Boletín de la Sociedad Geológica Mexicana ISSN: 1405-3322 Sociedad Geológica Mexicana, A.C.

Gómez-Espinosa, Catalina; Farinati, Ester; Aliotta, Salvador Predatory evidence in the high-latitude cold-water gastropod Buccinanops deformis (King, 1832) from the Holocene littoral sand ridges, Argentina, SW Atlantic Boletín de la Sociedad Geológica Mexicana, vol. 70, no. 2, 2018, May-August, pp. 293-305 Sociedad Geológica Mexicana, A.C.

DOI: https://doi.org/10.18268/BSGM2018v70n2a2

Available in: https://www.redalyc.org/articulo.oa?id=94362562003

How to cite Complete issue Scientific Information System Redalyc More information about this article Network of Scientific Journals from Latin America and the Caribbean, Spain and Journal's webpage in redalyc.org Portugal Project academic non-profit, developed under the open access initiative http://dx.doi.org/10.18268/BSGM2018v70n2a2 P. 293‒305 VOL. 70NO. 2 BOL. SOC.GEOL.MEX.2018 nos Aires, Argentina, 8000. Bue Blanca, Bahía 7.5, km Carrindanga La a Camino Instituto Argentino de Oceanografía, Salvador Aliotta nos Aires, Argentina, 8000. de Geología, San Juan 670,Bahía Blanca, Bue Universidad Nacional del Sur, Departamento Ester Farinati Guerrero, México, 40323. enda San Juan Bautista S/N, Taxco el Viejo, ExHaci Superior de de Cienciasla Tierra, Universidad Autónoma de Guerrero, Escuela [email protected] Catalina Gómez-Espinosa Catalina ridges, Argentina,SWAtlantic deformis Buccinanops Predatory evidence inthehigh-latitudecold-water gastropod Manuscript August accepted: 9,2017. Corrected manuscript received: May 24,2017. Manuscript received: January 25,2017. Gómez-Espinosa , Ester - - - (King,1832) from the Holocenelittoralsand Farinati Holocene, SWAtlantic. repair,high latitude, gastropod, Keywords: Predation, shell but effective predatory activity. alow because of maybe interpreted ate value for prey effectiveness (0.37) predation (0.16), andthe intermedi (0.06), the lowquency frequency of value observed for shell repair fre Argentinean low littoral coast. The deformis (A. Milne-Edwards, 1879)and tified between predatory-prey interaction was iden frequency andpreyeffectiveness. A predation, repair scar frequency of to evaluate durophagouspredation: ators. were Three parameters used activitybydurophagous pred tor of repaired shellasanindica traces of Sublethal damage was studied using as lethal and sublethal damage.agy duroph observed direct evidence of cold-waters andhighlatitudes. We hemisphere,a typicaltaxon is of 1832), endemicinthe southern Province. from the Argentinean Malacological activityona Holocene gastropodtory studyIn this investigates the preda ABSTRACT , Salvador duringthe Holocene on the Buccinanops deformis Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Danielethus crenulatus Aliotta (King, B. B. ------

1832) es una especie que habita en aguas 1832) esunaespecie que habita en aguas Argentina. del Holoceno de la Provincia Malacológica depredadora sobre unaespecie de gasterópodo la actividad En este estudio se investiga RESUMEN Holoceno, SOdelAtlántico. latitudes altas, gasterópodos, reparación deexoesqueletos, Palabras clave: Depredación, pero efectiva actividaddepredadora. se interpretaron comoresultado de unabaja medio dela efectividad de la presa (0.37) (0.06) yelvalor reparación del exoesqueleto de la frecuencia (0.16) y dedepredación las costas litorales argentinas. Elbajo grado 1879) y Danielethus interacción depredador-presa entre elcangrejo y laefectividad de lapresa. Se identificó la dación, la frecuencia decicatrices reparadas la frecuencia durófaga: dedepre depredación tres parámetros para de la la evaluación de este tipo de actividad. Se emplearon reparadas como un proxyexoesqueletos fue inferido de los utilizando la evidencia daño letal como subletal. El daño subletal directala evidencia dela durofagia tanto en Hemisferio Sur. Este estudio está basado en frías en latitudes altas, endémica del B .

