Palaeogeography, Palaeoclimatology, Palaeoecology · 1.2.3. Drilled Brachiopoda Drilling in Mesozoic brachiopods, especially those from the Triassic and Jurassic,is rare.Kowalewskietal

Palaeogeography, Palaeoclimatology, Palaeoecology 457 (2016) 342–359

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Palaeogeography, Palaeoclimatology, Palaeoecology

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Drill hole convergence and a quantitative analysis of drill holes inmollusks and brachiopods from the Triassic of Italy and Poland

Adiël A. Klompmaker a,b,⁎, Alexander Nützel c, Andrzej Kaim d

a Florida Museum of Natural History, University of Florida, 1659 Museum Road, Gainesville, FL 32611, USAb Department of Integrative Biology, Museum of Paleontology, University of California, Berkeley, 1005 Valley Life Sciences Building #3140, Berkeley, CA 94720, USAc SNSB-Bayerische Staatssammlung für Paläontologie und Geologie, Department für Geo- und Umweltwissenschaften, Paläontologie & Geobiologie, Geobio-Centre, Ludwig-Maximilians-Universität München, 80333 Munich, Germanyd Instytut Paleobiologii PAN, ul. Twarda 51/55, 00-818 Warsaw, Poland

⁎ Corresponding author at: Florida Museum of Natura1659 Museum Road, Gainesville, FL 32611, USA.

E-mail addresses: [email protected] ([email protected] (A. Nützel), kaim@twarda

http://dx.doi.org/10.1016/j.palaeo.2016.06.0170031-0182/© 2016 Elsevier B.V. All rights reserved.

a b s t r a c t
a r t i c l e i n f o
Article history:Received 11 March 2016Received in revised form 7 June 2016Accepted 9 June 2016Available online 14 June 2016

The history of predation is recorded primarily from drilling in Cenozoic invertebrates. Quantitative data are un-common from the Triassic, a period before the appearance and radiation of many known drill hole producerssuch as various gastropod families and octopods. We present quantitative evidence of drilling from the Late Tri-assic (Carnian) Cassian Formation of Italy and the Middle Triassic (Anisian) Lower Muschelkalk of Poland,documenting the first drill holes in Triassic brachiopods. A single brachiopod with a cylindrical, complete drillhole was found in a brachiopod sample from of Poland (drilling percentage = 0.3%, n = 365). The Cassian For-mation yielded drill holes in gastropods, bivalves, and brachiopods, indicating that more species are drilledthan was known previously. The minimum drilling percentage exclusive of incomplete drill holes of a samplefrom the Stuores Wiesen (Cassian Formation) is 1.7% (n = 116.5). Prey selectivity is evident: complete drillholes are primarily present in one gastropod species, Polygyrina lommeli (11.8% of specimens with a completedrill hole), whereas other common species were not drilled. Single drill holes in brachiopods are cylindricaland complete and may be predatory in origin. Multiple drill holes in mollusks are common, and drill holes areparabolic and often incompletewith a central boss, resembling the shape of drill holes produced by extant naticidgastropods. A survey of the Paleozoic literature showed that such drill holes are also present inDevonian and Car-boniferous brachiopods. However, naticids did not evolve until the Cretaceous sowe propose the term “drill holeconvergence” for similar-shaped drill holes produced by different organisms. The Triassic parabolic drill holes arenot caused by domicile-seeking or boring organisms. Instead, we favor a predatory origin of these drill holes, butwe cannot entirely rule out parasitism. Surveying other Triassic invertebrate assemblages should yield more evi-dence of drilling.

© 2016 Elsevier B.V. All rights reserved.

Keywords:BivalviaBrachiopodaGastropodaNaticidaeParasitismPredation

1. Introduction

1.1. Drilling predation and prey

The present study reports and discusses the occurrence of drill holesin marine invertebrate shells (gastropods, bivalves, and brachiopods)from the Upper Triassic (Carnian) Cassian Formation of Italy and fromthe Middle Triassic (Anisian) Lower Muschelkalk of Poland. Theserecords are one of the very few Triassic occurrences of drill holes, and,therefore, their analysis is pivotal for understanding the evolutionaryhistory of drilling predation.

l History, University of Florida,

Klompmaker),.pan.pl (A. Kaim).

Drilling predation is the best preserved and themost easily quantifi-able evidence of predation in the fossil record. Predatory drilling inmodern marine invertebrates is ascribed to flatworms, nematods,octopods, but most of all to gastropods (Kowalewski, 2002: Table 2).The shape of these drill holes can, by analogy, help to determine themost likely culprit of drill holes in fossil assemblages, especially duringthe Cenozoic when relatives of modern drillers were present. Giventhe abundance of shelly material in Cenozoic deposits, traces of gastro-pod predation have been studied extensively in a variety of marine in-vertebrate prey including bivalves and gastropods (e.g., Kowalewskiet al., 1998; Kelley and Hansen, 2003, 2006; Klompmaker, 2009;Chattopadhyay and Dutta, 2013; Klompmaker and Kelley, 2015),scaphopods (e.g., Yochelson et al., 1983; Klompmaker, 2011; Li et al.,2011), echinoids (e.g., Kowalewski and Nebelsick, 2003), ostracods(e.g., Reyment et al., 1987; Reyment and Elewa, 2003), annelids(e.g., Klompmaker, 2012; Martinell et al., 2012; Villegas-Martín et al.,

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2016), brachiopods (e.g., Leighton, 2003; Kowalewski et al., 2005;Baumiller et al., 2006), barnacles (Gordillo, 2013; Klompmaker et al.,2015), a chiton (Rojas et al., 2014), and decapod crustaceans (Pasiniand Garassino, 2012; Klompmaker et al., 2013). Cenozoic octopod drillholes have been encountered in bivalves and gastropods (Robba andOstinelli, 1975; Bromley, 1993; Harper, 2002; Todd and Harper, 2011),barnacles (Klompmaker et al., 2014, 2015), and decapods (Klompmakeret al., 2013). Ascribing traces to certain predators or parasites becomesmore difficult in the Mesozoic and older eras, when taxa with moderncounterparts that are known to drill are either absent or difficult to iden-tify (lack of taxonomic uniformitarism) unless the culprit is found at-tached to the host/prey (e.g., Baumiller, 1990, 1996, 2003; Baumilleret al., 1999, 2004a; Donovan and Webster, 2013, for platyceratid gastro-pods on Paleozoic echinoderms and brachiopods). The variety of drilledprey mentioned above is not known from the Mesozoic, at least in partbecause drilling appears to be less frequent.

1.2. Mesozoic drilling predation

Drilling predation in theMesozoic appears uncommon until the endof the Cretaceous (Kowalewski et al., 1998; Huntley and Kowalewski,2007), when its occurrence and the average frequency of drilling in-creased. This increase is coincident with the rise of muricid and naticidgastropods (Harper, 2006: fig. 3), although several biases may haveaffected the apparent pattern (see Kowalewski et al., 1998; Harperet al., 1998; Harper, 2003). The oldest undoubted neogastropod, aclade that includes known drillers such as muricids (see Bouchet et al.,2005), is recorded from the Valanginian (Kaim, 2004) and the samesamples provided a number of drilled gastropods (pers. obs. A.K.).Therefore, the rarity of Early Cretaceous drill holesmight be at least par-tially a sampling artifact. Drilled shell material needs to be very well-preserved to show drill holes clearly and traces of drilling predationmay have been obscured by diagenetic processes and corrosionwith in-creasing geological age.

1.2.1. Drilled BivalviaDrilling is rarely reported from the Triassic and Jurassic, unlike from

themid- to Late Cretaceous (e.g., Taylor et al., 1983; Kelley and Hansen,2006; Harries and Schopf, 2007). From the Jurassic, Kowalewski et al.(1998)mentioned only one drilled bivalve among ~20,000macrofossilsdominated by infaunal bivalves from the Middle Jurassic (Callovian) ofwestern India. They also indicated (their fig. 2) the possible presenceof drill holes in bivalves from the Late Jurassic (Kimmeridgian), MiddleJurassic (?Bajocian), and the Late Triassic (Norian). Early Jurassic shellshave yielded more drill holes. Harper et al. (1998) mentioned thepresence of drill holes in a variety of Early Jurassic bivalves from va-rious localities across the United Kingdom. A drilling percentage wasprovided for one well-preserved species: 20.4% (11/54) of articula-ted Pliensbachian Neocrassina gueuxi (d'Orbigny, 1850) was drilled.Aberhan et al. (2011) reported on a single drill hole in the samePliensbachian species from Germany in addition to drilled specimens(5/19, 26%) of Gresslya intermedia (Simpson, 1855), and one drilledspecimen each of Pholadomya (Pholadomya) ambigua (Sowerby, 1819)and Pleuromya costata (Young and Bird, 1828). Bardhan et al. (2012)found that 29.4–32.5% of Late Jurassic Neocrassina subdepressa Blakeand Hudleston, 1877, was drilled; other drilled taxa include Pinnamitis Phillips, 1829, and Grammatodon virgatus Sowerby, 1840. Fromthe Triassic, Fürsich and Jablonski (1984) reported on undisputeddrilled bivalves from the Upper Triassic (Carnian) Cassian Formationof Italy, but drilling percentages were not provided. Fürsich andWendt (1977: p. 283), however, mentioned for specimens from anassemblage from the same formation dominated by infaunal soft-bottom dwellers (nuculids, scaphopods) that “1.6% are bored bypredating gastropods”, referring to drilled nuculids and CassianellaBeyrich, 1862. Furthermore, Harper (2003: Appendix 1) noted (basedon a pers. comm. with Newton based on Newton et al., 1987) that the

Late Triassic (Norian) bivalve Mysidioptera williamsi (McLearn, 1941)from Oregon (USA) was drilled at a percentage of 40.0%. From theLate Triassic (Norian) of Alaska (USA), Newton (1983) reported ondrilled limid bivalves, and McRoberts and Blodgett (2000) showed twodrilled valves of Septocardia cf. S. peruviana (Cox, 1949). Lastly, VéghNeubrandt (1982) reported on drill holes in Late Triassic megalodontoidbivalve specimens from Hungary.

