IntJEarthSci(GeolRundsch)(2017)106:157–170
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DOI10.1007/s00531-016-1308-1 ORIGINALPAPER
IchnologicalrecordoftheFrasnian–
Famennianboundaryinterval:twoexamplesfromtheHolyCrossMts (CentralPoland)
MichałS t a c h a c z1·AlfredU c h m a n1·F r a n c i s c o J.R o d r í g u e z - Tovar2
Received:2October2015/Accepted:10February2016/Publishedonline:2March2016
©TheAuthor(s)2016.ThisarticleispublishedwithopenaccessatSpringerlink.com
AbstractT h e F r a s n i a n –
Famennian( L a t e D evonian)boundaryi n t e r valw i t h i n t h e c a r b o n a t e –
s i l i c i c l a s t i c s e r i e s intheKowalaandPłuckisection s(HolyCrossMts,Cen-
tralP o l a n d ) h a s b e e n a n a l y s e d t o evaluatet h e i n f l u e n c e oft h e Kellwasserevento n t h e m a c r o b e n t h i c t r a c e m a kercommunity.T h e U p p e r Kellwassereventh a s a l i t h o l o g i -
callyvariablerecord,ashorizonsofflints(Kowala)andasab edofbituminous,black,cephalopodlimestone(Płucki).Bot hs e c t i o n s s h owm o s t l y l a m i n a t e d , u n b i o t u r b a t e d b e d s ofm a r l s t o n e s o r s h a l e s j u s t a b ovet h e F r a s n i a n – Famenn-
ianb o u n d a r y,w h i c h p o i n t t o eventso f a n o x i a o n t h e s e a floor.H owever,t h e f i r s t a n o x i c h o r i z o n o c c u r s b e l owt h e Frasnian–
Famennianb o u n d a r y.T h e t r a c e f o s s i l s a n d b i o - turbationalstructuresareuncommonandpoorlydiversified.Tric hichnusandMultinaaretheonlyfrequenttracefossilsins omebeds.Moreover,onehorizonab ovetheFrasnian–
FamennianboundarycontainsnumerousMultinaandasin-gle?
Planolites.Suchpoorlydiversifiedtracefossilassem- blages u g g e s t s a n u n favourablee nvironmentf o r m o s t o f burrowingorganismsandfluctuationsinoxygenationfro manoxic,todysoxicconditions.Theoccurrenceofthetr acefossilsandbioturbationalstructuresasspottedandmot tledichnofabricsfromthe1.3mabovetheFrasnian–Famennian
*
MichałS t a c h a c z michal.st achacz@uj.edu.pl
AlfredUchmanalfred.uchma n@uj.edu.pl
FranciscoJ.Rodríguez- Tovarfjrtovar@ugr.es
1 InstituteofGeologicalSciences,JagiellonianUniversity,Ole andry2a,30-063Kraków,Poland
2 DepartmentofStratigraphyandPalaeontology,SciencesFac ulty,UniversityofGranada,Granada,Spain
2 IntJEarthSci(GeolRundsch)(2017)106:157–170
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boundaryisinterpretedasanimprovementinbottomwaterox ygenconditionsaftertheUpperKellwasserevent.
KeywordsTracefoss ils· Frasn ian –
Famennian·Devonian·Kellwasserevent·Trichichnus·
Multina·Poland
Introduction
Oneo f t h e l a rgerm a s s extinctionsi n E a r t h h i s t o r y h a p - penedd u r i n g t h e F r a s n i a n – Famennian( F – F ; L a t e D evo-
nian)transition.Itconcernedmostlyshall ow- waterorgan-
ismslivinginwarmerenvironments,foremostreef-
builders(RugosaandTabulatacapcorals,stromatoporoi ds)aswellammonoids,t e n t a c u l i t e s , c o n o d o n t s , p l a c o d e r m s a n d b r a -
chiopods(e .g .H al l a m a nd Wignall1 99 7;M cG he e20 1 3).TheLateDevonianextinctionhasbeenrecentlyinterprete dasa “ b i o d iversityc r i s i s ” c a u s e d b y s p e c i a t i o n r a t e c o l -
lapser a t h e r t h a n a c l a s s i c “ m a s s extinction”( e . g . R a c k i2005;B e c kere t al.2012;GerekeandSchindler20 12).Thiscrisisisrelatedmostlytosealevelchanges(e.g.