deformis Buccinanops crenulatus durante elHoloceno en (A.Milne-Edwards, /2018

deformis (King, 293 -

Durophagy in a high-latitude Holocene gastropod from Argentina ABSTRACT Durophagy in a high-latitude Holocene gastropod from Argentina INTRODUCTION / LOCATION 294 294 1. Introduction contributes tonatural selection (Vermeij, 1987; 2007) basedonitsrole as aputative factor that intensively(Skovstedthe 1980s studiedsince mechanismfor metazoan evolutionbeen has predation as the main importance of The nepee ae ntedge f earo the repair of of the degree based on interpreted predationUnsuccessful orsublethal damage is mechanical agents (Vermeij action, and(3)shellscattering bycurrents orother by shells non-predatory (2) the destruction of 2001; Zuschin either before or after burial (Zuschin and Stanton, forces taphonomic towards resistance the shell’s differential of effects the (1) of because pretation overlooked asit has been considered a misinter shellbreakageThe caused by durophagy hasbeen (see Kelley andHansen,2003). are several reportsthe literature in aboutthat topic on gastropodsby wasmainly drilling, and there In the fossil records, evidence shows that predation predation (Kowalewsky, 2002). durophagous used asanindicatorcommonly of that maybe considered asdirect evidence andis last twoDietl, 2003). The methodsleave traces to leave preservable evidence (Alexander and two are methods first unlikely The Skelton, 1993). pre-ingestive breakage, and (Harper andcrushing insertion andextraction, boring followed by the whole organism, methods: the ingestion of It isconsidered that there are four molluscivory ton isdefinedasdurophagy (Aronson, 2001). Predation protectedon organisms by a hard skele prey (Chattopadhyay andBaumiller, 2010). their leave notrace or destroythe hard partsof ton (Vermeij, 1987),whereas mostpredators either preya biomineralizedpossessing organisms skele on of predation evidence direct find to possible foundtogether (Molinaro because predatorsmainly and prey are rarely in the fossilrecord (Stuart and Greenstone, 1990), be estimated to interactions interspecific difficult the most 1996). Nevertheless, predationone of is Kelley andHansen,1993;Jablonski and Sepkoski, / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín et al. , 2003;Mapes et al. et al , 2013). It is only , 2013).Itis ., 1989). et al. , 2010), et al ., - - - /2018 2. Location (2003). gastropods waspublished by Alexander and Dietl shell-breaking predation in ing fossilrecords of and Peel, 2010). A comprehensive review regard and Dietl, 2003; Skovsted tropods’ shell breakage (Vermeij, 1987; Alexander focused onsublethal damage as they report gas publicationsMost regarding fossilrecords are al. (Staffordshell the on patterns ments,color as or surfaceorna growth lines, visible of as disruptions derived Scars 2011). from repair processesare visible of as disruptions Leighton, and (Stafford frequently used indicator for crushing-predation damaged shell. Recently this hasemerged as a gical studies indicate that the Holocene marine altitude shoals.and sand islands Several geolo​ tidal channels separated by low a dense net of byformed is thisestuary coastalregionThe of 1). (Figure W 62°20ʹ S, 38°45ʹ Location: Argentina. BuenosAires Province,Blanca estuary, southof the Bahia ridges located at theinnerregion of Samples were collected at the Holocene sand shell been estimated for fossilized repaired gastropods. not has effectiveness prey 2010), Baumiller, and Moreover, except for one report (Chattopadhyay to arctic latitudes (Alexander and Dietl, 2003). for bothfossiland recent locatedmollusks at Boreal describing repair frequencies is scarceInformation (Ogaya, 2004;Dietl breakage is distinguishedby apeculiar pattern ent regarding severity, size, and position, and the markedlydiffer is by crabs caused breakage Shell the Stafford described by evidence has been formally type of by other carnivorous gastropodsand crabs. This al be used to identify durophagous predation (Oji can lip shell external the on inflicted damage The ., 2003), although the lip may alsobe damaged , 2015). Caedichnus et al. (2015) on an ichnofossil assigned to assigned ichnofossil an on (2015) ichnogenera. et al. , 2010). et al ., 2007;Lindström et et - - - - 3. Targettaxa Figure 1 particular features onthe Bahía Blancaarea transgressive–regressive cycleimprinted its has Buccinanops deformis durophagous predation in the Holocene gastropod In thiswork we present documented evidence of pumice (AliottaandFarinati, 1990). quartzite pebbles, calcrete, siltstone, and tion of (approximately 30%) as well as a smaller propor whole shellsandtheir fragments percentage of high a having fine to medium from range which size sands Sand-shell ridges are composed of along thecoastline(Farinati andAliotta,1987). theyarea narrow in arranged sub-parallel strip are mainly found at the estuary’s inner region and ing 8–10mabove meansealevel. ridges These sand-shellridgesobserved at heightsrang series of produced highest Holocene transgression The a (Aliotta Mapof theBahiaBlancaEstuaryhighlighting thezone characterized bysand-shell ridgesand thelocationof thestudyarea. et al ., 2013). (King, 1832). Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín - - ered feeders, as carrion a proposal based on the Nassariidae familyhave been consid of Members 2A). (Figure the spire than larger times five almost is – 51 mm) having 5 to 5½ whorls. body The whorl size (24 intermediate depressed. Shell height is of a callus initsrear end whereas the outer lip is having inner lip isangled siphonal canal. The narrow opening. shell has awell-developed The with a low anddepressed spire and a subelliptical B. deformis Argentina (NuñezandNarosky, 1997). Santo, Brazil to Golfo SanMatías, Patagonia, the SW fromAtlantic Ocean, spanning Espiritu high latitudes endemic to it is a typical taxon of the Bahia Blanca estuary, andbecause ridges of mollusk foundand conspicuous inthe sand-shell specieswasselected because it is an abundantThis Recent (Aguirre, 1993). recordthe UpperMioceneto comprising (Caenogastropoda, Nassariidae) has a geological The Buccinanops possesses a smooth, wide,a smooth, possesses shell, andthick eu (’rin, 1841) (d’Orbigny, genus /2018 295 295 -