1.2.2. Drilled GastropodaNo drilled Jurassic or Triassic gastropods are mentioned in the over-

view papers by Kowalewski et al. (1998) and Harper (2003) nor havewe found any detailed reports [see Koken (1892), Kittl (1894), andZardini (1978), who reported drilled gastropods from the TriassicCassian Formation anecdotically, see Section 1.3.]. Conversely, drillholes in mid- to Late Cretaceous gastropods are well-known(e.g., Fischer, 1966; Dudley and Vermeij, 1978; Vermeij and Dudley,1982; Taylor et al., 1983; Pan, 1991; Kelley and Hansen, 2006; Li et al.,2011; Sørensen and Surlyk, 2011; Mallick et al., 2013, 2014).

1.2.3. Drilled BrachiopodaDrilling in Mesozoic brachiopods, especially those from the Triassic

and Jurassic, is rare. Kowalewski et al. (1998) reported the only quanti-tative data known then on drilled brachiopods from the Early Jurassic(Sinemurian) of Hungary in a fauna dominated by articulated brachio-pods (0.4%, 3/704, for all brachiopods combined). For drilled taxa,they found drilling percentages of 2.8% (2/71) for Rhapidothyris?beyrichi (Oppel, 1861) and 2.3% (1/43) for Calcirhynchia plicatissima(Quenstedt, 1852). They also referred to a possible drill hole in anEarly Jurassic (Pliensbachian) brachiopod (their fig. 2). In a recent arti-cle, Tyler et al. (2013: fig. 2B) did not report any new quantitative dataon drilling predation in Mesozoic brachiopods other than the values re-ported by Kowalewski et al. (1998). Harper and Wharton (2000: table1) showed that drill holes in brachiopods are present in Cretaceousand Jurassic assemblages, but absent from the Triassic based onEuropean collections (mostly from the UK). They reported that 31.4%(58/127 [sic], probably 58/(127 + 58)) of mid-Cretaceous (Albian/Cenomanian) brachiopods (probably mostly Rhynchonella grasianad'Orbigny, 1849) from Englandwas drilled. Sohl (1969) alsomentionedsome single brachiopod specimens from the Jurassic and Cretaceouscontaining drill holes, whereas Lee andMotchurova-Dekova (2007) re-ported on drilled Late Cretaceous specimens of Chathamirhynchiakahuitara Lee and Motchurova-Dekova, 2007. Leighton (2003) referredto Surlyk (1972) and Alexander, pers. comm., concerning Late Creta-ceous (Maastrichtian) drill holes from Denmark and New Jersey(USA), respectively, and Hiller (2014) recently showed that 16.2% (44/271) of the articulated shells of Wekarhynchia cataracta Hiller, 2011,from the Maastrichtian of New Zealand contained a drill hole. To ourknowledge, drilled Triassic brachiopods have not been reportedthus far, unlike for Permian brachiopods (Kowalewski et al., 2000;Hoffmeister et al., 2004).

1.2.4. SummaryMesozoic, especially Jurassic and Triassic, records on drilling preda-

tion available are often a) anecdotic, b) based on one invertebratetaxon, c) often qualitative in nature, and d) have focused on bivalvesprimarily. The Cassian Formation (Upper Triassic, Carnian) in northernItaly has yielded a well-preserved invertebrate fauna with a variety ofgroups that are suitable for quantitative analyses of drill holes.

1.3. History of research on drilling predation of the Cassian Formation

Drilled specimens from the Cassian Formation were first reportedin the late 19th century, but were not documented in detail prior tothe study of Fürsich and Jablonski (1984). Koken (1892: p. 473)was the first to mention that drill holes are common in shells fromthe Late Triassic Cassian Formation. “The following species could be

344 A.A. Klompmaker et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 457 (2016) 342–359

true naticids:N. substriataMü. (Lunatia sensu v. Zittel), pseudospirataMü., limnaeformis Mü., tyrolensis Lbe., subhybrida Mü., haudcarinataMü. They could be responsible for the numerous drill holes that I ob-served in shells from St. Cassian (especially Cheilotomona Blumi Mü.sp., Loxonema lommeli Mü. sp.)” (translated from German). Twoyears later, Kittl (1894) mentioned that the thousands of invariablyincomplete specimens of Polygyrina lommeli (zu Münster, 1841)from the Cassian fauna commonly show multiple perforations po-tentially made by drilling gastropods. Renewed interest in drillholes from the Cassian Formation was shown by Fürsich andWendt (1977), who mentioned a low predation percentage of 1.6%in nuculids and Cassianella. Subsequently, the works of Zardini(1978, 1981, 1985) figured various drill holes in bivalves and gastro-pods (see Appendix 1). Zardini (1985) collected N300 drilled bivalvespecimens (Nuculoidea and Cassianellidae) from the Cassian loca-tion Costalaresc near Cortina d'Ampezzo, co-occurring with nume-rous alleged members of the family Naticidae, i.e. Amauropsispaludinaris (zu Münster, 1841), A. sanctaecrucis Laube, 1868, andAmauropsis subhybrida (d'Orbigny, 1849). He alsomentioned that in-complete drill holes outnumber complete drill holes by far. Fürsichand Jablonski (1984) reported the same types of drill holes from se-veral bivalves and attributed them to naticids, without providingmuch quantitative data for the drill holes. These authors found drillholes in articulated specimens of Palaeonucula Quenstedt, 1930, (in-faunal deposit feeder) and in the buried valve of the reclining sus-pension feeder Cassianella, and less commonly in other infaunalbivalves such as Palaeocardita Conrad, 1867, and ProsoleptusBeushausen, 1895. Fürsich and Jablonski (1984) hypothesized thatAmpullina Férussac, 1822, could have been responsible for the drillholes.

1.4. Geology of the Cassian Formation

The sediments of the Triassic Cassian Formation, late Ladinian tomiddle Carnian in age, crop out in the Central Dolomites of the SouthernAlps. This region was situated at the western termination of the TethysOcean in the tropical Northern Hemisphere (Broglio Loriga et al., 1999;Keim et al., 2006). Sea surface temperatures were tropical with strongseasonal fluctuations (Nützel et al., 2010). The fossiliferous strata ofthe Upper Cassian Formation exposed at the Stuores Meadows are ofearly Carnian age (Urlichs, 1994). The formation consists of basin sedi-ments: clay- and marlstones and mass flow deposits. Beds of marly oroolithic limestone are intercalated in the basin marls. The Cassian For-mation represents the basin facies between carbonate platforms,which are now dolomitized and thus largely devoid of fossils, such asthe Cassian Dolomite (e.g., Wendt and Fürsich, 1980; Wendt, 1982;Bizzarini and Braga, 1987; Bosellini, 1998). Due to progradation of theplatfoms into the basins, the initially 300–500 m deep basins becameshallower during the Carnian and calcareous shallow-water materialwas increasingly transported into the basins. The Cassian fauna com-prises N1000 invertebrate species. There are more or less autochtho-nous basin assemblages and transported assemblages derived fromneighboring platforms (Fürsich and Wendt, 1977; Nützel and Kaim,2014; Hausmann and Nützel, 2015). The low grade of diagenetic andtectonic alteration resulted in a good preservation of fossils. Moreover,the low grade of lithification facilitates continuous weathering out ofeven small fossils and also allows for bulk sampling by wet sieving(Hausmann and Nützel, 2015).

1.5. Goals

The main goals of this paper are to (1) report and quantify data onthe drill holes found inmollusks and brachiopods from the Upper Trias-sic Cassian Formation of Italy, (2) document a drill hole in a Middle Tri-assic brachiopod sample from Poland, and (3) evaluate the cause ofthese drill holes.

2. Materials and methods

New and existing museum collections from the Upper Triassic(Carnian) Cassian Formation from northern Italy were checked fordrilled specimens. Specimens were collected from the locations inFigs. 1 and 2. New quantitative surface collections were made bycollecting all fossils weathered out frommarls at fossiliferous locations.Specimens were collected by kneeling or lying on the outcrop becausemost fossils of the Cassian Formation are small (mostly b10 mm). Aquantitative surface collection from the Stuores Wiesen (Stuores2010-3) was used to calculate a drilling percentage and documentdrill hole characteristics. Additional sediment bulk samples weretaken at fossiliferous spots and then disaggregated and wet-sieved at amesh size of 0.5mm.Drill hole characteristicswere reportedwhere pos-sible. Zardini's monographs on Cassian mollusks (1978, 1981, 1985)were examined for evidence of drill holes as well.

Additionally, a sample of 365 brachiopod shells of Coenothyrisvulgaris (Schlotheim, 1820) from the Middle Triassic (Anisian)Lower Muschelkalk of the Strzelce Opolskie Quarry in southwesternPoland (Trammer et al., 1996; Kaim, 1997: fig. 4C as “CV”) waschecked for drill holes. These shells were collected from a singleblock of marl by successive breaking of the rock and extracting thebrachiopods.

Drilled specimens from surface and museum collections werephotographed andmeasured using lightmicroscopy and SEM.Maximumspecimen height, width, and drill hole diameters were measured using astereomicroscope with calipers. Drill holes were rinsed under runningwater and cleaned with soft brushes to determine whether the drillhole was complete (i.e. completely penetrating the shell) or incomplete.

We intentionally use drilling percentage instead of the commonlyused drilling frequency because drilling frequencies are often incorrect-ly expressed as percentages (e.g., Chattopadhyay et al., 2015; Meadowset al., 2015; Klompmaker et al., 2015). Drilling percentages were calcu-lated by dividing the number of specimenswith a complete drill hole bythe total number of specimens times 100, corrected for the fact that bra-chiopods and bivalves consist of two valves (i.e. one specimen consistsof two valves) (e.g., Kowalewski, 2002).We refrain from using the com-monly used outer borehole diameter (OBD) as a measure of the size ofthe hole because we consistently use drill hole instead of bore hole forpredatory and/or parasitic holes.

Institutional abbreviations (repository of studied and illustratedma-terial): MWGUW: S.J. Thugutt Geological Museum, Faculty of Geology,University of Warsaw; NHM: Natural History Museum, London; PZO:Naturmuseum Bozen; NHMW: Naturhistorisches Museum Wien;MB.Ga: Museum für Naturkunde, Berlin.

