Drill Hole Convergence and a Quantitative Analysis of Drill Holes in Mollusks and Brachiopods from the Triassic of Italy and Poland

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

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Drill hole convergence and a quantitative analysis of drill holes in mollusks 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, USA b Department of Integrative Biology, Museum of Paleontology, University of California, Berkeley, 1005 Valley Life Sciences Building #3140, Berkeley, CA 94720, USA c SNSB-Bayerische Staatssammlung für Paläontologie und Geologie, Department für Geo- und Umweltwissenschaften, Paläontologie & Geobiologie, Geobio-Centre, Ludwig-Maximilians- Universität München, 80333 Munich, Germany d Instytut Paleobiologii PAN, ul. Twarda 51/55, 00-818 Warsaw, Poland article info abstract

Article history: The history of predation is recorded primarily from drilling in Cenozoic invertebrates. Quantitative data are un- Received 11 March 2016 common from the Triassic, a period before the appearance and radiation of many known drill hole producers Received in revised form 7 June 2016 such as various gastropod families and octopods. We present quantitative evidence of drilling from the Late Tri- Accepted 9 June 2016 assic (Carnian) Cassian Formation of Italy and the Middle Triassic (Anisian) Lower Muschelkalk of Poland, Available online 14 June 2016 documenting the ﬁrst drill holes in Triassic brachiopods. A single brachiopod with a cylindrical, complete drill hole was found in a brachiopod sample from of Poland (drilling percentage = 0.3%, n = 365). The Cassian For- Keywords: Bivalvia mation yielded drill holes in gastropods, bivalves, and brachiopods, indicating that more species are drilled Brachiopoda than was known previously. The minimum drilling percentage exclusive of incomplete drill holes of a sample Gastropoda from the Stuores Wiesen (Cassian Formation) is 1.7% (n = 116.5). Prey selectivity is evident: complete drill Naticidae holes are primarily present in one gastropod species, Polygyrina lommeli (11.8% of specimens with a complete Parasitism drill hole), whereas other common species were not drilled. Single drill holes in brachiopods are cylindrical Predation and complete and may be predatory in origin. Multiple drill holes in mollusks are common, and drill holes are parabolic and often incomplete with a central boss, resembling the shape of drill holes produced by extant naticid gastropods. A survey of the Paleozoic literature showed that such drill holes are also present in Devonian and Car- boniferous brachiopods. However, naticids did not evolve until the Cretaceous so we propose the term “drill hole convergence” for similar-shaped drill holes produced by different organisms. The Triassic parabolic drill holes are not caused by domicile-seeking or boring organisms. Instead, we favor a predatory origin of these drill holes, but we cannot entirely rule out parasitism. Surveying other Triassic invertebrate assemblages should yield more evidence of drilling. © 2016 Elsevier B.V. All rights reserved.

1. Introduction Drilling predation is the best preserved and the most easily quantiﬁ- able evidence of predation in the fossil record. Predatory drilling in 1.1. Drilling predation and prey modern marine invertebrates is ascribed to ﬂatworms, nematods, octopods, but most of all to gastropods (Kowalewski, 2002: Table 2). The present study reports and discusses the occurrence of drill holes The shape of these drill holes can, by analogy, help to determine the in marine invertebrate shells (gastropods, bivalves, and brachiopods) most likely culprit of drill holes in fossil assemblages, especially during from the Upper Triassic (Carnian) Cassian Formation of Italy and from the Cenozoic when relatives of modern drillers were present. Given the Middle Triassic (Anisian) Lower Muschelkalk of Poland. These the abundance of shelly material in Cenozoic deposits, traces of gastro- records are one of the very few Triassic occurrences of drill holes, and, pod predation have been studied extensively in a variety of marine in- therefore, their analysis is pivotal for understanding the evolutionary vertebrate prey including bivalves and gastropods (e.g., Kowalewski history of drilling predation. et 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., ⁎ Corresponding author at: Florida Museum of Natural History, University of Florida, 2011), echinoids (e.g., Kowalewski and Nebelsick, 2003), ostracods 1659 Museum Road, Gainesville, FL 32611, USA. E-mail addresses: [email protected] (A.A. Klompmaker), (e.g., Reyment et al., 1987; Reyment and Elewa, 2003), annelids [email protected] (A. Nützel), [email protected] (A. Kaim). (e.g., Klompmaker, 2012; Martinell et al., 2012; Villegas-Martín et al.,

2016), brachiopods (e.g., Leighton, 2003; Kowalewski et al., 2005; Late Triassic (Norian) bivalve Mysidioptera williamsi (McLearn, 1941) Baumiller et al., 2006), barnacles (Gordillo, 2013; Klompmaker et al., from Oregon (USA) was drilled at a percentage of 40.0%. From the 2015), a chiton (Rojas et al., 2014), and decapod crustaceans (Pasini Late Triassic (Norian) of Alaska (USA), Newton (1983) reported on and Garassino, 2012; Klompmaker et al., 2013). Cenozoic octopod drill drilled limid bivalves, and McRoberts and Blodgett (2000) showed two holes have been encountered in bivalves and gastropods (Robba and drilled valves of Septocardia cf. S. peruviana (Cox, 1949). Lastly, Végh Ostinelli, 1975; Bromley, 1993; Harper, 2002; Todd and Harper, 2011), Neubrandt (1982) reported on drill holes in Late Triassic megalodontoid barnacles (Klompmaker et al., 2014, 2015), and decapods (Klompmaker bivalve specimens from Hungary. et al., 2013). Ascribing traces to certain predators or parasites becomes more difﬁcult in the Mesozoic and older eras, when taxa with modern 1.2.2. Drilled Gastropoda counterparts that are known to drill are either absent or difﬁcult to iden- No drilled Jurassic or Triassic gastropods are mentioned in the over- tify (lack of taxonomic uniformitarism) unless the culprit is found at- view papers by Kowalewski et al. (1998) and Harper (2003) nor have tached to the host/prey (e.g., Baumiller, 1990, 1996, 2003; Baumiller we found any detailed reports [see Koken (1892), Kittl (1894),and et al., 1999, 2004a; Donovan and Webster, 2013, for platyceratid gastro- Zardini (1978), who reported drilled gastropods from the Triassic pods on Paleozoic echinoderms and brachiopods). The variety of drilled Cassian Formation anecdotically, see Section 1.3.]. Conversely, drill prey mentioned above is not known from the Mesozoic, at least in part holes in mid- to Late Cretaceous gastropods are well-known becausedrillingappearstobelessfrequent. (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., 1.2. Mesozoic drilling predation 2011; Sørensen and Surlyk, 2011; Mallick et al., 2013, 2014).