Durophagy in a high-latitude Holocene gastropod from Argentina LOCATION / TARGET TAXA Durophagy in a high-latitude Holocene gastropod from Argentina TARGET TAXA 296 296 bar 5mm. Side viewshowing Figure 2 /

Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Buccinanops deformis Caedichnus sp.repairedJ–K. Sideviewshowing shell.(PI-UNS3184 –3191); punctures 3192 –3193).Scale (PI-UNS (King, 1832)A1–A2. Non-preyed shell(PI-UNS 3183).B–K. Failed durophagous B–I. predation attempts. /2018 4. Materialsandmethods overlooked [exceptions are the studies conducted in coolandcoldwaters have remained largely Kowalewski,2002). Conversely, living mollusks 1967; Vermeij, 1977a;Kowalewski ing intemperate and tropical waters ( (whether living or fossil) isfocused onspeciesliv describing predationinformation on mollusks represent cold water inhabitants, whereas most Argentinean Malacological Province asit might the important to studyIt is an endemicgenus of (Pastorino,waters (2–20mdeep) 1993). toms onintertidal orinfralittoral zonesin shallow Regarding habitat, (King, 1832). ences describingthe bydiet consumed bivalves (Averbuj and predator) thatand living dead crabs fedon been reported as acarnivorousspecies (scavenger Only or undocumentedsources(González posal are scarce and obtained from unpublished relatively unknown asdata supporting this pro this familyare Nevertheless, the trophic aspects of the diet of examined underadissecting stereo-microscope. and sublethal predation (repairedwere scars) lethal After cleaning the shells, the evidences of out. carried the specimens corresponding to selective sampling of pod and24bivalve species (Aliotta gastro assemblageMollusk includes 23speciesof Farinati(1994). and (1993) Aguirre byestablished taxonomy the on based identified were shells All anddecapods.cirripedia, in thissamplewere chiton fragments, bryozoans, prised by bivalves andgastropods. Other remains assemblagesdried. The were almost entirely com sieves.mesh sampleswere The thenwashedand sievedwere mm samples Five throughbulk 4 field by Cadée(1999)and Bigatti Buccinanops deformis Buccinanops cochlidium Nassarius et al. Buccinanops genus (Harasewych, 1998). , 2012).There are norefer gastropod (King, 1832) was (Dillwyn,1817) has et al. dwell in soft bot dwell (2009)]. et al. et al. et al e.g. B. deformis B. deformis , 2013). A , Carter, ., 2011). , 1997; Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín ------sample (Equation 1). specimensinthe divided by the total number of is lethal injury (LTF)displayingimens evidencesof method Frequency spec (Kowalewski, 2002), where the number of Taxon Lower the using by calculated was predation of Frequency prey effectiveness. predation, repair scar frequency,frequency of and wereThree parameters used to evaluate predation: on repaired scars shells. occlude the observation of were considered as these featurestaphonomic predation. Neither extensively abraded nor corroded shells of effects the evaluating for selected Only those relatively intact specimens were thatsampled at was 38°45ʹS,dated62°20ʹ Wwas A the monospecific sample of Argentina. Nacional del Sur (PI-UNS), Bahía Blanca, Universidad The the Paleontological Collection of Specimens described in thiswork are housed in predation. evidence of displaying specimens shells. TT is the number of displaying specimens punctures andrepaired of number the is effectiveness. TF Prey is PE Where punctures andrepaired shells. of using Equation 2andby considering the number quantified was 2002), (Kowalewski, failure ator pred measure to parameter a effectiveness, Prey lation (Alexander andDietl,2003). sublethal attackswithin apopu the frequencyof per shell method because the latter underestimates sample.preferredmethod was This over thescar in the individuals divided by the total number of repaired is scars 2003) where the total number of the repairfrequency scar (Alexander andDietl, scarper shell method was usedto calculateThe specimensinthesample. N isthetotalnumber of specimens displaying lethal predation evidences. Where:the taxon Kis target. the numberis D of LTF =Dk/Nk(Equation 1) PE =TF/TT(Equation 2) B. deformis /2018 gastropod 297 297 - - -

