) and an unidentified one (UN1) were found in the same which was observed on the 31th May at Buol-2 and two new encountered again (Table 2: Buol-5) near a vertical wall at 145 surveys. Field surveys were conducted from the 19th to 30th May kilometers wide area (FRICKE et al., 1991). °C in a very short period of time (about 30 min according to our individuals were encountered above 200 m deep, in an Manado, Indonesia, for their generous supports to conduct BRUTON, M. N., CABRAL, A. J. P. and FRICKE, H. 1992. First daytime and come out and drift near the ocean floor for from L. chalumnae and a new species was erected (POUYAUD cave (Buol-1). The unidentified one immediately hid deeper in individuals, were observed in a vertical crack which was m depth (Table 2: E6). After a while, this individual (ID 3) The field survey in 2009 was carried out off Manado and 2015 off Lolak Island and from the 30th October to the 16th When individuals of L. menadoensis were encountered, recordings). The coelacanths however often stayed there environment that remains slightly bright. Latimeria expeditions. Also we express our thanks to Dr. SUHARSONO, Dr. capture of a coelacanth, Latimeria chalumnae (Pisces, hunting prey at night (FRICKE et al., 1991). However, since the et al., 1999). In 1999 two individuals were observed by the cave, where the ROV could not access, and only its lateral several hundred meters far from Buol-2 (Buol-3) for 2 hours moved upward about 18 m and stayed behind a large rock (Fig. around Talise and Bangka islands off the northern coast of November 2015 off Lolak and Bitung Islands (Table 1; R13 some of them stayed stationary at same place but some without moving away or significantly modifying their menadoensis seems to be less sensitive to the daylight than L. Zainal ARIFIN, Dr. DIRHAMSYAH and colleagues of the Indonesian Latimeriidae), off Mozambique. South African Journal of record of juvenile or small (below 80 cm long) individuals is submersible in a location 360 km away from Manado (FRICKE side of the body was recorded. The individual (ID 2) was and 14 minutes (Fig. 2C; Table 2: E4, ID 3, 4, 5). During this 3). The observation time was the longest of all surveys with Sulawesi Island from the 12th September to the 9th October and R14). Three new individuals (Table 2: ID 26–28) were individuals such as ID 6 on the 12th December 2006 (E5) and behavior. These water currents of different temperatures are chalumnae. Institute of Science for their advice and administrative work to Science, 88: 225–227. virtually absent, the growth, development and breeding et al., 2000). Today 8 individuals were officially caught in the observed for one minute before it hid inside the cave (Table 2: long period of time all individuals stayed near the rock wall four hours and 16 minutes of recording. The temperature (Table 1: R8). On the 24th September 2009, six individuals recognized at different places off Lolak Island (Lolak 1-4). All ID 24 on the 4th May 2012 (E22) swam away. It seemed to caused by the movement of the marine thermocline. During These first observations are potentially crucial since the obtain our research permit. Other thanks go to Mr. OPO, Mr. CUPELLO, C., BRITO, P. M., HERBIN, M., MEUNIER, F. J., biology of L. chalumnae remain poorly known. Our knowledge Indonesian seas. However, the details of their habitat have not E2). When a new individual could be identified by the white always keeping their head against the water flow. The water changed up and down several times (Fig. 4). Its difference (Table 2: E11, ID 9, 10, 11, 12, UN 2, 3) were found under an individuals were alone. ID 26 was observed in May and once avoid and escape from the brightness of the light or the sounds our surveys we observed in some area that as the thermocline habitats of the Indonesian coelacanths could be possibly not as REFRY and colleagues at Murex Dive Resort, and DAUD, AL, J ANVIER, P., DUTEL, H. and CLÉMENT, G. 2015. 54 IWATA et al. Field surveys on Latimeria menadoensis using ROV from 2005 to 2015 55 been described. The aim of this study is to clear their habitat Allometric growth in the extant coelacanth lung during capture of a coelacanth off Madagascar. South African and distribution. ontogenetic development. Nature Communications, 6: Journal of Science 92: 150–151. 8222. DOI: 10.1038/ncomms9222 POUYAUD, L., WIRJOATMODJO, S., RACHMATIKA, I., TJAKRAWIDJAJA, DE VOS, L. and OYUGI, D. 2002. First capture of a coelacanth, A., HADIATY, R. and HADIE, W. 1999. A new species of METHODS Latimeria chalumnae SMITH, 1939 (Pisces, Latimeriidae), coelacanth. Genetic and morphologic proof. C. R. off Kenya. South African Journal of Science, 98: 345–347. Academy of Science, 322: 261–267. Two remotely operated vehicles (ROVs) (Kowa; ERDMANN, M. V., CALDWELL, R. L. and MOOSA, M. K. 1998. SMITH, J. L. B. 1939. The living coelacanth fish from South VEGA300) were used for the surveys. The first ROV was Indonesian ‘king of the sea’ discovered. Nature, 395: 335. Africa. Nature, 143: 748–750. replaced with the second one in 2007. Our ROVs are able to overhang off Talise Island between 144 and 150 m deep. There again in October 2015 on a steep slope (Table 2: E25, E27). FRICKE, H., REINICKE, O., HOFER, H. and NACHTIGALL, W. SMITH, J. L. B. 1953. The second coelacanth. Nature, 171: dive up to 300 m depth. These had two vertical, two horizontal was almost no water current in this area. All individuals were The coelacanth ID 28 was found on the 1st November along 1987. Locomotion of the coelacanth Latimeria chalumnae 99–101. and two right-left propellers and these were controlled from a close to each other under the same overhang and most of them one of the large rocks scattered on a gentle slope in a bay of in its natural environment. Nature, 329: 331–333. SMITH, C. L., RAND, C. S., SCHAEFFER, B. and ATZ, J. W. 1975. boat on surface through a 400 m long tether cable. The stayed still with their head facing downward. Four days later, the Lolak Island by 125 m depth. FRICKE, H., HISSMANN, K., SCHAUER, J., REINICKE, O., KASANG, Latimeria, the living coelacanth, is ovoviviparous. Science, underwater operations were visualized on a screen with ID 9 was observed again, alone on a steep slope at 172.9 m The encountered Indonesian coelacanths during these L. and PLANTE, R. 1991. Habitat and population size of 190: 1105–1106. information including directions of ROVs, depth, water depth (Table 2: E12), which was located within several dozen surveys were in caves, alongside large rocks, under overhangs coelacanth Latimeria chalumnae at Grand Comoro. VENTER, P., TIMM, P., GUNN, G., LE ROUX, E., SERFONTEIN, E., temperature, date and time. All data were directly recorded on meters far from Talise-2 and 30 m deeper. or on steep slopes (Table 2; Fig. 2). Environmental Biology of Fishes, 32: 287–300. SMITH, P., SMITH, E., BENSCH, M., HARDING, D. and video-tapes on the boat. Since the 2007 survey, water Four new coelacanths were recorded from the 29th Most individuals were found under overhangs or FRICKE, H., HISSMANN, K., SCHAUER, J., ERDMANN, M., HEEMSTRA, P. 2000. Discovery of a viable population of temperature and depth were independently recorded with more September to the 6th October 2009 in Talise and Banggka alongside large rocks. Some individuals were encountered not MOOSA, M. K. and PLANTE, R. 2000. Biogeography of the coelacanths (Latimeria chalumnae SMITH, 1939) at accuracy by external measuring memories (Alec Electronics, Islands and off Manado (Table 2: E13 and 14, ID 13, 14, 15). to hide in any shade and stayed just on rocky slopes. In Indonesian coelacanth. Nature, 403: 38. Sodwana Bay, South Africa. South African Journal of later named JFE Alec; MDS-MKV/T, MDS-MKV/D). In ID 15 is the only juvenile coelacanth filmed so far. It was in a overhangs and cracks, all individuals had their ventral side HEEMSTRA, P. C., FREEMAN, A. L., WONG, H. Y., HENSLEY, D. Science, 96: 567–568. addition