Drilling predation in the Mesozoic appears uncommon until the end 1.2.3. Drilled Brachiopoda of the Cretaceous (Kowalewski et al., 1998; Huntley and Kowalewski, Drilling in Mesozoic brachiopods, especially those from the Triassic 2007), when its occurrence and the average frequency of drilling in- and Jurassic, is rare. Kowalewski et al. (1998) reported the only quanti- creased. This increase is coincident with the rise of muricid and naticid tative data known then on drilled brachiopods from the Early Jurassic gastropods (Harper, 2006: fig. 3), although several biases may have (Sinemurian) of Hungary in a fauna dominated by articulated brachio- affected the apparent pattern (see Kowalewski et al., 1998; Harper pods (0.4%, 3/704, for all brachiopods combined). For drilled taxa, et al., 1998; Harper, 2003). The oldest undoubted neogastropod, a they found drilling percentages of 2.8% (2/71) for Rhapidothyris clade that includes known drillers such as muricids (see Bouchet et al., ?beyrichi (Oppel, 1861) and 2.3% (1/43) for Calcirhynchia plicatissima 2005), is recorded from the Valanginian (Kaim, 2004) and the same (Quenstedt, 1852). They also referred to a possible drill hole in an samples provided a number of drilled gastropods (pers. obs. A.K.). Early Jurassic (Pliensbachian) brachiopod (their fig. 2). In a recent arti- Therefore, the rarity of Early Cretaceous drill holes might be at least par- cle, Tyler et al. (2013: fig. 2B) did not report any new quantitative data tially a sampling artifact. Drilled shell material needs to be very well- on drilling predation in Mesozoic brachiopods other than the values re- preserved to show drill holes clearly and traces of drilling predation ported by Kowalewski et al. (1998). Harper and Wharton (2000:table may have been obscured by diagenetic processes and corrosion with in- 1) showed that drill holes in brachiopods are present in Cretaceous creasing geological age. and Jurassic assemblages, but absent from the Triassic based on European collections (mostly from the UK). They reported that 31.4% 1.2.1. Drilled Bivalvia (58/127 [sic], probably 58/(127 + 58)) of mid-Cretaceous (Albian/ Drilling is rarely reported from the Triassic and Jurassic, unlike from Cenomanian) brachiopods (probably mostly Rhynchonella grasiana the mid- to Late Cretaceous (e.g., Taylor et al., 1983; Kelley and Hansen, d'Orbigny, 1849) from England was drilled. Sohl (1969) also mentioned 2006; Harries and Schopf, 2007). From the Jurassic, Kowalewski et al. some single brachiopod specimens from the Jurassic and Cretaceous (1998) mentioned only one drilled bivalve among ~20,000 macrofossils containing drill holes, whereas Lee and Motchurova-Dekova (2007) re- dominated by infaunal bivalves from the Middle Jurassic (Callovian) of ported on drilled Late Cretaceous specimens of Chathamirhynchia western India. They also indicated (their fig. 2) the possible presence kahuitara Lee and Motchurova-Dekova, 2007. Leighton (2003) referred of drill holes in bivalves from the Late Jurassic (Kimmeridgian), Middle to Surlyk (1972) and Alexander, pers. comm., concerning Late Creta- Jurassic (?Bajocian), and the Late Triassic (Norian). Early Jurassic shells ceous (Maastrichtian) drill holes from Denmark and New Jersey have yielded more drill holes. Harper et al. (1998) mentioned the (USA), respectively, and Hiller (2014) recently showed that 16.2% (44/ presence of drill holes in a variety of Early Jurassic bivalves from va- 271) of the articulated shells of Wekarhynchia cataracta Hiller, 2011, rious localities across the United Kingdom. A drilling percentage was from the Maastrichtian of New Zealand contained a drill hole. To our provided for one well-preserved species: 20.4% (11/54) of articula- knowledge, drilled Triassic brachiopods have not been reported ted Pliensbachian Neocrassina gueuxi (d'Orbigny, 1850) was drilled. thus far, unlike for Permian brachiopods (Kowalewski et al., 2000; Aberhan et al. (2011) reported on a single drill hole in the same Hoffmeister et al., 2004). Pliensbachian species from Germany in addition to drilled specimens (5/19, 26%) of Gresslya intermedia (Simpson, 1855), and one drilled 1.2.4. Summary specimen each of Pholadomya (Pholadomya) ambigua (Sowerby, 1819) Mesozoic, especially Jurassic and Triassic, records on drilling preda- and Pleuromya costata (Young and Bird, 1828). Bardhan et al. (2012) tion available are often a) anecdotic, b) based on one invertebrate found that 29.4–32.5% of Late Jurassic Neocrassina subdepressa Blake taxon, c) often qualitative in nature, and d) have focused on bivalves and Hudleston, 1877, was drilled; other drilled taxa include Pinna primarily. The Cassian Formation (Upper Triassic, Carnian) in northern mitis Phillips, 1829, and Grammatodon virgatus Sowerby, 1840. From Italy has yielded a well-preserved invertebrate fauna with a variety of the Triassic, Fürsich and Jablonski (1984) reported on undisputed groups that are suitable for quantitative analyses of drill holes. drilled bivalves from the Upper Triassic (Carnian) Cassian Formation of Italy, but drilling percentages were not provided. Fürsich and 1.3. History of research on drilling predation of the Cassian Formation Wendt (1977: p. 283), however, mentioned for specimens from an assemblage from the same formation dominated by infaunal soft- Drilled specimens from the Cassian Formation were first reported bottom dwellers (nuculids, scaphopods) that “1.6% are bored by in the late 19th century, but were not documented in detail prior to predating gastropods”, referring to drilled nuculids and Cassianella the study of Fürsich and Jablonski (1984). Koken (1892: p. 473) Beyrich, 1862.Furthermore,Harper (2003: Appendix 1) noted (based was the first to mention that drill holes are common in shells from on a pers. comm. with Newton based on Newton et al., 1987)thatthe the 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. substriata Mü. (Lunatia sensu v. Zittel), pseudospirata 2. Materials and methods Mü., limnaeformis Mü., tyrolensis Lbe., subhybrida Mü., haudcarinata Mü. They could be responsible for the numerous drill holes that I ob- New and existing museum collections from the Upper Triassic served in shells from St. Cassian (especially Cheilotomona Blumi Mü. (Carnian) Cassian Formation from northern Italy were checked for sp., Loxonema lommeli Mü. sp.)” (translated from German). Two drilled specimens. Specimens were collected from the locations in years later, Kittl (1894) mentioned that the thousands of invariably Figs. 1 and 2. New quantitative surface collections were made by incomplete specimens of Polygyrina lommeli (zu Münster, 1841) collecting all fossils weathered out from marls at fossiliferous locations. from the Cassian fauna commonly show multiple perforations po- Specimens were collected by kneeling or lying on the outcrop because tentially made by drilling gastropods. Renewed interest in drill most fossils of the Cassian Formation are small (mostly b10 mm). A holes from the Cassian Formation was shown by Fürsich and quantitative surface collection from the Stuores Wiesen (Stuores Wendt (1977), who mentioned a low predation percentage of 1.6% 2010-3) was used to calculate a drilling percentage and document in nuculids and Cassianella. Subsequently, the works of Zardini drill hole characteristics. Additional sediment bulk samples were (1978, 1981, 1985) figured various drill holes in bivalves and gastro- taken at fossiliferous spots and then disaggregated and wet-sieved at a pods (see Appendix 1). Zardini (1985) collected N300 drilled bivalve mesh size of 0.5 mm. Drill hole characteristics were reported where pos- specimens (Nuculoidea and Cassianellidae) from the Cassian loca- sible. Zardini's monographs on Cassian mollusks (1978, 1981, 1985) tion Costalaresc near Cortina d'Ampezzo, co-occurring with nume- were examined for evidence of drill holes as well. rous alleged members of the family Naticidae, i.e. Amauropsis Additionally, a sample of 365 brachiopod shells of Coenothyris paludinaris (zu Münster, 1841), A. sanctaecrucis Laube, 1868,and vulgaris (Schlotheim, 1820) from the Middle Triassic (Anisian) Amauropsis subhybrida (d'Orbigny, 1849). He also mentioned that in- Lower Muschelkalk of the Strzelce Opolskie Quarry in southwestern complete drill holes outnumber complete drill holes by far. Fürsich Poland (Trammer et al., 1996; Kaim, 1997: fig. 4Cas“CV”)was and Jablonski (1984) reported the same types of drill holes from se- checked for drill holes. These shells were collected from a single veral bivalves and attributed them to naticids, without providing block of marl by successive breaking of the rock and extracting the much quantitative data for the drill holes. These authors found drill brachiopods. holes in articulated specimens of Palaeonucula Quenstedt, 1930,(in- Drilled specimens from surface and museum collections were faunal deposit feeder) and in the buried valve of the reclining sus- photographed and measured using light microscopy and SEM. Maximum pension feeder Cassianella, and less commonly in other infaunal specimen height, width, and drill hole diameters were measured using a bivalves such as Palaeocardita Conrad, 1867,andProsoleptus stereomicroscope with calipers. Drill holes were rinsed under running Beushausen, 1895. Fürsich and Jablonski (1984) hypothesized that water and cleaned with soft brushes to determine whether the drill Ampullina Férussac, 1822, could have been responsible for the drill hole was complete (i.e. completely penetrating the shell) or incomplete. holes. We intentionally use drilling percentage instead of the commonly used drilling frequency because drilling frequencies are often incorrect- 1.4. Geology of the Cassian Formation ly expressed as percentages (e.g., Chattopadhyay et al., 2015; Meadows et al., 2015; Klompmaker et al., 2015). Drilling percentages were calcu- The sediments of the Triassic Cassian Formation, late Ladinian to lated by dividing the number of specimens with a complete drill hole by middle Carnian in age, crop out in the Central Dolomites of the Southern the total number of specimens times 100, corrected for the fact that bra- Alps. This region was situated at the western termination of the Tethys chiopods and bivalves consist of two valves (i.e. one specimen consists Ocean in the tropical Northern Hemisphere (Broglio Loriga et al., 1999; of two valves) (e.g., Kowalewski, 2002). We refrain from using the com- Keim et al., 2006). Sea surface temperatures were tropical with strong monly used outer borehole diameter (OBD) as a measure of the size of seasonal fluctuations (Nützel et al., 2010). The fossiliferous strata of the hole because we consistently use drill hole instead of bore hole for the Upper Cassian Formation exposed at the Stuores Meadows are of predatory and/or parasitic holes. early Carnian age (Urlichs, 1994). The formation consists of basin sedi- Institutional abbreviations (repository of studied and illustrated ma- ments: clay- and marlstones and mass flow deposits. Beds of marly or terial): MWGUW: S.J. Thugutt Geological Museum, Faculty of Geology, oolithic limestone are intercalated in the basin marls. The Cassian For- University of Warsaw; NHM: Natural History Museum, London; PZO: mation represents the basin facies between carbonate platforms, Naturmuseum Bozen; NHMW: Naturhistorisches Museum Wien; which are now dolomitized and thus largely devoid of fossils, such as MB.Ga: Museum für Naturkunde, Berlin. the Cassian Dolomite (e.g., Wendt and Fürsich, 1980; Wendt, 1982; Bizzarini and Braga, 1987; Bosellini, 1998). Due to progradation of the 3. Results platfoms into the basins, the initially 300–500 m deep basins became shallower during the Carnian and calcareous shallow-water material 3.1. Italian Cassian Formation was increasingly transported into the basins. The Cassian fauna com- prises N1000 invertebrate species. There are more or less autochtho- 3.1.1. Drill hole characteristics nous basin assemblages and transported assemblages derived from Nearly all drill holes are circular to slightly oval in outline and repre- neighboring platforms (Fürsich and Wendt, 1977; Nützel and Kaim, sent the ichnogenus Oichnus Bromley, 1981. None of the drill hole walls 2014; Hausmann and Nützel, 2015). The low grade of diagenetic and show obvious micro-rasp marks sensu Schiffbauer et al. (2008). All drill tectonic alteration resulted in a good preservation of fossils. Moreover, holes were inflicted starting from the outside of the shell and all are ori- the low grade of lithification facilitates continuous weathering out of ented approximately perpendicular to the shell surface. As in Fürsich even small fossils and also allows for bulk sampling by wet sieving and Jablonski (1984), most of the drill holes are parabolic, either (Hausmann and Nützel, 2015). representing the ichnospecies Oichnus paraboloides Bromley, 1981 (hole completely penetrating the shell) or Oichnus excavatus Donovan 1.5. Goals and Jagt, 2002 (incomplete hole with a central boss) (Figs. 3, 4). The central boss of incomplete drill holes, seemingly present in all incom- The main goals of this paper are to (1) report and quantify data on plete drill holes that are sufficiently clean to observe this feature, is the drill holes found in mollusks and brachiopods from the Upper Trias- circular to distinctly elongate. Whether drill holes are complete or in- sic Cassian Formation of Italy, (2) document a drill hole in a Middle Tri- complete is obscured by sediment matrix or cements in many cases. assic brachiopod sample from Poland, and (3) evaluate the cause of We were not able to assess whether incomplete drill holes showed ev- these drill holes. idence of healing or repair (blisters as in Smith et al., 1985) because the A.A. Klompmaker et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 457 (2016) 342–359 345