3. Results

3.1. Italian Cassian Formation

3.1.1. Drill hole characteristicsNearly all drill holes are circular to slightly oval in outline and repre-

sent the ichnogenus Oichnus Bromley, 1981. None of the drill hole wallsshow obviousmicro-rasp marks sensu Schiffbauer et al. (2008). All drillholes were inflicted starting from the outside of the shell and all are ori-ented approximately perpendicular to the shell surface. As in Fürsichand Jablonski (1984), most of the drill holes are parabolic, eitherrepresenting the ichnospecies Oichnus paraboloides Bromley, 1981(hole completely penetrating the shell) or Oichnus excavatus Donovanand Jagt, 2002 (incomplete hole with a central boss) (Figs. 3, 4). Thecentral boss of incomplete drill holes, seemingly present in all incom-plete drill holes that are sufficiently clean to observe this feature, iscircular to distinctly elongate. Whether drill holes are complete or in-complete is obscured by sediment matrix or cements in many cases.We were not able to assess whether incomplete drill holes showed ev-idence of healing or repair (blisters as in Smith et al., 1985) because the

Fig. 1.Map showing locations yielding drilled Triassic specimens that are figured below from the Cassian Formation (N Italy) and in the Lower Muschelkalk of Poland. Coordinates loca-lities: Lago Antorno-01: 46°35′39.97″ N/12°15′39.23″ E; Seelandalpe-01: 46°38′14.91″ N/12°11′51.08″ E; Stuores-03: 46°32′16.29″ N/11°55′24.28″ E; Stuores-06: 46°32′14.91″ N/11°55′26.94″ E; Stuores-07: 46°32′08.79″N/11°55′28.35″ E; Stuores-09: 46°31′44.98″N/11°56′16.25″ E; Strzelce Opolskie Quarry: 50°31′49.50″N/18°18′45.56″ E. At each Cassian location, bulksampling and quantitative surface collecting took place. Stuores Wiesen sample Stuores 2010-3 originates from Stuores-03.


inner surface of such shells was not exposed. However, we suspect thatvery few if any incomplete drill hole was healed because nearly all in-complete drill holes exhibit a central boss, a feature not known to beproduced by repairing of the shell. We found another type of drill holein two brachiopod specimens from the Seelandalpe (Alpe di Species)(Figs. 3E, F, 4Q, R). Both are complete and have walls perpendicular tothe shell surface (cylindrical), and are ascribed to the ichnospeciesOichnus simplex Bromley, 1981.

In the Stuores 2010-3 sample, drill hole diameters vary from0.3–1.1 mm (mean = 0.7 mm, n = 37) (Table 1). Most drill holes are

Fig. 2. Quantitative surface collecting at the Stuores Wiesen, where the Stuores 2010-3 sampl(~2000 m altitude) formed by the soft argillaceous basin sediments; fossiliferous locations occsurface collecting in 2013 by five persons including A.N. and A.K. at a typical fossil location of thmer carbonate platforms; deeper areas with forests and meadows represent argillaceous basin

incomplete (24/37, 65%). Whether the diameters of complete drillholes are significantly different from those of incomplete drill holescould not be assessed with only two complete drill holes present. Therelationship between drill hole size and shell size could not be investi-gated for this sample due to the low number of complete specimens.Multiple drilled specimens are common (9/14 or 64%).

Drill holes in twenty specimens from other samples from a variety oflocalities range from 0.33 to 1.4 mm (mean = 0.8 mm, n = 49)(Table 2). A third of the drill holes are incomplete (16/49, 33%). Al-though the mean outer diameter of complete drill holes appears higher

e originates from. A. Typical aspect of the smooth-hilled landscape of the Stuores Wiesenur where vegetation is absent e.g., location Stuores 2010-3 just above A.N. B. Quantitativee StuoresWiesen. Snow-covered DolomiteMountains in the background representing for-sediments of the Cassian and Wengen Formations.

Fig. 3.Circular drill holes (ichnotaxonOichnus spp.) from the Late Triassic Cassian Formation.A–D,G. Parabolic incomplete drill holeswith a central boss (Oichnus excavatus), all in the samespecimen of the gastropod Polygyrina lommeli, locality unknown, NHM 35406A. E, F. Cylindrical complete drill hole (Oichnus simplex) in an unidentified articulate brachiopod, Alpe diSpecie (Seelandalpe-01), PZO 4777. H, I. Parabolic complete drill hole (Oichnus paraboloides) in the bivalve Palaeocardita crenata, Alpe di Specie (Seelandalpe-01 surface), PZO 4778.


compared to incomplete drill holes (0.81 vs 0.66 mm), this difference isnot statistically significant (Mann-WhitneyU=81, p=0.12). Drill holesize is not correlated significantly with shell width (R2 = 0.433, p =0.16) and height (R2 = 0.478; p = 0.12). Specimens that are drilledmore than once are common (10/20 or 50%).

Drill holes in the shells of the Zardini (1978, 1981, 1985) mono-graphs are of similar size (0.4–1.6 mm, mean = 0.94, Appendix 1),and the diameters of complete and incomplete drill holes sizes are notstatistically different (Mann-Whitney U = 71, p = 0.28). Drill holesize is not correlated with shell width (R2 = 0.0007, p = 0.92) andheight (R2 = 0.003; p = 0.85).

3.1.2. The prey — drilled taxa from the Cassian FormationAt least 23 species have revealed drill holes so far, all but one of them

beingmollusks (Appendix 2), which is higher than previously assumedbecause several drilled gastropod taxa are reported for the first timehere (Appendix 2). Only two drilled brachiopod specimens (possiblyrepresenting a single species) are present in the investigated collectionsfrom the Cassian Formation. The drilled specimens have a height rangeof 3–27 mm (Tables 1, 2; Appendix 1), although they are commonly

fragmented, especially the high-spired gastropod P. lommeli that usuallyconsists of 2–5 whorls. Below, some taxa are discussed briefly, particu-larly thosewith quantitative data. For others, referral ismade to Tables 1and 2.

3.1.2.1. Bivalvia. Palaeonucula strigilata is one of the most abundant fos-sils of the Cassian Formation, especially in the soft-bottom communitiesof the Stuores Meadows. As is usual for nuculoids, it is interpreted as aninfaunal mobile deposit feeder. It is almost always preserved with arti-culated valves, which is also the case for drilled specimens. Drill holesof this species were illustrated by Zardini (1981, 1985) and Fürsichand Jablonski (1984) from the localities Staolin, Vervei, and Costalaresc(Appendices 1, 2). We also include Pa. strigilata f. jugulata sensuZardini (1981). We have examined an articulated specimen fromthe StuoresWiesen that has a single complete drill hole in the umbil-ical region (diameter = 1.1 mm).

An articulated specimen of the nuculoid Prosoleptus lineatus fromStuores (Stuores 7, coll. 2013) has two drill holes, one complete (dia-meter = 1.4 mm) near the umbo and one incomplete with centralboss (diameter = 1.14 mm) (Fig. 4N, O). The presence of drill holes in

Fig. 4. Drilled gastropods, bivalves, and brachiopods from the Late Triassic Cassian Formation. A. Polygyrina lommeli with three closely spaced incomplete drill holes, Stuores Wiesen(Stuores-03), PZO 4779. B, C, I. Polygyrina lommeli with eight drill holes, locality unknown, MB.Ga.44755. D. Kittliconcha obliquecostata, Stuores 2010-6, PZO 4790. E. Unidentifiedcaenogastropod, Lago Antorno-01, PZO 4780. F. Atorcula canalifera, locality unknown, NHMW 1899 V 406B. G. Anoptychia carinata, Stuores 03-07, PZO 4789. H. Neodonaldina sp.,Stuores Wiesen (Stuores-03), PZO 4781. J, K. Coelostylina conica, Lago Antorno-01, PZO 4782 and PZO 4783, resp. L, M. Prosoleptus lineatus, Stuores Wiesen (Stuores-07), PZO 4785 andPZO 4784, resp. N. Cassianella sp., Stuores Wiesen (Stuores-03), PZO 4786. O. Palaeonucula strigilata, Stuores Wiesen (Stuores-09), PZO 4787. P. cf. Spirigera wissmanni, Alpe di Specie(Seelandalpe-01), PZO 4788. Q. Unidentified articulate brachiopod, Alpe di Specie (Seelandalpe-01), PZO 4777. R. Palaeocardita crenata from Alpe di Specie (Seelandalpe-01), PZO 4778.


Table 1Characteristics ofmollusks and their drill holes from the StuoresWiesen (Stuores 2010-3 sample from locality Stuores-03) from the Cassian Formation. Non-drilled specimens of themostabundant species are added for comparison.

Gastropod (g)or bivalve (bi)

Taxon Preservation Height(mm)

Width orlength forbi (mm)

# drillholes

Outer drill holediameter completedrill hole (mm)

Outer drill holediameter incompletedrill hole(s) (mm)

Outer drill holediameter,completenessdrill hole(s)unknown (mm)

g Polygyrina lommeli A Fractured N10.5 ≥5.3 6 0.7 0.7 0.7 0.6 0.6 0.7g P. lommeli B Fractured N14.9 ≥5.4 1 0.7g P. lommeli C Fractured N9.0 ≥5.2 2 0.6 0.8g P. lommeli D Fractured N9.7 ≥3.4 2 1.0 0.7g P. lommeli E Fractured N9.4 ≥3.5 1 1.0g P. lommeli F Fractured N14.0 ≥6.0 6 0.7 0.7 0.7 0.7 0.5 0.3g P. lommeli G Fractured N6.0 ≥3.6 1 0.5g P. lommeli H Fractured N8.6 ≥3.8 5 0.8 1.0 1.0 0.7 0.6g P. lommeli I Fractured N5.2 ≥2.6 2 0.8 0.5g P. lommeli J Fractured N7.7 ≥3.8 1 0.9g P. lommeli K Fractured N10.7 ≥5.0 2 0.6 1.1g P. lommeli (non-drilled A) Fractured N14.5 N6.0g P. lommeli (non-drilled B) Fractured N10.0 N5.0g P. lommeli (non-drilled C) Fractured N10.0 N4.5g P. lommeli (non-drilled D) Fractured N7.0 N5.0g P. lommeli (non-drilled E) Fractured N6.0 N3.0g P. lommeli (non-drilled F) Fractured N6.3 N2.5g Kittliconcha obliquecostata

(Bronn in zu Münster, 1841)Fractured N10.8 ≥5.8 4 0.7 0.7 0.7 0.8

g Neodonaldina? n. sp. Apex complete ≥5.4 ≥2.8 1 0.7bi Cassianella sp. Nearly complete,

disarticulated5.2 8.6 3 0.5 0.9 0.7


specimens of Prosoleptus was also mentioned by Fürsich and Jablonski(1984). The life habit is the same as for Pa. strigilata.

