Echinoderm Holdfasts from the Carnic Alps / Ferretti, Annalisa; Ausich, William I.; Corradini, Carlo; Corriga, Maria Giovanna; Schönlaub, Hans Peter

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Stars in the Silurian sky: Echinoderm holdfasts from the Carnic Alps

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Original Stars in the Silurian sky: Echinoderm holdfasts from the Carnic Alps / Ferretti, Annalisa; Ausich, William I.; Corradini, Carlo; Corriga, Maria Giovanna; Schönlaub, Hans Peter. - In: GEOLOGICA ACTA. - ISSN 1695-6133. - STAMPA. - 14 (4)(2016), pp. 335-347.

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Published DOI:10.1344/GeologicaActa2016.14.4.1

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18 September 2017 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 , pgs-pgs t g p p i - r c s s g u n p a m y, º n n . a e u s s s i e . M e m u , l ? o v º r. a N e y a , t ? c A ? a . c l i g o o V l o , e G a / t 4 4 c 3 A 1 . 0 a 1 c : i I g O D o l o e G KEYWORDS non-echinoderm calcareousmaterial. other affected also has that substitution a documenting phase, iron-rich an in preserved are holdfasts distinctive The specimens. all on present not but common is depression central small ‘star-like’ holdfasts Aoutline. of sort the a create general, In to so disposition, adaptation. sub-radial a in environmental arranged processes several of with profile dome-like response massive and globous a a have as morphologies of range wide (Austria) a Alps Carnic contains the of Formation Cardiola Ludlow the from holdfasts echinoderm of collection small A INTRODUCTION with conchs commonly oriented parallel to bedding surfaces. parts of the Silurian may be missing. ofthearea, many successions areincomplete andsignificant owing to the complex sedimentological and tectonicexposedtheassignmentsections.ageof most Unluckily, of evolution biostratigraphical markers for the period, co-occurrencehasthefor graptolitesenabledof conodonts, and primarya precise blackgraptolitic shales.presenceThesuitable of conditions of black bituminous limestones, rich in cephalopod shellstheirforwell-preserved and fossilspeculiartheand associations  A.Ferretti, W.I. Ausich,C.Corradini,M.G.Corriga,H.P. Schönlaub,2016CCBY-SA Stars intheSiluriansky:EchinodermholdfastsfromCarnic Cephalopods are the most abundant elements in the fauna The Silurian deposits of the peri-Gondwana area are famous Pelmatozoans. Holdfast. Functional morphology. Palaeoecology. Iron. Silurian. Cardiola Formation. 1 Dipartimento diScienzeChimicheeGeologiche,UniversitàdegliStudiModenaReggioEmilia Dr. IgnazSeipel-Platz2,1010Vienna,Austria.SchönlaubE-mail:[email protected] A. FERRETTI 125 S.OvalMall,Columbus,OH43210,USA.AusichE-mail:[email protected] 2 via Trentino 51,I-09127Cagliari,Italy. CorradiniE-mail:[email protected] via Campi103,I-41125Modena,Italy. FerrettiE-mail:[email protected] Division ofEarthHistory, SchoolofEarthSciences,TheOhioStateUniversity 3 Dipartimento diScienzeChimicheeGeologiche,UniversitàCagliari 1 4 W.I. AUSICH Austrian AcademyofSciences,CommissionforGeosciences Corriga E-mail:[email protected] 2 C.CORRADINI Alps, Austria S B A T C A R T 3 content,including recently documented laminated structures palaeontologicalrichliteratureitsthe foralready inknown is locality Austria. The Rauchkofel in Mount the of south Siluriancalcareous occurrences Carnicthein Alps, exposed Iapetus Ocean during the Silurian, our attention has focused on facies (sensu Brett and at the Silurian/Devonian boundary. Ordovician, Silurian middle Late the rock-forming, in like remainsmay more commonly occur insuch masses asto be skeletal documented. Their poorly and rare extremely are Echinodermremainsabundant,are complete but specimens well. as present chitinozoans are and acritarchstrilobites, gastropods, brachiopods, corals, with associated Bivalves, M.G.CORRIGA n n tep t dcmn ad orlt time-specific correlate and document to attempt an In 3 H.P. Schönlaub et al. , 2012; Ferretti 4 et al. , 2012a) across the 1 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 i t t e r r e F . A GEOLOGICAL SETTING (Ferretti, 2005; Ferretti activity microbial Silurian of evidences interpreted as are discontinuity stromatolitic that along surfaces features and cephalopods echinoderms) some(mostly trilobites and and ferruginous the coatings skeletal around fragments as such to Corradini refer Carnic Alps, the a of geology the of description recent For 2015). Suttner, and (Corradini Alps Carnic the of Pre-Variscansequence the in discriminated formally now are formations 36 and recently, revised significantly was sequence this of lithostratigraphy The Variscansequence. Pre- so-called the constitute 2007) Forke, and Schönlaub during the late Bashkirian and Moscovian (Venturini, 1990; Variscanthe by affectedorogeny were that Pennsylvanian clear differences in litho- and biofacies along theDevonian thebeltCarnicin (Schönlaub, 