TARGET TAXA / MATERIALS AND Durophagy in a high-latitude Holocene gastropod from Argentina METHODS MATERIALS AND METHODS / Durophagy in a high-latitude Holocene gastropod from Argentina RESULTS 298 298 5. Results 3198 –3199).Scale bar 5mm. displaying twopunctures locatedatpenultimate (D1)and body(D2)whorl. E–FSideviewshowing apuncture onthebody whorl (PI-UNS Shell punctureD. by damage shows view Side D-F. 3196); – 3495 (PI-UNSwhorl body on the fracture an embayment shows view Side whorl and apartialmissing shellaperture. A2shows continuation asanembaymentfracture inthebodywhorl3194); B–C. (PI-UNS Figure 3 from Universidad NacionaldelaPlata (UNLP). Laboratorio de Tritio Radiocarbono(LATYR)y with C the samples. in predominant are height of mm 35 to mm 23 540 extractedfrom five from samples. Organisms bulk of amount total A 5690±70years BP.of datingThe process for the shell indicates a / Lethal damageasaconsequence of durophagous predation A–D. Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín 14 . Radiocarbon dating was performed at . Radiocarbondatingperformed was B. deformis shells were shells 14 C age /2018 Caedichnus trace fossil. evidence match the Praedichnia these types of agreement with Ekdalethe (1985), classification of In 3E–F). 2J–K, (Figures punctures exhibit shells al as identified is trace of with damage on the apertural lip and thiskind Most the observed predation evidence correlates 4B). of 3, (Figures 56 on lethal was damage whereas 4A) 2B–K, (Figures damages sublethal the latter show evidence of tion evidence, 33of preda display 89 examined, shells 540 the From ., 2015) (Figures 2B–I, 3A–D, 4A–B). Very few Very4A–B). 3A–D, 2B–I, (Figures 2015) ., isp.A1.Sideviewshowing puncture onthebody Caedichnus isp (Stafford isp et - 5.1. LOCATION ANDTYPEOFINJURIES bar 5mm. Scale (PI-UNS3201). the lip from back farbreakage shell deep a showinginjuryobservedas view lethal Side a B. 3200); (PI-UNS whorl body on the damage andrepair showingbreakage deep fracture asconsequence A.Sideview of crabpredatory activity Figure 4 h rqec f The predation frequency (LTF)of is 0.10 (95 % CI the whole-body whorl near the aperture (96.9%) Repairing observed on the shells mostlyoccuralong lar fracture (Figures 2B, 2D–F, 2H). irregu to 4A–B) 3A–C, 2C, (Figures embayment apertural lip region (94.3%) ranging from large outer the at is injuries Frequently,of location the (Figure 2D). the of repaired show Some zigzag-breaking scars patterns 2I–K). (Figure shells two only on observed were punctures and 4A) 2B–H, (Figures evidence withinthe sublethal damage category Repairedwereshells injurieson abundant the most (Figures 3E–F). 4B) andpunctures areobserved only ontwo shells 3A–C, (Figures category damage lethal the within Shell breakage bydurophagythe main evidence is (P is 0.06 (95 % CI ± 1.04) whereas prey effectiveness repaired scars ± 1.17). The observed frequency of E

) is0.37(95%CI±2.58). Caedichnus isp. shellsexhibiting alargeembayment Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín - 6. Discussion atory damage on the shell from abiotic damage 1998). However,possible itis pred to distinguish assemblage (Harper the paleoecology of dation intensity leading to the misinterpretation pre mayfossilpret mask recordeffects in some as inter to difficult is predation that recognized is It truncated apicalendontheshells. removed apex or 2D). There is noevidence of (Figure whorl second the on observed is 3.1%) to (equivalent scar one only and 2E–2I) 2C, (Figures tures toattack thickshellswithanarrow aperture. predator mustspecialized possess physicalstruc insertion andmolluscivoryextraction. Thus, the that the predatoron used amethod consisting mation on their interaction. In thiscaseit is shown the preytivity on skeletons provides useful infor site-selec Predators’ injuries. sublethal and lethal predation on of breakage evidenceis the most common Apertural to growth lines. are thinner thantheoriginalnotparallel and scars of the shell’s thickness. In the repaired area shells are modification subsequent the with continued growth and repair individual’s the breakage, by When shellsexhibit sublethal damage caused which correlate withlethalandsublethal attacks. predation, including breakage and puncture, The Klompmaker, 2009). important as well ( aperture (Ogaya, 2004). Compactionmaybe either a broken orunbroken shell the criterion of can be distinguished incrushedshellsbased on the remains. Similarly, breakage of the cause of are thissmooth, indicates transport or rework the edges damage caused by water currents and,if durophagy rather than ably the consequence of a disrupted apertural possesses margin, it is prob (Alexander and Dietl, 2003). For a instance, shell if B. deformis shells exhibit damage caused by B. deformis e.g. , Zuschin andStanton, 2001; asdemonstrated by /2018 et al 299 299 ., ------

