6 SPC Beche-de-mer Information Bulletin #20 – August 2004

Field observations of juvenile sea cucumbers Glenn Shiell1

Introduction The relative scarcity of knowledge obtained through direct observation of field based juvenile Recent advances in tropical mari- sea cucumbers is due possibly to two problems. culture have created scope for the rehabilitation of The first, as reported by Wiedemeyer (1994), is that populations affected by overfishing through the the calcareous spicule arrangement in juveniles production and release of hatchery-raised juve- could be different to that of adults. Hence, the iden- niles. Although the necessary technology required tification of juveniles based on keys developed for to implement rehabilitation programmes is pro- adults may lead to misidentification. Second, and gressing rapidly (Purcell 2004), there is some perhaps most importantly, juveniles are very rarely speculation as to the viability of such pro- encountered in sufficient numbers for study, if at grammes, given major shortfalls in knowledge of all. The fact that small juvenile sea cucumbers are important aspects of sea cucumber biology. (For a rarely encountered in the field (Seeto 1994) may be full review see Bell and Nash 2004.) One area of due to a number of scenarios. Juvenile holothuri- sea cucumber biology identified as being impor- ans have the potential to be misidentified given tant to the ultimate success of rehabilitation pro- their potential for morphological differences rela- grammes is a clear understanding of the habitat tive to the adult forms (Wiedemeyer 1994); they oc- and ecological requirements of juvenile sea cu- cupy habitats different to that of larger specimens cumbers (Wiedemeyer 1994; Mercier et al. 1999; (James et al. 1994; Lokani et al. 1996), and finally, Purcell 2004). An understanding of juvenile habitat they exist in the habitat occupied by the adult form preferences may help scientists to determine accu- but are obscured from view within sediments or rately the ultimate carrying capacity of a given sea crevices or beneath obscuring objects such as coral cucumber habitat (while acknowledging the space (Cameron and Fankboner 1989; Wiedemeyer 1994). occupied by juveniles) and additionally, enable the eventual release of juveniles to appropriate habi- The difficulty in locating juvenile sea cucumbers is tats with enhanced probability of survival (Bell and perhaps highlighted by the fact that studies rele- Nash 2004; Purcell 2004). vant to juvenile ecology often result from fortu- itous encounters (Conand 1983; Mercier et al. Much of the existing literature on juvenile 1999). For example, juvenile Actinopyga echinites holothurian biology has focused on observations or used in Wiedmeyer’s study (1994) were found on studies of hatchery-raised juveniles, particularly in the upper reef zone following a strong typhoon. the context of growth and mortality under labora- For this reason, and because juvenile sea cucum- tory or aquaculture conditions (e.g. Battaglene 1999; bers are rarely observed in large numbers, it Battaglene and Seymore 1998; Battaglene et al. 1999; seemed appropriate and useful to begin consoli- Engstrom 1980; Hamano et al. 1996; Hamel et al. dating anecdotal observations of juveniles made in 2003; Hatanaka 1996; Hatanaka et al. 1994; Ito et al. the field. This exercise may help to identify differ- 1994; James et al. 1994; Kobayashi and Ishida 1984; ences in habitat preferences between juveniles on Mercier et al. 1999, 2000a; Tanaka 2000). However, the inter- level and, additionally, between several authors have also made contributions to the adults and juveniles of the same species. Also, by understanding of juvenile biology based on studies consolidating these observations, it may be possi- of juveniles that were either captured or observed ble to discern future research directions to clarify in situ (e.g. Cameron and Fankboner 1989; Daud et some of the details of this little known but impor- al. 1993; Hamel and Mercier 1996; Mercier et al. tant life phase. 2000b; Muliani 1993; Purcell 2004; Purcell et al. Response to the questionnaire included in SPC’s 2002; Ramofafia et al. 1997; Tiensongrusmee and Beche-de-Mer Information Bulletin #19 Pontjoprawiro 1988; Wiedemeyer 1994; Young and Chia 1982). Additionally, observations of juvenile Holothuria scabra are reported from a variety of Questionnaires on observations of holothurian sources (Conand 1997; Gravely 1927; James 1976, spawning and fission have been published previ- 1983; Lokani et al. 1995; Long and Skewes 1997; ously in SPC’s Beche-de-Mer Information Bulletin. Shelley 1985). Following the success of these, a new questionnaire