Hal-
lamandWignall1999;Stigall2012)orclimatevariation s(Joachimskietal.2009);however,themainreasonsarestillunc lear.I n E u r o p e , t h e F –
F extinctioneventi s r e c o r d e d usuallyasaseriesofblac klimestonescalledtheKellwas-
ser,whichareassociatedwithanoxia(e.g.Buggisch1991
;Joachimskie t a l .2 0 0 1;M a r y n owskie t a l .2 0 1 1) .R e l a -
tionshipsb e t w e e n a n o x i a , s e a l evelc h a n g e s a n d t h e F –
F massextinctionw e r e p r o p o s e d f r o m t h e c i t e d p a p e r s , buttheya r e n o t u n iversallya c c e p t e d ( s e e d e t a i l e d r eviewi n Bonda n d Wignall2 0 0 8);i t s e e m s t h a t t h e extinctionh a s amoremulticausalnature(Racki2005;Ger ekeandSchin-
dler2 0 1 2).I n t h e l a t e s t y e a r s , i m p o r t a n t M e s o z o i c a n d Cenozoicanoxiceventshavebeenstudied bymeansofthe
ichnologicala p p r o a c h , a s t h o s e o c c u r r i n g a t t h e Toar cianOceanicAnoxicEvent(T-OAE;Rodríguez-
TovarandUch-man2010;Rodríguez-
TovarandReolid2013;Reolidetal.2014),theCenomanian–
Turonianboundary(Uchmanetal.2008,2013a,b;Rodrí gu ez-
Tovaret al .2009a,b;Monaco eta l .2 0 1 2),o r t h e O l i g o c e n e –
M i o c e n e ( Kotlarczyka n d Uchman2012).Muchlessiskn ownaboutichnologyoftheF–
Ftransition.Buggisch(1991)onlymentionedthelami -nated,u n b i o t u r b a t e d d e p o s i t s o n t h e F –
F b o u n d a r y.B o n d eta l .
(2004)r e p o r t e d b i o t u r b a t e d s e d i m e n t s , “ m i c r o bur-
rows”a n d “ l a rger,w i s pyburrows”i n t h e Famenniano f theKowalasectionintheHolyCrossMountainsinPoland(st udiedi n t h i s p a p e r ) . Wange t a l .
(2006)p r e s e n t e d a n assemblageoftracefossilasarec ordofrecoverythroughtheFamenniani n S o u t h C h i n a . H a d d a d e t a l . (2013)a n d Boyeretal.
(2014)focusedoni chnologyandgeochem is-
tryoftheboundaryintervalintheUSA.Thelatterauthorsd escribedatracefossilassemblagedominatedbydeep- tier,pyritizedS kolithosa n d C h o n d r i t e s ,b e s i d e a s l e s s a bun-
dantT h a l a s s i n o i d e s c l o s e t o t h e U p p e r Kellwasser i n t e r-
val.S t a c h a c z a n d U c h m a n (2012)p r e s e n t e d p r e l i m i n a r y resultso f i c h n o l o g i c a l s t u d i e s o f t h e F –
F i n t e r valo f t h e HolyC r os sM ou nt a i n si nP ol an d, w h i c ha re d evelopedi n thispaper.
TheHolyCrossMountainsinCentralPolando ffertw oimportants e c t i o n s o f t h e F r a s n i a n –
Famenniant r a n s i t i o n atKowalaa n d P ł u c k i ( F i g .1), w h e r e b l a c k , a n o x i c s e d i -
ments,referredtotheUpperKellwasser,cropout.Theyarethet a rgeto f a “ b e d - b y -
b e d ” i c h n o l o g i c a l a n a l y s i s , f r o m whichtheresu ltsanddiscussionarepresentedinthispaper.
Earlierg e o c h e m i c a l s t u d i e s o f t h e Kowalas e c t i o n s h owgeneraloxygendeficiencyneartheseafloorduringthe F–
Ftransition(Joachimskietal.2001;Bondetal.2004).Morec omplexoxygenationchangesarereferredtothehightotalorg aniccarbon(TOC)contentattheF–
Fboundaryinterval,whichisinterpretedasaresultofahigheri nputoforganicmatter(Rackietal.2002).