Durophagy in a high-latitude Holocene gastropod from Argentina RESULTS / DISCUSSION Durophagy in a high-latitude Holocene gastropod from Argentina DISCUSSION 300 300 EFFECTIVENESS 6.2. REPAIR SHELL FREQUENCY AND PREY 6.1. FREQUENCY OFPREDATION mligta lotnn fthe gastropods were implying that almost noneof predationobserved, is a relatively low frequency of the predationtion is frequency index. Inthisstudy indirectthe only approachout itsestima tocarry Predation intensity cannotbe directly inferred and that predatorswere abundantor and successful, Repaired shell frequency islow, implying either assuccessful(Kowalewski,interpreted 2002). tially their consume prey. Inthiscase, predation is predationpredation oras bythatorganisms par sublethal as failed damage may be interpreted crushing intensity (Vermeij Repairedmeasurecannot beusedas shells of defenses. prey’s the and effectiveness predator between link the reflects it as 2010), Peel, and (Lindström tors interaction between gastropodsand their preda an assemblage isa simple meansto estimate the repaired shell frequency within estimation of and Baumiller, 2010;Klompmaker, 2011). The ities (Alexander andDietl, 2003; Chattopadhyay used indicator for durophagouspredatory activ a commonly repairedis scars frequencyThe of (Allmon those reported for tropical Paleozoic communities lowThe predation values are also comparable with temperate latitudes (Vermeij, 1983). sure is greater in tropicswhen compared to more waters. that It has been suggested predation pres tion ongastropodslowis in coolandtemperate idea thattent with the generally accepted preda Nevertheless, alowpredation frequencyconsis is Molinaro attackssuccessful by predators (Kowalewski, 2002; increased attacks (predation) or decreased of because repairedmay also beinterpreted scars When studying predation, the frequency of predation does not have a fossil record. this type of the shells were crushed by predators, that some of attacked by we consider durophagy. Although, if / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín et al. et al. , 1990;Gordillo andArchuby, 2014). , 2013). et al. , 1980) because ------/2018 6.3. CENOZOICSHELL-BREAKINGPREDATORS oeta fpotential of al. out in tropicalstudies carried zones ( compared to that obtained on field and laboratory preyobtained The effectiveness low is value (0.37) potentialanti-predatory selection. measure of predationunsuccessful provides a conservative Vermeij (1983) considered that the incidence of as anti-predatorymechanism (Vermeij Prey effectiveness indicates the shells’ performance thatsuggest predators were scarce. is observed in the tropics) (Vermeij repair frequencies regarding latitude (the highest al (Schindel ineffective or scarce were predators elkon(emi,17b.Mmeso the well known(Vermeij, of 1977b).Members diet but their predatory activity ongastropods is Vermeij, 1987).Brachyuran crabshave a broad Buccinanops are the observedthe injurieson punctures unknown,some of shell’s the of causes the case this uncertain (Kowalewski,traces is 2002), andin through predator a of identification the Although (Alexander andDietl,2003). crustaceans, stromatopods,octopuses andsome palinurid lobsters, brachyuran crabs, pagurid seal, and walruses. Among the invertebrates are sea otters, birds, marine coastal some fishes, ost Vertebrate includespecialized shell crushers tele (2007, 2010). extensively byChattopadhyay and Baumiller nations for tropical environments were discussed activity. expla predatory Alternative effective but alow as the consequence of maybe interpreted Holocene sand shellridges in Bahia Blanca. This effectiveness(0.37) on prey of value intermediate an and we argue that a low shellrepair frequency (0.06) Although it is not possible to confirm this proposal, antipredatory system. gastropodsare notforced to develop acomplex the predation predation frequency.low, is of If case,this prey effectiveness may consequence a be ,18) osdrn h aito fgastropod ., 1982).Consideringthevariation of , 2010;Paul B. deformis shellsmaybe causedby crabs (see B. deformis et al ., 2013). This indicates the low., 2013).This is predominant is at least at the against itspredators. against In et al e.g. et al. ., 1980),we , Dietl, , 1980). et et - - traces were made by ascissor-like weapon, such that the skeleton suggests these of opposite sides on scars complementary presence2010). The of predation by crabs(Dietl the consequence of been observed damage has as whorl.type of This injuries lip the body observed as large embayed apertural fracture of exhibit 4B), 2C, (Figures on inflicted damage shells, either lethal (Figures 3A–C, 4A) or repaired the of Some distinguishable infossil records. caused bythem are well characterized andclearly al. Dietl 2004; Ogaya, 1985; Hughes, and Lawton Vermeij,1981; 1978;BertnessandCunningham, extensively described (Vermeij, 1977b; Zipserand Predation on gastropod shellsby crabshasbeen (Lawton andHughes, 1985). crushing, apertural breakage, andapex removal at least three attack gastropods: methods against use Crabs Feldman,2010). and Schweitzer 2008; as anadaptation todurophagy(Dietl and Vega, lipeds. larger The one contains a crushing tooth che right and left the between size in differences Platyxanthidae family are distinguishedbymarked lddta rb fsmaller size prefer thin shells, cluded that crabsof by Bertness andCunningham (1981). They con predationdefenses against by crabswasevaluated dation. Inrecent gastropods,and shellthickness maybean evolutionary as adaptation pre against Buccinanops such asmollusks(Walker andBrett, 2002). thattic feeders eatoccasionally hard-shelled preys, currently considered as generalist and opportunis Nevertheless, portunidsandxanthids crabsare Feldman,2010). and SchweitzerVega, 2008; and and crabs (Dietl crushing tips teeth typicalof molariform sharp display that 5), (Figure dactylus same sample. cheliped The remainscorrelate with Platyxanthidae) along with gastropods inthe chelipeds crab’sEdwards, 1879) species (Decapoda, Brachyura, of presence the fragments of the is crabs by Another fact supporting predation on 1997). (Kowalewskicrab’s3D–F) (Figuresa claw as , 2010) and injury types and breakage patterns typesandbreakage, 2010) and injury patterns possess a particularly possess thickshell, Danielethus crenulatus (A.Milne- Buccinanops Buccinanops et al et al Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín et ., ., - - - -