Specimens of Cassianella are abundant in the Cassian Formation. Thissoft-bottom recliner has a strongly convex lower (left) valve and a flatupper (right) valve as a cover. The drill holes are always located in thelower valve (Zardini 1981; Fürsich and Jablonski 1984; pers. obs.A.N.), but the flat upper valves are rarely preserved. Three drilledspecies have been reported so far (Cassianella ampezzana Bittner,1895, C. euglypha Laube, 1865, and C. beyrichi Laube, 1865). Most drillholes were reported in shells of C. ampezzana from Costaresc and Alpedi Specie, some of them with up to five drill holes on a single valve(Table 2). We report that Cassianella sp. from the Stuores Wiesen(Stuores 2010-3 sample), clearly distinct from theknowndrilled speciesin the Cassian Formation, shows three incomplete drill holes (Fig. 4L,M;Table 1).

Palaeocardita is an endobyssate suspension feeder present in someof the soft-bottom associations (see Fürsich and Wendt, 1977). Zardini(1981, 1985) reported a drilled specimen of Palaeocardita beneckei(Bittner, 1895) and P. pichleri (Bittner, 1895) from Alpe di Specie (Ap-pendix 1). We report a single valve of P. crenata (zu Münster, 1841)from Alpe di Specie that shows a single complete drill hole of 0.7 mmin diameter (Fig. 4K).

3.1.2.2. Gastropoda. Polygyrina lommeli is one of the most abundant gas-tropods of the Cassian Formation. It is a high-spired, largely smooth gas-tropod shell. Almost all known specimens are fractured as is alsocommon in other turreted shells from the Cassian Formation; fragmentsusually have two to seven whorls. Kittl (1894) mentioned thatP. lommeli commonly has multiple drill holes. Here, we documentthese drill holes for the first time. Specifically, the following specimenshave been examined. (A) A specimen from the NaturkundemuseumBerlin (MB.Ga.44755) contains eight drill holes, at least one of themcompletely penetrating the shell (Fig. 4B, C). (B) During a survey ofthe Klipstein (1843) collection (one of the earliest monographs of theCassian fauna) in the Museum of Natural History in London, a lot(35,406) with 14 well-preserved specimens was studied, whichcontained six specimens with multiple conspicuous drill holes(Table 2). Other lots of P. lommeli from the same collection with feweror no drilled specimen were available, but these specimens are

encrusted or poorly preserved. (C) A surface collection from the StuoresWiesen (Stuores 2010-3) yielded 17 specimens of P. lommeli, of which11 specimens contain drill holes and seven of them have multiple drillholes (Table 1). Drilled specimens are not of a statistically differentheight (Mann-Whitney U = 29.5, p = 0.75) or width (Mann-WhitneyU = 30.5, p = 0.84) compared to non-drilled specimens, although allspecimens are fractured.

3.1.2.3. Brachiopoda. Drill holes were found in an articulated specimen(cf. Spirigera wissmanni) and a shell fragment (clearly of brachiopod na-ture as is indicated by the presence of typical prisms as the shell micro-structure), both from the location Alpe di Specie (Seelandalpe) (Fig. 4Q,R). The holes are circular and have diameters of 0.34 and 0.50 mm. Thisis the first report of drilled brachiopods from the Cassian Formation andfrom the entire Triassic.

3.1.3. Drilling percentageThe drilling percentage in the various assemblages of the Cassian

Formation cannot be assessed comprehensively yet because there aretoo few quantitative data sets available. The only previous drilling per-centage was provided by Fürsich and Wendt (1977), who reportedthat ~6% of the 460 specimens from the Stuores marls was drilled.

Strong variations in drilling percentages exist within the Cassianbiota. Generally, transported shallow-water assemblages, as indicatedby the presence of oncoids, ooids, and encrusted fossils in a clayey ma-trix and shallow-water derived ooids and oncoids in calcareous turbi-dites (e.g., Misurina Skilift, parts of Alpe di Specie), have a low drillingpercentage (b1%). The large and diverse allochthonous assemblagefrom the Stuores Meadows (Hausmann and Nützel, 2015) containedno drilled specimens at all. However, specimens from this collectionare commonly encrusted so that drillingmay be obscured. Bulk samplesyielded small specimens mostly that are rarely drilled.

In contrast, autochthonous soft-bottom assemblages, as indicated by avery high percentage of articulated infaunal bivalves such as Pa. strigilataand Pr. lineatus in the quantitative surface collection from the StuoresWiesen (Stuores 2010-3), commonly yield a relatively high number ofdrilled specimens, although many show incomplete drill holes. The sam-ple Stuores 2010-3, comprising 119 benthic invertebrate specimens (plusseveral cidarid sea urchin spines not included in the analysis), yielded a

Table 2Characteristics of mollusks and brachiopods and their drill holes from various localities in the Cassian Formation.

Gastropod (g),bivalve (bi) orbrachiopod(br)

Taxon Preservation Museumnumber ifknown

Locality Height(mm)

Widthorlengthfor bi(mm)

#drillholes

Outerdrill holediametercompletedrillhole(s)(mm)

Outer drill holediameter incompletedrill hole(s) (mm)

Outer drill holediameter,completeness drillhole(s) unknown(mm)

g Polygyrina lommeli Fractured MB.Ga.44755 N9.5 ≥3.5 8 0.7 0.6 0.9 0.7 0.6 0.6 0.4 0.6g P. lommeli (specimen A) Fractured NHM 35406A 1 1.2g P. lommeli (specimen B) Fractured NHM 35406B 5 1.2 0.7 0.8 0.5 0.8g P. lommeli (specimen C) Fractured NHM 35406C 2 0.5 0.5g P. lommeli (specimen D) Fractured NHM 35406D 4 1.2 0.6 0.6 0.5g P. lommeli (specimen E) Fractured NHM 35406E 3 0.9 1.2 0.9g P. lommeli (specimen F) Fractured NHM 35406F 1 1.2g Atorcula canalifera (zu

Münster, 1841)Fractured NHMW

1899 V 406BN11.5 ≥5.0 2 1.0 0.67

g Atorcula? sp. Fractured Lago Antorno,near Misurinaa

N10.5 ≥5.0 1 1.0a 0.9

g Atorcula? sp. Fractured Lago Antorno,near Misurinaa

N9.5 ≥5.1 1 0.7

g Anoptychia carinata(zu Münster, 1841)

Fractured Stuores Wiesen2010-3-7,surface coll.2013b

N11.9 ≥7.5 7 0.6 0.7 0.5 0.4 0.4 1.0 0.4

g Kittliconcha obliquecoststa Fractured Stuores 2010-6b N14.2 ≥5.7 4 0.9 0.9 0.9 0.7g Coelostylina conica (zu

Münster, 1841)Apexcomplete,juvenile

Lago Antorno,near Misurinaa

4.3 2.3 1 0.8

g C. conica Apexcomplete,juvenile


4.8 2.9 1 0.33

g C. conica Apexcomplete,juvenile


3.5 2.0 1 0.8

bi Palaeonucula strigilata(Goldfuss, 1837)

Complete,articulated

Stuores Wiesenb 14.5 10.0 1 1.1

bi Prosoleptus lineatus(Goldfuss, 1840)


Stuores(Stuores 7, coll.2013)b

11.0 8.0 2 1.4 1.1

bi Palaeocardita crenata(zu Münster, 1841)

Fractured,disarticulated

Alpe di Specie ≥2.2 N3.2 1 0.7

br cf. Spirigera wissmannizu Münster, 1841


Alpe di Specie 5.5 4.3 1 0.34

br Brachiopod Fractured,disarticulated

Alpe di Specie N3.4 N7.0 1 0.5

a Lago Atorno-01 from Fig. 1.b Collected between Stuores 2010-3 and Stuores 2010-7 from Fig. 1.


complete drilling percentage of 1.7% (2/116.5) (Table 3). This must beconsidered a minimum percentage because many specimens arefragmented. Parts of the shell that were not preserved may havecontained drill holes. Including incompletely drilled specimens yields 14drilled shells and a drilling percentage of 12.0%. This collection is dominat-ed by the gastropods P. lommeli and Rhaphistomella radians, by thenuculoid bivalves Pa. strigilata and Pr. lineatus aswell as by the scaphopodPlagioglypta undulata. These are typical soft-bottom basin dwellers be-longing to the autochthonous R. radians/Pa. strigilata association (sensuFürsich and Wendt, 1977). Typically, the infaunal bivalves Pa. strigilataand Pr. lineatus have articulated valves, showing that this assemblagewasnot transported. The drilled specimens represent three gastropod spe-cies and the bivalveCassianella sp. Specimens of P. lommeli showmore drillholes than other species. The twomost abundant gastropod species in thiscollection (Stuores 2010-3) are P. lommeli (14%) and R. radians (12%).Whereas 11.8% (n = 17) of the P. lommeli specimens show a completedrill hole and 65% show a drill hole (complete + incomplete drill holes),no drill holes are present in R. radians (Fig. 5). Twenty-five percent ofthe gastropod specimens exhibit drill holes, showing that drilling in gas-tropods can be frequent at least locally. Of the 14 drilled specimens, ninehave multiple drill holes with 2–6 drill holes per specimen (Table 1).This high proportion (64%) of specimens with multiple drill holes is re-markable especially since these specimens are usually teleoconch

fragments of a few whorls, so that the number of drill holes in completespecimens could have been even higher. Although this collection issmall, its study shows that thedrillingpercentage in theCassian Formationcan exceed 10% of the specimens if incomplete specimens are includedand that gastropods appear to be the preferred prey.