1992).reflectedAlps.Thisis in the Late Ordovician to 35ºS in the Silurian and to the tropical northwardfaster than the main continent, from about 50ºS in the northern Gondwana margin in the Ordovician and moved terranes; von Raumer and Stampfli, 2008) that detached from Carnicthe terrainsgrouptheAlpswereof partof (Galatian Ordovician to the Late Permian. Middle During the from the early age Palaeozoic, in ranging world, the successions in border(Fig. 1),expose oneof the most complete Palaeozoic n nnw tikes f eietr rcs s locally is rocks sedimentary of thickness unknown the an to strata, due Silurian and and, Ordovician Llandovery, separating the disconformity of base overall the at The started (‘lydites’). cherts black 60m. exceed not does thickness transgression Silurian The and basinal or shales deep-shelf graptolitic to limestones, and shales interbedded limestones, bioclastic nautiloid-bearing water to shallow limestones from range and Chain Carnic illustrate and assess the significance of this fauna. Brett,2015). Thepurpose of the present paper is to describe, MacClintock,andZamora 2005; ( sediment-water interface the atconditions pelmatozoan palaeoecology on echinoderms the of and on calyx,holdfastmorphology provides importantinformation commonlyare preserved dissociated fromtheircolumnand al. othersFranzén,among(seeDonovanBrett,1981;1977; own their on studied been relatively rarely holdfasts have Pelmatozoan residue. acid the in recovered was holdfasts echinoderm of association small A acid. etched formic dilute and in collected were limestone of samples 1980), biostratigraphicalresolution based on conodonts (Schönlaub, .vol.num.nº manuscript p i r c s u n a m º n . m u n . ) l r o a v e r. y a ( e y s a g t p c - A s a g c p i g , o ) l m o u e n G ( / l 4 o 4 v 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G , 2007; Plotnick ok fo te ide roiin o h lower the to Ordovician Middle the from Rocks Italian-Austrian the across located Alps, Carnic The Silurian deposits are irregularly distributed within the within distributed irregularly are deposits Silurian . l a t e et al. (2015c,2016). et al. et al. , 2016). Despite the fact that holdfasts , 2012b). In order to improve the et al. et , 2010; Thomka2010;and , e.g. Seilacher , et Brett 1999; Histon, and (Schönlaub age Ludlow to Llandovery of zones conodont several to corresponds which missing, comprised of alternating black graptolitic shales, shales, graptolitic black Formation, Nölbling alternating beds. limestone and marlstones the of by comprised represented are shaley and facies calcareous between Intermediate conditions basin. the sedimentary of part deeper the and in deposited cherts were Formation interbedded Bischofalm the of shales with alum clay-rich shales graptolitic black Brett 2005; Ferretti, chitinozoans (Corradini and well as present are brachiopods ostracods, and, rarely, gastropods more crinoids, common; are wackestones-packstones. conodonts and bioclastic bivalves Trilobites, abundant. are cephalopods Nautiloid by the to represented mostly correspond Ludfordian-basal ‘ units (upper These (Ludfordian), Formation Lochkovian). Formation Alticola the Cardiola and the Ludfordian), (Telychian-lower Formation Kok the basin: the of parts FIGURE 1. Lake W Orthoceras 0 M. olayer (Hohe Three calcareous units are present in the proximal proximal the in present are units calcareous Three Coglians I 2780 500 et al. W 5k arte) m

Location mapoftheRauchkofelBodenSection. m A Pso , 2009;Corradini (Plockenpass) Carnico Timau limestones’ of earlier authors and are are and authors earlier of limestones’ di Silurian equinodermalholdfastsfromtheCarnicAlps Paluzza M. T I Croce Cuestalta (Hoher L A et al. et Koderhohe Paularo 2228 T rieb) 2198 Y , 2009; Histon, 2012a, b). The The b). 2012a, Histon, 2009; , et al. et al. et M. 2143 Pontebba * , 2015a). U A Zermula Passo , 2003, 2010, 2015a; 2015a; 2010, 2003, , T S di (Nassfeld Pramollo I R Hermagor ) A Ugovizza M. Osterni Cocco 1941 2228 2 g 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 i t t e r r e F . A .volume.issue.nº manuscript p i r c s u n a m º n . e u s s i . e m u l o ) v y r. n a a e ( y a s t g c p A - a s c g i p g o , l ) o º e n G ( / l 4 o 4 v 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G Section. The asterisk indicates the level from which the studied material was collected. Gor.: Gorstian; H.: Homerian; S.: Sheinwoodian; W.: Wenlock. 2. FIGURE ORDOVICIAN S I L U R I A N D E V O N I A N U p p e r W. L u d l o w P r i d o l i L o w e r Chronostratigraphy, lithostratigraphy, stratigraphical log (modified after Schönlaub, 1980) and selected views of the Rauchkofel Boden Rauchkofel the of views selected and 1980) Schönlaub, after (modified log Chronostratigraphy,lithostratigraphy,stratigraphical Katian/Hirnantian S.H. Gor. Ludfordian Lochkovian Pragian . l a t e

Wolayer Fm. Kok Fm. Cardiola Fm. Alticola Fm. Rauchkofel Fm. La Valute Fm. Findenig Fm.