Acknowledgements 7. Conclusions sublethal injuries. trace on breakagepredationcommon themost is Apertural predation isnotcompletelyexcluded. tropod small-size vulnerability against crabs, but by large-size crabs. Athick shell provides lessgas shells, andthepercentage similar foris predation predation bymiddle-size low crabsis onthick preyed onthinshells, whereas the percentage of also reported that small- and middle-size crabs but larger crabs have no preference. Palmer (1985) ral mrvdteqaiyo thepaper.greatly improved thequalityof and Gérard Breton for their comments, which ects. We expressour sincere to Greg thanks Dietl and PIP Nº5538 and Nº 00699 (CONICET) proj 24/H107 (Secretaría Técnica, UNS) de Ciencia y research This wassupported by PGI 24/H099 and (predator) andthe distinguished between the A Holocenepredatory-preyinteraction maybe crabs. whorl may be attributed to predatory activityby Caedichnus largeThe embayment fractures, corresponding to alow but effective predatory activity.the result of as shell ridges in Bahia Blanca may be interpreted tiveness (0.37) on effec prey for value intermediate the with along lowThe value for shellrepair frequency (0.06) predation pressure at temperate latitudes. consistent withproposalstion is lowsuggesting durophagous preda lowThe frequency (0.16) of lip.and specifically at the external by Most injuries, bothlethal and sublethal, displayed B. deformis isp., observed on the gastropod’s body gastropod’s the on observed isp., Buccinanops deformis shellsare located at the body whorl B. deformis B. deformis Danielethus crenulatus at the Holocene sand gastropod (prey). either for lethal or /2018 crab 301 301 - - - -

DISCUSSION / CONCLUSIONS / Durophagy in a high-latitude Holocene gastropod from Argentina ACKNOWLEDGEMENTS Durophagy in a high-latitude Holocene gastropod from Argentina REFERENCES 302 302 References Alexander, R.R.,Dietl, G.P., fossil 2003,The Aguirre, M.L.,1993,Typeof specimens Bigatti, G., Sánchez, C.J.M., Miloslavich, P., Bertness, M.,Cunningham,C.,1981,Crab Averbuj, A.,Palomo, G., Brogger, M.I., AronsonR.B., marine 2001, Durophagyin Allmon, W.D., Nieh, J.C., Norris, R.D., 1990, Aliotta, S., Ginsberg, S., Spagnuolo, J.O., Farinati, of Stratigraphy 1990, E., Farinati, S., Aliotta, / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín predatory neogastropod (Gastropoda: of behaviorFeeding P.E.,Penchaszadeh, 2009, andEcology,marine biology 50,213–230. experimental architectural defense: Journal of shell-crushing predation andgastropod Aquatic Biology, 17,261–268. Patagonia,1817) from Northern Argentina: eodo shell-breaking predation on record of Argentina: Ameghiniana,30,23–38. marine gastropods fromQuaternary the nassariid Penchaszadeh, P.E., 2012,Diet and feeding of Blackwell Science, 393–397. P.R. (eds.), PalaeobiologyII: Oxford, UK, organisms, Palaeontology, 33,595–611. Gastropods sincethe Late Cretaceous: Turritelline Drilling andPeeling of 45, Sciences, Earth 194–208. American South of coastalevolution: Journal indicators of and paleontologic system: Stratigraphic in a South Atlantic estuarine Quaternary E., Giagante, D., Vecchi, L.G., 2013,Late 94(4), 353–360. Blanca Estuary, Argentina: Marine Geology, the Bahia Holocene sand-shellridgesin Plenum Press, 141–176. Fossil the Record: NewYork, in Kluwer Academic/ Interactions Predator-Prey P.H., Kowalewski, M.,Hansen,T.A. (eds), marine bivalves andgastropods, Adelomelon ancilla in Buccinanops cochlidium Briggs, D.E.G, Crowther, (Lightfoot, 1786): A (Dillwyn, in Kelley, /2018 Chattopadhyay, D., Baumiller, T.K., 2007,Drilling Carter, and R.M.,1967,Onthe biology Cadée, G.C., 1999,Shell damage and shell repair aiai E, 94 Mcooucs (Gastropoda Micromoluscos 1994, the E., Farinati, of Paleoecology 1985, A.A., Ekdale, l’Amérique dans Voyages 1841, A., D’Orbigny, Dillwyn,L.W., descriptive 1817,A catalogue Dietl, G.P., Durham,S.R., Kelley, P.H., 2010, shell- Specialized 2008, F.J., Vega, G.P., Dietl, Chattopadhyay,Effect Baumiller,T.K.,D.,2010, Experimental Marine Biology and Ecology, Experimental Marine Biology apredator: Journal of in the presence of drilling behavior of of under threat: An experimental assessment Paleoecology, 4,29–65. Mollusca: Paleogeography, Paleoclimatology, bivalved somepredators of of paleontology 41, 149–161. Sea Research, King George Island: Journal of in the Antarctic limpet communities: Nautilus, The 123,159–165. Volutidae) in Patagonian benthic 303–315. Blanca, Argentina: Ameghiniana,31, y Bivalvia) del Holoceno del Área de Bahía Palaeoecology, Palaeoclimatology, 50, 63–81. marine endobenthos: Palaeogeography, Paris, France, P. Mollusques: Berdtan. 3, Part 5, méridionale. 581–1092. to the synonymy:J.London, Arch, andA. Linnæan method;withparticular attention recent shells, according arranged to the of 174–184. Paleoclimatology, Paleoecology, 296, thein fossil record: Paleogeography, predation by shell-wedging gastropods bivalve Shell repair asareliable indicator of Biological Letters, 4,290–293. breaking crab claws inCretaceous seas: America: HistoricalBiology, 22(4),367–379. Cenozoic molluscsfrom North studyof durophagy ondrilling predation: A case of 352, 257–266. Nacella concinna Nucella lamellosa from