3.2. Polish Lower Muschelkalk

The sample from the LowerMuschelkalk of Poland contains one drillhole in a single specimen (Fig. 6) of articulated valves of C. vulgaris,resulting in a drilling percentage of 0.3% (1/365). The drill hole with adiameter of 1.9 mm is located in the dorsal or brachial valve (30.7 mmlong; 24.4mmwide) to the right-posterior of the umbo. The subcircular,cylindrical drill hole (O. simplex) completely penetrates the shell.

4. Discussion

4.1. Naticids as drill hole producers in the Triassic?

Most of the drill holes found in the Cassian Formation representOichnus paraboloides and O. excavatus, whereas only two drill holescan be ascribed to O. simplex. The parabolic drill holes closely resemblethe drill holes of modern naticids (see also Fürsich and Jablonski,

Table 3Total number of specimens, number of drilled specimens, and the drilling percentage per taxon from the Stuores Wiesen 2010-3 sample (locality Stuores-03).

Gastropod (g),bivalve (bi),brachiopod (br)or scaphopod(sc)

Taxon # specimens # specimensdrilled

# specimenswith completedrill holes

Preservation ofvalves

Corrected# specimens

Drilling percentagefor taxa with ≥10specimens

g Polygyrina lommeli (zu Münster, 1841) 17 11 2 17 11.8g Rhaphistomella radians (Wissmann in zu

Münster, 1841)14 14 0

g Neritid, strongly encrusted 3 3g Pseudoclanculcus cassianus (Wissmann in zu

Münster, 1841)2 2

g Worthenia sp. 1 1g Cheilotomona blumi (zu Münster, 1841) 1 1g Zygopleura tenuis (zu Münster, 1841) 1 1g Zygopleura sp. 1 1g Kittliconcha obliquecostata (Bronn in zu

Münster, 1841)1 1 0 1

g cf. Neritaria mandelslohi (Klipstein, 1843) 1 1g Neodonaldina? n. sp. 1 1 0 1g Promathidia subornata (zu Münster, 1841) 1 1g Schizogonium sp. 1 1g Coelostylina conica (zu Münster, 1841) 1 1g cf. Spirostylus subcolumnaris (zu Münster, 1841) 1 1g Cylindrobullina scalaris (zu Münster, 1841) 1 1g Indet. varia 4 4sc Plagioglypta undulata (zu Münster, 1841) 17 17 0?sc Unknown tube 2 2bi Palaeonucula strigilata (Goldfuss, 1837) 22 All articulated

(i.e. 2 valvesper specimen)

22 0

bi Prosoleptus lineatus (Goldfuss, 1837) 14 All articulated 14 0bi Cassianella tenuistria (zu Münster, 1841) 1 Disarticulated

(i.e. 1 valve)0.5

bi Cassianella sp. 1 1 0 Disarticulated 0.5bi Paleocardita sp. 1 Disarticulated 0.5bi Costatoria harpa (zu Münster in Goldfuss, 1838) 1 Disarticulated 0.5bi cf. Nuculana sulcellata (Wissmann in Münster,

1841)1 Articulated 1

bi Astartid 1 Articulated 1br Koninckina leonhardi (Wissmann in zu

Münster, 1841)1 Disarticulated 1

br cf. Spirigera wissmanni (zu Münster, 1841) 3 2 articulated,1 disarticulated

2.5

br Brachiopod 1 1 Articulated 1br Brachiopod 2 1 Articulated 1

Total 119 14 2 116.5 1.7

0

10

20

30

40

50

60

70 Drilling percentage[complete drill holes]

Drilling percentage[all drill holes]

Per

cent

age

Fig. 5.Drilling percentages of species represented by ≥10 specimens in the Stuores 2010-3sample. G = gastropod; Sc = scaphopod; Bi = bivalve.


1984). Fürsich and Jablonski (1984) implied that Naticidae (orNaticoidea) were present as early as the Late Triassic and were the pro-ducers of these drill holes. Naticids gained the ability to drill, lost it sub-sequently, and regained it by the Cretaceous according to Fürsich andJablonski (1984). However, there are numerous fossil bulbous largelysmooth gastropods resembling modern naticoids (i.e. naticiform)known since the Paleozoic. Currently, 16 Triassic gastropod species areformally assigned to Natica Scopoli, 1777, which is based on a recenttype species (Database A.N. based on the Fosslilium Katalogus byDiener (1926) and Kutassy (1940)). For most if not all of the Paleozoicand Early Mesozoic “naticids” the assignment to Naticoidea, Naticidae,or even to modern naticid genera may reflect a lack of recent revisions,shell convergence, and/or poor preservation. The status of the Triassic-Jurassic “naticids” is doubtful because they may be referrable to theNeritoidea or to extinct Mesozoic families instead (Kabat, 1991).

“Natica” is the secondmost common (after Turritella Lamarck, 1799)gastropod genus in the Paleobiology Database (see Plotnick andWagner, 2006), and belongs to one of the extremely long-ranging,species-rich, potentially wastebasket taxa. Similarly, Nützel (2005) sta-ted that Natica, Turritella, Trochus Linnaeus, 1758, or Turbo Linnaeus,1758, are names for general, archaetypic gastropodmorphologies ratherthan names for real biological entities, except for themodern represen-tatives of these genera that are closely related to the type species.

Fig. 6. TheMiddle Triassic (Anisian) brachiopod Coenothyris vulgaris, MWGUWZI/74/001,from the Strzelce Opolskie Quarry in Polandwith a cylindrical drill hole (Oichnus simplex).


Koken (1892: p. 473) noted that several gastropod species couldrepresent “real naticids” that drill. Fürsich and Wendt (1977: p. 299)stated that occasional naticid borings in the umbonal region of nuculidsand in some convex valves of Cassianella indicated the presence of an in-faunal carnivorous gastropod and they suggested that Natiria de Koninck,1881, or Naticopsis M'Coy, 1844, could have been responsible. However,both genera are not naticoids, but rather neritimorphs, a group of whichmodern representatives are unable to drill. Fürsich and Jablonski (1984:p. 79) mentioned that the naticids (ampullospirids) potentially responsi-ble for the drill holes are abundant in the Cassian Formation, notably“Ampullina sanctaecrucis,A. paludinaris, andA. subhydridica”. These specieshave been reassigned to different genera, however: Ptychostomasanctaecrucis (Laube, 1868) and Prostylifer paludinaris (zu Münster,1841). For “A. subhydridica”, Amauropsis subhybrida was probablymeant, and this taxon is a synonym of P. paludinaris (Bandel, 1992a).P. paludinaris, an abundant caenogastropod species in the Cassian Forma-tion, has a tiny larval shell with strong spiral cords (Bandel, 1992a;Hausmann and Nützel, 2015: fig. 7G). This protoconch differs stronglyfrom naticid protoconchs. The same type of protoconch is also presentin P. sanctaecrucis (pers. obs. A.N.). Thus, none of the “naticid” taxa men-tioned by Fürsich and Jablonski (1984) can be regarded as naticids anylonger.

Bandel (1988, 1999) stated that all assumed naticids from theCassian Formation representmembers of the Neritimorpha. In addition,some of the assumed naticids (“Ampullina”) (i.e. Prostylifer Koken, 1889,and Ptychostoma Laube, 1868) are caenogastropods unrelated tonaticids based on the size and morphology of their protoconchs.Modern representatives of naticids are characterized by relativelylarge (up to 1 mm) globular protoconchs (Bandel, 1999). Theseprotoconchs are weakly ornamented with weak collabral lirae. Bandel(1999) concluded that the earliest confirmed naticids are of mid-Cretaceous age (see also Kollmann, 1982; Tracey et al., 1993). Traceyet al. (1993: p. 148) suggested a late Early Cretaceous first occurrenceof the Naticidae (Aptian/Albian) and aMiddle Jurassic (Aalenian) originfor theAmpullospiridae, another family thatwas included inNaticoidea,but is now considered unrelated to the Naticidae (Kase and Ishikawa,2003; Bouchet et al., 2005).

In contrast, Bouchet and Warén (1993: p. 753) noted that theproblems of the classification of modern naticids could originate fromtheir long geological history back to the Triassic and they referred toillustrations of possible Naticoidea by Zardini (1978). Without a de-tailed explanation, they stated: “We do not share the view of Bandel(1988: p. 277) that none of the Cassian gastropods belong to thefamily Naticidae”. They seemed, however, unaware that there is an-other group of naticiform gastropods present in the early Mesozoic(Ampullospiridae=Ampullinidae), a group that is unrelated to naticids

and apparently herbivorous (Kase and Ishikawa 2003). Bandel (1999)mentioned that the Naticidae originated in the mid-Cretaceous, but hedid not refer to drill holes and especially not to pre-Cretaceous reportsof drill holes. Furthermore, Bandel (1999) characterized the membersof the modern Naticoidea, but he did not try to identify their pre-Cretaceous relatives and their sister-group.

Kase and Ishikawa (2003) pointed out that the sole living represen-tative of the naticiformAmpullospiridae, a group that is widespread anddiverse in the Mesozoic, is not a carnivorous driller, but an herbivo-rous grazer. The Ampullospiridae are treated as a synonym of theAmpullinidae and are placed in the non-drilling Campaniloidea byBouchet et al. (2005). These authors only listed the family Naticidae inNaticoidea. The shell morphology of some Ampullinidae (respectivelyAmpullospiridae) is most probably convergent. Whereas we agree thatnaticiform shells evolved repeatedly in unrelated groups (very similarshells are also classified as neritimorphs), the claim by Kase andIshikawa (2003) that all fossil species placed in Ampullospiridae wereherbivorousmaybepremature (see also Aronowsky and Leighton, 2003).

In conclusion, we agree with Bandel (1999) that true naticids wereabsent in the Late Triassic Cassian Formation. Instead, convergentcaenogastropods and neritimorphs are present, which implies that thedrill holes, very reminiscent of naticid drill holes, are caused by a diffe-rent organism.