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 localities: 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, bulk sampling 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 that incomplete (24/37, 65%). Whether the diameters of complete drill very few if any incomplete drill hole was healed because nearly all in- holes are signiﬁcantly different from those of incomplete drill holes complete drill holes exhibit a central boss, a feature not known to be could not be assessed with only two complete drill holes present. The produced by repairing of the shell. We found another type of drill hole relationship between drill hole size and shell size could not be investi- in two brachiopod specimens from the Seelandalpe (Alpe di Species) gated for this sample due to the low number of complete specimens. (Figs. 3E, F, 4Q, R). Both are complete and have walls perpendicular to Multiple drilled specimens are common (9/14 or 64%). the shell surface (cylindrical), and are ascribed to the ichnospecies Drill holes in twenty specimens from other samples from a variety of Oichnus simplex Bromley, 1981. localities range from 0.33 to 1.4 mm (mean = 0.8 mm, n = 49) In the Stuores 2010-3 sample, drill hole diameters vary from (Table 2). A third of the drill holes are incomplete (16/49, 33%). Al- 0.3–1.1 mm (mean = 0.7 mm, n = 37) (Table 1). Most drill holes are though the mean outer diameter of complete drill holes appears higher

Fig. 2. Quantitative surface collecting at the Stuores Wiesen, where the Stuores 2010-3 sample originates from. A. Typical aspect of the smooth-hilled landscape of the Stuores Wiesen (~2000 m altitude) formed by the soft argillaceous basin sediments; fossiliferous locations occur where vegetation is absent e.g., location Stuores 2010-3 just above A.N. B. Quantitative surface collecting in 2013 by ﬁve persons including A.N. and A.K. at a typical fossil location of the Stuores Wiesen. Snow-covered Dolomite Mountains in the background representingfor- mer carbonate platforms; deeper areas with forests and meadows represent argillaceous basin sediments of the Cassian and Wengen Formations. 346 A.A. Klompmaker et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 457 (2016) 342–359

Fig. 3. Circular drill holes (ichnotaxon Oichnus spp.) from the Late Triassic Cassian Formation. A–D, G. Parabolic incomplete drill holes with a central boss (Oichnus excavatus), all in the same specimen of the gastropod Polygyrina lommeli, locality unknown, NHM 35406A. E, F. Cylindrical complete drill hole (Oichnus simplex) in an unidentified articulate brachiopod, Alpe di Specie (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 is fragmented, especially the high-spired gastropod P. lommeli that usually not statistically significant (Mann-Whitney U = 81, p = 0.12). Drill hole consists of 2–5 whorls. Below, some taxa are discussed briefly, particu- size is not correlated significantly with shell width (R2 = 0.433, p = larly those with quantitative data. For others, referral is made to Tables 1 0.16) and height (R2 = 0.478; p = 0.12). Specimens that are drilled and 2. more than once are common (10/20 or 50%). Drill holes in the shells of the Zardini (1978, 1981, 1985) mono- 3.1.2.1. Bivalvia. Palaeonucula strigilata is one of the most abundant fos- graphs are of similar size (0.4–1.6 mm, mean = 0.94, Appendix 1), sils of the Cassian Formation, especially in the soft-bottom communities and the diameters of complete and incomplete drill holes sizes are not of the Stuores Meadows. As is usual for nuculoids, it is interpreted as an statistically different (Mann-Whitney U = 71, p = 0.28). Drill hole infaunal mobile deposit feeder. It is almost always preserved with arti- size is not correlated with shell width (R2 = 0.0007, p = 0.92) and culated valves, which is also the case for drilled specimens. Drill holes height (R2 = 0.003; p = 0.85). of this species were illustrated by Zardini (1981, 1985) and Fürsich and Jablonski (1984) from the localities Staolin, Vervei, and Costalaresc 3.1.2. The prey — drilled taxa from the Cassian Formation (Appendices 1, 2). We also include Pa. strigilata f. jugulata sensu At least 23 species have revealed drill holes so far, all but one of them Zardini (1981). We have examined an articulated specimen from being mollusks (Appendix 2), which is higher than previously assumed the Stuores Wiesen that has a single complete drill hole in the umbil- because several drilled gastropod taxa are reported for the first time ical region (diameter = 1.1 mm). here (Appendix 2). Only two drilled brachiopod specimens (possibly An articulated specimen of the nuculoid Prosoleptus lineatus from representing a single species) are present in the investigated collections Stuores (Stuores 7, coll. 2013) has two drill holes, one complete (dia- from the Cassian Formation. The drilled specimens have a height range meter = 1.4 mm) near the umbo and one incomplete with central of 3–27 mm (Tables 1, 2; Appendix 1), although they are commonly boss (diameter = 1.14 mm) (Fig. 4N, O). The presence of drill holes in A.A. Klompmaker et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 457 (2016) 342–359 347

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. Unidentiﬁed caenogastropod, 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 and PZO 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. 348 A.A. Klompmaker et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 457 (2016) 342–359

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

Gastropod (g) Taxon Preservation Height Width or # drill Outer drill hole Outer drill hole Outer drill hole or bivalve (bi) (mm) length for holes diameter complete diameter incomplete diameter, bi (mm) drill hole (mm) drill hole(s) (mm) completeness drill hole(s) unknown (mm)