Geologicalsetting Generalremarks
ThestudyareaislocatedinthesouthernpartoftheHol yCrossMountainscalledtheKielceR egion(Fig.1),whi chisasurfacepartoftheMałopolskaBlock(e.g.Buła200 0;CocksandTorsvik2005).LateDevoniansedimentationtook placehereonanextensiveshelfinthemarginalpartoftheLaurus siapalaeocontinent(e.g.Joachimskietal.2001;Rackieta l .2002;Maryn owskietal.2011andreferences citedt h e r e i n ) . T h e Kowalas e c t i o n i s i n a n a c t ivel a rge quarrylocatedsouthofKielce,whilethePłuckisectioni savailableinasmall ,abandonedquarryl ocat ednort h- west ofPłucki(Figs.1,2).
TheU p p e r-
MiddleD evonian( F r a s n i a n ) i n t h e K i e l c e Region o f t h e H o l y C r o s s M o u n t a i n s i s d evelopeda s a s erieso f s h a l l owwater,s t r o m a t o p o r o i d –
c o r a l r e e f l i m e - stones(DyminyReef),whichare330–
800mthick(Narkie-
wiczetal.1990).Thereefsrimmedthenorthernmarginofthec arbonateshelfoftheMałopolskaBlockandformeda
Fig.1L o c a l i t y maps.a,bGenerallocation.cSketchdrawoftheKowalaquarry.dLocationofthePłuckiexposurenearŁagów
159 IntJEarthSci(GeolRundsch)(2017)106:157–170
distinctp o s i t iver e l i e f i n t h i s p a r t o f t h e b a s i n , w h i l e t h e ŁysogóryBasinlocatedtothenorthwasmuchdeeper(
Narkiewicz1988).Generally,theLateDevonianreefsstret chedalongthesouthernmarginofLaurussia(e.g.Cop-
per2002).ThereefswerealmosttotallyfloodedattheendofF rasni an(R acki1990).Asaresul t,th eyarecappedby p elagicl i m e st on es ,m ar l s t o ne sa nd sh al es of t h e Fame nn-
ian,whichareinterpretedasturbiditefedfromerodedpartsofthe carbonateplatformlocatedbetweenthesouthernChe˛ciny- ZbrzaBasinandtheŁysogóryBasintothenorth (Joachi mskie t a l .2 0 0 1).The F – F b o u n d a r y i n t e r valu s u - allyoccursinthisareaasathickseriesoflimestonesinter- beddedbymarlstones,marlyorsiltyshales(e.g.Joachi m-
skietal.2001;Rackietal.2002;DeVleeschouweretal.
2 0 1 3a n d r e f e r e n c e s c i t e d t h e r e i n ) . T h e s o - c a l l e d L owerKellwasserandUpperKellwassercanbedis tinguishedwithins u c h l i t h o l o g y i n t h e P ł u c k i s e c t i o n , w h i l e i n t h e Kowalaq u a r r y s e c t i o n t h e s e h o r i z o n s a r e d i f f i c u l t t o r e c -
ognize(cf.Rackietal.2002).TheboundaryintervalintheKo walaquarrymainlyconsistsofgrainstonesandmicritic limestonesw h i c h c o n t a i n t woo r t h r e e l a y e r s o f n o d u l a r
cherts( J o a c h i m s k i e t a l .2 0 0 1;F i g s .5,6).G e o c h e m i c a l recordsfromthislocality,especiallyTOC,showt womax-
imat h a t c o u l d b e r e l a t e d t o t h e t woKellwassere p i s o d e s (Joachimskietal.2001).Accordingtothecitedauthors,thestrat igraphicallyl owerTOCm a x i m u m c o r r e s p o n d s w i t h theLowerKellwasserhori zon,whil et hebasalpart of theupperTOCm a x i m u m c o i n c i d e s w i t h t h e U p p e r K ellwas-
serh o r i z o n . T h e u p p e r p a r t o f t h e u p p e r TOCm a x i m u m isabout8mabovetheF–Fboundary,withintheH- 3unit(Joachimskietal.2001).InthesameKowalasection,Racki andBalin´ski(1998)andRackietal.