2 m * Alticola Fm. Findenig Fm. Cardiola

Fm. La Valute Fm.

Rauchkofel Fm. Wolayer Fm. Silurian equinodermalholdfastsfromtheCarnicAlps

Kok Fm. Alticola Fm. Kok Fm. 3 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 i t t e r r e F . A .vol.num.nº manuscript p i r c s u n a m º n . m u n . ) l r o a v e r. y a ( e y s a g t p c - A s a g c p i g , o ) l m o u e n G ( / l 4 o 4 v 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G The RauchkofelBodenSection FIGURE 3. Valute formations. formations. Valute La the and Wolayer the both of Section stratotype the Boden represents Rauchkofel The Formation (uppermost Valute Lochkovian-Eifelian). Formation Findenig La the and Formation the (Lochkovian) (Lochkovian), Alticola Rauchkofel the Formation the Lochkovian), Ludfordian-basal (upper (Ludfordian), Cardiola Formation the Ludfordian), Kok (Telychian-lower the Formation Hirnantian), ?basal to Katian (Late Formation Wolayer the Section: Boden Rauchkofel the in exposed authors the of paper. this of some by (2001) revision under Schönlaub by currently and is and provided Ferretti and was (1980) frame Schönlaub biostratigraphical Corradini to conodont refer and chronostratigraphical and description litho- up-to-date dacryoconarids an for and corals, Ferretti see summary a For Problematica. conodonts, bivalves, trilobites, cephalopods, nautiloid on section, concentrate the papers of geochemistry, and descriptions stratigraphy sequence detailed sedimentology, numerous a to and out. addition Alps, carried In been have Carnic works monographic whole and studies the of sections known best the of the one is Section in Boden Rauchkofel The present is Llandovery. whole gap the almost embracing Silurian, significant lower a are but facies here, Wolayer in Upper exposed Devonian from Lower at rocks to of Ordovician 65m Rauchkofel, Approximately altitude Mt. an 2175m. and of of 12º52’30” E slope 46º36’54”, N coordinates southwestern the on cephalopod association.Scalebar corresponds to5cm. Seven formations, formally introduced recently, are are recently, introduced formally formations, Seven located is 2) (Fig. Section Boden Rauchkofel The

Slab of Cardiola Fm. showing the peculiar cardiolid bivalve- . l a t e t al. et (2015b). The The (2015b). et al. et (1999), (1999), The CardiolaFormation Material andmethods STARS INTHESILURIANSKY true brachiopod and trilobite dominated communities communities dominated present. are trilobite no Oxygen and as 1999). limited brachiopod true apparently 1998, however, 1979, was, (Kříž, concentration substrate cephalopod firm to a representing and attach bottom the to on deposited order the conchs in conic at triangle a distributed of strategy, points vertix feeding prominent and three morphology developing their bivalves adapted Cardiolid empty had region. with the over along shells bivalve, larvae cephalopod fauna temporarily distributing other and and was gastropods base sea-bottom the wave ventilating normal below present current surface a (1995), Kříž and Ferretti to According speculations. different attracted have smell epibyssate bituminous of peculiar the and matter organic mostly abundant the and forms, comprised the Group, of Community communities presence the with bivalve-dominated coupled of fauna shelly benthic real of a absence The debate. strong of peri-Gondwana matter a been the has area along limestone deposited cephalopod were black the occurrences general, in and, Fm. 4m. to up reach may unit the Alps, Carnic the of parts other In 30-40cm. to limited is thickness its and the section Boden in the World exposed Fm. is poorly Cardiola War Rauchkofel I trench, the In beds calcisiltites. calcareous and fine-grained hummocky-laminated, to to planar thin as gray intercalations dark limestone with by shales black constituted 3), (Fig. (Fm.) Formation specimens were coated with gold and were observed observed were and gold few with A coated 990. were Coolpix specimens Nikon M8 a Heerbrugg microscope using Wild a Optical stereomicroscope under 4). taken were (Fig. photographs microscopy optical in were revealed was specimens colour reddish the Apeculiar observed. of complete. rarely and cleavage and preserved, in well Microfracturing were common generally were methods residue, holdfasts the net concentration echinoderm 100μm The further a applied. No with sieved dried. acid, and formic diluted with processed been had which limestone a from specimens binocular microscope 11 SV The Stemi Zeiss a Alps. under picked Carnic hand were Austrian the in Boden Silurian