We examine the pervasiveness of alternative modes of predation employed by naticids reported in the literature and offer recommendations regarding the terminology used in referring to such mechanisms. 3A), and in the absence of it (Fig. (2) Prey abundance should be controlled and monitored, and naticid predators should be isolated from one another unless testing for effects of multiple predators. Neverita duplicata (Say, 1822) is an abundant moon snail inhabiting shallow intertidal to subtidal environments along the eastern coast of the United States. Although TTX is found mostly in the muscle or digestive glands of naticids, Natica lineata demonstrated a capacity to release seawater yielding acute paralytic toxicity in response to external stimulation, i.e. 3 Comments Mark Ridgway a year ago. Jackson, 1973) in support of his speculation. However, as long as some sediment was present, regardless of depth, variation in drilling position did not impact predation mode, frequency of feeding or stereotypy of drillholes, as nearly all penetrated in the vicinity of the umbo (Fig. Nondrilling predation by moon snails has been linked to suffocation for nearly a century (Agersborg, 1920). (1) Tank space and substrate depths should be considered with respect to predator and prey sizes, life habits, and any attack, burrowing or escape behaviours. Moreover, the differences in health of the resident population of several parts of Europe seem to be related with their varied traditional diet (Cencic & Chingwaru, 2010 ). Examining literature accounts of alternative modes of naticid predation is challenging, because potential confounding variables have often not been reported (e.g. Funding for writing of the dissertation was provided by a Ford Foundation Fellowship and Association for Women Geoscientists Chrysalis Scholarship. Such studies typically define predation intensity as percent of prey individuals with complete drillholes. The present work focuses on deaths due to suffocation in which entry through the commissure is permitted via forced gaping before or during the drilling process, rather than through an existing permanent gape, which may allow feeding without prior suffocation of prey. Reid & Gustafson (1989) stated that bivalve prey were limp and unresponsive after being drilled, leading them to investigate pharmacological properties of oesophageal gland secretions of Lunatia lewisii. One specimen, in a 20-cm aquarium, was discovered empty on the sediment surface after only 3 d. The other, in a 2-cm tank, was found on the surface at 3 d but with signs of gaping, perhaps indicating that weakness prevented it from burrowing. peer-reviewed journal articles) and grey literature (e.g. Abbreviations: as in Table 2; SL, sand layer provided but precise depth not given. Nearly all drillholes in this investigation were located at umbo, as seen in these specimens. Members of Family Naticidae have wide, globose shells with short coiled spires (total of 5-6 whorls) and a diameter nearly equal to the height, no siphonal notch, interior of the shell is not pearly (photo), there is a horny or calcified operculum. Secondly, dates of entry into aquaria were recorded on all prey as a way to monitor how long individuals remained in experiments; average duration of occupancy in aquaria was quantified. Drilling captive prey commenced only upon relocation to a setup containing 9 cm of sand, in which they immediately burrowed. In addition, predator size may influence the depth of sediment required for normal feeding behaviour; Kabat (1990) noted that most experiments offer only slightly more sand than the size of the predators or prey under observation. Our results provide insight as to appropriate sediment depths for laboratory work on N. duplicata, alleviating prior concerns that minimal sediment leads to suffocation of prey. due to suffocation); (2) restrict use of ‘suffocation’ to situations in which mortality is attributed to respiratory distress; (3) promote the more appropriate phrase ‘alternative modes of predation’ as encompassing all feeding by naticids that is not accomplished using a completed drillhole and (4) abandon the term ‘smothering’ as it is not employed consistently or clearly in the literature, in part because multiple mechanisms may be executed by naticids in achieving apparent suffocation. accepted. All dietary recommendations show the need to increase Omega-3 long-chain fatty acids (LC n-3 PUFA) in the diet (Leventakou et al., 2014). The decision to use M. mercenaria as prey was based on several factors. To our knowledge, smothering, as an attack behaviour executed by gastropods, was used first by Morton (1958) to describe predation by members of the Cassididae, Harpidae, Olividae, Tonnidae and Volutidae. The number of predators confined to the same aquarium setup is denoted as single (Si), multiple (Mu) or both (B). For full access to this pdf, sign in to an existing account, or purchase an annual subscription. In our study, incomplete drilling, typically due to interruption during experimental checks, was monitored carefully. Species Parent. The third clam showed no indication of decay afterward or weakness, yet was discovered in the 20-cm setup within the substrate and completely empty at the next experimental check. Several authors have noted that only larger prey were suffocated in their experiments (e.g. Predators often attempted to bury themselves at least partially in the sediment, however, indicating that more substrate is preferred. Kitchell et al. Our experiments, however, show that most modern accounts of suffocation in tightly closing bivalves can be discounted as a result of weak prey in laboratory settings, alleviating concerns regarding the interpretation of the frequency of complete and incomplete drillholes. Overall, our experiments indicate that suffocation by Neverita duplicata is not linked to insufficient sediment. 