two laser beam irradiators, that provide line lasers of small crack between 164.6 and 170.9 m deep in a temperature alongside and close to a rock wall, but without touching it. A. and RABESANDRATANA, H. D. 1996. First authentic 20-cm distance, were attached to the second ROV and have between 14.5 and 15 °C. Estimated total lengths are shown in Table 3. Most been used to register the size of encountered animals. The The shallowest record of a coelacanth was at 115.6 m individuals are more than 1 m in total length. ID 15 is a Fig. 3. Water temperature and depth during observations of coelacanths in each locality. The orange box shows depth. The top and Fig. 4. Change of the water temperature at Buol-5 off Buol of Sulawesi Island. surveys were conducted off northern parts of Sulawesi Island depth off the southern part of Manado (Table 2: E16, juvenile individual of 31.5 cm and ID 23 is 90 cm in total bottom of the box mean a minimum depth and maximum depth of coelacanths stay. The blue bar shows a maximum and and along the Biak Island located in northwestern New Guinea Manado-3) on the 9th October in 2009. ID 16 (Table 2) was length. No individual beyond 140 cm has been recorded in minimum water temperatures. The open circle shows an average of temperature during observation. All individuals of L. Island, Indonesia. The areas and time periods of each survey found at the end of the morning, alone alongside a large rock Indonesia so far. menadoensis were observed in depth between 115.6 and 218.9 m and water temperature between 12.4 and 21.5 ºC. restricted as those of the African coelacanths, and so a broader and other staffs for our work on the research boat. Dr. BATUNA are shown in Table 1 and Fig. 1. on the edge of a shelf. The water temperature was about 20.0 distribution of the Indonesian coelacanth could be expected. and the Murex staffs supported our daily life during the ºC. Occurrences of coelacanths along the eastern coast of the expeditions. We would like to express our sincere thanks to From the 6th to the 16th November in 2010, the field of the propellers of the ROV. level changed, the water flow was reversed. Coelacanths African continent were recorded in South Africa (SMITH, colleagues at Aquamarine Fukushima, Japan, for their RESULTS survey was conducted around Biak Island (Table 1: R9). Adult DISCUSSION Temperature and depth data for each Indonesian always kept their head against the water flow, like other 1939), Comoros (SMITH, 1953), Madagascar (HEEMSTRA et al., understanding and supports. We would like to thank Dr. Gaël coelacanths were found at two different sites (Table 2: coelacanth record are shown in Fig. 3. Though an average of observed actinopterygian fishes did. It means that as the 1996), Kenya (DE VOS and OYUGI, 2002), Tanzania (BENNO et CLÉMENT of Muséum national d’Histoire naturelle and Dr. The field survey of the Indonesian coelacanth, Latimeria E17–19). Two individuals (ID 17 and 18) were found under an During the field surveys by underwater ROV recording water temperature on the ocean surface was about 30 °C, the direction of the water flow changed, the swimming direction of al., 2006), and Mozambique (BRUTON et al., 1992). In Camila CUPELLO of Universidade do Estado do Rio de Janeiro menadoensis, was conducted 14 times from 2005 to 2015 with overhang between 212.5 and 218.9 m deep (Table 2: Biak-1) for Indonesian coelacanths, Latimeria menadoensis, from 2005 temperature often dropped into about 11 °C at 300 m water depth. each coelacanth individual changed accordingly. Indonesia, coelacanths have only been found off the northern as a referee for their critical reading of the manuscript and their 1173 underwater operations around Sulawesi and Biak islands on the 11th November. Three other ones (ID 19, 20, 21) were to 2015, 30 different individuals were identified and three The temperatures changed widely even though the changes of On the 24th September 2009 (Table 2: E11) six coast of the Sulawesi Island and the southern coast of the Biak comments. in Indonesia (Fig. 1, Table 1). No coelacanth was encountered registered under an overhang between 193.2 and 195.9 m deep others were unidentified. Among those, six individuals were depth were small at several places. All Indonesian coelacanths, individuals were recognized under an overhang (Talise-2). The Island in northern New Guinea. Unfortunately one of our authors, Dr. Djoko Hadi KUNARSO during the first surveys in 2005 although 452 underwater (Table 2: Biak-2) on the 13th November. Two days later, observed more than twice: ID 3 at Buol-2, 3, 5, 6 (Table 2: E3, L. menadoensis, were encountered during day time at a depth all had about the same size. There was almost no water flow, Field surveys of the Indonesian coelacanths should be has passed away during the process of this contribution. We operations were conducted around Manado-tua Island and spots pattern on its body, an ID number was allocated. The temperature changed up and down by five degrees during the between the maximum and the minimum was 6.7 °C, which Biak-2 was observed again and only ID 20 stayed there. E4, E6 and E7); ID 8 at Talise-1 (E9 and E10), ID 9 at Talise 2 between 115.6 and 218.9 m and a temperature between 12.4 and each individual was head downward. They were close to continued in all area of Indonesia and expanded to all pray for the response of his soul. other close Indonesian islands from the 17th to 30th of April individual which could not be distinguished was allocated an observation due to water flow reversals. When the water flow was the largest range of all surveys. Two new coelacanths were recorded off Manado in and 3 (E11 and E12), ID 20 at Biak-2 (E18 and E19), ID 24 at and 21.5 ºC. Indonesian coelacanths have exceptionally been each other but no direct contacts or interactions between Southeast Asia in order to increase our knowledge of the This study was supported in part by funding from (Table 1, R1). The second field survey was held along the UN (unknown) + number. was reversed some times, all individuals always changed their On the next day, on the 15th December, the ID 3 was December 2010 (Table 1: R10; Table 2: E 20 and 21; ID 22 Manado-6 and 8 (E22 and E24), ID 26 at Lolak-1 and 3 (E25 recorded in an environment with a temperature exceeding 20 coelacanths were observed. No evidence of a social behavior distribution of this rare species and to define, in a close future, Interdisciplinary Collaborative Research program of Atmosphere northern coast of Sulawesi from the 6th to 19th May in 2006, In the end of the morning of the same day, a third directions against the water flow. observed again under an overhang (Table 2: E7, Buol-6) near and 23). and E27). ID 8 was observed on the 14th September 2009 and ºC, regardless of the depth. The African coelacanth, L. has been recognized in the Comorian populations of the most relevant conservation policies of the different known and Ocean Research Institute, the University of Tokyo. but once again despite of 107 underwater operations no individual (Table 2: E3, ID 3) was found in another cave During the field survey carried on December of the same other sites where it was previously found at Buol-2 , Buol-3 On the 4th May 2012, an individual (Table 2: E22, ID 24) was encountered again at the same place (Talise-1) the day chalumnae, was observed by submarine vehicles in steep coelacanths (FRICKE et al., 1991). populations. coelacanth was recorded. (Buol-2), larger and located 20 m deeper (Buol-2) than the first year, in the same Buol area (Table 1: R4) the sixth individual and Buol-5. Its posture was upside down with its ventral side was found on a steep slope at 169.4 m depth off Manado after. ID 24 was observed on the 4th May 2012 and then 13 slopes between 150 m and 253 m (FRICKE et al., 1991). Divers According to the observations of the African coelacanths, INTRODUCTION of the reproduction biology of the coelacanths is