Section Rauchkofel the the of Fm. from Cardiola residue Ludlow) (early conodont a from holdfasts The depositional environment in which the Cardiola Cardiola the which in environment depositional The Cardiola the from collected was material studied The We have obtained a small fauna of echinoderm echinoderm of fauna small a obtained have We Silurian equinodermalholdfastsfromtheCarnicAlps Cardiola 4 - 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 Mineralogical composition Biostratigraphical assignment Other faunalelements 28992-29022. IPUM numbers repository under Emilia, Reggio e Modena di Studi degli e Università Chimiche Geologiche, Scienze di Dipartimento the of Palaeontological the Collections in reposited is based is which paper and on this material Modena The Italy). of (Modena, Emilia University Reggio the of (CIGS) Strumenti’ Grandi at were the performed Interdipartimentale ‘Centro detector. X’Celerator an with equipped diffractometer PRO randomly X’Pert Philips on a using samples conditions powdered room oriented at samples selected on Oxford collected was (XRPD) an diffraction powder spectrometer X-ray system. with dispersive energy X-ray equipped 300 INCA EDX 200, ESEM-Quanta Microscope FEI Electron Scanning Environmental an with picked in the residue has a peculiar rusty appearance, appearance, rusty peculiar a has residue the in picked Slavik, 2010; Corriga, and Corradini (Corradini processes branched with elements unicostatus excavata ichnusae variabilis 5A), (Fig. 1933 Mehl, and Branson the topofCardiolaFm.,havenotbeenfound. above immediately level the in documented 2013) al., et (Ferretti rings phosphatic unassigned and 2008) Serpagli, enigmatic the Eurytholia or and Muellerisphaerida sorted well appears equidimensional. material all the general in and complete, rarely are Trilobites holdfasts. echinoderm for the detected phase iron-rich same the in preserved been have elements faunal Surprisingly,the 5). all (Fig. well as echinoderm gastropods, ossicles and brachiopods, nautiloid ostracodes, embryonic chambers were recovered bivalves, spiculae, sponge cardiolid trilobites, Abundant elements. faunal other included also that residue conodont 2009, 2014; Slavík Slavík 2014; Corriga 2009, 1999; Corradini, and (Serpagli Ludlow) (Ludfordian, Zone the of part lower the in extinct became the of of LADs the before part lower the to level the of et al. i t t e r r e F . A .volume.issue.nº manuscript p i r c s u n a m º n . e u s s i . e m u l o ) v y r. n a a e ( y a s t g c p A - a s c g i p g o , l ) o º e n G ( / l 4 o 4 v 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G Optical microscopy, XRPD, Optical microscopy, and analyses ESEM-EDX As previously remarked, the faunal association association faunal the remarked, previously As includes fauna The conodont a from recovered been have holdfasts echinoderm The , 2014). This association allows a precise assignment assignment a allows precise , association This 2014). (Branson and Mehl, 1933), 1933), Mehl, and (Branson Serpagli and Corradini, 1998, 1998, Corradini, and Serpagli Walliser, 1957 (Fig. 5B), 5B), (Fig. 1957 Walliser, . l a t e lts (Ferretti plates (Branson and Mehl, 1933) and anomalous anomalous and 1933) Mehl, and (Branson et al. et K. K. v. variabilis , 2010; Corradini Corradini 2010; , 20; erti and Ferretti 2006; al., et siluricus Polygnathoides and and Kockelella variabilis variabilis Kockelella Kockelella variabilis variabilis Kockelella et al. et . siluricus P. K. K. v. ichnusae , 2010; Corriga Corriga , 2010; et al. et Panderodus Panderodus et al. et Wurmiella Wurmiella , 2015a). , , 1996; , 1996; Zone, Zone, t al. et that that , (Fe oxyhydr-/oxides goethite (α-Fe by goethite oxyhydr-/oxides Analyses iron the of consists material composition. red the that revealed XRPD mineralogical and aspect usual their keep conodonts as such elements Phosphatic preservation. rich’ ‘iron- this have organisms originally residue calcareous the only the Apparently, magnetic from magnet. a of picked have use the easily by be elements can the and properties brown Furthermore, deep a 4). to red dark (Fig. a from ranging colour with either during later diagenesis or during the formic acid acid formic processing. sample the during or away diagenesis later dissolved during was either it and J-K) 7C-D, (Fig. not is preserved specimens the of centre The phases. iron-rich the by replaced was specimen original the of layer outler the that appears it preserved, are 7H) (Fig. facet column the stereomic of the details the of some and 