2223, Stereotypic and size-selective predation in, The evolution of predator-prey behavior: naticids gastropods and their molluscan prey, Evolution of animal behavior: paleontological and field approaches, Prey selection by naticid gastropods: experimental tests and application to the fossil record, Anomalies in naticid predatory behavior: a critique and experimental observations, Drill hole predation on fossil serpulid polychaetes, with new data from the Pliocene of the Netherlands, Drill holes produced by the predatory gastropod, Yale University Reprographics & Imaging Services, New example of Devonian predatory boreholes and the influence of brachiopod spines on predator success, Inferring predation intensity in the marine fossil record, Trial control of the greater clam drill (, Journal of the Fisheries Research Board of Canada, The marine fauna and flora of Moreton Bay, Queensland, Memoirs of the Queensland Museum – Nature 54, Proceedings of the Thirteenth International Marine Biological Workshop, Brisbane. These moon snails were stressed in the absence of sediment and showed no interest in available prey. Use of ‘suffocation’ (SU) vs ‘smothering’ (SM) in the literature in reference to alternative predation by naticids. These examples highlight the challenges in assessing how undrilled prey perish in the field; concerns regarding prey health are not limited to laboratory experiments in attempting to recognize alternative modes of predation by naticids. Three replicate trials of 48 days each were conducted at the University of North Carolina Wilmington's Center for Marine Science during September–October 2010, October–December 2010 and June–July 2011, in part due to limited availability of specimens during the winter and concerns regarding suppressed feeding rates in cooler months. They found no paralysing effect in placing these secretions on the heart of Tresus nuttallii and concluded that prey must be suffocated as suggested by others. (1986) recognized the issue of prey health in laboratory studies and proposed that Medcof & Thurber (1958) incorrectly ascribed Mya arenaria deaths without drillholes to naticid predation instead of considering background mortality of experimental prey after being placed in the field. To minimize concerns regarding prey health in our study, we eliminated unhealthy prey from our study, i.e. Rank. Although sediment likely offers greater stability in handling of prey items, one of our N. duplicata regularly pursued and drilled clams in the absence of supportive sediment; however, fewer prey were consumed relative to most moon snails in aquaria with sand (22 prey compared with an average of 32 prey per predator in tanks with sediment). During the dieta, the spirit of the plant presents itself and asks the purpose or the intentions that one has for working with “his” plant. Neverita Didyma or moon shell isolated on black background. Although his comments are based on only 11 observations, he cited high anaerobic capacities of the Lucinidae (e.g. Burrowing through soft substrates, most naticids forage at or below the sediment surface and remain submerged while in pursuit of their prey. A thorough investigation of the term's nuanced history and semantic connotations requires a wide diversity of information sources, including primary literature (e.g. Thanks to G. Vermeij for input on valve closure. Can naticid gastropod predators be discriminated by the holes they drill? Drilling occurred beneath the sediment, on the sediment (Fig. Because it is unclear if predatory behaviours such as suffocation are common in natural settings or are mostly artefacts of laboratory conditions such as insufficient substrate, we examined experimentally the influence of different sediment depths on drilling vs suffocation of Mercenaria mercenaria prey by Neverita duplicata. Finally, following Visaggi (2012), empty shells recovered every 6 d were analysed for signs of decay following natural mortality, including odour and whether any soft parts remained, as naticids typically consume the entire soft tissues of prey (Kitchell et al., 1981). predator-prey sizes, frequency of monitoring, density of predator and prey individuals and prey health). 601 S. College Road, Wilmington, NC 28403. Turner, 1950; Edwards & Huebner, 1977). Christy C. Visaggi, Gregory P. Dietl, Patricia H. Kelley, Testing the influence of sediment depth on drilling behaviour of Neverita duplicata (Gastropoda: Naticidae), with a review of alternative modes of predation by naticids, Journal of Molluscan Studies, Volume 79, Issue 4, November 2013, Pages 310–322, https://doi.org/10.1093/mollus/eyt023. The latter interpretation is further supported by the observation that a naticid repeatedly ignored a decaying Mercenaria mercenaria on the sediment surface in a 6-cm setup. Didyma was an ancient Ionian shrine which is today known as Didim in Turkey . D. The recurrent behavioural display characterized as upside-down and foot extended by a naticid in a 0-cm setup. (2002) A compendium of fossil marine animal genera, Bulletins of American Paleontology 363, 1-560: Martin G. C. (1904) Gastropoda, Maryland Geological Survey Miocene Text, 131 … Gastropods that can be attacked through the aperture may also be suffocated more easily. Related Products Mercenaria mercenaria has tightly closing valves and a moderate metabolic rate and may be able to withstand lower oxygen conditions (Savage, 1976), perhaps contributing to the low incidence of suffocation observed here. Despite exhibiting stereotypic behaviours useful for studying ecological and evolutionary aspects of predator-prey interactions, naticid gastropods have been reported as utilizing alternative modes of predation, such as suffocation. For example, Bayliss (1986) found that Euspira pulchella was unable to drill prey in aquaria containing only a few millimetres of sand; although victims could be captured, moon snails were unable to burrow and merely moved in circles, dragging their prey with them. Taxon names for naticids are updated as per Torigoe & Inaba (2011). We were able to minimize prey health as a concern by assessing the condition of prey before, during and after experimentation and monitoring the duration of prey used in aquaria. Our literature review also highlights that different causal mechanisms may allow moon snails to feed in the absence of a completed drillhole; research is needed on alternative naticid predation modes that may be a concern for studying predator-prey interactions using drillholes. Regarded as models of stereotypy in their experiments ( e.g after removal of yielded! Didyma ) adds brilliant color to the same setup upon experimental checks and possibly water! 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