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Drilling Predation Traces on Recent Limpets from Northern Patagonia, Argentina

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Drilling Predation Traces on Recent Limpets from Northern Patagonia, Argentina Palaeontologia Electronica palaeo-electronica.org Drilling predation traces on recent limpets from northern Patagonia, Argentina Fernando M. Archuby and Sandra Gordillo ABSTRACT In this study we analyze the predator-prey relationship between muricid gastro- pods (mainly Trophon geversianus) and the limpets Nacella magellanica, Fissurella radiosa, Diodora patagonica and Siphonaria lessoni from recent mollusk death assem- blages at Puerto Lobos (41°59’54.2’’ S), on the Atlantic coast of northern Patagonia. The majority of the drill holes fit the ichnospecies Oichnus simplex Bromley, 1981. In five cases the holes resemble Oichnus ovalis Bromley (1983), attributable to octo- puses. In analyses of O. simplex, the drilling frequency of the whole mollusk assem- blage was 4.72%, which is higher than for N. magellanica (3.97%) and for all limpet specimens together (1.57%). Drillings were site-selective in the apical sector of the shell, with a notable and statistically significant preference for the U-shaped muscle scar. There is also some evidence of selective predation with respect to size, although the pattern is not strong. The low drilling frequencies on limpets compared to mussels coincide with the general idea that muricid gastropods have a strong preference for sessile prey, due to their inability to handle mobile items. The quantitative incidence of drilling predation on limpets varies considerably between published examples and within the different species of prey studied. The absence of records of drilling predation on fossil limpets remains partially unexplained, except for the fact that limpets are rarely preserved in the fossil record, thus reducing the probability of finding drilled specimens, plus the low frequency in which muricids choose limpet prey for drilling. This is the first study in South America centered on this biotic interaction. Fernando M. Archuby. Instituto de Investigación en Paleobiología y Geología, Universidad Nacional de Río Negro - Consejo Nacional de Investigaciones Científicas y Técnicas (IIPG, UNRN-CONICET), Av. Roca nº1242 (8332) General Roca (Río Negro), Argentina [email protected] Sandra Gordillo. Centro de Investigaciones en Ciencias de la Tierra, Consejo Nacional de Investigaciones Científicas y Técnicas - Universidad Nacional de Córdoba (CICTERRA, CONICET-UNC), Avenida Vélez Sarsfield 1611, X5016GCA Córdoba, Argentina [email protected] Keywords: Drilling predation; Limpets; Trophon geversianus; Octopuses; Argentine Patagonia, Death assemblages Archuby, Fernando M. and Gordillo, Sandra. 2018. Drilling predation traces on recent limpets from northern Patagonia, Argentina. Palaeontologia Electronica 21.3.36A 1-23. https://doi.org/10.26879/620 palaeo-electronica.org/content/2018/2246-limpets-predation-in-patagonia Copyright: September 2018 Palaeontological Association. This is an open access article distributed under the terms of Attribution-NonCommercial-ShareAlike 4.0 International (CC BY-NC-SA 4.0), which permits users to copy and redistribute the material in any medium or format, provided it is not used for commercial purposes and the original author and source are credited, with indications if any changes are made. creativecommons.org/licenses/by-nc-sa/4.0/ ARCHUBY & GORDILLO: LIMPETS PREDATION IN PATAGONIA Submission: 16 December 2015 Acceptance: 16 August 2018 INTRODUCTION tern of drill hole features for the sample (Lever et al., 1961; Chattopadhyay et al., 2013a and b). Predator-prey interactions constitute a major Previous studies of drilling predation are avail- ecological process which affects individuals, popu- able, in the form of in situ surveys and/or experi- lations and communities, and links various organ- ments with living fauna (e.g., Palmer, 1984; 1988; isms within and across ecosystems (Kowalewski Kabat, 1990; Kowalewski, 2004; Chattopadhyay and Leighton, 2007). Studying predator-prey inter- and Baumiller, 2007; Pio, 2010; Tyler and Schiff- actions in deep time also helps evaluate its effect bauer, 2012; Gordillo and Archuby, 2012; Archuby on the diversity, evolution and distribution of a and Leighton, 2014), recent death assemblages group (Paine, 1966; Vermeij, 1977, 1987; Roy, (Yanes and Tyler, 2009; Martinelli et al., 2013; Gor- 1996; Kelley and Hansen, 2003; Leighton and dillo and Archuby, 2014) and fossil assemblages Aronowsky, 2003; Chattopadhyay and Baumiller, (Taylor et al., 1983; Leighton, 2003; Ceranka and 2010; among others). Predator-prey systems Złotnik, 2003; Klompmaker et al., 2015, and many involving drilling predation on mollusk shells are others). There are also studies, which deal with the especially relevant for paleontologists because