7C) Because 6D; (Figs. microstructure material. ferruginous of layer Description columnals makes any taxonomic identification beyond identification pelmatozoan echinodermspeculative. taxonomic any makes columnals and holdfasts than other remains pelmatozoan of material our in absence The are echinoderm. pelmatozoan holfasts some from these that indicates columnals holdfasts isolated these and of presence The 7H. pentagonal Figure the on and opening 7F Figure in opening facet column pentalobate the 6B, Figure on opening facet column the of outline subpentagonal coarse a include These specimens. few a in preserved aspects symmetry pentameral the do as echinoderm, an from are holdfasts these that demonstrates Section, Austria. Note the reddish colour of the material. Scale bar Scale material. the of colour reddish the Note Austria. Section, Boden Rauchkofel Fm., Cardiola the from 28992a-i IPUM holdfasts IUE 4. FIGURE corresponds to1mm. 3+ The pelmatozoan holdfasts are preserved as a thin thin a as preserved are holdfasts pelmatozoan The The presence of stereomic microstructure (Figs. 6D, 7C) 2 Fe 2+

O pia seemcocp iae o echinodermal of images stereomicroscope Optical 4 ). ). Silurian equinodermalholdfastsfromtheCarnicAlps 3+ O(OH)) and magnetite magnetite and O(OH)) 5 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 i t t e r r e F . A 29000. I-K)Echinodermossicles,specimensIPUM29001-29003.Scale barcorrespondsto500μm. specimen IPUM 28998. G) Ostracode with overgrowth of pyrite crystals, specimen IPUM 28999. H) Articulated cardiolidid bivalve, specimen IPUM views of trilobite fragments, specimen IPUM 28995 and IPUM 28996. E) nautiloid embryonic chamber, specimen IPUM 28997. F) Sponge spicula, B) 28993. IPUM element P2 of view upper 1933, FIGURE 5. .vol.num.nº manuscript p i r c s u n a m º n . m u n . ) l r o a v e r. y a ( e y s a g t p c - A s a g c p i g , o ) l m o u e n G ( / l 4 o 4 v 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G (here regarded as the upper part). Therefore, in lateral lateral in Therefore, part). upper the as regarded (here dome opposite bulbous a and one) lower the as regarded (here surface flat a with body globous a have apparently specimens the feature, general a As 425μm. to 290 from size in ranges holdfasts the on diameter facet column the from 1 dimension to 4mm, with a mean of value 2mm, and Echinoderm holdfasts of our material range in in range material our of holdfasts Echinoderm Scanning electron micrographs of main faunal elements from the Cardiola Fm. of Austria. A) . l a t e B H E A Walliser, 1957, upper view of P1 element IPUM 28994. C-D) oral C-D) 28994. IPUM element P1 of view Walliser,upper variabilis 1957, variabilis Kockelella penetration into a soft sediment were recognized. were sediment soft a into penetration indicating possibly branches oriented vertically No 6H). (Fig. profile discoidal flat distinctive a have others Some 8B, F). (Fig. directions in opposite and growing departing basal curved more branches with profile, curved lateral a a thinner and surface have systems holdfast Some 6J). (Fig. shapes conical truncate nearly are specimens view, I D F J Silurian equinodermalholdfastsfromtheCarnicAlps Polygnathoides siluricus Branson and Mehl, G C K 6 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 i t t e r r e F . A .volume.issue.nº manuscript p i r c s u n a m º n . e u s s i . e m u l o ) v y r. n a a e ( y a s t g c p A - a s c g i p g o , l ) o º e n G ( / l 4 o 4 v 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G of specimenIPUM29005.Scalebars correspondto500μminallframesexceptC-Dwherebarsrepresent 100μm. views lateral and Oral I-J) 29007. IPUM specimen of views lateral and oral Sub-lateral, F-H) 29006. IPUM specimen of view Oral E) echinoderms. and oral views of specimen IPUM 29004. C-D) Details of specimen IPUM 29004 and specimen IPUM 29005 showing the typical stereom pattern of 6. FIGURE Scanning electron micrographs of Silurian pelmatozoan holdfasts. All specimens are from the Cardiola Fm. of Austria. A-B) Sub-lateral A-B) Austria. of Fm. Cardiola the from are specimens All holdfasts. pelmatozoan Silurian of micrographs electron Scanning . l a t e E A G B H F C D Silurian equinodermalholdfastsfromtheCarnicAlps J I 7 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 i t t e r r e F . A .vol.num.nº manuscript p i r c s u n a m º n . m u n . ) l r o a v e r. y a ( e y s a g t p c - A s a g c p i g , o ) l m o u e n G ( / l 4 o 4 v 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G Aboral view of specimen