g Polygyrina lommeli A Fractured N10.5 ≥5.3 6 0.7 0.7 0.7 0.6 0.6 0.7 g P. lommeli B Fractured N14.9 ≥5.4 1 0.7 g P. lommeli C Fractured N9.0 ≥5.2 2 0.6 0.8 g P. lommeli D Fractured N9.7 ≥3.4 2 1.0 0.7 g P. lommeli E Fractured N9.4 ≥3.5 1 1.0 g P. lommeli F Fractured N14.0 ≥6.0 6 0.7 0.7 0.7 0.7 0.5 0.3 g P. lommeli G Fractured N6.0 ≥3.6 1 0.5 g P. lommeli H Fractured N8.6 ≥3.8 5 0.8 1.0 1.0 0.7 0.6 g P. lommeli I Fractured N5.2 ≥2.6 2 0.8 0.5 g P. lommeli J Fractured N7.7 ≥3.8 1 0.9 g P. lommeli K Fractured N10.7 ≥5.0 2 0.6 1.1 g P. lommeli (non-drilled A) Fractured N14.5 N6.0 g P. lommeli (non-drilled B) Fractured N10.0 N5.0 g P. lommeli (non-drilled C) Fractured N10.0 N4.5 g P. lommeli (non-drilled D) Fractured N7.0 N5.0 g P. lommeli (non-drilled E) Fractured N6.0 N3.0 g P. lommeli (non-drilled F) Fractured N6.3 N2.5 g Kittliconcha obliquecostata Fractured N10.8 ≥5.8 4 0.7 0.7 0.7 0.8 (Bronn in zu Münster, 1841) g Neodonaldina? n. sp. Apex complete ≥5.4 ≥2.8 1 0.7 bi Cassianella sp. Nearly complete, 5.2 8.6 3 0.5 0.9 0.7 disarticulated specimens of Prosoleptus was also mentioned by Fürsich and Jablonski encrusted or poorly preserved. (C) A surface collection from the Stuores (1984). The life habit is the same as for Pa. strigilata. Wiesen (Stuores 2010-3) yielded 17 specimens of P. lommeli,ofwhich Specimens of Cassianella are abundant in the Cassian Formation. This 11 specimens contain drill holes and seven of them have multiple drill soft-bottom recliner has a strongly convex lower (left) valve and a flat holes (Table 1). Drilled specimens are not of a statistically different upper (right) valve as a cover. The drill holes are always located in the height (Mann-Whitney U = 29.5, p = 0.75) or width (Mann-Whitney lower valve (Zardini 1981; Fürsich and Jablonski 1984; pers. obs. U = 30.5, p = 0.84) compared to non-drilled specimens, although all A.N.), but the flat upper valves are rarely preserved. Three drilled specimens are fractured. species have been reported so far (Cassianella ampezzana Bittner, 1895, C. euglypha Laube, 1865, and C. beyrichi Laube, 1865). Most drill 3.1.2.3. Brachiopoda. Drill holes were found in an articulated specimen holes were reported in shells of C. ampezzana from Costaresc and Alpe (cf. Spirigera wissmanni) and a shell fragment (clearly of brachiopod na- di Specie, some of them with up to five drill holes on a single valve ture as is indicated by the presence of typical prisms as the shell micro- (Table 2). We report that Cassianella sp. from the Stuores Wiesen structure), both from the location Alpe di Specie (Seelandalpe) (Fig. 4Q, (Stuores 2010-3 sample), clearly distinct from the known drilled species R). The holes are circular and have diameters of 0.34 and 0.50 mm. This in the Cassian Formation, shows three incomplete drill holes (Fig. 4L, M; is the first report of drilled brachiopods from the Cassian Formation and Table 1). from the entire Triassic. Palaeocardita is an endobyssate suspension feeder present in some of the soft-bottom associations (see Fürsich and Wendt, 1977). Zardini 3.1.3. Drilling percentage (1981, 1985) reported a drilled specimen of Palaeocardita beneckei The drilling percentage in the various assemblages of the Cassian (Bittner, 1895)andP. pichleri (Bittner, 1895)fromAlpediSpecie(Ap- Formation cannot be assessed comprehensively yet because there are pendix 1). We report a single valve of P. crenata (zu Münster, 1841) too few quantitative data sets available. The only previous drilling per- from Alpe di Specie that shows a single complete drill hole of 0.7 mm centage was provided by Fürsich and Wendt (1977), who reported in diameter (Fig. 4K). that ~6% of the 460 specimens from the Stuores marls was drilled. Strong variations in drilling percentages exist within the Cassian 3.1.2.2. Gastropoda. Polygyrina lommeli is one of the most abundant gas- biota. Generally, transported shallow-water assemblages, as indicated tropods of the Cassian Formation. It is a high-spired, largely smooth gas- by the presence of oncoids, ooids, and encrusted fossils in a clayey ma- tropod shell. Almost all known specimens are fractured as is also trix and shallow-water derived ooids and oncoids in calcareous turbi- common in other turreted shells from the Cassian Formation; fragments dites (e.g., Misurina Skilift, parts of Alpe di Specie), have a low drilling usually have two to seven whorls. Kittl (1894) mentioned that percentage (b1%). The large and diverse allochthonous assemblage P. lommeli commonly has multiple drill holes. Here, we document from the Stuores Meadows (Hausmann and Nützel, 2015) contained these drill holes for the first time. Specifically, the following specimens no drilled specimens at all. However, specimens from this collection have been examined. (A) A specimen from the Naturkundemuseum are commonly encrusted so that drilling may be obscured. Bulk samples Berlin (MB.Ga.44755) contains eight drill holes, at least one of them yielded small specimens mostly that are rarely drilled. completely penetrating the shell (Fig. 4B, C). (B) During a survey of In contrast, autochthonous soft-bottom assemblages, as indicated by a the Klipstein (1843) collection (one of the earliest monographs of the very high percentage of articulated infaunal bivalves such as Pa. strigilata Cassian fauna) in the Museum of Natural History in London, a lot and Pr. lineatus in the quantitative surface collection from the Stuores (35,406) with 14 well-preserved specimens was studied, which Wiesen (Stuores 2010-3), commonly yield a relatively high number of contained six specimens with multiple conspicuous drill holes drilled specimens, although many show incomplete drill holes. The sam- (Table 2). Other lots of P. lommeli from the same collection with fewer ple Stuores 2010-3, comprising 119 benthic invertebrate specimens (plus or no drilled specimen were available, but these specimens are several cidarid sea urchin spines not included in the analysis), yielded a A.A. Klompmaker et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 457 (2016) 342–359 349

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

Gastropod (g), Taxon Preservation Museum Locality Height Width # Outer Outer drill hole Outer drill hole bivalve (bi) or number if (mm) or drill drill hole diameter incomplete diameter, brachiopod known length holes diameter drill hole(s) (mm) completeness drill (br) for bi complete hole(s) unknown (mm) drill (mm) hole(s) (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 0.6 g P. lommeli (specimen A) Fractured NHM 35406A 1 1.2 g P. lommeli (specimen B) Fractured NHM 35406B 5 1.2 0.70.80.5 0.8 g P. lommeli (specimen C) Fractured NHM 35406C 2 0.5 0.5 g P. lommeli (specimen D) Fractured NHM 35406D 4 1.2 0.60.60.5 g P. lommeli (specimen E) Fractured NHM 35406E 3 0.9 1.2 0.9 g P. lommeli (specimen F) Fractured NHM 35406F 1 1.2 g Atorcula canalifera (zu Fractured NHMW N11.5 ≥5.0 2 1.0 0.67 Münster, 1841) 1899 V 406B g Atorcula? sp. Fractured Lago Antorno, N10.5 ≥5.0 1 1.0a 0.9 near Misurinaa g Atorcula? sp. Fractured Lago Antorno, N9.5 ≥5.1 1 0.7 near Misurinaa g Anoptychia carinata Fractured Stuores Wiesen N11.9 ≥7.5 7 0.6 0.7 0.5 0.4 0.4 1.0 0.4 (zu Münster, 1841) 2010-3-7, surface coll. 2013b g Kittliconcha obliquecoststa Fractured Stuores 2010-6b N14.2 ≥5.7 4 0.9 0.9 0.9 0.7 g Coelostylina conica (zu Apex Lago Antorno, 4.3 2.3 1 0.8 Münster, 1841) complete, near Misurinaa juvenile g C. conica Apex Lago Antorno, 4.8 2.9 1 0.33 complete, near Misurinaa juvenile g C. conica Apex Lago Antorno, 3.5 2.0 1 0.8 complete, near Misurinaa juvenile bi Palaeonucula strigilata Complete, Stuores Wiesenb 14.5 10.0 1 1.1 (Goldfuss, 1837) articulated bi Prosoleptus lineatus Complete, Stuores 11.0 8.0 2 1.4 1.1 (Goldfuss, 1840) articulated (Stuores 7, coll. 2013)b bi Palaeocardita crenata Fractured, Alpe di Specie ≥2.2 N3.2 1 0.7 (zu Münster, 1841) disarticulated br cf. Spirigera wissmanni Complete, Alpe di Specie 5.5 4.3 1 0.34 zu Münster, 1841 articulated br Brachiopod Fractured, Alpe di Specie N3.4 N7.0 1 0.5 disarticulated

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 be fragments of a few whorls, so that the number of drill holes in complete considered a minimum percentage because many specimens are specimens could have been even higher. Although this collection is fragmented. Parts of the shell that were not preserved may have small, its study shows that the drilling percentage in the Cassian Formation contained drill holes. Including incompletely drilled specimens yields 14 can exceed 10% of the specimens if incomplete specimens are included drilled shells and a drilling percentage of 12.0%. This collection is dominat- and that gastropods appear to be the preferred prey. ed by the gastropods P. lommeli and Rhaphistomella radians,bythe nuculoid bivalves Pa. strigilata and Pr. lineatus as well as by the scaphopod 3.2. Polish Lower Muschelkalk Plagioglypta undulata. These are typical soft-bottom basin dwellers be- longing to the autochthonous R. radians/Pa. strigilata association (sensu The sample from the Lower Muschelkalk of Poland contains one drill Fürsich and Wendt, 1977). Typically, the infaunal bivalves Pa. strigilata hole in a single specimen (Fig. 6) of articulated valves of C. vulgaris, and Pr. lineatus have articulated valves, showing that this assemblage resulting in a drilling percentage of 0.3% (1/365). The drill hole with a was not transported. The drilled specimens represent three gastropod spe- diameter of 1.9 mm is located in the dorsal or brachial valve (30.7 mm cies and the bivalve Cassianella sp. Specimens of P. lommeli show more drill long; 24.4 mm wide) to the right-posterior of the umbo. The subcircular, holes than other species. The two most abundant gastropod species in this cylindrical drill hole (O. simplex)completelypenetratestheshell. collection (Stuores 2010-3) are P. lommeli (14%) and R. radians (12%). Whereas 11.8% (n = 17) of the P. lommeli specimens show a complete 4. Discussion drill hole and 65% show a drill hole (complete + incomplete drill holes), no drill holes are present in R. radians (Fig. 5). Twenty-ﬁve percent of 4.1. Naticids as drill hole producers in the Triassic? the gastropod specimens exhibit drill holes, showing that drilling in gastropods can be frequent at least locally. Of the 14 drilled specimens, nine Most of the drill holes found in the Cassian Formation represent have multiple drill holes with 2–6 drill holes per specimen (Table 1). Oichnus paraboloides and O. excavatus, whereas only two drill holes This high proportion (64%) of specimens with multiple drill holes is re- can be ascribed to O. simplex. The parabolic drill holes closely resemble markable especially since these specimens are usually teleoconch the drill holes of modern naticids (see also Fürsich and Jablonski, 350 A.A. Klompmaker et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 457 (2016) 342–359