(2002)notedbenthicbiotaextinctiona n d b i o t i c s w i t c h f r o m m o st l y c a l c a r e o u s benthosa n d p l a n k t o n t o s i l i c i s p o n g e s a n d r a d i o l a r i a n s a t thetransitionfromth eunitH-2totheunitH-3,justbelowtheF–
Fboundar y.AccordingtoFilipiak(2002),thereare noi mportantchangesinassemblagesofmarinepalynoflorabelowan dabovetheF–
Fboundary;however,morediversepalynofaciesaroundthis boundarymaybearesultofeutrophication.TheUpperKell wasserhorizoncorrespond-ingt o t h e F –
F b o u n d a r y c r o p s o u t i n t h e P ł u c k i s e c t i o n , wherei twasexaminedforthisstudy(Fig.2).Thesection
IntJEarthSci(GeolRundsch)(2017)106:157–170
6Fig.2O u t c r o p s s t u d i e d . a U p p e r Kellwasserc e p h a l o p o d l i m e s t o n e bedi n t h e P łu ck i s e c t i o n . b ,c F r a s n i a n –
Famenniant r an s i ti o n i n t e r-val,n o r t h -
e a s t wallo f t h e Kowalaq u a r r y ; b G e n e r a l v i ewo f wall,taluslim esto nesarevisibleo n th erig ht,F– Fb ou nd aryinter valrep -
resentedbymicriticlimestoneswithhorizonsofflintsincentralpart,an dFamennianmicriticandnodularlimestonesandmarlsontheleft;cMic ritic,laminatedlimestoneswithoneoftheflinthorizonofthe F –Fboundaryinterval.Thehammeris40cmlong
iscomposedofmicriticlimestonesandmarls,inwhichtheF–
Fboundaryisrecognizedwithinamediumbedofcepha-
lopodlimestone(Rackietal.2002;SzrekandGinter2007).Adisti nctbenthicfaunacrisiswasdocumentedinthissec-
tionbyRackiandBalin´ski(1998)andOlempska(2002).
Outcropsstudied Płuckioutcrop
Ina s m a l l p i t , 1 k m N W o f Ł a g ó w ( F i g .1),a n a p p r o x i - mately5 - m - t h i c k s e c t i o n o f c a r b o n a t e – s i l i c i c l a s t i c r o c k s isexposed( F i g .3).T h e l owerp a r t o f t h e s e c t i o n , a b o u t
1.5mthick,mostlyconsistsofmarlstonesinterbeddedwithr arebedsofsiltyshale.Primaryhorizontallaminationisd o m i n a n t b o t h i n m a r l s t o n e s a n d i n s i l t y s h a l e s ; h o w-
ever,som em ar l s t o ne be ds a r es l i gh t l yb i o t u rb at ed , s h ow-
ings p o t t e d a n d l e s s c o m m o n l y m o t t l e d fabricsa n d t h e n partlydi st urbedl am i nat i ons. Thebi ot urbat i o nal st ruct ures areu s u a l l y t i ny,u p t o 3 m m . Al a rgen u m b e r o f b e d s i n thispartofthesectioncontainthetrace fossilTrichichnusisp.,whichisvisibleincrosssectiona snumerousminute ferruginouss p o t s w i t h a h a l o ( F i g . 6).T h e u p p e r p a r t o f thes e c t i o n i s c o m p o s e d m o s t l y o f l a m i n a t e d o r m a s s ive,micriticlimestonesandles scommonlyslightlybiotur-
batedlimestones.Thebaseofthispartcontainstwobedsofcorrod edl i m e s t o n e s : t h e h i g h e r o n e r e p r e s e n t s t h e U p p e r Kellwasserh o r i z o n ( J a n i s z ewskae t a l .2 0 0 7;S z r e k a n d Ginter2007).Thisbedcontainsnumerouspoorlypr eservedgoniatites,clymenids,nautiloids,brachiopodsa ndraretri-lobites.Rackietal.
(2002)distinguishedapurelimestone unit(UnitC)whi chincludesstronglybituminouscephalo-
podlimestonelayeratthebottompart,foll owedbyathin coquinaintercalation,andthenrhythmicallybedded,unfos- siliferousmarlylimestonesandshaleswithbrachiopods atthetopmostpart.Moreover,Bondetal.
(2004)recognizedthebivalveaff.Buchiolasp.,entomozoa ceanostracodaandh o m o c t e n i d s . O n l y s o m e b e d s o f l i m e s t o n e s c o n t a i n Trichichnusisp.Thebedsabove andbelowtheUpperKell-
wasserhorizonareusuallyalsopartlycorrodedandcontainnum erouscorrodedclastsandnodules(Fig.6d).Thelime- stoneb e d s a r e r a r e l y i n t e r b e d d e d w i t h t h i n b e d s o f s i l t y shales(Fig.4).Onespecimenofthetracefossil?