4.2. Ecology prey and the drill hole producer

Because the bivalve prey was predominantly infaunal, Fürsich andJablonski (1984) stated that the predator was infaunal as is the casein Recent naticids, although naticids can hunt epifaunally as well(e.g., Reyment, 1999; Dietl, 2002; Huelsken, 2011). If the Triassic drillerwas infaunal, it may imply that P. lommeli and Atorcula Nützel, 1998,were infaunal or recliners too. Polygyrina is one of the most abundantgastropods in the marly deposits of the Stuores Meadows nearSt. Cassian. These outcrops consist of dark marls that suggest asoft-bottom environment. However, Polygyrina is relatively rare inthe outcrops in the area of Cortina d'Ampezzo. Zardini (1978) tabulatedabout 22,000 gastropod specimens representing ~360 species from 17fossil locations in the Cortina area. P. lommeli is present in six localitieswith a total of only 11 specimens. In contrast, thousands of specimenshave been collected from the Stuores Meadows near San Cassiano(pers. obs. A.N. in the Natural History Museums of Vienna andLondon; Fürsich and Wendt, 1977).

So far, all drilled gastropods are high-spiredwith a smooth or axiallyribbed teleoconch, which could point to an infaunal (or facultatively in-faunal) mode of life. However, Fürsich and Wendt (1977) interpretedP. lommeli as feeding on algal meadows and living on algae. The pre-sence of algal meadows as ancient analogs of modern seagrassmeadows is, however, speculative because photosynthetic benthossuch as calcareous algae or hermatypic corals are absent in the autoch-thonous basin assemblages near San Cassiano. Thus, there is no indica-tion that the sea bottom was in the photic zone. There are also noother indications for very shallow water conditions. In summary, theautecology of P. lommeli remains enigmatic. Its abundance argues thatthis species is relatively low in the food chain. It was either an epifaunalgrazer or detritivore ormobile or stationary infaunal inhabitant (the lat-ter is the case for modern Turritella, a commonly drilled taxon). An in-faunal life habit is supported by the fact that the shells are rarelyencrusted and the shell is smooth and elongated. However, the autecol-ogy of gastropods in deep time is speculative because taxonomic unifor-mitarianism usually does not work and functional morphology is lessobvious than in other invertebrates such as bivalves (Todd, 2001; pers.obs. A.N.).

We found that Cassian parabolic drill holes are most abundant inmore or less autochthonous soft-bottom assemblages dominated bymollusks such as nuculoid bivalves (mostly with articulated valves),the reclining bivalve Cassianella, the gastropods P. lommeli and

Fig. 7. Two compressed specimens of the agglutinating foraminifer Bathysiphon sp. A, D.PZO 4792. B, C, E. PZO 4791.


R. radians, and the scaphopod Plagioglypta. They are rare or absent intransported or parautochthonous shallow water assemblages. Thus,there is a habitat and ecological preference of the drill hole producer(s).Although we found N20 drilled species (gastropods and bivalves), mostdrill holes occur in P. lommeli and in the bivalves Cassianella spp., Pr.lineatus, and Pa. strigilata. Although most drill holes are incomplete, it isremarkable that a high percentage of specimens of P. lommeli from an as-semblage from the Stuores Wiesen is drilled, whereas the abundantR. radians is not drilled. This result suggests that the producer(s) hadclear preferences if both prey taxa were either infaunal or epifaunal in-habitants. The presence of multiple incomplete drill holes suggest thatthe ability of the producer to fully penetrate the shell was low, that theproducer was interrupted frequently, or that it was not the goal to fullypenetrate the shell. In addition, the ability of the gastropod prey tomove may have made it difficult to finish a drill hole. Infrequently, aspecimen was found with more than one complete drill hole (at least2/18 or 11% for multiply drilled specimens, Tables 1 and 2) and thesame observation was made for Cassian bivalves (Fürsich andJablonski, 1984). Thus, the producer was most likely able to separatedead shells from living individuals.

Although many Cassian shells are well-preserved (Fig. 4), micro-rasp marks have not been found thus far. Such marks are reportedfrom the Pleistocene (Chiba and Sato, 2016), the Miocene (Pek et al.,1997; Schiffbauer et al., 2008), and the Late Cretaceous (Tapanilaet al., 2015), but such marks appear rare (two, three, three, and onespecimen, resp.). Thus, their absence from the Triassic may reflect dia-genetic alteration. Alternatively, the drill holes may not be producedby rasping with a proboscis-like apparatus, but principally caused byacids dissolving the calcium carbonate shell instead. The latter scenariowould slow down drilling substantially and explain the high incidenceof incomplete drill holes and shells drilled more than once. An ineffi-cient proboscis-like apparatus, no use of acids, a poor grip of the produ-cer on its prey so that escape was common, and/or effective defencemechanisms may also explain this result. Defence mechanisms mayconsist of a high agility/manoeuvrability of the prey and chemical/toxic repellents to reduce palatability, the latter having been discountedfor fossil brachiopods (Tyler et al., 2013). Secreting repellents as a de-fence mechanism is known for modern gastropods from the familyPleurotomariidae (e.g., Harasewych, 2002).

4.3. The cause of the drill holes

Smith et al. (1985) documented drill holes with a similarmorpholo-gy in Devonian brachiopods and interpreted them to be predatory in

Table 4Characteristics of drilled specimens from the Triassic Cassian Formation and how well they fit

Characteristics Type of producer

Post-mortem domicilorganism(domichnia)

Shell always penetrated from outside NDrill holes perpendicular to shell surface NSpecificity for culprit NHealing blisters absent YMultiple holes very common YMany incomplete holes YMultiple complete drill holes uncommon NNo attachment scars YNo attached specimens that may have served as culprit Y/NPreferred drill hole site (Fürsich and Jablonski, 1984, only;not testable herein)

Y/N

Hole size and culprit size not correlated YHoles parabolic with central boss for incomplete ones ?Shape drill holes circular Y/NCertain size class culprit preferred (not testable with completespecimens herein)

origin (even though 8/12 incomplete specimens were repaired), butdid not discuss other potential causes. Likewise, the drill holes in bi-valves from the Cassian Formation were only discussed in the contextof predators by Fürsich and Jablonski (1984). Here, four possibilitiesare discussed for the drill holes from the Cassian Formation: (1) thedrill holes are caused by organisms seeking a home (domichnia) inshells, (2) the holes are part of bore holes penetrating hardened sedi-ments and the embedded shells, (3) drill holes are produced by carniv-orous predators such as gastropods, and (4) the drill holes are caused byparasites. Characteristics of the drilled specimens are summarized inTable 4.

4.3.1. Domicile-seeking organisms (domichnia)The Stuores Wiesen sample yielded also numerous (several hun-

dreds) fragments of the large agglutinating foraminifer BathysiphonSars, 1872. This foraminifer has the shape of a tube (mostly collapsednow due to compression) and its diameter is within the range of thedrill hole diameter (0.3–1.3 mm) (Fig. 7). This raises the possibilitythat O. paraboloides, especially the incomplete ones, represent dwellingtraces or attachment scars of Bathysiphon. This hypothesis is rejected forseveral reasons. This species is also abundant in other Cassian locations,especially in autochthonous soft-bottom assemblages. However, insome of the assemblages (Lago Antorno, pers. obs. A.N.) drill holes arepresent, but Bathysiphon is absent, which argues against Bathysiphoncausing O. paraboloides. The taxon selectivity further argues against

with each type of possible producer.

e-seeking Pre-mortem domicile-seekingorganism(domichnia)

Borer Parasite Predator

Y N Y YN N Y YN N Y Y? Y N YY Y/N Y NY N N NN Y/N Y YY Y N YY/N Y Y/N YY/N N Y Y

Y Y Y N? N ? YY/N Y/N Y/N Y


domichnia. Finally, the low percentage of specimens with more thanone complete drill hole for multiply drilled specimens (at least 11%,see Section 4.2.), suggests that post-mortem domichnia are an unlikelyexplanation. The same applies to pre-mortem domichnia (Table 4).

4.3.2. Boring organismsCircular perforations can also be made by boring organisms in hard

substrates, thereby completely penetrating shells. Paleozoic brachiopodshells were bored this way (Richards and Shabica, 1969; Wilson andPalmer, 2001). Such bore holes tend to be cylindrical and complete,may penetrate the shell at an oblique angle, and should not besite and prey-specific. Hardgrounds are unknown from the Cassian For-mation and their existence in the clayey soft-bottom environment isalso unlikely. Thus, such bore holes cannot explain the holes seen inthe Cassian shells.

4.3.3. Predators

4.3.3.1. Parabolic drill holes. The fact that multiple drill holes and incom-plete drill holes are so frequent with extremes of eight drill holes on asingle gastropod fragment of about four whorls (another example is afragment of two whorls having six drill holes), sheds doubt as to whe-ther these drill holes are predatory in origin. Such an inefficient predatorcould hardly evolve and survive. Moreover, such a high percentage ofincomplete drill holes (40/86 or 47%, for all drilled Cassian shells) andmany specimens with multiple drill holes (19/34 or 56%) is unknownto us from modern drilling predators. The fact that the majority (24/37 or 65%) of the drill holes from the Stuores 2010-3 sample is incom-plete and the low percentage of specimens with a single hole (5/14,36%) may argue against predation as the cause. Moreover, and contraryto the previous assumption, there are no naticids present in the CassianFormation.

If the traces were produced by a predator, its producer remains un-known at present. It may be a non-naticid gastropod because there arevarious modern gastropod groups that drill similar-sized (sub)circularholes today (Kowalewski, 1993, 2002). Moreover, gastropods arethe only motile benthic group that is abundant at the locations withreported drill holes. If the drill holes were predatory in origin, theproducer may be identified by a careful analysis of co-occurrences atthe various Cassian locations if the predator possessed a mineralizedskeleton, but our current data are too sparse for this purpose.

4.3.3.2. Cylindrical drill holes. The two cylindrical drill holes (O. simplex)found in two brachiopod shells from the Cassian Formation and the sin-gle drill hole in a brachiopod from the Lower Muschelkalk may be pre-datory in origin. Both are completely penetrating the shell, orientedperpendicular to the shell surface, no attachment scars are found, andmultiple drill holes of this type in the same shell are unknown thusfar. More drilled brachiopod specimenswith such drill holes are neededto further test this hypothesis.

4.3.4. ParasitesThe drill holes referred to O. paraboloides may also have a parasitic

origin. This hypothesis is supported by a combination of several linesof evidence (cf. Daley, 2008: Table 1): the presence of multiple drillholes for many specimens, drill holes are placed perpendicular to theshell surface, and taxon selectivity. Parasites are often very smallin comparison to their host (Warén, 1981, for eulimid gastropods;Klompmaker and Boxshall, 2015, for crustaceans), but the diameter ofO. paraboloides in the high-spired gastropod may suggest a relativelylarge producer.