almost The first individual of Indonesian coelacanth (Table 2, ID cave (Buol-1). After 29 minutes of recording, this individual (Table 2: E5, ID 6) was recognized at 14:34 on the 12th facing the ceiling of the overhang. Such posture is also known (Table 2: Manado-6). It swam up and down 5.4 m between months later, on the 9th June 2013 at the same locations of recorded the presence of this species at 104 m depth at the species L. chalumnae is nocturnal and hides in cave in REFERENCES restricted to an ovoviviparity strategy since some caught large 1) was recorded during the third field survey, in the morning of hid inside the cave. Although the water temperature increased December alongside a large rock. This individual stayed in the in coelacanths from the Comorian Archipelago (FRICKE et al., 168.1 and 173.5 m deep for 17 minutes. Thirteen months later Manado area within several kilometers. Sodwana Bay, South Africa (VENTER et al., 2000). Latimeria daytime but comes out at night, supposedly in search of food ACKNOWLEDGMENTS The first living coelacanth, Latimeria chalumnae, was females carried unfertilized eggs or embryos in their oviduct the 30th May 2006 (Table 1, R3). The coelacanth was of more than five degrees, from 14.8 to 20.4 ºC, it stayed at the beginning of the observation and then moved ahead to deeper 1987). ID 3 was found totally four times in this area in two (the 9th June 2003), this individual (ID 24) was observed again All individuals observed more than twice were chalumnae seem to prefer a temperature range of 15–20 °C, (FRICKE et al., 1991). The living geomorphologic environment BENNO, B., VERHEIJI, E., STAPLEY, J., RUMISHA, C., NGATUNGA, discovered in South Africa in the Indian Ocean in 1938 (SMITH et al., 1975; CUPELLO et al., 2015). observed in a cave of 165 m depth (Table 2, Boul-1) for 10 same place, not moving away. Despite it disappeared from the slowly. Eventually it reached 12 m deeper during one hour and surveys during the seven months and seems to live in this area. near a cliff at 152.3 m depth (Table 2: E24, Manado-8), where encountered in the same area, at locations close from each where they tend to choose their daytime habitats (FRICKE et al., of L. menadoensis is similar to that of L. chalumnae, with We are grateful to Dr. Teruya UYENO for his valuable B., ABDALLAH, A. and KALOMBO, H. 2006. Coelacanth (SMITH, 1939). The first observation of the living coelacanth The Indonesian coelacanth, L. menadoensis, was first minutes (Table 2, E1). The adult individual was in stationary camera frame we kept setting the ROV there in front of the 42 minutes. During the 2007 survey (Table 1: R5) a coelacanth (Table located on the same slope of Manado-6. other within hundreds of meters. However, in the Comorian 1991). The suitable temperature for L. menadoensis seems to be rocky steep slopes with undermarine caves or overhangs. advice, guidance and encouragement throughout the present (Latimeria chalumnae SMITH, 1939) discoveries and habitats using submersibles was reported in Comoros (FRICKE discovered in Manado, Sulawesi Island, Indonesia, in 1997. position in a cave (Buol-1) slowly moving its pectoral, pelvic, cave until the next morning, however it did not appear again. On the 14th December, the individual ID 3 previously 2: ID 7) was recorded off Manado by 195 m depth, with a Another new coelacanth (Table 2: E23, ID 25) was Archipelago, same individuals of L. chalumnae were observed, almost the same as L. chalumnae. However, Latimeria menadoensis was also observed alongside study. Many thanks go to colleagues of the Faculty of Fisheries conservation in Tanzania. South African Journal of et al., 1987). Latimeria chalumnae hide in submarine caves The first specimen was captured in 1998 (ERDMANN et al., second dorsal and anal fins. The day after, another individual On the 4th June in 2006, three individuals, including ID 3 recorded at Buol-2 in a cave and at Buol-3 in a crack, was temperature of 12.4 °C, the coolest one registered during all recorded alone on a Manado site (Manado-7). within a two week period, in several caves distributed in a 8 Water temperature can increase or drop for more than 5 large rocks or along vertical wall in daytime. Some of these and Marine Sciences of the Sam Ratulangi University at Science, 102: 486–490. along steep slopes between 100 m and 200 m depth during 1998). According to a genetic study, the specimen differed (ID 2) and an unidentified one (UN1) were found in the same which was observed on the 31th May at Buol-2 and two new encountered again (Table 2: Buol-5) near a vertical wall at 145 surveys. Field surveys were conducted from the 19th to 30th May kilometers wide area (FRICKE et al., 1991). °C in a very short period of time (about 30 min according to our individuals were encountered above 200 m deep, in an Manado, Indonesia, for their generous supports to conduct BRUTON, M. N., CABRAL, A. J. P. and FRICKE, H. 1992. First daytime and come out and drift near the ocean floor for from L. chalumnae and a new species was erected (POUYAUD cave (Buol-1). The unidentified one immediately hid deeper in individuals, were observed in a vertical crack which was m depth (Table 2: E6). After a while, this individual (ID 3) The field survey in 2009 was carried out off Manado and 2015 off Lolak Island and from the 30th October to the 16th When individuals of L. menadoensis were encountered, recordings). The coelacanths however often stayed there environment that remains slightly bright. Latimeria expeditions. Also we express our thanks to Dr. SUHARSONO, Dr. capture of a coelacanth, Latimeria chalumnae (Pisces, hunting prey at night (FRICKE et al., 1991). However, since the et al., 1999). In 1999 two individuals were observed by the cave, where the ROV could not access, and only its lateral several hundred meters far from Buol-2 (Buol-3) for 2 hours moved upward about 18 m and stayed behind a large rock (Fig. around Talise and Bangka islands off the northern coast of November 2015 off Lolak and Bitung Islands (Table 1; R13 some of them stayed stationary at same place but some without moving away or significantly modifying their menadoensis seems to be less sensitive to the daylight than L. Zainal ARIFIN, Dr. DIRHAMSYAH and colleagues of the Indonesian Latimeriidae), off Mozambique. South African Journal of record of juvenile or small (below 80 cm long) individuals is submersible in a location 360 km away from Manado (FRICKE side of the body was recorded. The individual (ID 2) was and 14 minutes (Fig. 2C; Table 2: E4, ID 3, 4, 5). During this 3). The observation time was the longest of all surveys with Sulawesi Island from the 12th September to the 9th October and R14). Three new individuals (Table 2: ID 26–28) were individuals such as ID 6 on the 12th December 2006 (E5) and behavior. These water currents of different temperatures are chalumnae. Institute of Science for their advice and administrative work to Science, 88: 225–227. virtually absent, the growth, development and breeding et al., 2000). Today 8 individuals were officially caught in the observed for one minute before it hid inside the cave (Table 2: long period of time all individuals stayed near the rock wall four hours and 16 minutes of recording. The temperature (Table 1: R8). On the 24th September 2009, six individuals recognized at different places off Lolak Island (Lolak 1-4). All ID 24 on the 4th May 2012 (E22) swam away. It seemed to caused by the movement of the marine thermocline. During These first observations are potentially crucial since the obtain our research permit. Other thanks go to Mr. OPO, Mr. CUPELLO, C., BRITO, P. M., HERBIN, M., MEUNIER, F. J., biology of L. chalumnae remain poorly known. Our knowledge Indonesian seas. However, the details of their habitat have not E2). When a new individual could be identified by the white always keeping their head against the water flow. The water changed up and down several times (Fig. 4). Its difference (Table 2: E11, ID 9, 10, 11, 12, UN 2, 3) were found under an individuals were alone. ID 26 was observed in May and once avoid and escape from the brightness of the light or the sounds our surveys we observed in some area that as the thermocline habitats of the Indonesian coelacanths could be possibly not as REFRY and colleagues at Murex Dive Resort, and DAUD, AL, J ANVIER, P., DUTEL, H. and CLÉMENT, G. 2015. 