1. School of Biology (MO92), University of Western Australia, 35 Stirling Hwy, Nedlands, WA 6009. Email: [email protected]

SPC Beche-de-mer Information Bulletin #20 – August 2004 7 aimed at consolidatng anecdotal observations of ju- Smaller specimens of H. scabra (> 10–40 mm) are venile sea cucumbers in the field, was included in also reported to remain hidden for the majority of issue #19 (p. 41). Many questionnaire respondents daylight hours. Larger H. scabra juveniles (> 40–140 provided observations of juveniles ranging between mm) emerged from sediments in the middle of the 0.3 and 21.0 cm in length. Given the inter-species day at around 13:30 hours (Mercier et al. 1999). variability in size at first maturity among holothuri- ans, all of the observations were retained in the final Additional results presented in Table 1 indicate a results. Twenty-six responses to the questionnaire preference for juveniles of selected species to oc- have been received at the time of writing. cupy habitats slightly different to those of adults. Results and discussion In these cases, juvenile Holothuria fuscogilva and H. nobilis were located in shallower water adjacent to the deeper water habitats occupied by adults The results to date (Table 1) are organised under (Conand 1981). A similar phenomenon was re- the following sub-headings: species observed, cor- ported in Cucumaria frondosa (Hamel and Mercier responding observed habitat, date and time of ob- 1996) that were observed to undergo a process of servation, proximity of adults in relation to ob- progressive migration from shallower protected served juvenile(s) and the name and affiliation of reef areas to deeper more exposed sandy areas as the observer. they matured sexually. The reported observation of H. fuscogilva juveniles (Table 1) in very shallow Results presented in Table 1 indicate a broad range water is suggestive of a similar pattern of migra- of habitat preferences among juvenile holothuri- tion in this species. Evidence for this notion was ans. In most cases, juvenile holothurians were ob- more recently provided by Ramofafia et al. (2000) served within close proximity to adult holothuri- who obtained sexually mature individuals of ans of the same species. These observations in- H. fuscogilva in water between 25 and 30 m depth. cluded representatives of the following genera: Size related migration from shallow water to deep Stichopus (2 species), Actinopyga (3 species), water has also been reported in Stichopus variegatus Isostichopus (2 species), Astichopus, Thelenota, (now S. hermanni) (Conand 1993). Cucumaria, Chiridota, and (1 species each). Conclusions Habitat preferences of juvenile Holothuria spp. var- ied, but in most cases, juveniles of this genus were also observed in the presence of adults (e.g. H. From the observations provided in Table 1 and the scabra, H. leucospilota, H. mexicana, H. atra and in additional evidence provided in the literature, it some instances, H. nobilis). Concurrent occupation seems reasonable to conclude that juvenile of similar habitat between both juvenile and adult holothurians display some degree of cryptic be- H. scabra was reported by Mercier et al. (1999, haviour in the earliest stages of their life cycle fol- 2000b) who found newly settled and smaller juve- lowing settlement. Cryptic behaviour most likely niles in the same general area as adults. Both continues until juveniles are large enough to avoid Cameron and Fankboner (1989) and Young and most forms of (Cameron and Fankboner Chia (1982) provide similar evidence, where 1989). Further migration and hence greater habitat Parastichopus californicus and Psolus chitonoides ju- separation between juveniles and adults of some veniles were also regularly observed within habi- species may then occur as holothurians mature tats occupied by the adult form. Although it seems (Hamel and Mercier 1996), but the degree to which common for juveniles and adults to occupy habi- this process is common among all holothurians re- tats simultaneously, there is evidence that most mains unclear. small juveniles retain some degree of cryptic be- haviour that appears to diminish with size. An additional observation is that juvenile and Therefore, it is likely that juveniles may be ob- adult sea cucumber habitat preferences consis- scured from view even when present in large num- tently appear to differ at some level of scale. For bers within an adult habitat. For example, example, although H. scabra juveniles and adults Isostichopus fuscus juveniles (< 6 cm) maintained in have been observed in the same general area, aquaculture ponds remain hidden in the rocky sub- clear differences in habitat preferences at the mi- strate during the day, but begin to emerge follow- crohabitat level have been identified (Mercier et ing nightfall; larger specimens of the same species al. 2000b). Microhabitat requirements of juvenile were otherwise visible throughout the day (Mr holothurians are likely to vary both between Roberto Ycaza, pers. comm.). Similarly, Cucumaria stages of development and between species. To frondosa and Actinopyga echinites juveniles dis- further develop viable restocking programmes played a progressive tendency to leave protected using hatchery-raised juveniles, it would be desir- locations within the substrate as they grew larger able to investigate microhabitat requirements in (Hamel and Mercier 1996, Wiedemeyer 1994). more detail. Specifically, future research direc- 8 SPC Beche-de-mer Information Bulletin #20 – August 2004