The drill holes produced by parasitic capulids can be up to ~5mm indiameter (e.g., Orr, 1962; Kosuge and Hayashi, 1967; Matsukuma,1978). Capulidae have been mentioned from the Cassian Formation(zu Münster, 1841; Böhm, 1895), but this caenogastropod family origi-nated in the Cenozoic or Late Cretaceous (Bandel, 1993; Tracey et al.,

1993). All pre-Cretaceous references to Capulus Montfort, 1810, orcapulids are outdated. Three cap-shaped gastropod species from theCassian Formation have been assigned to the modern genus Capulus:of those, Capulus alatus Laube, 1869, and C. fenestratus Laube, 1869,are now assigned to the neritimorph genera Marmolatella (Bandel,2007) and Pseudorthonychia Bandel and Fryda, 1999. Capulus muensteriGiebel, 1852, has not been revised yet. These gastropods are generallyrare in the Cassian Formation and are absent from the Stuores 2010-3sample. Morever, modern capulids produce a different drill hole mor-phology with an attachment scar and they prefer pectinid bivalves(Orr, 1962; Kosuge and Hayashi, 1967; Matsukuma, 1978).

Modern parasitic eulimid gastropods can also drill (e.g.,Wáren et al.,1994). Eulimids parasitizing echinoderms, especially echinoids, producemultiple drill holes (Kowalewski and Nebelsick, 2003), and theirdrill holes can show a circular groove around it (e.g., Neumann andWisshak, 2009). Eulimids originated in the Cretaceous (Dockery,1993) and, as for Naticidae and Capulidae, Triassic references (Kittl,1891) are outdated.

As for a predatory origin of the drill holes, there is ample evidenceagainst a parasitic producer: the inferred absence of healing blisters,the presence of many incomplete drill holes, and the absence of attach-ment scars.

4.3.5. A clue about the culprit from the Paleozoic?As naticids, capulids, and eulimids cannot have been the drill hole

producers of O. paraboloides and O. excavates from the Cassian For-mation, another culprit is must be responsible. Instead of lookingfor a late Mesozoic and/or Cenozoic producer, detailed analyses ofdrill holes from the Paleozoic may provide more insight (Fig. 8,Table 5).

Brett (2003) divided Paleozoic drill holes into four categories, and heclassified the drill holes in Middle Devonian brachiopods from NewYork as presented in Smith et al. (1985) as type 3: conical, chamfered,1–3 mm in diameter, and incomplete drill holes have a raised centralboss or raised knob. Brett (2003: table 4) interpreted type 3 drill holesto be caused by predatory gastropods. Drill holes in Middle Devonian(Buehler, 1969) and Mississippian (Ausich and Gurolla, 1979) brachio-pods from New York and Indiana, respectively, were also ascribed tothis type. The Middle Devonian brachiopods may not be ascribed tothis type because Buehler (1969) indicated that incomplete drill holesare flat-bottomed and most drill holes are not beveled or tapered, al-though some possess a slight flange near the bottom. More drilled spe-cimens can be assigned to type 3. Fenton and Fenton (1931) describeddrilled Devonian brachiopods; one of the incomplete ones contained araised central boss (see Carriker and Yochelson, 1968: B15). Brett andBordeaux (1990) mentioned Middle Devonian brachiopods withbeveled holes, including incomplete drill holes with a raised centralboss. Brett (2003), via pers. comm. with S. Felton in 2001, brieflymentioned Upper Ordovician brachiopods, gastropods, and crinoidsfrom Ohio with type 3 drill holes, a result that would be well worth de-scribing further. Some years later, Daley (2008) mentioned and figured(her figs. 7.5, 7.6) some incomplete, parabolic drill holes in Devonianbrachiopods from Canada with much more pronounced raised centralbosses that may or may not be ascribed to this type. Finally, we as-cribe the drill holes from the Cassian Formation classified asO. paraboloides and O. excavates to type 3 drill holes, includingthose in Fürsich and Jablonski (1984). Many of them are smallerthan 1 mm, but drill hole size can differ substantially for moderndrill hole producers (e.g., Kowalewski, 1993) depending on the sizerange of the producer. Some other drill holes have central raisedbosses, but are not parabolic: Miller and Sundberg (1984)mentionedstraight-sided drill holes in Upper Cambrian brachiopods with raisedcentral bosses. Thus, type 3 drill holes seem to be present in at leastthe Devonian, Mississippian, and Triassic. Naticiform drill holesoccur at least in these geological periods, in addition to the Creta-ceous and Cenozoic occurrences, when naticids were present. We

0

5

10

15

20

25

30

35

40

45

50

55

60

65

Percentage of drilled specimens

with multiple drill holes

Percentage of drill holes that is

incomplete

Pe

rce

nta

ge

Fig. 8. The percentage of drilled specimenswithmultiple drill holes and the percentage of drill holes that is incomplete for taxa from the Paleozoic and the Stuores 2010-3 sample from theTriassic Cassian Formation (see Table 5 for details).


call these Triassic and Paleozoic naticiform drill holes an example of“drill hole convergence” because they must have been produced byorganisms other than naticids.

These Paleozoic and Triassic occurrences have more in commonthan the parabolic drill hole shape and raised central bosses for in-complete drill holes. The percentage of multiple drill holes per shelland incomplete drill holes of all drill holes are strikingly high withvalues in excess of 40%, an almost unique combination for PaleozoicandMesozoic drill holes (Fig. 8, Table 5). This combination is not mu-tually exclusive because specimens can be drilled again when an (in-complete) drill hole is abandoned. At least two complete drill holesper shell are uncommon (b20% of shells with multiple drill holes,Table 5), which is to be expected because a complete predatorydrill hole is often lethal and/or there is less space for a second para-site to attach to the shell.

The similarity in drill hole characteristics justifies the questionwhether they could be produced by the same group or, alternatively,whether members of different hole-producing lineages caused thesedrill holes. In the first case, the responsible group should have a variedor changing diet: from various brachiopods in the Devonian to bivalvesand mostly gastropods in the Triassic (Smith et al., 1985; Tables 1, 2).

Platyceratids, an extinct gastropod family, are the only gastropodsthat are known to drill their victims in the Paleozoic (although faculta-tively). Drill holes by platyceratids are most frequently reported incrinoids and blastoids that show cylindrical to conical drill holes(e.g., Baumiller, 1990: fig. 2H, 1993, 1996; Baumiller and Macurda,1995). Drill holes in brachiopods and platyceratid gastropods werealso attributed to platyceratids (Baumiller et al., 1999, 2004b; Brett,2003), indicating that platyceratids, as a group, drilled more prey thanpreviously assumed. Platyceratids have also been found attached tocystoids (Clarke, 1908; Bowsher, 1955; Kluessendorf, 1983) and deadremains of orthocone nautiloid cephalopods plus other substrates onthe ocean bottom (Horný, 2000), but drill holes were not reported inthis case. Platyceratid drilling in echinoderms and brachiopods is pre-sumed to have been kleptoparasitic (Baumiller, 1990, 2003; Baumilleret al., 1999, but see Brett, 2003, who suggested predatory drilling),whereas a coprophagous and gametophagous mode of life, not associa-ted with drilling, has also been suggested (e.g., Bowsher, 1955; Lane,1984; Sutton et al., 2006). Additionally, algal grazing and filter-feeding

strategies were mentioned (Rollins and Brezinski, 1988; Horný, 2000).Baumiller et al. (1999) argued that host switching from echinoderm tobrachiopod hosts may have been possible. After the switch to brachio-pods, the platyceratids still may have been parasitic because the drillholes were mostly positioned near the lophophore, an ideal locationto steal food (Baumiller et al., 1999).

Platyceratids were present in the Middle Devonian of New York,from where type 3 drill holes in brachiopods have been found (Smithet al., 1985: Naticonema Perner, 1903, and Platyceras Conrad, 1840;Brett and Bordeaux, 1990; Bezusko, 2001: Platyceras; Brett et al., 2007:Platyceras), as is the case for drilled Misssissippian brachiopodsfrom Indiana (Ausich and Gurrola, 1979: Platyceras (Platyceras) andPlatyceras (Orthonychia)). This co-occurrence is further corroboratedby Leighton (2003), who observed that platyceratids are presentwherever type 3 drill holes have been reported; Fenton and Fenton(1931), who put forward Platyceras spp. as drill hole producers inDevonian brachiopods; and Brett (2003) noted that drill holes inUpper Ordovician invertebrates are associated with the platyceratidgastropod Cyclonema Hall, 1852. Platyceras has been interpreted as acoprophagous feeder (Smith et al., 1985; Bezusko, 2001), but Baumiller(1990) unequivocally showed that Platyceras (Orthonychia) can drill.For Naticonema, Brett (2003) interpreted this platyceratid, found inclose association with the drilled Middle Devonian brachiopods, torepresent a non-specialized platyceratid gastropod thatmay have beena predatory driller. Platyceratids can leavemuscle scars on the animal towhich they attach (Rollins and Brezinski, 1988, and references),but how often this occurs is unknown. In the case of predatoryplatyceratids, no muscle scars are expected.

Although platyceratids were thought to have gone extinct withcamerate and inadunate crinoids in the Permian (Bowsher, 1955),platyceratids have been claimed from the Triassic. Bandel (1992b)assigned Orthonychia alata (Laube, 1869) from the Cassian Formationto this family and proposed, without any evidence, that this taxonwould have parasitized a crinozoan host. This species became the typespecies of Pseudorthonychia Bandel and Fryda, 1999, and was placed inPseudorthonychiidae, however. To our knowledge, this rare specieshas never been found attached to a possible victim. Nevertheless,Hausmann and Nützel (2015) referred to this species as parasitic, citingBandel (1992b). Could it be possible that the drill holes from the Cassian

Table 5Characteristics of drill holes of Paleozoic and early Mesozoic assemblages with at least seven drilled specimens, ordered by age from young to old. Drill holes with similar cha-racteristics to drill holes from the Cassian Formation in gray.