56 IWATA et al. Bull. Kitakyushu Mus. Nat. Hist. Hum. Hist., Ser. A, 17: 57–65, March 31, 2019 been described. The aim of this study is to clear their habitat Allometric growth in the extant coelacanth lung during capture of a coelacanth off Madagascar. South African Observation of the first juvenile Indonesian coelacanth, Latimeria menadoensis from of juveniles of coelacanth is discussed. In addition, the detail et al., 2019) was found on the southern coast of Manado Bay the pectoral fin and 26 rays of the pelvic fin. Allometric growth in the extant coelacanth lung during Indonesian coelacanth, Latimeria menadoensis using and distribution. measurements and counts of the juvenile based on video of Sulawesi, Indonesia, on the 6th October in 2009 (Fig. 1). ontogenetic development. Nature Communications, 6: Journal of Science 92: 150–151. Indonesian waters with a comparison to embryos of ontogenetic development. Nature Communications, 6: remotely operated vehicles from 2005 to 2015. Bulletin of 8222. DOI: 10.1038/ncomms9222 POUYAUD, L., WIRJOATMODJO, S., RACHMATIKA, I., TJAKRAWIDJAJA, Latimeria chalumnae footages are made and compared to embryos of L. chalumnae, The individual stayed in a narrow overhang at 164.6 m depth 8222. DOI 10.1038/ncomms9222 Kitakyushu Museum of Natural History and Human DE VOS, L. and OYUGI, D. 2002. First capture of a coelacanth, A., HADIATY, R. and HADIE, W. 1999. A new species of which are from a large female (CCC no. 162) caught off (Fig. 4A). The overhang was long and progressively became DISCUSSION FOREY, P. L. 1998. History of the Coelacanth Fishes. Chapman History, Series A, Natural History, 17: 49–56. METHODS Latimeria chalumnae SMITH, 1939 (Pisces, Latimeriidae), coelacanth. Genetic and morphologic proof. C. R. Masamitsu IWATA 1, Yoshitaka YABUMOTO2, Toshiro SARUWATARI3,4, Shinya YAMAUCHI1, Kenichi FUJII1, Mozambique water in 1991 (BRUTON et al., 1992). This deeper (Fig. 4B). The height of the overhang seemed narrower and Hall, London, xiii+419 pp. NULENS, R., SCOTT, L. and HERBIN, M. 2011. An updated off Kenya. South African Journal of Science, 98: 345–347. Academy of Science, 322: 261–267. 1 1 5 5 6 morphological data, based on extant coelacanths, should than the observed juvenile standard length (Fig. 3). The Latimeria chalumnae is known to be ovoviviparous (SMITH FRICKE, H., REINICKE, O., HOFER, H. and NACHTIGALL, W. 1987. inventory of all known specimens of the coelacanth Rintaro ISHII , Toshiaki MORI , Frensly D. HUKOM , DIRHAMSYAH ,Teguh PERISTIWADY , Augy Two remotely operated vehicles (ROVs) (Kowa; ERDMANN, M. V., CALDWELL, R. L. and MOOSA, M. K. 1998. SMITH, J. L. B. 1939. The living coelacanth fish from South provide invaluable information to the study of fossil and extant juvenile was observed for 17 minutes from 11:34. It stayed et al., 1975). To date, no females of L. menadoensis with Locomotion of the coelacanth Latimeria chalumnae in its Latimeria spp. South African Institute for Aquatic 7 8,9 8,9 8,9 1 VEGA300) were used for the surveys. The first ROV was Indonesian ‘king of the sea’ discovered. Nature, 395: 335. Africa. Nature, 143: 748–750. SYAHAILATUA , Kawilarang W. A. MASENGI , Ixchel F. MANDAGI , Fransisco PANGALILA and Yoshitaka ABE coelacanths. below the overhang and moved slowly, between the depth of matured eggs or embryos has been captured. Reproductive natural environment. Nature, 329: 331–333. Biodiversity, 3: 1–52. replaced with the second one in 2007. Our ROVs are able to overhang off Talise Island between 144 and 150 m deep. There again in October 2015 on a steep slope (Table 2: E25, E27). FRICKE, H., REINICKE, O., HOFER, H. and NACHTIGALL, W. SMITH, J. L. B. 1953. The second coelacanth. Nature, 171: 164.6 and 170.9 m. The water current was gentle and the water organs of L. chalumnae and L. menadoensis are almost the FRICKE, H. and HISSMANN, K. 1992. Locomotion, fin SECRETARIAT AT THE J. L. B. SMITH INSTITUTE OF ICHTHYOLOGY. dive up to 300 m depth. These had two vertical, two horizontal was almost no water current in this area. All individuals were The coelacanth ID 28 was found on the 1st November along 1987. Locomotion of the coelacanth Latimeria chalumnae 99–101. 1Aquamarine Fukushima, Marine Science Museum, 50 Tatsumi-cho, Onahama, Iwaki, Fukushima, 971-8101, Japan temperature during the observation was stable, from 14.5 ºC to same anatomically (our observation) and, consequently, L. coordination and body form of the living coelacanth, 1993. Coelacanth conservation council newsletter no.4. and two right-left propellers and these were controlled from a close to each other under the same overhang and most of them one of the large rocks scattered on a gentle slope in a bay of in its natural environment. Nature, 329: 331–333. SMITH, C. L., RAND, C. S., SCHAEFFER, B. and ATZ, J. W. 1975. 2Kitakyushu Museum of Natural History and Human History, 2-4-1 Higashida, Yahata Higashi-ku, MATERIALS AND METHODS 15.0 ºC, the average temperature was 14.8 ºC. menadoensis is here considered to be ovoviviparous as well. A Latimeria chalumnae. Environmental Biology of Fishes, Environmental Biology of Fishes, 36: 395–406. boat on surface through a 400 m long tether cable. The stayed still with their head facing downward. Four days later, the Lolak Island by 125 m depth. FRICKE, H., HISSMANN, K., SCHAUER, J., REINICKE, O., KASANG, Latimeria, the living coelacanth, is ovoviviparous. Science, Kitakyushu, Fukuoka, 805-0071, Japan Coelacanth has six lobed fins (two pectoral, two pelvic, large female of L. chalumnae that held juveniles from 30.8 cm 34: 329–356. SMITH, C. L., RAND, C. S., SCHAEFFER, B. and ATZ, J. W. 1975. 3 underwater operations were visualized on a screen with ID 9 was observed again, alone on a steep slope at 172.9 m The encountered Indonesian coelacanths during these L. and PLANTE, R. 1991. Habitat and population size of 190: 1105–1106. Atmosphere and Ocean Research Institute, The University of Tokyo, 5-1-5 Kashiwanoha, Kashiwa, Materials are the video footage of the juvenile of one dorsal, and one anal fins). It exhibited a unique swimming to 35.8 cm inside its body was caught off Mozambique water FRICKE, H., HISSMANN, K., FROESE, R., SCHAUER, J., PLANTE, Latimeria, the living coelacanth, is ovoviviparous. Science, Chiba, 277-8564, Japan information including directions of ROVs, depth, water depth (Table 2: E12), which was located within several dozen surveys were in caves, alongside large rocks, under overhangs coelacanth Latimeria chalumnae at Grand Comoro. VENTER, P., TIMM, P., GUNN, G., LE ROUX, E., SERFONTEIN, E., Indonesian coelacanth L. menadoensis (Fig. 1) and photos of style by paddling and twisting the lobed fins (FRICKE and in 1991 (BRUTON et al., 1992). Latimeria chalumnae embryo R. and FRICKE, S. 2011. The populations biology of the : 1105–1106. 