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Defensive Strategies of European Spiny Lobster Palinurus Elephas During Predator Attack

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Defensive Strategies of European Spiny Lobster Palinurus Elephas During Predator Attack Vol. 423: 143–154, 2011 MARINE ECOLOGY PROGRESS SERIES Published February 10 doi: 10.3354/meps08957 Mar Ecol Prog Ser OPENPEN ACCESSCCESS Defensive strategies of European spiny lobster Palinurus elephas during predator attack G. Buscaino1,*, F. Filiciotto1, M. Gristina2, G. Buffa1, A. Bellante1, V. Maccarrone1, B. Patti1, S. Mazzola1 1Istituto per l’Ambiente Marino Costiero U.O. S. di Capo Granitola — Consiglio Nazionale delle Ricerche, Via del Faro No. 3, 91021 Granitola, TP, Italy 2Istituto per l’Ambiente Marino Costiero U.O. S. di Mazara del Vallo — Consiglio Nazionale delle Ricerche, Via L. Vaccara no. 61, 91026 Mazara del Vallo, TP, Italy ABSTRACT: The aim of the present work was to reveal the behaviour of the European spiny lobster Palinurus elephas (Fabricius, 1787) during encounters with predators. The study was conducted in a tank, exposing lobsters individually or in groups to a single specimen of the European conger eel or the common octopus. The behaviours of the lobsters, their sounds and the behaviours of the predators were recorded using a digital audio-video underwater acquisition system and analysed. A behav- ioural event not found in literature, dubbed ‘Alert’ by the authors, was described. The results showed that the most recurrent events were Alert, Point and Whip, which have the function of controlling the predator with the antennae, while maintaining the predator–prey distance. Moreover, a strong asso- ciation between Tail Flip and Lunge (events that were stimulated in response to an imminent danger to life) with the sounds emitted by lobsters was observed. Significantly more sounds were emitted during the tests with single lobsters compared to grouped lobsters. As a result of the present study, the European conger eel may also be considered a potential natural predator of the European spiny lobster. KEY WORDS: European spiny lobster · Behaviour · Antipredatory strategies · Common octopus · European conger eel Resale or republication not permitted without written consent of the publisher INTRODUCTION ster before an attack to reduce the risk of predator en- counters (Barshaw et al. 2003, Briones-Fourzán et al. Benthic stages of palinurid lobsters are subjected to 2006, Gristina et al. 2009) and (2) ‘antipredator mecha- a wide array of predators: groupers (Eggleston et al. nisms’ (e.g. escape, aggregation, cooperative defence, 1997, Quetglas et al. 2001), triggerfish (Barshaw et al. weaponry, clinging to the substrate) adopted during an 2003, Briones-Fourzán et al. 2006, Weiss et al. 2008, attack to reduce the risk of capture and killing (Vermeij Lavalli & Herrnkind 2009), sharks (Berry 1971, Eggle- 1982, Sih 1987, Brodie et al. 1991, Spanier et al. 1991, ston & Lipcius 1992), stingrays and octopus (Berger & Barshaw & Spanier 1994, Hazlett et al. 2000, Herrnkind Butler 2001). et al. 2001, Seitz et al. 2001, Barshaw et al. 2003, Bouw- To minimize the predation risk, which represents the ma & Herrnkind 2005, Lind & Cresswell 2005, Briones- primary source of natural population mortality (Butler & Fourzán et al. 2006, Lavalli & Herrnkind 2009). With Herrnkind 2000, Butler et al. 2006), lobsters have devel- regards to the ‘antipredator mechanisms’, several stud- oped effective defensive strategies for each life-history ies have described a range of defensive actions adopted stage. Defence mechanisms are generally mediated by by spiny lobsters to counteract diurnal predators in the 2 major strategies (Sih 1985, Brodie et al. 1991): (1) ‘pre- open (Herrnkind et al. 2001, Barshaw et al. 2003, dator-avoidance mechanisms’ (e.g. crypticity, immo- Bouwma 2006, Briones-Fourzán et al. 2006, Parsons & bility, nocturnal activity, sheltering) adopted by a lob- Eggleston 2006, Lavalli & Herrnkind 2009). These stud- *Email: [email protected] © Inter-Research 2011 · www.int-res.com 144 Mar Ecol Prog Ser 423: 143–154, 2011 ies, carried out both in the field and in mesocosms, have and natural protective holes are numerous from ap- identified a wide set of behavioural actions adopted by proximately 10 to 200 m in depth (Ceccaldi & Latrouite spiny lobsters during predator encounters. Within 2000). This species, primarily active at night, repre- these, the use of antennae probably represents the most sents one of the main targets of Mediterranean arti- effective antipredator weapon for spiny lobsters. In fact, sanal fisheries, and, while catches are now reduced the spinous antennae, with their powerful basal muscu- and sporadic, this fishery has a long history (Goñi & lature, can be used in a wide array of defensive tactics: Latrouite 2005, Groeneveld et al. 2006). As a result, to keep the predator at