Table 1. In situ observations of juvenile holothurians.

Species Approx. size Location Habitat Time Date Adults Observers’ name(s) and affiliation / observed and number present? Source of further information

Astichopus 8–21 cm North at Isla de la Seagrass 15:40 April 2002 Irma Alfonso Hernández, María del multifidus Juventud Pilar Frías; Fishery Research Centre of Cuba Actinopyga 5–18 cm North central Seagrass 14:30 May 2001 Yes Irma Alfonso Hernández, María del agassizii region of Cuba Pilar Frías; Fishery Research Centre of Cuba A. mauritiana 2–3 cm Unia Reef, New Reef flat Daytime 1989 Ye s Chantal Conand; Caledonia Université de La Réunion A. echinites 4 cm Ricaudy Reef, New Coral rubble Daytime Sept. 1981 Ye s Chantal Conand; Université de La Caledonia Réunion (unpub. photo available) Chiridota laevis 0.3–1 cm Bic Provincial Park At low tide Summer Ye s J.-F. Hamel & A. Mercier; (Quebec), Canada below the rocks 1994 Society for the Exploration & Valuing on sand flat of the Environment, Canada Cucumaria 1–3 cm Passamaquody Bay, Within tide Summer Ye s J-F. Hamel & A. Mercier; frondosa New Brunswick, pools near the 2000 Society for the Exploration & Valuing Canada. lowest tide mark; of the Environment, Canada below rocks and in crevices Holothuria 5–18 cm Pingües Channel, Sand 11:20 and May 2000 No Irma Alfonso Hernández, María del mexicana south eastern 14:25 July 2001 Pilar Frías; region of Cuba, Fishery Research Centre of Cuba Baie des Baraderes, Haití H. leucospilota 1–5 cm Guadalcanal, At low tide on 1998 Ye s J-F. Hamel & A. Mercier; Solomon Islands reef flat; below Society for the Exploration & Valuing rocks and in of the Environment, Canada crevices H. scabra 1–5 cm Ambandjoa, Muddy intertidal Daytime 1997 Nearby, but Chantal Conand - Photo published in: Madagascar in deeper Conand, C. 1999. Manuel de qualité water des holothuries commerciales du Sud-Quest de l’ Océan Indien. Commission Océan Indien: 39 p. H. scabra 8 cm Crab Island, Seagrass / algae Daytime 2002 Yes,within Grant Leeworthy; (1≈ individual) Moreton Bay, Shallow water 50 m Tasmanian Seafoods, Australia Queensland. H. scabra 10 cm long, Pouangué, Nor- Seagrass on a About Oct. 2002 Yes,same Steve Purcell; 2 cm in thern Province, muddy inshore 15:00 habitat, but WorldFish Center, New Caledonia diameter New Caledonia reef flat (shallow Low tide, juveniles water) when generally little burrowed in water mud, adults present foraging on on reef surface H. fucogilva 1–5 cm Fiji barrier reef Halimeda field, Daytime 1979 Yes,nearby, Chantal Conand; very shallow but deeper Université de La Réunion water on coral Photo published in Bull. Mar. Sci. 1981 reefs 31(3):523–543 H. nobilis 5 cm, Fiji barrier reef Seagrass bed, Daytime 1979 Yes,nearby, Chantal Conand; colour and shallow water but deeper Université de La Réunion different New Caledonia on coral Photo published in Bull. Mar. Sci. 1981 from adults, reefs 31(3):523–543 cream pat- ches on black background H. nobilis 14–21 cm Raine Island, Seagrass bed, Daytime Dec. 2003 Ye s Sven Uthicke; (300–625 g), Great Barrier Reef, shallow water AIMS,Townsville Australia colour Queensland, different Australia from adults, 3 specimens H. nobilis 12 cm Michaelmas Reef, Shallow water, Daytime Mar. 2004 Ye s Sven Uthicke; (estimated), Great Barrier Reef, lagoonal reef AIMS,Townsville Australia colour Queensland, area different Australia from adults, 3 specimens H. atra 2–3 cm Ilôt Maitre reef flat, Coral rubble Ye s Chantal Conand; (a few New Caledonia; Université de La Réunion specimens) Planch' Alizés, La Réunion H. atra 1–4 cm Likiep Atoll, At low tide on May 2001 Ye s J.-F. Hamel & A. Mercier; Marshall Islands reef flat; in Society for the Exploration & Valuing crevices of the Environment, Canada SPC Beche-de-mer Information Bulletin #20 – August 2004 9