IPUM 29015, showing again the hollow central part. Scale bars correspond to 500μm in all frames except C where bar where C except frames represents 100μm. all in 500μm to correspond bars Scale part. central hollow the again showing 29015, IPUM specimen of view Aboral empty.is body main the that reveals part K) apical the in holdfast the of breakage The 29014. IPUM specimen of view Sub-lateral J) 29013. IPUM E-F) Lateral and oral views of IPUM 29010. G) ?Oral view of specimen IPUM 29011. H) Oral view of specimen IPUM 29012. I) Oral view of specimen 29009. IPUM specimen in process hollow a showing D of Detail C) 29009. IPUM specimen of views lateral and Oral D) B, 29008. IPUM specimen 7. FIGURE Scanning electron micrographs of Silurian pelmatozoan holdfasts. All specimens are from the Cardiola Fm. of Austria. A) Oral view of view Oral A) Austria. of Fm. Cardiola the from are specimens All holdfasts. pelmatozoan Silurian of micrographs electron Scanning . l a t e G D E A J B H C F K Silurian equinodermalholdfastsfromtheCarnicAlps I 8 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 DISCUSSION including hardgrounds (Brett stem holdfasts. holdfastsunder investigation thewould beincluded 348),interminal p. 1, tab. (1981, environmental terminology specific his In conditions. to response a as substrate the to Brett(1981,1984)identified fixationthree major modes of be very helpful for understanding the paleoecological setting. depression( median the lackelements other Some 7H). Fig. inpentagonal and 7F Fig. (pentalobatein detected be may designgeometric aspecimens, few a On has a general circular outline with conical borders (Fig. 6A). lumencentral The part.basal thereaching depression,not 7C). (Fig. processes the inside and 7J-K) (Fig. body basal the in both hollow possibly to 7B) (Fig. are to internally adhesion Holdfasts the increase substrate. enlargement terminal bulbous in a expanding or 7F) (Fig. end proximal its at bifurcated and section in circular are Processes 8K). 7F, 6G, (Figs. partly processes is between at holes merging circular delineating or sometimes length, mid 8E) (Fig. and one other following the the overlapping from branch separate Each 6B). is (Fig. arrangement net-like a dendritic creating or elongate and slender rarely more massive, and short growth commonly are Processes 6F-G). as (Fig. processes infilled secondarily be may which 8C), symmetry (Fig. quadrilobate a developed and processes four have specimens of couple A 8I-J). (Fig. direction that in growth prevented has that the holdfast nearby of a of response presence a as possibly side, unbranched straight a have specimens few a keeping Just signal. but radial directions, subordinate a all in arrayed be to appear that processes with material, our in detected be may patterns Eucalyptocrinites mid-Palaeozoic the of holdfast in pattern distribution branch geometric in preferred distinctive a described has (Plotnick paper recent A number. in 12 the to from up be may departing Branches star. of sort a delineating and irregularly centre branches several with original architecture of the holdfast. ossicles was detected in any specimen, suggesting a massive noevidence of segmentation and/or articulation of different that has occurred and possibly obliterated primary structure, substitutionmineral the the of of spite Finally,in insteadlumen.central 8C-D) I-J, 6D, (Figs. area a raised of median evidence clear is there specimens, of number a On i t t e r r e F . A .volume.issue.nº manuscript p i r c s u n a m º n . e u s s i . e m u l o ) v y r. n a a e ( y a s t g c p A - a s c g i p g o , l ) o º e n G ( / l 4 o 4 v 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G emtza hlfss r kon rm settings from known are holdfasts Pelmatozoan Attachment structures in pelmatozoan echinoderms may central a have holdfasts echinoderm the of Most outline subcircular a has holdfast the above, From . l a t e . However, no precise symmetric symmetric precise no However, . et al., 1983), skeletal substrata e.g. t al. et , Fig. 7G). Fig. , , 2016) 2016) , (e.g., Hollisand Ausich, 2009). crinoids fossil many for and isocrinoids living among true autotomized at least the juvenile, attached holdfast. This is stalked holdfast cemented a than other holdfast a had that crinoids Adult animal. free-living a yield to autotomizes pentacrinoid stage. In living comatulid crinoids, the crown grow, the post-larval crinoid is considered to be part of the to begin arms initial the When stage. cystidean the as to referred echinoderm. is and stemmed arms lacks