Stratigraphic age Taxon Country (state/province)

Size drill holes (mm): range

(mean)

General drill hole shape (circular in outline unless stated otherwise)

Percentage of drilled specimens with multiple drill holes and the number of specimens involved

Percentage of drill holes that is incomplete and the number of specimens involved

Incomplete drill holes with central boss

Percentage of shells with ≥2 complete drill holes of shells with ≥2 parabolic drill holes

Reference

Triassic (Carnian) Mollusca Italy 0.3–1.1 (0.7) parabolic 64.3 (9/14) 64.9 (24/37) Y≥11.1 (≥2/18)

herein (Stuores 2010-3 sample)

Permian (Upper) Bivalvia Brazil 0.4–5.8 (2.0) cylindrical 12.5 (2/16) 0.0 (0/16) Kowalewski et al. (2000)

Carboniferous (Pennsylvanian)

brachiopod Cardiarina cordata Cooper, 1956

USA (New Mexico) 0.03–0.19 (0.1) slightly beveled 5.3 (7/131) 0.0 (0/131) Hoffmeister et al. (2003)

Carboniferous (Mississippian) Brachiopoda USA (Indiana) 1.6–3.2 parabolic rare rare

Y (sometimes) 0.0

Ausich and Gurrola (1979)

Carboniferous (Mississippian) Brachiopoda USA (Indiana) 0.1–1.6 cylindrical common

Ausich and Gurrola (1979)

Carboniferous (Mississippian) Brachiopoda USA (Kentucky) 1.1–4.7 (2.8)

cylindrical to slightly tapered 4.3 (2/46) Baumiller et al. (1999)

Carboniferous (Mississippian)

brachiopod Perditocardinia cf. P. dubia (Hall in Hall and Whitney, 1858) USA (Missouri) 0.13–0.80 (0.40) ?cylindrical 0.0 (0/67) Deline et al. (2003)


brachiopod Rhipidomella Oehlert, 1890

USA (Montana, Utah) cylindrical/parabolic ~5.2 (1/~19) N

Rodriguez and Gutschik (1970)

Carboniferous (Mississippian) Brachiopoda Northern Ireland 0.1–1.0 ?cylindrical 4.3 (2/46) Brunton (1966)


brachiopod Crurithyris George, 1931 Belgium 0.16–1.08 (0.57)

cylindrical or weakly conical 0.0 (0/35) 0.0 (0/60)

Mottequin and Sevastopulo (2009)


blastoid Orophocrinus von Seebach, 1865 0.5–2.5 (1.3)

cylindrical to slightly tapered 17.4 (4/23) ≥10.7 (≥3/28)

Baumiller and Macurda (1995)

Devonian (Frasnian) Brachiopoda USA (Iowa) (0.8) cylindrical14.9 (21/141)

~55.6 (≥90/≥162) N Leighton (2001)

Devonian (Givetian)

brachiopod Pholidostrophia Hall and Clarke, 1892 USA (Ohio) 0.5–1.3

cylindrical, unbeveled

4.2-8.3 (1-2/24)

N Leighton (2003)

Devonian (Middle) Brachiopoda USA (New York) 0.2–3.08 (1.18) parabolic 47.6 (10/21) 46.7 (21/45) Y (6/48) 4.8 (1/21) Smith et al. (1985)

Devonian (Middle)

brachiopod Spinocyrtia Fredericks, 1916 USA (New York) 2.5–3.0 ?parabolic 45.4 (5/11) 45.4 (5/11) Y 18.2 (2/11)

Brett and Bordeaux (1990)

Devonian (Middle) colonial metazoans

USA (Ohio, New York) and Canada (Ontario) 0.15–0.30 (0.24) beveled 14.3 (3/21) Wilson and Taylor (2006)

Devonian (Middle)

hyolithid Hallothecacf. H. aclis (Hall, 1876) Canada (Ontario) 0.3–0.9 (0.6)

conical or cylindrical 0.0 (0/7) 0.0 (0/7) Baumiller et al. (2010)

Devonian (Middle)

blastoid Heteroschisma Wachsmuth, 1883 0.3–2.4 (0.9)

conical or cylindrical 5.0 (7/139) Baumiller (1996)

Devonian (Lower) Brachiopoda Canada (Quebec) 0.8–1.7 non-beveled 0.0 (0/13)

Lespérance and Sheehan (1975), Sheehan and Lespérance (1978)

Devonian BlastoideaUSA (Ohio, Indiana) 1.1–2.4 (1.7)

conical or cylindrical 2.0 (1/51) 1.9 (1/52) Baumiller (1993)

Silurian (Wenlock)

brachiopod Dicaelosia King, 1850 Canada (Nunavut) (0.32) cylindrical 2.9 (2/68) 2.9 (2/68) N Rohr (1976)

Silurian

brachiopod Artiotreta Ireland, 1961 USA (Oklahoma) 0.08–0.13 countersunk 1.4 (1/71) ≥1.4 (≥1/72) N

Chatterton and Whitehead (1987)

Ordovician (Upper)

brachiopod Dalmanella Hall and Clarke, 1892 USA (Ohio) 1.2–3.1 (2.05) cylindrical/beveled 8.3 (2/24)

Bucher (1938), Carriker and Yochelson (1968)

Ordovician (Middle) Brachiopoda USA (New York, Michigan) 0.4–2.0 beveled 0.0 (0/9) Cameron (1967)

Ordovician (Middle) Brachiopoda Sweden 0.07–0.22 countersunk 3.3 (1/30) ≥3.2 (≥1/31) Holmer (1989)

Ordovician, Silurian, Devonian Brachiopoda

Canada (Eastern provinces) (≤0.48) cylindrical mostly 0-62.8 0-100

N (few exceptions) Daley (2008)

Cambrian (Upper) Brachiopoda USA (California, Nevada) 0.05–0.2 (0.11) cylindrical 0.0 (0/13) ≥7.7 (≥1/13) Y

Miller and Sundberg (1984)

Cambrian (Middle-Upper) Brachiopoda

USA (South Dakota)

0.066–0.130 (0.170) non-beveled 1.6 (1/63) Robson and Pratt (2007)

Cambrian (Middle-Upper) Brachiopoda

USA (South Dakota)

0.137–0.675 (0.342) irregular 0.0 (0/62) Robson and Pratt (2007)

Cambrian (Middle)

brachiopod Linnarssonia Walcott, 1885 Sweden

0.01–0.24 (0.0925) cylindrical mostly 36.7-58.8

Conway Morris and Bengtson (1994)

Cambrian (Middle) Brachiopoda Morocco0.05–0.17

(0.097) various 13.2 (7/53) 47.0 (31/66) N Streng (1999)

Cambrian (Lower)MobergellaHedström, 1923 Sweden 0.07–0.24 cylindrical 12.5 (1/8) 55 (5/9) N

Conway Morris and Bengtson (1994)



Formationwere caused by P. alata? This species appears too rare to pro-duce that many drill holes. It was, however, reported from the StuoresWiesen (where the Stuores 2010-3 sample was also collected), whichyielded a shallow-water assemblage that was transported to the basinby mass flows (Hausmann and Nützel, 2015). The shell shape ofP. alata is reminiscent of species that can attach to substrates such asplatyceratids and capulids. If specimens of Pseudorthonychia are thepro-ducers of the drill holes, this is yet another example of a lineage produ-cing naticiform drill holes.

4.3.6. SynthesisThe cylindrical drill holes (O. simplex) in brachiopods from the

Cassian Formation may have been caused by a predator. The producerremains unknown, however.

Most observations with regard to the parabolic drill holes(O. paraboloides and O. excavatus) from the Cassian Formation arein line with either a predatory or parasitic origin (Table 4). Choosingbetween the two is difficult as some observations argue against bothpossibilities. We hypothesize that the drill holes are predatory in ori-gin because not a single attachment scar is found on any of the drilledspecimens and healing blisters are inferred to be absent, both ofwhich may be expected in the case of parasitic drill holes. The com-mon presence of multiple drill holes and incomplete drill holesmay be explained by the use of chemical repellents in multipletaxa, effective escape strategies, and/or a high rate of disturbance ofthe attacker. The fact that drill hole size and culprit size are not correlat-ed may be related to the low number of complete specimens involvedand the narrow size range of the drilled specimens. The producer ofthe parabolic drill holes remains enigmatic. True platyceratids, sug-gested to have produced similar-shaped drill holes in Paleozoic in-vertebrates, are absent from the Cassian Formation, although onespecies is reminiscent of this family and may have caused at leastsome drill holes.

5. Conclusions

1. Drill holes are reported in bivalves, gastropods, and brachiopodsfrom the Late Triassic (Carnian) Cassian Formation of Italy, increasingthe number of drilled species known from this formation and theTriassic.

2. A single, cylindrical drill hole is reported from a Middle Triassic(Anisian) brachiopod from the Lower Muschelkalk of Poland.

3. Drill holes are reported from Triassic brachiopods for the first time.4. The minimum drilling percentages exclusive of incomplete drill

holes are generally low with 1.7% for a sample from the CassianFormation and 0.3% for a brachiopod species from the LowerMuschelkalk.

5. Single, cylindrical, complete drill holes in brachiopods from theCassian Formation and the Lower Muschelkalk may be predatory inorigin.

6. Multiple drill holes in mollusks from the Cassian Formation are com-mon, and drill holes are parabolic and often incomplete and with acentral boss. These parabolic drill holes are probably caused by preda-tors, although equally ineffective parasites cannot be ruled out entirely.

7. Similar-shaped drill holes are also present in Devonian and Carboni-ferous brachiopods and resemble drill holes produced by extantnaticid gastropods, despite the absence of naticids in pre-Cretaceousrocks. This is an example of drill hole convergence.

Supplementary data to this article can be found online at http://dx.doi.org/10.1016/j.palaeo.2016.06.017.

Acknowledgements

We are grateful to Patricia Kelley (University of North CarolinaWilmington) and Lydia Tackett (North Dakota State University) for

very useful comments that improved this article. A.A.K. was in partsupported by the Jon L. and Beverly A. Thompson Endowment Fund.A.N. acknowledges the Deutsche Forschungsgemeinschaft (project NU96/13-1). Funding sources had no role in the study design, data collec-tion, analysis and interpretation of the data, and during the writingand submission process. This is University of Florida Contribution toPaleobiology 819.

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