Curvoli-thusisp.hasbeenfoundabout60cmundertheF–
Fbound-ary(Fig.7f).
Kowalaquarry
Thisi s a n a c t iveq u a r r y i s l o c a t e d 5 k m S W o f K i e l c e (Fig.1).TheFrasnian–
Famennianboundaryintervaliscroppedoutonthenorth- easternwallofthequarry,onthetwol owermostexploitationl e vels( F i g .2),w h e r e l a t e r a l changesi nlit hologyh aveb eenobser vedduri ngthestudy
int he ye ar s2 01 1, 20 12 an d2 01 5 ( c f. R a c ki et al .20 0 2).Thel owerp a r t o f t h e i n t e r vali s c o m p o s e d o f t a l u s r u d -
stoneb e d s c o n t a i n i n g m o s t l y f r a g m e n t s o f s t r o m a t o p o r o -
idsa n d R u g o s a c o r a l s . I n p r eviouss t u d i e s o f t h i s l owerpart,amostlyr hythmic,marlysuccession wass ubdividedintotheH1andH2units(Rackietal.2002;Vish nevskayaetal.2002)ortheunits4and5(Bondetal.2004 ).Theseunitsw e r e r e c o g n i z e d —
M a y 2 0 1 2 a n d S e p t e m b e r 2 0 1 5 , whicharerece ntlyrepresentedbythickbedsofbioclastic limeston es.Thisisexplainedbythelateral facieschangeswhichar eexposedbytheprogressinexploitationandremovalofthep reviouslyobservedrockswithintheactive,largeq u a r r y.Ab ovet h e l owerp a r t , a n a p p r o x i m a t e l y 1-m-
thicks e r i e s o f l a m i n a t e d , i n p l a c e s “ s p o t t e d ,”l i m e -
stones,whichcontainabundantTrichichnusisp.andother ,sparsenon-
i dent i fi ed b i ot ur ba t i on al st ru ct ur es (F i g s.4,7,8), i s o b s e r ved.Th e l i m e s t o n e s , i n t h e i n t e r vall o c a t e d 150–
260c m a b ovet h e r u d s t o n e s , c o n t a i n t h r e e b e d s o f micriticlimestoneswitha3or4horizonsofflints(Fig .5).Thethirdbedcontainstwoflinthorizons(Fig.6d), whereaccordingt o bi b l i o g r a p h i c a l d a t a i n p r eviouss t u d i e s ( e . g . Joachimskietal.2001),theFrasnian–
Famennianisrecog-
nized.Abovethisbed,laminated,non-
bioturbatedbedsof limestonesandmarlstonescropout.
Acharacteristic,lami-
natedbedofmarlstonewithcorroded,bioturbatednodu lesandpyriteconcretionsandimpregnationoccurabout50cm abovetheF–
Fboundary.Ametreabovetheboundary,bedsofmarlstone sdisplayprimarylamination,partlydisturbed byuni denti fi ed bi ot urbat i onal st ruct ures(F i g.6g).Hi gh er up( f ewde ci m e t r es t o 1. 7m ), bi ot ur ba t e dm a rl st o ne sa nd limestonescont ai nnum eroust rac efossi l s
—sm al l Mul ti na isp.and?Planolites(Figs.6h,7g–
l).Fluctuatingcontentofcarbonatesandstronglaterallitho logicalchangesarechar-
acteristicf e a t u r e s o f t h i s p a r t o f t h e s e c t i o n . Af ewh o r i - zons,e s p e c i a l l y 0 . 5 – 1 m a b ovet h e F –
F b o u n d a r y,s h ownumerouscorrodedcarbonateclastsa ndnoduleswithinshales,marlstonesorsiltyshale,whic harelaterallypartlyreplacedbydissolvedlimestone.