phase attached, post-larval earliest The small, a as benthic a existence begins it where surface hard a substratum to attaches the and to settles eventually which larva, free- a living is crinoid made. a of be phase now developmental earliest can The juveniles fossil and living between (Amemiya documented crinoid, from theCardiolaFormation. reported pelmatozoans juvenile undoubtedly small, very the with size in sharply contrast and pelmatozoans adult However, from mostly are 2016). above mentioned studies the Wilson,in holdfasts and Ausich 2015; Brett, and Sumrall 2009; Ausich, and Hollis and have been attributed to many groups (e.g., Brett, 1978; localities many from reported been have Fm. Cardiola the encrusting Dendritic, 1978). holdfasts with a similar morphology to the specimens from (Brett, America North of America North and 2005), Brower, (e.g., from crinoids Ordovician 2010), al., et (Zamora Spain northern of Cambrian are the from eocrinoids examples Three preserved. are specimens complete cases rare relatively in however, known; not are holdfasts isolated most of affinities The references). many (among 2010), and soft substrata (Seilacher (Zamora and MacClintock, firmgrounds 2005) 2016), Wilson, and (Ausich Kammer (see Slate Hunsrück Devonian the from reported crinoids juvenile larger and smallest the between individuals from probably were pelmatozoans These forms. juvenile somewhat older or stage pentacrinoid late to middle the from be to interpreted are holdfasts Cardiola the pelmatozoans, can be drawn between parallels that assuming Thus, 425μm. to 290 from ranges Fm. holdfasts have a facet for the distal-most columnal that (Amemiya stage pentacrinoid early the by 215μm approximately was diameter column this whereas 130μm, approximately of holdfast the above immediately diameter column had stage cystidean late to life. Because organisms in this fauna with original original with fauna this in during organisms attached were Because they life. which to surface the from In the living crinoid living the In Recently, the complete development of a living stalked All of the holdfast specimens are preserved isolated isolated preserved are specimens holdfast the of All apne, 85 hs been has 1885, Carpenter, rotundus Metacrinus et al. Silurian equinodermalholdfastsfromtheCarnicAlps , 2015,fig.2versus1B). Metacrinus rotundus and Palaeozoic , the middle the rotundus, Metacrinus 21) s comparisons so 2014), al., et Caryocrinites , 2014). The Cardiola The 2014). al., et , 2015; Thompka 2015; al., et from the Silurian the from t al., et 9 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 i t t e r r e F . A .vol.num.nº manuscript p i r c s u n a m º n . m u n . ) l r o a v e r. y a ( e y s a g t p c - A s a g c p i g , o ) l m o u e n G ( / l 4 o 4 v 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G to 500μminallframesexceptDwhere barrepresents100μm. J) Oral view of specimen IPUM 29021. Note the pyrite crystal overgrowth on the left. K) Oral view of specimen IPUM 29022. Scale bars correspond E-F) Oral ridge. and lateral views apical of specimen IPUM 29018. central G-H) Lateral and the oral views of in specimen IPUM 29019. pattern I) Oral view of stereom specimen IPUM 29020. the illustrating C of detail D) 29017. IPUM specimen of view Oral C) 29016. IPUM specimen of views FIGURE 8. Scanning electron micrographs of Silurian pelmatozoan holdfasts. All specimens are from the Cardiola Fm. of Austria. A-B) Oral and lateral . l a t e G A B F H J C E D Silurian equinodermalholdfastsfromtheCarnicAlps K I 10 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 Middle Palaeozoic Revolution - Bridging the Gap between the the between Gap the Bridging - to Revolution Early Palaeozoic ‘The Middle 591 Project IGCP the to Relations Environments’ and AF) Anoxic (Responsible in ‘ROSAE: Sediments and Project Organism between UNIMORE-FAR2014 the Malferrari contribution fundamental a to is for paper Daniele This Tonelli investigations. SEM draft, Massimo in expertise and the analysis of X-Ray for preparation the remarks during critical for Papazzoni Cesare thank also Zamora. authors been The Samuel and have Donovan Stephen by not manuscript comments the useful on would acknowledge We paper why. knows He this possible. support whose without CONCLUSIONS identification isnotpossible. more than one of clade of pelmatozoan echinoderm, a adults more specific in morphologies with overlaps juveniles these of morphology the Because pelmatozoan. juvenile a object withrelief. an of side the to attached was holdfast this