Table 3 Total 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), Taxon # specimens # specimens # specimens Preservation of Corrected Drilling percentage bivalve (bi), drilled with complete valves # specimens for taxa with ≥10 brachiopod (br) drill holes specimens or scaphopod (sc)

g Polygyrina lommeli (zu Münster, 1841) 17 11 2 17 11.8 g Rhaphistomella radians (Wissmann in zu 14 14 0 Münster, 1841) g Neritid, strongly encrusted 3 3 g Pseudoclanculcus cassianus (Wissmann in zu 22 Münster, 1841) g Worthenia sp. 1 1 g Cheilotomona blumi (zu Münster, 1841)1 1 g Zygopleura tenuis (zu Münster, 1841)1 1 g Zygopleura sp. 1 1 g Kittliconcha obliquecostata (Bronn in zu 110 1 Münster, 1841) g cf. Neritaria mandelslohi (Klipstein, 1843)1 1 g Neodonaldina? n. sp. 1 1 0 1 g Promathidia subornata (zu Münster, 1841)1 1 g Schizogonium sp. 1 1 g Coelostylina conica (zu Münster, 1841)1 1 gcf.Spirostylus subcolumnaris (zu Münster, 1841)1 1 g Cylindrobullina scalaris (zu Münster, 1841)1 1 g Indet. varia 4 4 sc Plagioglypta undulata (zu Münster, 1841)17 170 ?sc Unknown tube 2 2 bi Palaeonucula strigilata (Goldfuss, 1837) 22 All articulated 22 0 (i.e. 2 valves per specimen) bi Prosoleptus lineatus (Goldfuss, 1837) 14 All articulated 14 0 bi Cassianella tenuistria (zu Münster, 1841) 1 Disarticulated 0.5 (i.e. 1 valve) bi Cassianella sp. 1 1 0 Disarticulated 0.5 bi Paleocardita sp. 1 Disarticulated 0.5 bi Costatoria harpa (zu Münster in Goldfuss, 1838) 1 Disarticulated 0.5 bi cf. Nuculana sulcellata (Wissmann in Münster, 1 Articulated 1 1841) bi Astartid 1 Articulated 1 br Koninckina leonhardi (Wissmann in zu 1 Disarticulated 1 Münster, 1841) br cf. Spirigera wissmanni (zu Münster, 1841) 3 2 articulated, 2.5 1 disarticulated br Brachiopod 1 1 Articulated 1 br Brachiopod 2 1 Articulated 1 Total 119 14 2 116.5 1.7

1984). Fürsich and Jablonski (1984) implied that Naticidae (or

70 Naticoidea) were present as early as the Late Triassic and were the pro- Drilling percentage ducers of these drill holes. Naticids gained the ability to drill, lost it sub- [complete drill holes] 60 sequently, and regained it by the Cretaceous according to Fürsich and Drilling percentage Jablonski (1984). However, there are numerous fossil bulbous largely 50 [all drill holes] smooth gastropods resembling modern naticoids (i.e. naticiform) 40 known since the Paleozoic. Currently, 16 Triassic gastropod species are formally assigned to Natica Scopoli, 1777, which is based on a recent 30 type species (Database A.N. based on the Fosslilium Katalogus by Percentage 20 Diener (1926) and Kutassy (1940)). For most if not all of the Paleozoic and Early Mesozoic “naticids” the assignment to Naticoidea, Naticidae, 10 or even to modern naticid genera may reﬂect a lack of recent revisions,

0 shell convergence, and/or poor preservation. The status of the Triassic- Jurassic “naticids” is doubtful because they may be referrable to the Neritoidea or to extinct Mesozoic families instead (Kabat, 1991). “Natica” is the second most common (after Turritella Lamarck, 1799) gastropod genus in the Paleobiology Database (see Plotnick and Wagner, 2006), and belongs to one of the extremely long-ranging, species-rich, potentially wastebasket taxa. Similarly, Nützel (2005) stated that Natica, Turritella, Trochus Linnaeus, 1758,orTurbo Linnaeus, 1758, are names for general, archaetypic gastropod morphologies rather Fig. 5. Drilling percentages of species represented by ≥10 specimens in the Stuores 2010-3 than names for real biological entities, except for the modern represen- sample. G = gastropod; Sc = scaphopod; Bi = bivalve. tatives of these genera that are closely related to the type species. A.A. Klompmaker et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 457 (2016) 342–359 351

and apparently herbivorous (Kase and Ishikawa 2003). Bandel (1999) mentioned that the Naticidae originated in the mid-Cretaceous, but he did not refer to drill holes and especially not to pre-Cretaceous reports of drill holes. Furthermore, Bandel (1999) characterized the members of 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 naticiform Ampullospiridae, a group that is widespread and diverse in the Mesozoic, is not a carnivorous driller, but an herbivorous grazer. The Ampullospiridae are treated as a synonym of the Ampullinidae and are placed in the non-drilling Campaniloidea by Bouchet et al. (2005). These authors only listed the family Naticidae in Naticoidea. The shell morphology of some Ampullinidae (respectively Ampullospiridae) is most probably convergent. Whereas we agree that naticiform shells evolved repeatedly in unrelated groups (very similar shells are also classified as neritimorphs), the claim by Kase and Ishikawa (2003) that all fossil species placed in Ampullospiridae were Fig. 6. The Middle Triassic (Anisian) brachiopod Coenothyris vulgaris, MWGUW ZI/74/001, herbivorous may be premature (see also Aronowsky and Leighton, 2003). from the Strzelce Opolskie Quarry in Poland with a cylindrical drill hole (Oichnus simplex). In conclusion, we agree with Bandel (1999) that true naticids were absent in the Late Triassic Cassian Formation. Instead, convergent Koken (1892: p. 473) noted that several gastropod species could caenogastropods and neritimorphs are present, which implies that the represent “real naticids” that drill. Fürsich and Wendt (1977: p. 299) drill holes, very reminiscent of naticid drill holes, are caused by a diffe- stated that occasional naticid borings in the umbonal region of nuculids rent organism. and in some convex valves of Cassianella indicated the presence of an infaunal carnivorous gastropod and they suggested that Natiria de Koninck, 4.2. Ecology prey and the drill hole producer 1881,orNaticopsis M'Coy, 1844, could have been responsible. However, both genera are not naticoids, but rather neritimorphs, a group of which Because the bivalve prey was predominantly infaunal, Fürsich and modern representatives are unable to drill. Fürsich and Jablonski (1984: Jablonski (1984) stated that the predator was infaunal as is the case p. 79) mentioned that the naticids (ampullospirids) potentially responsi- in Recent naticids, although naticids can hunt epifaunally as well ble for the drill holes are abundant in the Cassian Formation, notably (e.g., Reyment, 1999; Dietl, 2002; Huelsken, 2011). If the Triassic driller “Ampullina sanctaecrucis, A. paludinaris,andA. subhydridica”. These species was infaunal, it may imply that P. lommeli and Atorcula Nützel, 1998, have been reassigned to different genera, however: Ptychostoma were infaunal or recliners too. Polygyrina is one of the most abundant sanctaecrucis (Laube, 1868) and Prostylifer paludinaris (zu Münster, gastropods in the marly deposits of the Stuores Meadows near 1841). For “A. subhydridica”, Amauropsis subhybrida was probably St. Cassian. These outcrops consist of dark marls that suggest a meant, and this taxon is a synonym of P. paludinaris (Bandel, 1992a). soft-bottom environment. However, Polygyrina is relatively rare in P. paludinaris, an abundant caenogastropod species in the Cassian Forma- the outcrops in the area of Cortina d'Ampezzo. Zardini (1978) tabulated tion, has a tiny larval shell with strong spiral cords (Bandel, 1992a; about 22,000 gastropod specimens representing ~360 species from 17 Hausmann and Nützel, 2015: fig. 7G). This protoconch differs strongly fossil locations in the Cortina area. P. lommeli is present in six localities from naticid protoconchs. The same type of protoconch is also present with a total of only 11 specimens. In contrast, thousands of specimens in P. sanctaecrucis (pers. obs. A.N.). Thus, none of the “naticid” taxa men- have been collected from the Stuores Meadows near San Cassiano tioned by Fürsich and Jablonski (1984) can be regarded as naticids any (pers. obs. A.N. in the Natural History Museums of Vienna and longer. London; Fürsich and Wendt, 1977). Bandel (1988, 1999) stated that all assumed naticids from the So far, all drilled gastropods are high-spired with a smooth or axially Cassian Formation represent members of the Neritimorpha. In addition, ribbed teleoconch, which could point to an infaunal (or facultatively in- some of the assumed naticids (“Ampullina”)(i.e.Prostylifer Koken, 1889, faunal) mode of life. However, Fürsich and Wendt (1977) interpreted and Ptychostoma Laube, 1868) are caenogastropods unrelated to P. lommeli as feeding on algal meadows and living on algae. The pre- naticids based on the size and morphology of their protoconchs. sence of algal meadows as ancient analogs of modern seagrass Modern representatives of naticids are characterized by relatively meadows is, however, speculative because photosynthetic benthos large (up to 1 mm) globular protoconchs (Bandel, 1999). These such as calcareous algae or hermatypic corals are absent in the autoch- protoconchs are weakly ornamented with weak collabral lirae. Bandel thonous basin assemblages near San Cassiano. Thus, there is no indica- (1999) concluded that the earliest confirmed naticids are of mid- tion that the sea bottom was in the photic zone. There are also no Cretaceous age (see also Kollmann, 1982; Tracey et al., 1993). Tracey other indications for very shallow water conditions. In summary, the et al. (1993: p. 148) suggested a late Early Cretaceous first occurrence autecology of P. lommeli remains enigmatic. Its abundance argues that of the Naticidae (Aptian/Albian) and a Middle Jurassic (Aalenian) origin this species is relatively low in the food chain. It was either an epifaunal for the Ampullospiridae, another family that was included in Naticoidea, grazer or detritivore or mobile or stationary infaunal inhabitant (the lat- but is now considered unrelated to the Naticidae (Kase and Ishikawa, ter is the case for modern Turritella, a commonly drilled taxon). An in- 2003; Bouchet et al., 2005). faunal life habit is supported by the fact that the shells are rarely In contrast, Bouchet and Warén (1993: p. 753) noted that the encrusted and the shell is smooth and elongated. However, the autecol- problems of the classification of modern naticids could originate from ogy of gastropods in deep time is speculative because taxonomic unifor- their long geological history back to the Triassic and they referred to mitarianism usually does not work and functional morphology is less illustrations of possible Naticoidea by Zardini (1978).Withoutade- obvious than in other invertebrates such as bivalves (Todd, 2001;pers. tailed explanation, they stated: “We do not share the view of Bandel obs. A.N.). (1988: p. 277) that none of the Cassian gastropods belong to the We found that Cassian parabolic drill holes are most abundant in family Naticidae”. They seemed, however, unaware that there is an- more or less autochthonous soft-bottom assemblages dominated by other group of naticiform gastropods present in the early Mesozoic mollusks such as nuculoid bivalves (mostly with articulated valves), (Ampullospiridae = Ampullinidae), a group that is unrelated to naticids the reclining bivalve Cassianella, the gastropods P. lommeli and 352 A.A. Klompmaker et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 457 (2016) 342–359