distance (‘Point’ or ‘Parry’) or to almost all P. elephas populations in the Mediterranean slap against the predator’s body (‘Whip’ or ‘Lunge’) are clearly overexploited (Goñi & Latrouite 2005, Gal- (Lavalli & Herrnkind 2009). The use of antennae in hardo et al. 2006). Notwithstanding the economic and combination with rapid contractions of the abdomen ecological importance of P. elephas in most parts of the musculature (‘Tail Flip’, ‘Rear Back’) (Lavalli & Her- Mediterranean coastal area, experimental data on the rnkind 2009), with movement of walking legs (‘Pirou- antipredator mechanisms of P. elephas are scanty or ette’, ‘Walk’) (Lavalli & Herrnkind 2009) and with coop- absent (Hunter 1999, Goldman & Patek 2002, Barshaw erative defence strategies (‘queue’, ‘rosette’, ‘phalanx’) et al. 2003, Patek & Oakley 2003, Gristina et al. 2009). (Herrnkind et al. 2001, Briones-Fourzán et al. 2006), The aim of the present study was to experimentally confers to spiny lobster a useful tool with which to reveal the antipredator behaviours of European spiny counteract predator attack during different life-history lobster Palinurus elephas in the presence of potential stages and in several ecological conditions. Further- predators with different swimming abilities and attack more, in combination with antennae sweeping, stridu- strategies: the common octopus Octopus vulgaris latory spiny lobsters are able to produce a characteristic which employs an ambush strategy (Hanlon & Hixon loud sound (Moulton 1957, Smale 1974, Mercer 1975, 1980) and the European conger eel Conger conger Mulligan & Fischer 1977, Patek 2001, 2002, Bouwma & which mainly attacks prey in the open using the accel- Herrnkind 2005, 2009). eration of its undulatory swimming (Sfakiotakis et al. Despite the wide scientific interest in this topic, the 1999). Both common octopus and European conger eel sound-production mechanism of stridulatory decapod co-occur with common lobster in the same coastal, crustaceans within the general framework of ‘anti- rocky habitat, sheltering in natural holes during the predator mechanisms’ is not completely understood. day and mainly active at night (Morato et al. 1999). Some authors have assumed that lobsters only produce Many octopuses are natural predators of spiny lobsters sounds in an antipredatory context (Lindberg 1955, (Joll 1977, Harrington et al. 2006, Bouwma & Her- Moulton 1957, Hazlett & Winn 1962, Meyer-Rochow & rnkind 2009, Butler & Lear 2009) and O. vulgaris preys Penrose 1974, 1976, Patek 2001, 2002, Patek & Oakley on the common spiny lobster P. elephas (Quetglas et al. 2003, Bouwma & Herrnkind 2009), while others sug- 2001, Goñi & Latrouite 2005). When threatened by gest that sounds may be used in social context as well octopus in the open, lobsters attempt to escape by (Moulton 1957, Mercer 1973, 1975). In particular, using tailflips, darting backwards away from the at- Bouwma & Herrnkind (2005) and Bouwma (2006) de- tacker. As a result, octopus predation on lobsters in the monstrated that Caribbean spiny lobsters Panulirus open may be a rare occurrence, while, using their long argus and P. guttatus in which the sound-production and flexible arms, octopuses mainly prey on lobsters mechanism had been disabled were attacked and inside their dens (Weiss et al. 2006, Bouwma & Her- killed more frequently by Octopus briareus than spiny rnkind 2009). The European conger eel is considered lobsters with intact sound-producing structures. Sound a voracious predator, eating a variety of species — emanates from a specialized stridulating organ com- Clupeidae (especially sardines), cephalopods (cuttle- posed of 2 parts: a movable plectrum attached to the fish, squid, octopus) and crustaceans such as shrimps last segment of the antennal peduncle and a rigid file and small crabs (Cau & Manconi 1984, Morato et al. (Patek 2002, Patek & Oakley 2003, Patek & Baio 2007). 1999, O’Sullivan et al. 2004) — and is also known to This organ is located at the base of the long second enter lobster pots (O’Sullivan et al. 2003). Conger eel antennae, which is also the lobster’s primary defensive predation on lobster is mostly anecdotal and lacking weapon (Patek 2002). To produce sound, the lobster experimental evidence; however, we believe that, draws the plectrum up the file by moving the antenna inhabiting the same habitat, conger eels probably base posteriorly. compete with lobster for limited available shelter and The European spiny lobster Palinurus elephas simultaneously could represent a menacing predator. (Fabricius, 1787) is common along the Mediterranean We chose to carry out the experiments without any and northeastern Atlantic coasts (Hunter 1999, Ceccal- dens, in order to prevent the presence of available di & Latrouite 2000). It is a temperate species living on shelters from affecting the attack strategies of the 2 rocky and coralligenous
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