Kelley, P.H., Hansen,T.A., fossil 2003,The Kelley, P.H., Hansen,T.A., the 1993,Evolution of Jablonski,D., J.J. Sepkoski, jr., 1996,Paleobiology, T., Palmer, G.T.W., Forsythe, E.M., Harper, Harper, E.M.,Skelton, P.W., Mesozoic 1993,The Harasewych, M.G., 1998,Infraorder and Archuby,Live-Live S.,F.M., 2014, Gordillo, De F., Scarabino, A., Carranza, S., González, aiai E, lot, . 18, Malacofauna 1987, S., Aliotta, E., Farinati, Paleontologia: Santa Cruz, Bolivia, 574–585. Actas 4 de BuenosAires, Argentina (abstract), proximidades de Punta Alta, Provincia litoral en las holocena de un depósito M., Hansen,T.A. (eds.), Predator-Prey gastropods, drilling predationbivalves on and record of Palaios, 8,358–375. Escalation: of theHypothesis Evaluation of Naticid Gastropod Predator-PreySystem: An Ecology,pattern: 77,1367–1378. ecological communityecology, andscalesof 352–360. boring predators: Palaios, 13, importance of revolution: preservation statethe masks 1998, Taphonomy andthe Mesozoic marine Scripta Geologica,SpecialIssue2,128–153. Marine Revolution bivalves: andepifaunal Melbourne, AU, Csiro Publishing, 819–845. Nassariidae, Synthesis: (eds.), Mollusca:the Southern Family in Neogastropoda, Polonica, 59,429–442. Clam the Patagonian Assemblages: Case of The Live-Dead Interactions inMarineDeath Sciences, 6,247–259. Aquatic Pan-Americanestuary: Journal of andthe Río de la Plata continental shelf Buccinanops benthic gastropods: the genus of patterns Mello, C.,Ligróne, A.,2011,Coexistence Beesley, L.P., Ross, G.J.B., Wells, A. Venus antiqua th Congreso Latinoamericano Congreso de (Nassariidae) the inner Uruguay in Kelley, P.H., Kowalewski, : ActaPaleontolologica Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín in

Klompmaker, A.A.,2009,Taphonomy ondrill- King, P.P., Cirrhipeda, 1832,Descriptionof Mapes, R.H.,Landman,N.H., Cochran, K., Lindström, A.,Peel, J.S., 2010,Shell repair and Foraging 1985, R.N., Huges, P., Lawton, Kowalewski, M., K.W.,Flessa, Marcot, J.D., 1997, Kowalewski, fossil M., 2002, The record of Klompmaker, A.A.,2011, Drilling and crushing The Netherlands:The Palaios, 24,772–779. fromPliocene mollusks Langenboom(Mill), hole predationof intensitiesandpaleoecology Broderip”]: W.J. Zoological Journal, 5,332–349. by [“assisted Fuego del Tierra andthe coast of Magalhaens of South America,includingtheStraits of and 1830insurveying coasts the southern employedand Beagle between the years 1826 Adventure H.M.S. of officers the by formed Conchifera and Mollusca,inacollection Kluwer Academic/Plenum Press, 113–139. York,FossilRecord:New the in Interactions h Nw aeoi Rgo: aao, 25, Palaios, 597–610. Region: Caledonia of New the significance and naturally submerged taphonomy Early Goiran, C., Richer, B., Renfro, A.,2010, Geosciences, 8,541–550. gastropods from Bulletin of England: in Jurassicshell form pleurotomarioid 143–154. Progress Series, Marine Ecology theory: 27, littorea on the gastropods thecrab behaviour of Polonica, 42,497–532. ecological implications: Acta Palaeontologica lingulid brachiopod: Paleontological and a recent Predatory inthe shells of scars Society Papers, 8,3–42. Predation: Paleontological fossilRecord of in analytical methods, predation: An overview of theNetherlands: Lethaia,44,429–439. of predation fromon scaphopods the Miocene Kowalewski, M.,Kelley, P.H. (eds.), The : comparisons with optimalforaging: comparisons Nucella lapillus Cancer pagurus Nautilus /2018 shellsfrom and feeding Littorina 303 303