4Seikei Education and Research Center for Sustainable Development, 3-3-1 Kichijoji-Kitamachi, 190 temperature, date and time. All data were directly recorded on meters far from Talise-2 and 30 m deeper. or on steep slopes (Table 2; Fig. 2). Environmental Biology of Fishes, 32: 287–300. SMITH, P., SMITH, E., BENSCH, M., HARDING, D. and Musashino-shi, Tokyo, 180-8633, Japan the embryos of the African coelacanth L. chalumnae, which HISSMANN, 1992). The swimming of the juvenile was similar to CCC no. 29.5 found inside CCC no. 29 is 32.3 cm TL with a living coelacanth studied over 21 years. Marine Biology, UYENO, T. 1991. Observations on locomotion and feeding of video-tapes on the boat. Since the 2007 survey, water Four new coelacanths were recorded from the 29th Most individuals were found under overhangs or FRICKE, H., HISSMANN, K., SCHAUER, J., ERDMANN, M., HEEMSTRA, P. 2000. Discovery of a viable population of 5Research Center for Oceanography, Indonesian Institute of Science, Jakarta, Indonesia are CCC no. 162.16 housed in the J. L. B SMITH Institution, those of adults. During the observation of the juvenile, one yolk sac, and CCC no, 162.21 is a late embryo without a yolk 158: 1511–1522. DOI 10.1007/s00227-011-1667-x released coelacanths, Latimeria chalumnae. Environmen- temperature and depth were independently recorded with more September to the 6th October 2009 in Talise and Banggka alongside large rocks. Some individuals were encountered not MOOSA, M. K. and PLANTE, R. 2000. Biogeography of the coelacanths (Latimeria chalumnae SMITH, 1939) at 6Division of Marine Resources Research, Indonesian Institute of Sciences, Jakarta, Indonesia South Africa (Fig. 2A) (SECRETARIAT AT THE J. L. B. SMITH cycle of paddling was five seconds and the juvenile swam sac of 35.6 cm TL found inside CCC no. 126 captured in 1991 HOLDEN, C. 2009. Infant fossil. Science, 326: 1327. tal Biology of Fishes, 32: 267–273. 7 accuracy by external measuring memories (Alec Electronics, Islands and off Manado (Table 2: E13 and 14, ID 13, 14, 15). to hide in any shade and stayed just on rocky slopes. In Indonesian coelacanth. Nature, 403: 38. Sodwana Bay, South Africa. South African Journal of Centre for Deep Sea Research,Indonesian Institute of Sciences, Jakarta, Indonesia INSTITUTE OF ICHTHYOLOGY, 1993, fig. 3), CCC no. 162.12 forward and backward. The paddling pattern and the cycle (CUPELLO et al., 2015). CCC no. 29.5 is housed in Muséum HENSEL, K. and BALON, K. 2001. The sensory canal system of WOURMS, J. P., ATZ, J. W. and STRIBLING, M. D. 1991, 8 later named JFE Alec; MDS-MKV/T, MDS-MKV/D). In ID 15 is the only juvenile coelacanth filmed so far. It was in a overhangs and cracks, all individuals had their ventral side HEEMSTRA, P. C., FREEMAN, A. L., WONG, H. Y., HENSLEY, D. Science, 96: 567–568. International Coelacanth Research Center and Marine Museum, Sam Ratulangi University, Manado 95115, Indonesia (Fig. 2B) and CCC no. 162.22 (Fig. 2C) housed in University were the same both the forward and backward swimming. national d’Histoire naturelle, Paris (France) and CCC no. the living coelacanth, Latimeria chalumnae: a new Viviparity and meternal-embryonic relationship in the 9Faculty of Fisheries and Marine Science, Sam Ratulangi University, Manado 95115, Indonesia addition two laser beam irradiators, that provide line lasers of small crack between 164.6 and 170.9 m deep in a temperature alongside and close to a rock wall, but without touching it. A. and RABESANDRATANA, H. D. 1996. First authentic of Guelph (HENSEL and BALON, 2001, fig. 2a, b). CCC stands The dorso-posterior edge of the first dorsal fin, the dorsal 162.21 is housed in Zoologisches Staatsaammling, München instalment. Environmental Biology of Fishes, 61: 117–124. coelacanth Latimeria chalumnae. Environmental Biology 20-cm distance, were attached to the second ROV and have between 14.5 and 15 °C. Estimated total lengths are shown in Table 3. Most for the Coelacanth Conservation Council. The video footages edge of the dorsal lobe of the caudal fin (the third dorsal fin), (Germany). These specimens suggest that L. chalumnae IWATA, M., YABUMOTO, Y., SARUWATARI, T., YAMAUCHI, S., of Fishes, 32: 225–248. been used to register the size of encountered animals. The The shallowest record of a coelacanth was at 115.6 m individuals are more than 1 m in total length. ID 15 is a (Received August 24, 2018; accepted November 9, 2018) including this finding were taken by a remotely operated and the ventral edge of the ventral lobe of the caudal fin (the delivers juveniles of around 30 cm in total length. Since the FUJII, K., ISHII, R., MORI, T., HUKOM, F., DIRHAMSYAH, YABUMOTO, Y. 2008. A new Mesozoic coelacanth from Brazil surveys were conducted off northern parts of Sulawesi Island depth off the southern part of Manado (Table 2: E16, juvenile individual of 31.5 cm and ID 23 is 90 cm in total vehicle (ROV) (Kowa; HDTV VEGA-300) in the north of second anal fin) are white. The dorsal margin of the head is smaller individuals of L. chalumnae observed in their habitat PERISTIWADY, T., SYAHAILATUS, A., MASENGI, K. W. A., (, Actinistia). Paleontological Research, 12: and along the Biak Island located in northwestern New Guinea Manado-3) on the 9th October in 2009. ID 16 (Table 2) was length. No individual beyond 140 cm has been recorded in Sulawesi Island and Biak Island in Indonesia from 2005 to convex and angular, where seems to be the intracranial joint were from 50 to 60 cm long, the ecology of juveniles remain MANDAGI, I. F. and ABE, Y. 2019. Field Survey on the 329–343. Latimeria menadoensis Island, Indonesia. The areas and time periods of each survey found at the end of the morning, alone alongside a large rock Indonesia so far. restricted as those of the African coelacanths, and so a broader and other staffs for our work on the research boat. Dr. BATUNA ABSTRACT − The juvenile of Indonesian coelacanth, is here described for the first 2015 (IWATA et al., 2019). The video was recorded with two-line (Fig. 1B). Scales have many minute spines, which reflect the poorly known (FRICKE et al., 2011). According to CCC are shown in Table 1 and Fig. 1. on the edge of a shelf. The water temperature was about 20.0 distribution of the Indonesian coelacanth could be expected. and the Murex staffs supported our daily life during the time in detail with comparison to embryos of Latimeria chalumnae. The juvenile was found in free swimming at lasers placed 20 cm apart. These lines were applied by two light of the ROV. records, the size of captured small coelacanths are: 50 cm for ºC. Occurrences of coelacanths along the eastern coast of the expeditions. We would like to express our sincere thanks to 164.6 m depth off Manado, Indonesia on the 6th October in 2009. Because the total length of the juvenile is 31.5 laser beam irradiators attached to the ROV. The measurements CCC no. 84 caught in Comoros in 1973; 42.5 cm TL for CCC From the 6th to the 16th November in 2010, the field of the propellers of the ROV. level changed, the water flow was reversed. Coelacanths African continent were recorded in South Africa (SMITH, colleagues at Aquamarine Fukushima, Japan, for their cm, which is smaller than the embryos of L. chalumnae, it is speculated that not much time has passed from its of the juvenile were calculated based on the beams (Fig. 3; Comparative description of the juvenile of L. menadoensis no. 94 caught in Comoros in 1974; 60 cm TL for CCC no. 116 birth. The depth at which the juvenile was found is within the range of the depth where adult L. menadoensis RESULTS survey was conducted around Biak Island (Table 1: R9). Adult DISCUSSION Temperature and depth data for each Indonesian always kept their head against the water flow, like other 1939), Comoros (SMITH, 1953), Madagascar (HEEMSTRA et al., understanding and supports. We would like to thank Dr. Gaël Table 1) and the fin ray counts are made from the video and embryos of L. chalumnae caught in Comoros in 1979; and 62 cm for CCC no. 160 coelacanths were found at two different sites (Table 2: coelacanth record are shown in Fig. 3. Though an average of observed actinopterygian fishes did. It means that as the 1996), Kenya (DE VOS and OYUGI, 2002), Tanzania (BENNO et CLÉMENT of Muséum national d’Histoire naturelle and Dr. were observed, hidden in a narrow and long overhang where large predators could not enter. The juvenile has a footages (Fig. 1). Among the embryos, the total length of CCC The total length of the juvenile is 31.5 cm. The standard caught in Comoros in 1989 (NULENS et al., 2011). The field survey of the Indonesian coelacanth, Latimeria E17–19). Two individuals (ID 17 and 18) were found under an During the field surveys by underwater ROV recording water temperature on the ocean surface was about 30 °C, the direction of the water flow changed, the swimming direction of al., 2006), and Mozambique (BRUTON et al., 1992). In Camila CUPELLO of Universidade do Estado do Rio de Janeiro more slender body, smaller orbit, shorter and deeper posterior part of the body (caudal peduncle) between the no. 162.12, 162.16 and 162.22 are 33.3 cm, 34.8 cm and 34.4 length is 26.1 cm. It is smaller than the embryos of L. The underwater observation at 164.6 m depth off menadoensis, was conducted 14 times from 2005 to 2015 with overhang between 212.5 and 218.9 m deep (Table 2: Biak-1) for Indonesian coelacanths, Latimeria menadoensis, from 2005 temperature often dropped into about 11 °C at 300 m water depth. each coelacanth individual changed accordingly. Indonesia, coelacanths have only been found off the northern as a referee for their critical reading of the manuscript and their second dorsal and the anal fins and anterior ends of the dorsal and ventral lobes of the caudal fin (the third dorsal cm respectively (SECRETARIAT AT THE J. L. B. SMITH INSTITUTE chalumnae used here for comparison, which are 30.8–35.8 cm Manado, Indonesia on the 6th October in 2009 is the first 1173 underwater operations around Sulawesi and Biak islands on the 11th November. Three other ones (ID 19, 20, 21) were to 2015, 30 different individuals were identified and three The temperatures changed widely even though the changes of On the 24th September 2009 (Table 2: E11) six coast of the Sulawesi Island and the southern coast of the Biak comments. and second anal fins), longer dorsal and ventral lobes of the caudal fin (the third dorsal and second anal fins), OF ICHTHYOLOGY, 1993). The body part lengths and proportion (BRUTON et al., 1992) and it is smaller than the smallest account of a live juvenile of L. menadoensis in its natural in Indonesia (Fig. 1, Table 1). No coelacanth was encountered registered under an overhang between 193.2 and 195.9 m deep others were unidentified. Among those, six individuals were depth were small at several places. All Indonesian coelacanths, individuals were recognized under an overhang (Talise-2). The Island in northern New Guinea. Unfortunately one of our authors, Dr. Djoko Hadi KUNARSO broader peduncles of broader lobed fins, larger first dorsal fin and longer supplementary lobe of the caudal fin to the total length of the three embryos are calculated by specimen (CCC no. 94) of L. chalumnae that has been collected habitat. The juvenile is 31.5 cm in total length. The estimated during the first surveys in 2005 although 452 underwater (Table 2: Biak-2) on the 13th November. Two days later, observed more than twice: ID 3 at Buol-2, 3, 5, 6 (Table 2: E3, L. menadoensis, were encountered during day time at a depth all had about the same size. There was almost no water flow, Field surveys of the Indonesian coelacanths should be has passed away during the process of this contribution. We (caudal fin) than embryos of L. chalumnae. This indicates clear differences in the first ontogenetic stages of the measuring the figures (Table 1). In the present study, names of is 42.5 cm TL.. Drawings of the L. menadoensis juvenile and dorsal lobe of the caudal fin (the third dorsal fin) is shorter size at newborn of L. chalumnae is around 30 cm as mention operations were conducted around Manado-tua Island and spots pattern on its body, an ID number was allocated. The temperature changed up and down by five degrees during the between the maximum and the minimum was 6.7 °C, which Biak-2 was observed again and only ID 20 stayed there. E4, E6 and E7); ID 8 at Talise-1 (E9 and E10), ID 9 at Talise 2 between 115.6 and 218.9 m and a temperature between 12.4 and each individual was head downward. They were close to continued in all area of Indonesia and expanded to all pray for the response of his soul. two species, although adults have almost the same morphological features. Latimeria menadoensis appears to fins follow that of FOREY (1998) which differ from UYENO L. chalumnae embryos (CCC no. 162.16) are shown in Fig. 5. than that of L. chalumnae (Fig. 5). Positions of fins are almost earlier. This suggests that not much time has passed since the other close Indonesian islands from the 17th to 30th of April individual which could not be distinguished was allocated an observation due to water flow reversals. When the water flow was the largest range of all surveys. Two new coelacanths were recorded off Manado in and 3 (E11 and E12), ID 20 at Biak-2 (E18 and E19), ID 24 at and 21.5 ºC. Indonesian coelacanths have exceptionally been each other but no direct contacts or interactions between Southeast Asia in order to increase our knowledge of the This study was supported in part by funding from reproduce in a rather confined area, because both the juvenile and adults have been found within the same area (1991). Dorsal lobe of caudal fin, ventral lobe of caudal fin and The body of the Indonesian coelacanth juvenile is slender than the same in both species, except for the dorsal lobe of the observed juvenile L. menadoensis was born. About 30 adult (Table 1, R1). The second field survey was held along the UN (unknown) + number. was reversed some times, all individuals always changed their On the next day, on the 15th December, the ID 3 was December 2010 (Table 1: R10; Table 2: E 20 and 21; ID 22 Manado-6 and 8 (E22 and E24), ID 26 at Lolak-1 and 3 (E25 recorded in an environment with a temperature exceeding 20 coelacanths were observed. No evidence of a social behavior distribution of this rare species and to define, in a close future, Interdisciplinary Collaborative Research program of Atmosphere inside of Manado Bay. supplementary lobe of caudal fin in FOREY (1998) are 3rd that of the L. chalumnae embryos (Table 1). The body depth is caudal fin (the third dorsal fin). The anterior end of the dorsal coelacanths of about 1 m long were found at the depth between ACKNOWLEDGEMENTS and her comments. This study was supported in part by funding northern coast of Sulawesi from the 6th to 19th May in 2006, In the end of the morning of the same day, a third directions against the water flow. observed again under an overhang (Table 2: E7, Buol-6) near and 23). and E27). ID 8 was observed on the 14th September 2009 and ºC, regardless of the depth. The African coelacanth, L. has been recognized in the Comorian populations of the most relevant conservation policies of the different known and Ocean Research Institute, the University of Tokyo. dorsal fin, 2nd anal fin and caudal fin of UYENO (1991), 6.8 cm, 26 % of the standard length in the L. menadoensis lobe of the caudal fin base is located more anteriorly than that 115.6 m and 218.9 m in Indonesia (IWATA et al. 2019). The fins, longer fins, larger principal caudal fin (the third dorsal menadoensis and L. chalumnae are scarce when compared from Interdisciplinary Collaborative Research Program of but once again despite of 107 underwater operations no individual (Table 2: E3, ID 3) was found in another cave During the field survey carried on December of the same other sites where it was previously found at Buol-2 , Buol-3 On the 4th May 2012, an individual (Table 2: E22, ID 24) was encountered again at the same place (Talise-1) the day chalumnae, was observed by submarine vehicles in steep coelacanths (FRICKE et al., 1991). populations. KEY WORDS: habitat, heterochrony, Indonesian coelacanth, juvenile, Latimeria menadoensis respectively. The terminology of fins by UYENO (1991) is juvenile and 8.7 cm and 28 % in L. chalumnae (Table 1). The of L. chalumnae (Fig. 5). The bases of the dorsal lobe of the depth of 164.6 m at which the juvenile was recorded is well and second anal fins), and longer supplementary lobe of the between adults. However, there are several differences We are grateful to Dr. Teruya