Tracefossilsassemblagesandbioturbational structures
Both,theKowalaandPłuckisectionsshownumerousbeds whichcontainabundantTrichichnusisp.,characteristica
llyintheuppermostpartoftheFrasnian(Figs.3,4,5,6,7,8).The largestnumberofTrichichnusisp.occursinlaminatedlimest onesusual l ywi t hout ot hert ra cefossi l sorbi ot urba- tionalst ruct ures(F igs.6b,7a–
e,8).Trichichnusis visibl e asreddishorbrownish,ferruginou sorpyritizedthreadsandspots.Theyaregenerallyisolated,0.5–
1.2mmindiameter,int h e P ł u c k i s e c t i o n w i t h d i s t i n c t b r i g h t e r h a l o , w h i c h i s upt o 5 m m i n d i a m e t e r.Th e o u t e r m a rgino f t h e h a l o i s
IntJEarthSci(GeolRundsch)(2017)106:157–170 16 1
Fig.3L o g oftheFrasnian–
FamennianboundaryintervalwithKellwasserbedinthePłuckisection.Thecurvesoftheoxygenationontheseafloorbasedonpresenceorabsenceof thetracefossilsandbioturbationstructures,nottoscale
Fig.4S e d i m e n t a r y structuresandtracefossils,Płuckisection.a,bTrichichnusisp.inmicriticlimestone;aHorizontalcut;bVerticalcut;c–e partlycorrodednodulesoflimestonecontainspecimensofTrichichnusisp.withinlaminatedmarls;verticalcuts
diffusive.D e t e r m i n a t i o n o f t h e l e n g t h o f Trichichnusv i s -
ibleinsurfacesofcutslabsisdifficult;thelongestobservedfragme ntattainsalengthofabout20mm.Othertracefos-
silsaregenerallyabsentortheirstateofpreser vationdoes notpermitacloserdetermination;h owever,someofthe mcanbereservedlyassignedtoMultinaisp.and?
Planolitesisp.( F i g .7g–
l).B o t h m e n t i o n e d t r a c e f o s s i l s a r e v i s i b l e onl yinfracturedorpolishedsurfaces.Multinaincrosssec- tionisvisibleasspotsandshort,branchedandunbranchedst ripes.S e r i e s o f c u t s a l l owa n i n s i g h t i n t h e i r 3 - D m o r-
phology.Theyareendichnial,horizontal,obliqueand v er-
ticalthin,tubular,branchedstrings,whichareunlined,about 0.3mmindiameterandfilledwithmaterialofdiffer-
entcol oursthanthehostrock. Thebranchedst ri ngsf orm irregularp o l y g o n s , w h i c h s h owm a nyc r o s s i n g s a t d i ffer-entl evelst h a t f o r m e d a 3 -
D n e t work( F i g .7g–l).S p e c i - mensof?
Planolitesisp.arevisibleassubcylindricalrollersof0.4mm in diameter(Fi g.7k–l).Onlyonespeci menof
?
Curvolithusi s p . f r o m t h e P ł u c k i s e c t i o n i s v i s i b l e a s a n epichnialribbon,about40mmwide,withadistinctb e velontheoneside(Fig.7f).Itsstateofpreservationdoesno
tpermita c l o s e r d e t e r m i n a t i o n . C l e a r l y v i s i b l e butu n d e -
terminedb i o t u r b a t i o n a l s t r u c t u r e s o c c u r o n l y w i t h i n a veryf ewb e d s . Theya r e r e p r e s e n t e d b y i r r egular,s t r a i g h t orslightlycurvedstripesupto5mmlongonthecutsu r -
faces(Fi g.6g).However,m ost oft hesl i ght l ybi ot urbat e d
bedss h owo n l y i n d i s t i n c t s p o t t e d o r m o t t l e d i c h n o fabric(Fig.6c,g,h).