R. radians,andthescaphopodPlagioglypta. They are rare or absent in transported 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), most drill holes occur in P. lommeli and in the bivalves Cassianella spp., Pr. lineatus,andPa. strigilata. Although most drill holes are incomplete, it is remarkable that a high percentage of specimens of P. lommeli from an assemblage from the Stuores Wiesen is drilled, whereas the abundant R. radians is not drilled. This result suggests that the producer(s) had clear preferences if both prey taxa were either infaunal or epifaunal in- habitants. The presence of multiple incomplete drill holes suggest that the ability of the producer to fully penetrate the shell was low, that the producer was interrupted frequently, or that it was not the goal to fully penetrate the shell. In addition, the ability of the gastropod prey to move may have made it difficult to finish a drill hole. Infrequently, a Fig. 7. Two compressed specimens of the agglutinating foraminifer Bathysiphon sp. A, D. specimen was found with more than one complete drill hole (at least PZO 4792. B, C, E. PZO 4791. 2/18 or 11% for multiply drilled specimens, Tables 1 and 2) and the same observation was made for Cassian bivalves (Fürsich and Jablonski, 1984). Thus, the producer was most likely able to separate dead shells from living individuals. origin (even though 8/12 incomplete specimens were repaired), but Although many Cassian shells are well-preserved (Fig. 4), micro- did not discuss other potential causes. Likewise, the drill holes in bi- rasp marks have not been found thus far. Such marks are reported valves from the Cassian Formation were only discussed in the context from the Pleistocene (Chiba and Sato, 2016), the Miocene (Pek et al., of predators by Fürsich and Jablonski (1984). Here, four possibilities 1997; Schiffbauer et al., 2008), and the Late Cretaceous (Tapanila are discussed for the drill holes from the Cassian Formation: (1) the et al., 2015), but such marks appear rare (two, three, three, and one drill holes are caused by organisms seeking a home (domichnia) in specimen, resp.). Thus, their absence from the Triassic may reflect dia- shells, (2) the holes are part of bore holes penetrating hardened sedi- genetic alteration. Alternatively, the drill holes may not be produced ments and the embedded shells, (3) drill holes are produced by carniv- by rasping with a proboscis-like apparatus, but principally caused by orous predators such as gastropods, and (4) the drill holes are caused by acids dissolving the calcium carbonate shell instead. The latter scenario parasites. Characteristics of the drilled specimens are summarized in would slow down drilling substantially and explain the high incidence Table 4. of incomplete drill holes and shells drilled more than once. An inefficient proboscis-like apparatus, no use of acids, a poor grip of the produ- 4.3.1. Domicile-seeking organisms (domichnia) cer on its prey so that escape was common, and/or effective defence The Stuores Wiesen sample yielded also numerous (several hun- mechanisms may also explain this result. Defence mechanisms may dreds) fragments of the large agglutinating foraminifer Bathysiphon consist of a high agility/manoeuvrability of the prey and chemical/ Sars, 1872. This foraminifer has the shape of a tube (mostly collapsed toxic repellents to reduce palatability, the latter having been discounted now due to compression) and its diameter is within the range of the for fossil brachiopods (Tyler et al., 2013). Secreting repellents as a de- drill hole diameter (0.3–1.3 mm) (Fig. 7). This raises the possibility fence mechanism is known for modern gastropods from the family that O. paraboloides, especially the incomplete ones, represent dwelling Pleurotomariidae (e.g., Harasewych, 2002). traces or attachment scars of Bathysiphon.Thishypothesisisrejectedfor several reasons. This species is also abundant in other Cassian locations, 4.3. The cause of the drill holes especially in autochthonous soft-bottom assemblages. However, in some of the assemblages (Lago Antorno, pers. obs. A.N.) drill holes are Smith et al. (1985) documented drill holes with a similar morpholo- present, but Bathysiphon is absent, which argues against Bathysiphon gy in Devonian brachiopods and interpreted them to be predatory in causing O. paraboloides. The taxon selectivity further argues against

Table 4 Characteristics of drilled specimens from the Triassic Cassian Formation and how well they ﬁt with each type of possible producer.

Characteristics Type of producer

Post-mortem domicile-seeking Pre-mortem domicile-seeking Borer Parasite Predator organism organism (domichnia) (domichnia)