UYENO for his valuable Atmosphere and Ocean Research Institute, The University of coelacanth was recorded. (Buol-2), larger and located 20 m deeper (Buol-2) than the first year, in the same Buol area (Table 1: R4) the sixth individual and Buol-5. Its posture was upside down with its ventral side was found on a steep slope at 169.4 m depth off Manado after. ID 24 was observed on the 4th May 2012 and then 13 slopes between 150 m and 253 m (FRICKE et al., 1991). Divers According to the observations of the African coelacanths, given in parentheses to avoid confusion of names. Fin names head is slightly shorter than that of L. chalumnae. It is 5.5 cm caudal fin (the third dorsal fin) and the ventral lobe of the within the range of adult habitat. The individual of 113 cm caudal fin (caudal fin) than those of the embryos of African between juveniles and embryos of these species (Fig. 5). This advice, guidance and encouragement throughout the present Tokyo. of the reproduction biology of the coelacanths is almost INTRODUCTION The first individual of Indonesian coelacanth (Table 2, ID cave (Buol-1). After 29 minutes of recording, this individual (Table 2: E5, ID 6) was recognized at 14:34 on the 12th facing the ceiling of the overhang. Such posture is also known (Table 2: Manado-6). It swam up and down 5.4 m between months later, on the 9th June 2013 at the same locations of recorded the presence of this species at 104 m depth at the species L. chalumnae is nocturnal and hides in cave in REFERENCES are abbreviated in the tables as follows. A1: Anal fin. A2: and 22 % of the standard length in L. menadoensis and 8.0 cm caudal fin (the second anal fin) are longer than those of L. total length (ID 16 in Iwata et al., 2019), was observed at coelacanth. indicates possibility of the ontogenetic difference of the two study. Many thanks go to colleagues of the Faculty of Fisheries REFERENCES restricted to an ovoviviparity strategy since some caught large 1) was recorded during the third field survey, in the morning of hid inside the cave. Although the water temperature increased December alongside a large rock. This individual stayed in the in coelacanths from the Comorian Archipelago (FRICKE et al., 168.1 and 173.5 m deep for 17 minutes. Thirteen months later Manado area within several kilometers. Sodwana Bay, South Africa (VENTER et al., 2000). Latimeria daytime but comes out at night, supposedly in search of food ACKNOWLEDGMENTS Ventral lobe of caudal fin. C: Supplementary lobe of caudal and 26 % in L. chalumnae. The depth of the head is slightly chalumnae (Table 1). The base of the dorsal lobe of the caudal 115.6 m depth in the same area three days after the juvenile One individual of L. menadoensis, 90 cm of total length species. and Marine Sciences of the Sam Ratulangi University at The first living coelacanth, Latimeria chalumnae, was females carried unfertilized eggs or embryos in their oviduct beginning of the observation and then moved ahead to deeper 1987). ID 3 was found totally four times in this area in two RICKE ENNO ERHEIJI TAPLEY UMISHA GATUNGA ENNO ERHEIJ TAPLEY GATUNGA the 30th May 2006 (Table 1, R3). The coelacanth was of more than five degrees, from 14.8 to 20.4 ºC, it stayed at the (the 9th June 2003), this individual (ID 24) was observed again All individuals observed more than twice were chalumnae seem to prefer a temperature range of 15–20 °C, (F et al., 1991). The living geomorphologic environment B , B., V , E., S , J., R , C., N , fin. D1: First dorsal fin. D2: Second dorsal fin. D3: Dorsal lobe smaller than that of L. chalumnae. The mouth is slightly fin (the third dorsal fin) is longer than that of the ventral lobe was found. Presence of both the juvenile and adults of L. and the second smallest one among the coelacanths observed The juvenile of L. menadoensis has a long supplemental Manado for their generous supports to conduct expeditions. B , B., V , E., S , C. R., N , B., discovered in South Africa in the Indian Ocean in 1938 (SMITH et al., 1975; CUPELLO et al., 2015). observed in a cave of 165 m depth (Table 2, Boul-1) for 10 same place, not moving away. Despite it disappeared from the slowly. Eventually it reached 12 m deeper during one hour and surveys during the seven months and seems to live in this area. near a cliff at 152.3 m depth (Table 2: E24, Manado-8), where encountered in the same area, at locations close from each where they tend to choose their daytime habitats (FRICKE et al., of L. menadoensis is similar to that of L. chalumnae, with We are grateful to Dr. Teruya UYENO for his valuable B., ABDALLAH, A. and KALOMBO, H. 2006. Coelacanth INTRODUCTION ovoviviparous because captured female specimens carried of caudal fin. P1: Pectoral fin. P2: Pelvic fin. Measurements are smaller than that of L. chalumnae. The gape of the mouth is of the caudal fin (the the second anal fin) in both species. The menadoensis within the same area inside of Manado Bay, in Indonesian water from 2005 to 2015 (IWATA et al., 2019), lobe of the caudal fin, which is seen in adult and often young Also, we express our thanks to Dr. SUHARSONO, Dr. Zainal ABDALLAH, A. and KALOMBOM, H. 2006. Coelacanth (SMITH, 1939). The first observation of the living coelacanth The Indonesian coelacanth, L. menadoensis, was first minutes (Table 2, E1). The adult individual was in stationary camera frame we kept setting the ROV there in front of the 42 minutes. During the 2007 survey (Table 1: R5) a coelacanth (Table located on the same slope of Manado-6. other within hundreds of meters. However, in the Comorian 1991). The suitable temperature for L. menadoensis seems to be rocky steep slopes with undermarine caves or overhangs. advice, guidance and encouragement throughout the present (Latimeria chalumnae SMITH, 1939) discoveries and developing embryos (SMITH et al., 1975, WOURMS et al., 1991). abbreviated as follows. HL: Head length. SL: Standard length. probably smaller than that of L. chalumnae. The eye is smaller supplementary lobe of the caudal fin (caudal fin) is longer than suggests that the species reproduce in a rather confined area. was observed at 187.4 m depth on the 6th December 2012 specimens of some fossil species and early stages of embryos ARIFIN, and colleagues of the Indonesian Institute of Science (Latimeria chalumnae SMITH, 1939) discoveries and habitats using submersibles was reported in Comoros (FRICKE discovered in Manado, Sulawesi Island, Indonesia, in 1997. position in a cave (Buol-1) slowly moving its pectoral, pelvic, cave until the next morning, however it did not appear again. On the 14th December, the individual ID 3 previously 2: ID 7) was recorded off Manado by 195 m depth, with a Another new coelacanth (Table 2: E23, ID 25) was Archipelago, same individuals of L. chalumnae were observed, almost the same as L. chalumnae. However, Latimeria menadoensis was also observed alongside study. Many thanks go to colleagues of the Faculty of Fisheries conservation in Tanzania. South African Journal of The first individual of the extant coelacanth, Latimeria Aquamarine Fukushima has conducted field surveys for TL: Total length. than that of L. chalumnae. The orbit diameter of the L. that of L. chalumnae. All fins are larger and broader than those Comparisons of the juvenile L. menadoensis and the (Fig. 6: ID 23 in IWATA et al., 2019). Although ID 23 is the of L. chalumnae (BENNO et al., 2006, fig. 2; CUPELLO et al., for advices and administrable work to obtain our research conservation in Tanzania. South African Journal of et al., 1987). Latimeria chalumnae hide in submarine caves The first specimen was captured in 1998 (ERDMANN et al., second dorsal and anal fins. The day after, another individual On the 4th June in 2006, three individuals, including ID 3 recorded at Buol-2 in a cave and at Buol-3 in a crack, was temperature of 12.4 °C, the coolest one registered during all recorded alone on a Manado site (Manado-7). within a two week