Table 1(continued). In situ observations of juvenile holothurians.

Species Approx. size Location Habitat Time Date Adults Observers name and affiliation / observed and number present? Source of further information

Holothuria atra 1–4 cm Likiep Atoll, At low tide on May 2001 Ye s J.-F. Hamel and A. Mercier; Marshall Islands reef flat; in Society for the Exploration & Valuing crevices of the Environment, Canada Isostichopus 1–3 cm Along the coast of Between 5 and Autumn Ye s J.-F. Hamel and A. Mercier; fuscus mainland Ecuador 10 m water 2000. Society for the Exploration & Valuing depth of the Environment, Canada Isostichopus 5–14 cm North at Isla de la Seagrass Various April to Yes Irma Alfonso Hernández, María del badionotus Juventud, Banes daylight Nov. 2002 Pilar Frías; Bay, northern hours Fishery Research Centre of Cuba region of Cuba; Pilón Inlet, south eastern region of Cuba; Baie des Baraderes, Haití Psolus fabricii 0.5–3 cm Les Escoumins 3–10 m depth; Summer Ye s J.-F. Hamel and A. Mercier; (Quebec), Canada below rocks and 1991 Society for the Exploration & Valuing in crevices of the Environment, Canada

Stichopus 9 cm Sainte Marie Bay, Seagrass Sept. 1981 Ye s Chantal Conand; Université de La hermanni New Caledonia Réunion. For more information see: Bull. Mar. Sci. 52(3):970–981 S. chloronotus 3–4 cm (two Coral Bay, Ningaloo Reef flat near Daytime, Aug. 2003 Ye s Glenn Shiell; specimens) Reef,Western crevice afternoon University of Western Australia Australia

S. chloronotus 2–3 cm (very Great Palm Island, On Sargassum Daytime Yes,dense Sven Uthicke; rare, Great Barrier Reef, populations AIMS,Townsville Australia observed Queensland, only 3–5 in a Australia 2-year field study at that site) S. chloronotus 2–3 cm Several Reunion Rubble and sand Daytime Ye s For more information see: Conand Island reefs C., Uthicke S. and Hoareau T. 2002 Invert. Reprod. Develop. 41(1–3):235–242 Thelenota 12 cm Uitoe Pass, Coral 1981 Ye s Chantal Conand ananas New Caledonia Photo available in: Bull. Mar. Sci. 1981 31(3):523–543

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10 SPC Beche-de-mer Information Bulletin #20 – August 2004

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PIMRIS is a joint project of five international organi- the availability of information on marine resources sations concerned with fisheries and marine re- to users in the region, so as to support their ratio- source development in the Pacific Islands region. nal development and management. PIMRIS activi- The project is executed by the Secretariat of the ties include: the active collection, cataloguing and Pacific Community (SPC), the South Pacific Forum archiving of technical documents, especially ephe- Fisheries Agency (FFA), the University of the South mera (‘grey literature’); evaluation, repackaging Pacific (USP), the South Pacific Applied Geoscience and dissemination of information; provision of lit- Commission (SOPAC), and the South Pacific erature searches, question-and-answer services and Regional Environment Programme (SPREP). This bibliographic support; and assistance with the de- bulletin is produced by SPC as part of its commit- Pacific Islands Marine Resources velopment of in-country reference collections and ment to PIMRIS. The aim of PIMRIS is to improve Information System databases on marine resources.