Hermit Crab Population Ecology on a Shallow Coral Reef (Bailey’S Cay, Roatan, Honduras): Octopus Predation and Hermit Crab Shell Use
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Memoirs of Museum Victoria 60(1): 35–44 (2003) ISSN 1447-2546 (Print) 1447-2554 (On-line) http://www.museum.vic.gov.au/memoirs Hermit crab population ecology on a shallow coral reef (Bailey’s Cay, Roatan, Honduras): octopus predation and hermit crab shell use SANDRA L. GILCHRIST New College of Florida, 5700 N. Tamiami Trail, Sarasota, Florida 34243, USA ([email protected]) Abstract Gilchrist, S.L. 2003. Hermit crab population ecology on a shallow coral reef (Bailey’s Cay, Roatan, Honduras): octopus predation and hermit crab shell use. In: Lemaitre, R., and Tudge, C.C. (eds), Biology of the Anomura. Proceedings of a symposium at the Fifth International Crustacean Congress, Melbourne, Australia, 9–13 July 2001. Memoirs of Museum Victoria 60(1): 35–44. Shells can be a limiting factor in allowing hermit crab populations to increase. Predators of gastropod molluscs and of hermit crabs release shells into reef environments where hermit crabs find and cycle them within their populations. Predators also play a role in distributing shells among hermit crab species. To highlight how octopuses influence shell availability to hermit crabs, observations were made on members of Octopus vulgaris Cuvier, 1797 and O. briareus Robson, 1929 at Bailey’s Cay Reef (Roatan, Honduras) during July and August each of three years, 1999–2001. In addi- tion to feeding while foraging, Octopus vulgaris and O. briareus individuals create shell and debris middens outside of their temporary dens. These middens concentrate shells and food for hermit crabs in the reef environment where locat- ing an empty shell could be difficult. However, because hermit crabs are prey items for octopuses, hermit crabs using the middens risk predation from the den occupant. Relatively small hermit crab species such as Pagurus brevidactylus (Stimpson, 1858) and P. criniticornis (Dana, 1852) were found commonly in dens and among middens, opening the possibility that the den functions as a refugium for some species as well. Keywords Crustacea, Anomura, hermit crab, predation, shell use, octopus Introduction even intact shells begin deterioration upon entering the hermit crab use cycle. Hermit crabs generally do not procure shells directly from live The cycle of shells among gastropods and hermit crabs is molluscs (Hazlett, 1981; for an exception see Rutherford, not well known in coral reef environments. Octopuses consume 1977). Recycling of postmortem shells from gastropods and both crustaceans and molluscs, making their potential impact from live or postmortem hermit crabs is common (Bertness, on hermit crab shell cycling complex. They can prey upon her- 1982; Wilber and Herrnkind, 1984). Because shell availability mit crabs as well as crustaceans that are hermit crab predators. has been shown to be important in determining hermit crab In addition, they not only consume gastropods that can provide population size, hermit crab shell use has been widely investi- shells for hermit crabs, they also carry prey from various parts gated both in the lab and in the field (Benvenuto and Gherardi, of the reef back to a den location. Mather et al. (1997) specul- 2001; Elwood et al., 1979; Garcia and Mantelatto, 2001; Hahn, ated that hermit crabs are sometimes associated with octopus 1998; Hazlett, 1996; Osorno et al., 1998; Siu and Lee, 1992; dens as scavengers opportunistically feeding on remains of Vance, 1972 inter alia). Predators on gastropods and hermit prey left in middens and dens. Some workers have indicated crabs provide a variety of shells for habitation by hermit crabs hermit crabs as prey items for octopuses in field studies (Carikker, 1981; Mather, 1991; Tirelli et al., 2000). Sustainable (Iribarne et al., 1993) while other researchers have used hermit recycling requires hermit crabs to assess continually the quali- crab zoeae as prey in food searching studies of octopus par- ty of resources within the recycle pool. Some predators leave alarvae (Navarro and Villanueva, 2000; Villanueva et al., shells intact with little apparent damage (Gilchrist, 1984; Jory 1996). Octopuses can crush or drill their prey. They may also and Iversen, 1983; Ray and Stoner, 1995) while other predators extract gastropod or hermit crab prey through shell apertures, crush or smash the shells beyond use for hermit crabs (Brown leaving a relatively intact shell. The fact that octopuses not only et al., 1979; Hsueh et al., 1992; Hughes, 2000; Seed and feed on hermit crabs, but can also concentrate shell and food Hughes, 1995; Vermeij, 1977; Yamada and Boulding, 1998). resources in middens formed outside of their dens suggested However, LaBarbera and Merz (1992) noted decreases in shell that their role in the cycling process should be examined more strength after removal of the living gastropod, suggesting that 36 S.L. Gilchrist closely in