the temporal patterns of drilling predation (e.g., Kow- evidence of predation can be analyzed quantita- alewski et al., 1998; Harper, 2003; Kelley and Han- tively. Along the extensive Atlantic coast of Argen- sen, 2003; Dietl et al., 2004; Klompmaker and tine Patagonia, recent shells are naturally Kelley, 2015, among others). However, there are concentrated on the beaches by the action of tides, only a few articles, mostly recent, which provide a waves and wind. These concentrations are com- quantitative study of drilling predation in death mon, and allow a variety of studies centered on the assemblages from southern South America analysis of ecological attributes, such as drilling (namely, Gordillo, 1994, 1998; Gordillo and predation at local or regional level (Gordillo and Archuby, 2012, 2014; Martinelli et al., 2013). Archuby, 2012, 2014; Martinelli et al., 2013). The In an attempt to characterize the association death assemblages constitute appropriate samples between muricid drilling intensity and latitude, Mar- of the shelled portion of living communities and are tinelli et al. (2013) sampled dead shells along more affected mainly by the action of time (Tomašových than 1,000 km of the Patagonian Atlantic coast and Kidwell, 2009, 2010, 2011; Kidwell and (Argentina). The authors found drilling frequencies Tomašových, 2013; Archuby et al., 2015). The between 3 and 19% (with an anomalous figure of effect of time, i.e., the addition of shells over days, 36 in an anthropogenically impacted locality) at the years, centuries or even a few thousand years, cre- assemblage level, but no correlation with water ates time-averaged skeletal assemblages that, temperature or latitude. In this study the predator although lacking time resolution, rules out the bias- was most probably the muricid Trohpon gever- ing effect of short-term sudden changes in species sianus. A similar lack of correlation was found for composition and abundance, generated by abiotic the drilling frequency on the mytilid Brachidontes or biotic phenomena. An example of this kind of purpuratus, which was the species with the highest change would be the local extinction of the yellow number of drilled specimens (Martinelli et al., clam Mesodesma mactroides in the southwestern 2013). In a survey developed from recent dead Atlantic in 1995 (Fiori and Cazzaniga, 1999; shells in Bahía Golondrina, southern Tierra del Dadon, 2005). These time-averaged samples have Fuego (southernmost Argentina), estimated preda- been shown to be good proxies of the living com- tion frequencies were 16.2% for Mytilus chilensis munities, from deep marine to continental habitats and 11.7% for Aulacomya atra (Gordillo and (Kidwell and Tomašových, 2013). In addition, sam- Archuby, 2012). Visaggi and Kelley (2015) investi- pling or experimenting with living marine communi- gated naticid drilling frequency along 4,000 km of ties is normally complex and expensive (Warwick Brazilian coast, and found a significant increase in and Light, 2002) and does not necessarily guaran- drilling frequency towards the Equator, which they tee that the results can be extrapolated to the eco- related with temperature (although other factors systems (Cleland, 2001). Nevertheless, a possible might be involved, such as predator diversity and drawback of using death assemblages as proxies predator-prey size distribution). In a similar study of living communities is the effect of transportation, developed in the northern hemisphere, drilling fre- which might bias species composition and the pat- quency was highest at the mid-latitude in the Caro- 2 PALAEO-ELECTRONICA.ORG linian Province (29 to 34º N; Kelley and Hansen, interaction in the fossil record is due to a tapho- 2007). Drilling predation on grazing gastropods nomic bias. such as limpets from the shallow rocky shores of The Potential Drilling Predators Patagonia is interesting for biologists because of the ecological role that the consumers play in regu- Trophon geversianus Pallas is the most com- lating community structure (Bazterrica et al., 2007; mon predator sea snail inhabiting both rocky and Adami, 2008). The study of drilling predation on soft shallow bottoms along the Argentine Patago- limpets has attracted little attention, as shown by nian coast. Its area of distribution extends along the scarce literature available, despite the fact that the Atlantic coast from Buenos Aires province to limpets and drilling predators are major compo- Tierra del Fuego and along the Pacific coast of nents of rocky shallow marine environments every- Chile to at least the Chonos Archipelago (Griffin where (Black, 1978; Palmer, 1980, 1984, 1988; and Pastorino, 2005). In southern Patagonia, this Visaggi and Kelley, 2007; Yanes and Tyler, 2009; muricid gastropod mainly preys upon mytilids and Yamamoto, 1993, 2004; Chim and Ong, 2012; venerid clams (Gordillo, 1994,
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