Durophagy in a high-latitude Holocene gastropod from Argentina REFERENCES Durophagy in a high-latitude Holocene gastropod from Argentina REFERENCES 304 304 Molinaro, D.J., Stafford, E.S., Collins, B.M.J., Collins, E.S., Stafford, D.J., Molinaro, Skovsted, C.B., Brock, G.A., Lindström, A.,Peel, The 2010, R.M., Feldman, C.E., Schweitzer, Schindel, D.E., Vermeij, D.E., Zipser, E.,1982, Pastorino, G., taxonomic 1993,The status of Paul, G., Das,Bardhan, A., S., Mondal,S., shell of value Adaptive 1985, A.R., Palmer, Oji, T., Ogaya, C.,Sato, T., shell- 2003,Increase of the Ogaya, C.,2004,Presence or absenceof Nuñez, C., Narosky, T., 1997, Cien caracoles / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín with Programs: Denver, USA,45,756. AmericaAbstracts Geological Society of predation?understanding (Abstract), repair frequency be a usefulproxy towards 2013, Peeling out predation intensity:can Barclay, K.,Tyler, C.L.,Leighton, L.R., Letters, 3, 314–317. mollusc:Biology shell repair in anepibenthic failed durophagy and Cambrian record of S.J.,J.R.,Paterson,EarlyFuller, 2007, R.K., 167–182. 25, Palaios, time: geologic trough Mollusca Decapoda (Crustacea) as predators on 729–740. Paleontology,56, of JournalTexas: Central North- the Pennsylvanian gastropods of among fractures shell repaired Frequencyof Veliger,Nassariidae): The 36,160–165. Buccinanops Database: Malacologia,56,193–213. and ComparisonwithaRevisedGlobal Gastropodsfrom the Indian Subcontinent 2013, PredationRecenton Turritelline crabs: Veliger, 27,349–356. shells onvulnerability to and preference by variation in fragmentation: Paleobiology, 29,520–526. crushing predation recorded in fossil shell Paleontological Research, 8,311–324. (Mollusca: Gastropoda: Trochidae): breakage induced bydurophagy in shell aperture: a criterion to identify shell 158 p. argentinos, BuenosAires, Editorial Albatros, ’rin, 81 (Gastropoda: 1841 d’Orbigny, Thais lamellosa Thais efcs fthick of effects : Umbonium in /2018

Stuart, M.K., Greenstone, M.H.,1990,Beyond M.P., Gingras, G.P., Dietl, E.S., Stafford, Vermeij 2011, L.R., Leighton, E.S., Stafford, Zuschin,M., Stanton, R.J., 2001,Experimental Walker, S.E., Brett, C.E., 2002,Post-Paleozoic Vermeij, G.J., Zipser, E.,Dudley, E.C.,1989, Vermeij, G.J., Zipser, E.,Dudley, E.C.,1980, Vermeij, G.J., 1987,Evolution andEscalation: An Vermeij, G.J., 1983,Traces andtrends of Vermeij, G.J., 1977b, Patterns incrabclaw size: the Vermeij, G.J., Mesozoic 1977a,The Marine otns naso the Entomological contents: of Annals stomach predator of identification for assay ELISA: a rapid, sensitive, specific immunodot 87–102. on the Gastropod Ichnos, Shell Aperture: 22, Ichnogenus Representing Predatory Attack 2015, L.R., Leighton, 305, 123–137. Palaeoecology, Palaeoclimatology, gastropod assemblages: Palaeogeography, for estimatingpredation crushing in A newoldtechniqueanalysis: crushing 161–170. Taphonomic Palaios, Interpretation: 16, Shell Strength and its Measurement of Paleontological SocietyPapers, 8,119–194. The Predation: of RecordFossil The (eds.), revolution?, Mesozoic andCenozoic marine predatory inmarine predation:patterns Was there a in recent pelecypods:Veliger, 32,266–273. drilling predationand unsuccessful Successful 6, 352–364. drilling in terebrid gastropods: Paleobiology, Predation intime and space:peeling and Princeton University Press, 527p. life: New Jersey,of ecological history USA, animals: Paleontology, 26,455–465. predation, withspecialreference tobivalved 26, 138–151. Systematic crushing: Zoology, geographyof and Grazers: Paleobiology, 3,245–258. Revolution: Evidence from Snails, Predators America,83,1101–1107. Society of in Kowalewski, M.,Kelley, P.H. Caedichnus , aNew Zuschin, M.,Stachowitsch, M.,Stanton, R.J., 2003, Earth ScienceReviews, 63,33–82. and ancientmarineenvironments: modern in shellfragmentation Patternsand processes of Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín /2018 305 305

Durophagy in a high-latitude Holocene gastropod from Argentina REFERENCES