Shell always penetrated from outside N Y NY Y Drill holes perpendicular to shell surface N N NY Y Specificity for culprit N N NY Y Healing blisters absent Y ? YN Y Multiple holes very common Y Y Y/N Y N Many incomplete holes Y Y NN N Multiple complete drill holes uncommon N N Y/N Y Y No attachment scars Y Y YN Y No attached specimens that may have served as culprit Y/N Y/N Y Y/N Y Preferred drill hole site (Fürsich and Jablonski, 1984, only; Y/N Y/N NY Y not testable herein) Hole size and culprit size not correlated Y Y YY N Holes parabolic with central boss for incomplete ones ? ? N? Y Shape drill holes circular Y/N Y/N Y/N Y/N Y Certain size class culprit preferred (not testable with complete specimens herein) A.A. Klompmaker et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 457 (2016) 342–359 353 domichnia. Finally, the low percentage of specimens with more than 1993). All pre-Cretaceous references to Capulus Montfort, 1810,or one complete drill hole for multiply drilled specimens (at least 11%, capulids are outdated. Three cap-shaped gastropod species from the see Section 4.2.), suggests that post-mortem domichnia are an unlikely Cassian Formation have been assigned to the modern genus Capulus: explanation. The same applies to pre-mortem domichnia (Table 4). of those, Capulus alatus Laube, 1869,andC. fenestratus Laube, 1869, are now assigned to the neritimorph genera Marmolatella (Bandel, 4.3.2. Boring organisms 2007)andPseudorthonychia Bandel and Fryda, 1999. Capulus muensteri Circular perforations can also be made by boring organisms in hard Giebel, 1852, has not been revised yet. These gastropods are generally substrates, thereby completely penetrating shells. Paleozoic brachiopod rare in the Cassian Formation and are absent from the Stuores 2010-3 shells were bored this way (Richards and Shabica, 1969; Wilson and sample. Morever, modern capulids produce a different drill hole mor- Palmer, 2001). Such bore holes tend to be cylindrical and complete, phology with an attachment scar and they prefer pectinid bivalves may penetrate the shell at an oblique angle, and should not be (Orr, 1962; Kosuge and Hayashi, 1967; Matsukuma, 1978). site and prey-specific. Hardgrounds are unknown from the Cassian For- Modern parasitic eulimid gastropods can also drill (e.g., Wáren et al., mation and their existence in the clayey soft-bottom environment is 1994). Eulimids parasitizing echinoderms, especially echinoids, produce also unlikely. Thus, such bore holes cannot explain the holes seen in multiple drill holes (Kowalewski and Nebelsick, 2003), and their the Cassian shells. drill holes can show a circular groove around it (e.g., Neumann and Wisshak, 2009). Eulimids originated in the Cretaceous (Dockery, 4.3.3. Predators 1993) and, as for Naticidae and Capulidae, Triassic references (Kittl, 1891) are outdated. 4.3.3.1. Parabolic drill holes. The fact that multiple drill holes and incom- As for a predatory origin of the drill holes, there is ample evidence plete drill holes are so frequent with extremes of eight drill holes on a against a parasitic producer: the inferred absence of healing blisters, single gastropod fragment of about four whorls (another example is a the presence of many incomplete drill holes, and the absence of attach- fragment of two whorls having six drill holes), sheds doubt as to whe- ment scars. ther these drill holes are predatory in origin. Such an inefficient predator could hardly evolve and survive. Moreover, such a high percentage of 4.3.5. A clue about the culprit from the Paleozoic? incomplete drill holes (40/86 or 47%, for all drilled Cassian shells) and As naticids, capulids, and eulimids cannot have been the drill hole many specimens with multiple drill holes (19/34 or 56%) is unknown producers of O. paraboloides and O. excavates from the Cassian For- to us from modern drilling predators. The fact that the majority (24/ mation, another culprit is must be responsible. Instead of looking 37 or 65%) of the drill holes from the Stuores 2010-3 sample is incom- for a late Mesozoic and/or Cenozoic producer, detailed analyses of plete and the low percentage of specimens with a single hole (5/14, drill holes from the Paleozoic may provide more insight (Fig. 8, 36%) may argue against predation as the cause. Moreover, and contrary Table 5). to the previous assumption, there are no naticids present in the Cassian Brett (2003) divided Paleozoic drill holes into four categories, and he Formation. classified the drill holes in Middle Devonian brachiopods from New If the traces were produced by a predator, its producer remains un- York as presented in Smith et al. (1985) as type 3: conical, chamfered, known at present. It may be a non-naticid gastropod because there are 1–3 mm in diameter, and incomplete drill holes have a raised central various modern gastropod groups that drill similar-sized (sub)circular boss or raised knob. Brett (2003: table 4) interpreted type 3 drill holes holes today (Kowalewski, 1993, 2002). Moreover, gastropods are to be caused by predatory gastropods. Drill holes in Middle Devonian the only motile benthic group that is abundant at the locations with (Buehler, 1969)andMississippian(Ausich and Gurolla, 1979)brachio- reported drill holes. If the drill holes were predatory in origin, the pods from New York and Indiana, respectively, were also ascribed to producer may be identified by a careful analysis of co-occurrences at this type. The Middle Devonian brachiopods may not be ascribed to the various Cassian locations if the predator possessed a mineralized this type because Buehler (1969) indicated that incomplete drill holes skeleton, but our current data are too sparse for this purpose. are flat-bottomed and most drill holes are not beveled or tapered, although some possess a slight flange near the bottom. More drilled spe- 4.3.3.2. Cylindrical drill holes. The two cylindrical drill holes (O. simplex) cimens can be assigned to type 3. Fenton and Fenton (1931) described found in two brachiopod shells from the Cassian Formation and the sin- drilled Devonian brachiopods; one of the incomplete ones contained a gle drill hole in a brachiopod from the Lower Muschelkalk may be pre- raised central boss (see Carriker and Yochelson, 1968:B15).Brett and datory in origin. Both are completely penetrating the shell, oriented Bordeaux (1990) mentioned Middle Devonian brachiopods with perpendicular to the shell surface, no attachment scars are found, and beveled holes, including incomplete drill holes with a raised central multiple drill holes of this type in the same shell are unknown thus boss. Brett (2003), via pers. comm. with S. Felton in 2001, briefly far. More drilled brachiopod specimens with such drill holes are needed mentioned Upper Ordovician brachiopods, gastropods, and crinoids to further test this hypothesis. from Ohio with type 3 drill holes, a result that would be well worth describing further. Some years later, Daley (2008) mentioned and figured 4.3.4. Parasites (her figs. 7.5, 7.6) some incomplete, parabolic drill holes in Devonian The drill holes referred to O. paraboloides may also have a parasitic brachiopods from Canada with much more pronounced raised central origin. This hypothesis is supported by a combination of several lines bosses that may or may not be ascribed to this type. Finally, we as- of evidence (cf. Daley, 2008: Table 1): the presence of multiple drill cribe the drill holes from the Cassian Formation classified as holes for many specimens, drill holes are placed perpendicular to the O. paraboloides and O. excavates to type 3 drill holes, including shell surface, and taxon selectivity. Parasites are often very small those in Fürsich and Jablonski (1984). Many of them are smaller in comparison to their host (Warén, 1981, for eulimid gastropods; than 1 mm, but drill hole size can differ substantially for modern Klompmaker and Boxshall, 2015, for crustaceans), but the diameter of drill hole producers (e.g., Kowalewski, 1993) depending on the size O. paraboloides in the high-spired gastropod may suggest a relatively range of the producer. Some other drill holes have central raised large producer. bosses, but are not parabolic: Miller and Sundberg (1984) mentioned The drill holes produced by parasitic capulids can be up to ~5 mm in straight-sided drill holes in Upper Cambrian brachiopods with raised diameter (e.g., Orr, 1962; Kosuge and Hayashi, 1967; Matsukuma, central bosses. Thus, type 3 drill holes seem to be present in at least 1978). Capulidae have been mentioned from the Cassian Formation the Devonian, Mississippian, and Triassic. Naticiform drill holes (zu Münster, 1841; Böhm, 1895), but this caenogastropod family origi- occur at least in these geological periods, in addition to the Creta- nated in the Cenozoic or Late Cretaceous (Bandel, 1993; Tracey et al., ceous and Cenozoic occurrences, when naticids were present. We 354 A.A. Klompmaker et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 457 (2016) 342–359

60 Percentage of drilled specimens 55 with multiple drill holes 50 Percentage of drill holes that is 45 incomplete 40 35 30

Percentage 25 20 15 10 5 0

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

call these Triassic and Paleozoic naticiform drill holes an example of strategies were mentioned (Rollins and Brezinski, 1988; Horný, 2000). “drill hole convergence” because they must have been produced by Baumiller et al. (1999) argued that host switching from echinoderm to organisms other than naticids. brachiopod hosts may have been possible. After the switch to brachio- These Paleozoic and Triassic occurrences have more in common pods, the platyceratids still may have been parasitic because the drill than the parabolic drill hole shape and raised central bosses for in- holes were mostly positioned near the lophophore, an ideal location complete drill holes. The percentage of multiple drill holes per shell to steal food (Baumiller et al., 1999). and incomplete drill holes of all drill holes are strikingly high with Platyceratids were present in the Middle Devonian of New York, values in excess of 40%, an almost unique combination for Paleozoic from where type 3 drill holes in brachiopods have been found (Smith and Mesozoic drill holes (Fig. 8, Table 5). This combination is not mu- et al., 1985: Naticonema Perner, 1903,andPlatyceras Conrad, 1840; tually exclusive because specimens can be drilled again when an (in- Brett and Bordeaux, 1990; Bezusko, 2001: Platyceras; Brett et al., 2007: complete) drill hole is abandoned. At least two complete drill holes Platyceras ), as is the case for drilled Misssissippian brachiopods per shell are uncommon (b20% of shells with multiple drill holes, from Indiana (Ausich and Gurrola, 1979: Platyceras (Platyceras)and Table 5), which is to be expected because a complete predatory Platyceras (Orthonychia)). This co-occurrence is further corroborated drill hole is often lethal and/or there is less space for a second para- by Leighton (2003), who observed that platyceratids are present site to attach to the shell. wherever type 3 drill holes have been reported; Fenton and Fenton The similarity in drill hole characteristics justifies the question (1931), who put forward Platyceras spp. as drill hole producers in whether they could be produced by the same group or, alternatively, Devonian brachiopods; and Brett (2003) noted that drill holes in whether members of different hole-producing lineages caused these Upper Ordovician invertebrates are associated with the platyceratid drill holes. In the first case, the responsible group should have a varied gastropod Cyclonema Hall, 1852.Platycerashas been interpreted as a or changing diet: from various brachiopods in the Devonian to bivalves coprophagous feeder (Smith et al., 1985; Bezusko, 2001), but Baumiller and mostly gastropods in the Triassic (Smith et al., 1985; Tables 1, 2). (1990) unequivocally showed that Platyceras (Orthonychia)candrill. Platyceratids, an extinct gastropod family, are the only gastropods For Naticonema, Brett (2003) interpreted this platyceratid, found in that are known to drill their victims in the Paleozoic (although faculta- close association with the drilled Middle Devonian brachiopods, to tively). Drill holes by platyceratids are most frequently reported in represent a non-specialized platyceratid gastropod that may have been crinoids and blastoids that show cylindrical to conical drill holes a predatory driller. Platyceratids can leave muscle scars on the animal to (e.g., Baumiller, 1990: fig. 2H, 1993, 1996; Baumiller and Macurda, which they attach (Rollins and Brezinski, 1988, and references), 1995). Drill holes in brachiopods and platyceratid gastropods were but how often this occurs is unknown. In the case of predatory also attributed to platyceratids (Baumiller et al., 1999, 2004b; Brett, platyceratids, no muscle scars are expected. 2003), indicating that platyceratids, as a group, drilled more prey than Although platyceratids were thought to have gone extinct with previously assumed. Platyceratids have also been found attached to camerate and inadunate crinoids in the Permian (Bowsher, 1955), cystoids (Clarke, 1908; Bowsher, 1955; Kluessendorf, 1983) and dead platyceratids have been claimed from the Triassic. Bandel (1992b) remains of orthocone nautiloid cephalopods plus other substrates on assigned Orthonychia alata (Laube, 1869) from the Cassian Formation the ocean bottom (Horný, 2000), but drill holes were not reported in to this family and proposed, without any evidence, that this taxon this case. Platyceratid drilling in echinoderms and brachiopods is pre- would have parasitized a crinozoan host. This species became the type sumed to have been kleptoparasitic (Baumiller, 1990, 2003; Baumiller species of Pseudorthonychia Bandel and Fryda, 1999,andwasplacedin et al., 1999, but see Brett, 2003, who suggested predatory drilling), Pseudorthonychiidae, however. To our knowledge, this rare species whereas a coprophagous and gametophagous mode of life, not associa- has never been found attached to a possible victim. Nevertheless, ted with drilling, has also been suggested (e.g., Bowsher, 1955; Lane, Hausmann and Nützel (2015) referred to this species as parasitic, citing 1984; Sutton et al., 2006). Additionally, algal grazing and filter-feeding Bandel (1992b). Could it be possible that the drill holes from the Cassian A.A. Klompmaker et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 457 (2016) 342–359 355

Table 5 Characteristics 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 characteristics to drill holes from the Cassian Formation in gray.