Discussi on
TracefossilassemblageattheFrasnian–
FamenniantransitioninCentralPoland ComparativeanalysisoftheFrasnian–
Famennianboundaryintervali n t h e s t u d i e d s e c t i o n s a t t h e KowalaQ u a r r y a n d thePłuckioutcropshowssimilar features,suchasvariablecarbonatecontent,laterallithologi calchangesandthepres-
enceofhorizonswiththeabundantferruginousorpyritizedtra cefossilTrichichnusisp.However,detailedichnologicalan alysisthroughtheF–
Ftransitionrevealssignificantdif- ferences,whichareasfollows:
1. int h e U p p e r F r a s n i a n o f t h e Kowalas e c t i o n , b e l owtheF–
Fboundary,numerouslaminated,unbioturbated,limes tonebedsarepunctuatedonlybythepresenceofseve ralinter valswithabundantTrichichnus,whilethePłu ckis e c t i o n i s d o m i n a t e d b y b i o t u r b a t e d b e d s ( f r e -
quentTrichichnusandrare,undeterminedbioturba- tionalstructures),withafewunbioturbatedintervals;
2. att heF–
Fboundar y,simi lari chnologi calfeat uresare pr esenti n b o t h s e c t i o n s , g e n e r a l l y m a n i f e s t e d b y t h e
Fig.5L o g oftheFrasnian–FamennianboundaryintervalintheKowalaquarry.Thecurvesoftheoxygenationontheseafloorbasedonichno- logicaldata,nottoscale.aDatafromtheyears2011–2012andbdatafromtheyear2015
disappearanceo f t r a c e f o s s i l s . H owever,t h i s a b s e n c e isregisteredseveralcentimetresbelowtheboundary intheKowalasect ion,andj ustbel owtheF –
Fboundary inthePłuckisection;
3. abovet h e F –
F b o u n d a r y,t r a c e f o s s i l a s s e m b l a g e s a r e signific antlyd i fferentb e t w e e n t h e s e c t i o n s , w i t h t h e a ppearanceo f l a rgers t r u c t u r e s ( Multinaa n d ?
Plano-lites),somecentimetresabovetheF–
Fboundaryinthe
Fig.6S e d i m e n t a r y s t r u c t u r e s , d i a g e n e t i c s t r u c t u r e s a n d t r a c e f o s -
silsi n p o l i s h e d s u r faces,Kowalaq u a r r y.a B e d o f t a l u s r u d s t o n e s . bLaminated bedso flimestoneswith Trichichnusisp.wh itearr ows.cMixedlayercontainnumerousTrichichnusisp.dFrasni an–Famen-
nianboundary,bedoflaminatedlimestonewithtwohorizonsofflint.
eB e d o f l a m i n a t e d m a r l . f Partlyl a m i n a t e d b e d o f m a r l w i t h c o r-
roded,bioturbatednodules(cn)andpyrite.gBedoflaminated,partlybiot urbatedmarl.hBedofbioturbatedlimestonethatcontainsmainlyMultinai sp.(M)and?Planolitesisp.(P)
Kowalas e c t i o n , w h i l e u n b i o t u r b a t e d b e d s w i t h o n l y rarer e c o r d o f Trichichnus,?
Curvolithuso r u n d e t e r-
minedb i o t u r b a t i o n a l s t r u c t u r e s a r e p r e s e n t i n a t h i c k intervalabovetheboundaryinthePłuckisection.
Thesignificantdifferencesincompositionanddistri- butionoftracefossilsandbioturbationalstructuresintheF–
Fboundaryintervalsofthetwostudiedsectionscouldberelat edtoadifferentincidenceoftheUpperKellwasser
Fig.7Tracef o s s il s f r o m t h e F–
F b o u n d a r y i n t e r valo f Kowalaan d Płuckis e c t i o n s . a – eTrichichnusi s p . i n t h i n s e c t i o n s ; a h o r i z o n t a l cut;b–
fverticalcut;a,b,dKowala;c,ePłucki.f?Curvolithusisp.,
epichnionon med iu mb ed o fmarls. g–
lMultina (M)an d Plan olites
(P)insections,Kowala;g–jverticalcut;k,lhorizontalsections;dis- tancesbetweenneighbouringcutca.0.3mm
Fig.8S ketchdrawandexamplesoftherecordofdepositoxygena- tion.aLaminated,non-
bioturbateddeposit,blaminateddepositwithTrichichnusisp ., cm i xedlayerwith Trichichnusisp ., dbio turba ted,
spottedlayerwithMultinaisp.andPlanolitesisp.Theimproveme ntofoxygenationfollowstherecoverytrendaftertheF–
Fmassextinc-tion,interpretedbyWangetal.(2006)
eventintheregionaldepositionalsetting.TheDevoniansedime ntsoftheHolyCrossMountainsweredepositedina600-km- wideshelf,showingpalaeogeographicandtopo-
graphicvariations.TheKowalasectioncorrespondstoa shallo wer,intrashelfdepositionalsetting(Che˛ciny-
ZbrzaBasin)neartheDyminyReef,whilesedimentsofthePłuck isectionweredepositedinthedeeperŁysogóryBasin(e.g.Racki etal.2002).Thesedifferentdepositionalsettingscoulddetermin eavariableincidenceofthepalaeoenviron-
mentalchangesaffectingmacrobenthictracemakercom- munityprior,duringandaftertheUpperKellwasserphe- nomena,andthusthedifferentichnologicalrecord.Thus,intheK owalasettingdeteriorationofthemacrobenthictrace-
makerenvironmenttookplaceearlier(priortotheUpperKellwa sserevent),butalsotherecoverywascomparativelyearlieraftert heevent,whileinthePłuckisection,theich-
nologicaldatapointtoadepauperatedenvironmentthatappeare dnearlysimultaneouslywiththeUpperKellwassereventandper sistedforlongeraftertheevent.