the reef system. The types of shells entering the her- observed most often in early evening and at night; they were mit crab shell cycle for this study were noted at Bailey’s Cay rarely active outside of the den during daylight hours. Each reef by observing octopuses in the field to determine their role animal was observed for a minimum of 20 minutes and a max- in the hermit crab shell cycle. imum of 30 minutes as it emerged from the den and proceeded with foraging for each of 3 consecutive days. Civil dusk in the Methods area occurs around 7:30 CST during July and August. Lights were used after civil dusk. These lights were not directly Description of sampling area. The eastern reef area surround- applied to octopuses as they foraged. Indirect lighting did not ing Bailey’s Cay provides an opportunity to examine the hermit result in colour changes by the octopuses and the animals con- crab shell cycle under field conditions. The Cay is located with- tinued to forage for several hours. The observations of foraging in the Roatan Marine Preserve, so no spearfishing is allowed in allowed establishment of general feeding areas, feeding the area and only artisanal line fishing from traditional canoes duration and habitats visited. Remains discarded by foraging is permitted. Bailey’s Cay is part of a collapsed volcanic ridge O. vulgaris and O. briareus individuals were collected and surrounded by patch reefs to the east ending with a reef wall categorised by shell type, organism consumed. Characteristic that drops nearly vertically to about 30 m. There is a wall that behaviours exhibited by octopuses described by Hanlon and drops vertically to 35 m with a narrow shelf to the north. A boat Messenger (1996) and Mather (1991) were used to suggest channel about 15 m deep between Bailey’s Cay and Roatan items eaten away from the den, though specific numbers from bounds the southern part of the reef. Thus, because of the water foraging were not determined over the entire foraging time of depth surrounding the reefs of the cay, once hermit crabs and octopuses in this study. Postmortem gastropod shells were molluscs metamorphose from the plankton, they are surround- identified by remains of muscles or flesh attached to shells. ed by deeper water, restricting movements from the area. Gastropod shells recovered with no remaining flesh were cate- gorised as formerly occupied by hermit crabs. Gastropod shell General octopus foraging observations. Octopuses were active fragments were noted separately because former occupant both day and night. They were observed using focal animal and could not be determined clearly for most fragments. A shell was focal area techniques (Altman, 1974) during July and August of considered a fragment for this study if at least two whorls 1999–2001 at the same area of the eastern reef of Bailey’s Cay. beyond the protoconch were intact. The sampling area was approximately 5000 m2, extending from shallow seagrass through the top of the forereef. Hermit crab observations at octopus dens. After establishing Though observations were made on octopuses and their den areas and general foraging patterns for O. vulgaris, remains dens in the same area for three years, it is unlikely that the in middens were collected and catalogued daily for seven con- octopuses were the same individuals each year. The life span of secutive days. Octopus briareus individuals generally con- these animals is limited and they grow rapidly (Hanlon, 1983). sumed prey while hunting (about 80% eaten away from the The size of each octopus observed was small (3–5 cm head den), however, the small middens formed by these animals diameter; estimated using the methods described by Aronson, were also sampled for seven consecutive days. Two active dens 1982), suggesting that they were juveniles. To facilitate obser- of O. vulgaris and one of O. briareus were selected in August vations, dens of octopuses in the sampling area were identified 2001 for more focused hermit crab observations. and marked discreetly. Observation areas were chosen where Active den sites for both octopus species were observed for the minimum distance between conspecific dens was 5 m while hermit crab activity every 30 minutes for a two-hour period minimum distance for different species was approximately 12 starting an hour before sunset. A circular area of one-metre m for octopuses observed in 1999–2001. Observers using diameter was outlined around each den using plastic tent pegs SCUBA or snorkeling remained at 2–3 m from foraging octo- driven into the substrate. The 1-m area was considered the den puses. Only animals that habituated to presence of observers area while the den was the physical shelter used by the octopus. were used for collecting foraging data. (The area is frequented As background information, two circles with 1 m diameters during the day by recreational snorkelers and swimmers, thus were marked elsewhere in the seagrass and in the coral rubble octopuses are not in an isolated habitat).