that suggesting illustrated in Figure 6I was at a high angle to the substratum, the column was attached. However, when the column of the holdfast substratum the to vertical near or surface) vertical lower was holdfast the of orientation the on (based column the of posture the examples, most In substrate. the floating to algae. or shell cephalopod organisms swimming the a to into pseudoplanktonic attached as introduced were assemblages they Cardiola that possible also However, is facies. it adjacent in absent are holdfasts these because these setting Fm. that Cardiola the likely in is lived It pelmatozoans algae. as such soft- a to organism, or bodied shell holdfasts aragonitic an these either to alive, attached when were that probable is it oxides, oxyhydr-/ iron in preserved are material skeletal calcite ACKNOWLEDGMENTS substrate. bottom firm planar a to adhesion suggest ( al. holdfasts et juvenile for expected relatively be and may shorter which occurrences, Silurian are other to compared if material thicker our on branching. of Processes pattern simple Carnic general a with Austrian appearance the discoidal massive a in design holdfasts Fm. Documented Alps. Cardiola Silurian recovered the been has in structures attachment echinoderm i t t e r r e F . A .volume.issue.nº manuscript p i r c s u n a m º n . e u s s i . e m u l o ) v y r. n a a e ( y a s t g c p A - a s c g i p g o , l ) o º e n G ( / l 4 o 4 v 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G The Cardiola Fm. holdfasts are interpreted to be from be to interpreted are holdfasts Fm. Cardiola The above elevated moderately was stem the of base The Our special thanks to Juan Carlos Gutiérrez-Marco Gutiérrez-Marco Carlos Juan to thanks special Our pelmatozoan ferruginous of number large A , 2016). Their dominant flat lower surface would would surface lower flat dominant Their 2016). , . l a t e e.g. , Plotnick Plotnick , Corradini, C., Olivieri, R., Serpagli, E., 1996. Possible relationship Corradini, C., Suttner, T.J. (eds.), 2015. The Pre-Variscan sequence lowermost and Silurian 2010. M.G., Corriga, C., Corradini, of Species New Three On 1885. 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55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 hma JR, rt, .. 21. aeeooy f pelmatozoans of Paleoecology 2015. C.E., Brett, J.R., Thomka, Zamora, S.L., Sheffield, Colmenar,J., B., Deline, C.D., Surmall, mid- the of Reflection 2010. P., Carls, J., Kříž, L., Slavík, of evolution and Taxonomy 1999. C., Corradini, E., Serpagli, of taxa New 1998. C., Corradini, E., Serpagli, i t t e r r e F . A .volume.issue.nº manuscript p i r c s u n a m º n . e u s s i . e m u l o ) v y r. n a a e ( y a s t g c p A - a s c g i p g o , l ) o º e n G ( / l 4 o 4 v 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G ide iuin ase omto, otesen Indiana. southeastern Palaeogeography, the Formation, Palaeoclimatology, Palaeoecology, in Massie 420, 1-2. surface Silurian hardground middle a from structures attachment de España,CuadernosdelMuseoGeominero,19,175-180. Minero y Geológico Instituto Paleobiology. Echinoderm in from Sardinia, Italy. In: Zamora, S., Rábino, I. (eds.). Progress S., 2015. New data on Late Ordovicican (Katian) echinoderms Bohemia. of faunas conodont Bulletin ofGeosciences,85,395-414. in Event Lau Ludfordian della SocietàPaleontologicaItaliana,38,275-298. Kockelella of (Silurian) Wenlock-Ludlow Sardinia. GiornalediGeologia,60,79-83. Late from (Conodonta) . l a t e (Conodonta) from Silurian of Sardinia. Bollettino Sardinia. of Silurian from (Conodonta) Kockelella

aoa S, lue, . Ávr, .. Sih AB, 2010. A.B., Smith, J.J., Álvaro, S., Clausen, S., Zamora, Gotlandium oberen dem aus Conodonten 1957. Walliser,O.H., Rheic the of birth The 2008. G.M., Stampfli, J.F.,Raumer, von Venturini, C., 1990. Geologia delle Alpi Carniche centro-orientali. published OnlineOctober2016. revision acceptedOctober2016; Manuscript receivedSeptember2016; Cambrian. Palaios,25,764-768. substrates in high energy environments ground since the basal Middle firm carbonat colonized echinoderms Pelmatozoan 85, 28-52. des Notizblatt Alpen. zu Wiesbaden,Bodenforschung für Landesamtes Hessischen Karnischen der und Deutschlands tectonic platescenarios. Tectonophysics, 461,9-20. Ocean – Early Palaeozoic subsidence patterns and subsequent 36, 222pp. Naturale, Storia di Friulano Museo del Pubblicazioni Udine, Silurian equinodermalholdfastsfromtheCarnicAlps 13 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 10 11 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1