Stratigraphic age Taxon Country Size drill holes General drill hole Percentage of Percentage of Incomplete Percentage Reference (state/province) (mm): range shape (circular in drilled drill holes that drill holes of shells (mean) outline unless specimens with is incomplete with central with ≥2 stated otherwise) multiple drill and the number boss complete holes and the of specimens drill holes of number of involved shells with specimens ≥2 parabolic involved drill holes

≥11.1 herein (Stuores 2010-3 Triassic (Carnian) Mollusca Italy 0.3–1.1 (0.7) parabolic 64.3 (9/14) 64.9 (24/37) Y (≥2/18) sample) Permian (Upper) Bivalvia Brazil 0.4–5.8 (2.0) cylindrical 12.5 (2/16) 0.0 (0/16) Kowalewski et al. (2000) brachiopod Carboniferous Cardiarina cordata USA (New (Pennsylvanian) Cooper, 1956 Mexico) 0.03–0.19 (0.1) slightly beveled 5.3 (7/131) 0.0 (0/131) Hoffmeister et al. (2003) Carboniferous Y Ausich and Gurrola (Mississippian) Brachiopoda USA (Indiana) 1.6–3.2 parabolic rare rare (sometimes) 0.0 (1979) Carboniferous Ausich and Gurrola (Mississippian) Brachiopoda USA (Indiana) 0.1–1.6 cylindrical common (1979) Carboniferous cylindrical to (Mississippian) Brachiopoda USA (Kentucky) 1.1–4.7 (2.8) slightly tapered 4.3 (2/46) Baumiller et al. (1999) brachiopod Perditocardinia cf. P. dubia (Hall in Carboniferous Hall and Whitney, (Mississippian) 1858) USA (Missouri) 0.13–0.80 (0.40) ?cylindrical 0.0 (0/67) Deline et al. (2003) brachiopod Carboniferous Rhipidomella USA (Montana, Rodriguez and Gutschik (Mississippian) Oehlert, 1890 Utah) cylindrical/parabolic ~5.2 (1/~19) N (1970) Carboniferous (Mississippian) Brachiopoda Northern Ireland 0.1–1.0 ?cylindrical 4.3 (2/46) Brunton (1966) brachiopod Carboniferous Crurithyris George, cylindrical or Mottequin and (Mississippian) 1931 Belgium 0.16–1.08 (0.57) weakly conical 0.0 (0/35) 0.0 (0/60) Sevastopulo (2009) blastoid Carboniferous Orophocrinus von cylindrical to Baumiller and Macurda (Mississippian) Seebach, 1865 0.5–2.5 (1.3) slightly tapered 17.4 (4/23) ≥10.7 (≥3/28) (1995) ~55.6 14.9 (21/141) Devonian (Frasnian) Brachiopoda USA (Iowa) (0.8) cylindrical (≥90/≥162) N Leighton (2001) brachiopod Pholidostrophia 4.2-8.3 (1-2/24) Hall and Clarke, cylindrical, Devonian (Givetian) 1892 USA (Ohio) 0.5–1.3 unbeveled 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) brachiopod Spinocyrtia Brett and Bordeaux Devonian (Middle) Fredericks, 1916 USA (New York) 2.5–3.0 ?parabolic 45.4 (5/11) 45.4 (5/11) Y 18.2 (2/11) (1990) USA (Ohio, New York) and Canada Devonian (Middle) colonial metazoans (Ontario) 0.15–0.30 (0.24) beveled 14.3 (3/21) Wilson and Taylor (2006) hyolithid Hallotheca cf. H. aclis (Hall, conical or Devonian (Middle) 1876) Canada (Ontario) 0.3–0.9 (0.6) cylindrical 0.0 (0/7) 0.0 (0/7) Baumiller et al. (2010) blastoid Heteroschisma conical or Devonian (Middle) Wachsmuth, 1883 0.3–2.4 (0.9) cylindrical 5.0 (7/139) Baumiller (1996) Lespérance and Sheehan (1975), Sheehan and Devonian (Lower) Brachiopoda Canada (Quebec) 0.8–1.7 non-beveled 0.0 (0/13) Lespérance (1978) USA (Ohio, conical or Devonian Blastoidea Indiana) 1.1–2.4 (1.7) cylindrical 2.0 (1/51) 1.9 (1/52) Baumiller (1993) brachiopod Dicaelosia King, Silurian (Wenlock) 1850 Canada (Nunavut) (0.32) cylindrical 2.9 (2/68) 2.9 (2/68) N Rohr (1976) brachiopod Artiotreta Ireland, Chatterton and Silurian 1961 USA (Oklahoma) 0.08–0.13 countersunk 1.4 (1/71) ≥1.4 (≥1/72) N Whitehead (1987) brachiopod Dalmanella Hall Bucher (1938), Carriker Ordovician (Upper) and Clarke, 1892 USA (Ohio) 1.2–3.1 (2.05) cylindrical/beveled 8.3 (2/24) and Yochelson (1968) USA (New York, Ordovician (Middle) Brachiopoda 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, Canada (Eastern N (few Silurian, Devonian Brachiopoda provinces) (≤0.48) cylindrical mostly 0-62.8 0-100 exceptions) Daley (2008) USA (California, Miller and Sundberg Cambrian (Upper) Brachiopoda Nevada) 0.05–0.2 (0.11) cylindrical 0.0 (0/13) ≥7.7 (≥1/13) Y (1984) Cambrian (Middle- USA (South 0.066–0.130 Upper) Brachiopoda Dakota) (0.170) non-beveled 1.6 (1/63) Robson and Pratt (2007) Cambrian (Middle- USA (South 0.137–0.675 Upper) Brachiopoda Dakota) (0.342) irregular 0.0 (0/62) Robson and Pratt (2007) brachiopod Linnarssonia 0.01–0.24 Conway Morris and Cambrian (Middle) Walcott, 1885 Sweden (0.0925) cylindrical mostly 36.7-58.8 Bengtson (1994) 0.05–0.17 Cambrian (Middle) Brachiopoda Morocco (0.097) various 13.2 (7/53) 47.0 (31/66) N Streng (1999)

Mobergella Conway Morris and Cambrian (Lower) Hedström, 1923 Sweden 0.07–0.24 cylindrical 12.5 (1/8) 55 (5/9) N Bengtson (1994) 356 A.A. Klompmaker et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 457 (2016) 342–359

Formation were caused by P. alata? This species appears too rare to pro- very useful comments that improved this article. A.A.K. was in part duce that many drill holes. It was, however, reported from the Stuores supported by the Jon L. and Beverly A. Thompson Endowment Fund. Wiesen (where the Stuores 2010-3 sample was also collected), which A.N. acknowledges the Deutsche Forschungsgemeinschaft (project NU yielded a shallow-water assemblage that was transported to the basin 96/13-1). Funding sources had no role in the study design, data collec- by mass ﬂows (Hausmann and Nützel, 2015). The shell shape of tion, analysis and interpretation of the data, and during the writing P. alata is reminiscent of species that can attach to substrates such as and submission process. This is University of Florida Contribution to platyceratids and capulids. If specimens of Pseudorthonychia are the pro- Paleobiology 819. ducers of the drill holes, this is yet another example of a lineage producing naticiform drill holes. References

4.3.6. Synthesis Aberhan, M., Scholz, A., Schubert, S., 2011. Das Ober-Pliensbachium (Domerium) der The cylindrical drill holes (O. simplex) in brachiopods from the Herforder Liasmulde - Teil 3 - Taxonomie und Paläoökologie der Bivalvia aus der Cassian Formation may have been caused by a predator. The producer Amaltheenton-Formation (Unterjura) der Herforder Liasmulde. Geol. Paläont. in Westfalen 80, 61–109. remains unknown, however. Aronowsky, A., Leighton, L.R., 2003. Mystery of naticid predation history solved: evidence Most observations with regard to the parabolic drill holes from a “living fossil” species: comment and reply COMMENT. Geology 31, e34–e35. (O. paraboloides and O. excavatus) from the Cassian Formation are Ausich, W.I., Gurrola, R.A., 1979. Two boring organisms in a Lower Mississippian community of southern Indiana. J. Paleontol. 53, 335–344. in line with either a predatory or parasitic origin (Table 4). Choosing Bandel, K., 1988. 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Similar-shaped drill holes are also present in Devonian and Carboni- der Aviculiden. Abh. der königlich-Jahrb. Preuss. Geol. Landesanst., neue Serie 17, ferous brachiopods and resemble drill holes produced by extant 1–514 (pls. 1–38). Beyrich, E., 1862. Zwei aus dem deutschen Muschelkalk noch nicht bekannte Avicula- naticid gastropods, despite the absence of naticids in pre-Cretaceous artige Muscheln. Z. Dtsch. Geol. Ges. 14, 9–10. rocks. This is an example of drill hole convergence. Bezusko, K.M., 2001. Biotic Interaction Versus Abiotic Response as Mediators of Biodiver- sity in the Middle Devonian (Givetian) Upper Hamilton Group of New York State (MS Supplementary data to this article can be found online at http://dx. Thesis) University of Cincinnati. doi.org/10.1016/j.palaeo.2016.06.017. Bittner, A., 1895. Lamellibranchiaten der alpinen Trias. 1. Theil: Revision der Lamellibranchiaten von St. Cassian. Abh. K.-Kg. Geol. Reichsanstalt 18, 1–235. Bizzarini, F., Braga, G., 1987. 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