period, in several caves distributed in a 8 Water temperature can increase or drop for more than 5 large rocks or along vertical wall in daytime. Some of these and Marine Sciences of the Sam Ratulangi University at Science, 102: 486–490. chalumnae, was discovered in South Africa in 1938 (SMITH, Indonesian coelacanth, Latimeria menadoensis from 2005 to menadoensis juvenile is almost half that of L. chalumnae. The of L. chalumnae. Peduncles of all lobed fins are also broader embryos of L. chalumnae show some interesting facts about second smallest individual among the coelacanths observed in 2015, fig. 1a; FOREY, 1998; YABUMOTO, 2008), indicating L. permit. Thanks also to Dr. BATUNA and the Murex Dive Resort Science, 102: 486–490. along steep slopes between 100 m and 200 m depth during 1998). According to a genetic study, the specimen differed (ID 2) and an unidentified one (UN1) were found in the same which was observed on the 31th May at Buol-2 and two new encountered again (Table 2: Buol-5) near a vertical wall at 145 surveys. Field surveys were conducted from the 19th to 30th May kilometers wide area (FRICKE et al., 1991). °C in a very short period of time (about 30 min according to our individuals were encountered above 200 m deep, in an Manado, Indonesia, for their generous supports to conduct BRUTON, M. N., CABRAL, A. J. P. and FRICKE, H. 1992. First 1939). The first observation of the living coelacanth habitats 2015 (IWATA et al., 2019). During the surveys a small coelacanth part (caudal peduncle) between the second dorsal and the anal than those of L. chalumnae. The juvenile of L. menadoensis ontogeny of the congeneric species. The embryos of L. Indonesian water, it is considered to be an adult because its menadoensis retained more juvenile characters during its staffs. We would like to express our sincere thanks to BRUTON, M. N., CABRAL, A. J. P. and FRICKE, H. 1992. First daytime and come out and drift near the ocean floor for from L. chalumnae and a new species was erected (POUYAUD cave (Buol-1). The unidentified one immediately hid deeper in individuals, were observed in a vertical crack which was m depth (Table 2: E6). After a while, this individual (ID 3) The field survey in 2009 was carried out off Manado and 2015 off Lolak Island and from the 30th October to the 16th When individuals of L. menadoensis were encountered, recordings). The coelacanths however often stayed there environment that remains slightly bright. Latimeria expeditions. Also we express our thanks to Dr. SUHARSONO, Dr. capture of a coelacanth, Latimeria chalumnae (Pisces, using submersible was reported in Comoros (FRICKE et al., was observed in 2009. This discovery was simply reported as a fins and the anterior ends of the dorsal lobe of the caudal fin has 8 spines of the first dorsal fin; 26 rays of the secod dorsal chalumnae are larger than the juvenile L. menadoensis and morphological feature is almost identical to other adults. Also, development than L. chalumnae. This indicates presence of colleagues at Aquamarine Fukushima for their understanding capture of a coelacanth, Latimeria chalumnae (Pisces, hunting prey at night (FRICKE et al., 1991). However, since the et al., 1999). In 1999 two individuals were observed by the cave, where the ROV could not access, and only its lateral several hundred meters far from Buol-2 (Buol-3) for 2 hours moved upward about 18 m and stayed behind a large rock (Fig. around Talise and Bangka islands off the northern coast of November 2015 off Lolak and Bitung Islands (Table 1; R13 some of them stayed stationary at same place but some without moving away or significantly modifying their menadoensis seems to be less sensitive to the daylight than L. Zainal ARIFIN, Dr. DIRHAMSYAH and colleagues of the Indonesian Latimeriidae), off Mozambique. South African Journal of 1987). Juveniles or small individuals of Latimeria are rarely scientific news (HOLDEN, 2009). Heterochrony of recent RESULTS (the third dorsal fin) and the ventral lobe of the caudal fin (the fin; 23 rays of the anal fin; 25 rays of the dorsal lobe of the have almost completely absorbed their yolk reserves according juvenile characters, observed in the juvenile here described, heterochrony in the development of Latimeria, with L. and supports. We thank Dr. Rik NULENS to provide information Latimeriidae), off Mozambique. South African Journal of record of juvenile or small (below 80 cm long) individuals is submersible in a location 360 km away from Manado (FRICKE side of the body was recorded. The individual (ID 2) was and 14 minutes (Fig. 2C; Table 2: E4, ID 3, 4, 5). During this 3). The observation time was the longest of all surveys with Sulawesi Island from the 12th September to the 9th October and R14). Three new individuals (Table 2: ID 26–28) were individuals such as ID 6 on the 12th December 2006 (E5) and behavior. These water currents of different temperatures are chalumnae. Institute of Science for their advice and administrative work to Science, 88: 225–227. obsereved or caught, and the growth and reproductive biology coelacanths was discussed based on this discovery. However, second anal fin) is shorter and deeper than that of L. caudal fin (the third dorsal fin); 23 rays of the ventral lobe of to CCC newsletter (SECRETARIAT AT J. L. B. SMITH INSTITUTE are not observed in ID 23. The body proportion of the juvenile menadoensis being paedomorphic during its juvenile stage. of CCC records. We would like to thank Dr. Camila CUPELLO Science, 88: 225–227. virtually absent, the growth, development and breeding et al., 2000). Today 8 individuals were officially caught in the observed for one minute before it hid inside the cave (Table 2: long period of time all individuals stayed near the rock wall four hours and 16 minutes of recording. The temperature (Table 1: R8). On the 24th September 2009, six individuals recognized at different places off Lolak Island (Lolak 1-4). All ID 24 on the 4th May 2012 (E22) swam away. It seemed to caused by the movement of the marine thermocline. During These first observations are potentially crucial since the obtain our research permit. Other thanks go to Mr. OPO, Mr. CUPELLO, C., BRITO, P. M., HERBIN, M., MEUNIER, F. J., of extant species of Latimeria remain mostly unknown. The the detail of findings was not described. In the present paper, ROV observations chalumnae. The distance between the posterior end of the base the caudal fin (the second anal fin); 26 rays of the OF ICHTHYOLOGY, 1993), just before delivery. The juvenile of is very different from that of the adult in L. menadoensis (Figs. of Departamento de Zoologia, Universidade do Estado do Rio CUPELLO, C., BRITO, P. M., HERBIN, M., MEUNIER, F. J., biology of L. chalumnae remain poorly known. Our knowledge Indonesian seas. However, the details of their habitat have not E2). When a new individual could be identified by the white always keeping their head against the water flow. The water changed up and down several times (Fig. 4). Its difference (Table 2: E11, ID 9, 10, 11, 12, UN 2, 3) were found under an individuals were alone. ID 26 was observed in May and once avoid and escape from the brightness of the light or the sounds our surveys we observed in some area that as the thermocline habitats of the Indonesian coelacanths could be possibly not as REFRY and colleagues at Murex Dive Resort, and DAUD, AL, J ANVIER, P., DUTEL, H. and CLÉMENT, G. 2015. only knowledge of its reproduction is that Latimeria is this finding will be described in detail and the probable habitat The juvenile of Latimeria menadoensis (ID 15 in IWATA of the second dorsal fin peduncle and the anterior end of the supplementary lobe of the caudal fin (caudal fin) ; 29 rays of L. menadoensis has a smaller eye, larger peduncles of lobed 1 and 6). The morphological differences between L. de Janeiro as a referee for her critical reading of the manuscript J ANVIER, P., DUTEL, H. and CLÉMENT, G. 2015.