Oxygenation/productivitychangesattheUpperKellwasserev ent
Previousi n t egrativea n a l y s e s o f t h e F r a s n i a n – Famenn-
ianboundaryintervalintheHolyCrossMountainsmainly focusedattheKowalasection,referredtochangesinoxy- genationa n d p r o d u c t ivitya s t h e m a i n e nvironmentalfac -
torsd u r i n g t h e U p p e r Kellwasserevent( e . g . J o a c h i m s k i eta l .2 0 0 1;G i r a r d a n d L é c u y e r2 0 0 2;P u j o l e t a l .2 0 0 6;
BorcuchandRakocin
´ski2007).TheanalysisofV/Crratios,withvalueso f 0 a b o u t 2 . 5 m b e l owt h e F – F b o u n d a r y,6 attheF–
Fboundary,and18about8mab ovethisbound- ary(Joachimskietal.2001),clearlypointstoanoxiaaf terthisboundary.TheV/Crratios>5inafewspotsbelowth eF–
Fb o u n d a r y p o i n t a n o x i a a l s o b e f o r e t h i s c r i s i s . H ow-
ever,a l a t e r s t u d y s h owsd i fferentvalueso f V / C r r a t i o s
(0.25a t t h e F –
F b o u n d a r y a n d 2 . 2 5 t e n s o f c e n t i m e t r e s abo vetheboundary)andsuggestsgeneraldysoxiaandthe absenceofapermanentanoxiaattheF–Fboundarytransi- tion(Rackietal.2002).Theloweringinoxygenationju stabovetheF–
FboundaryintheKowalasectionisindicatedbyt h e C e a n o m a l y ( G i r a r d a n d L é c u y e r2 0 0 2).H owever,thea n o x i a d u r i n g t h e Kellwassere p i s o d e s wasi n t e r m i t -
tent,p e r h a p s s e a s o n a l , n o t p e r m a n e n t ( R a c k i e t a l .2 0 0 2andc i t e d r e f e r e n c e s t h e r e i n ) . A cycli cm o d e l o f a n o x i c – dysoxic–
weaklyoxicconditionsduringtheLateDevo-
niana c c o m p a n i e d b y m a s s o c c u r r e n c e s o f cyanob acterialmatsw h i c h c o l o n i z e d t h e o x y g e n -
d e f i c i e n t s e a f l o o r wasproposedb y K a z
´mierczake t a l . (2012).S i m i l a r,a n o x i c – suboxic–
oxice nvironmentsb a s e d o n pyritef r a m b o i d s i z e andcy anobacterialm a t s o n t h e s e a f l o o r a r o u n d t h e F – F boundaryatKowalawereinterpretedbyMarynowskietal.
(2011).C o m p l e t e a b s e n c e o f b i o t u r b a t i o n a l s t r u c t u r e s i n theL owera n d U p p e r Kellwasserb e d s i n G e r m a nys u g -
gesta n o x i a ( B u g g i s c h1 9 9 1).I c h n o l o g i c a l d a t a o b t a i n e d inthepresentstudygenerallyagreewiththe interpretationsbyBuggisch(1991)andJoachimskietal.
(2001),buttheybringa m o r e p r e c i s e p i c t u r e o f t h e o x y g e n a t i o n c h a n g e s .
Trichichnus,p r o b a b l y p r o d u c e d b y o p p o r t u n i s t i c c h e m o -
symbionts(Uchman1995),istypicalofpoorlyoxygenatedde posit(McBrideandPicard1991),especiallyifitoccurs ast h e o n l y t r a c e f o s s i l ( Kotlarczyka n d U c h m a n 2 0 1 2).Recently,Ke˛dzierskietal.
(2015)proposedthattheichno-
genusTrichichnusi s a s t r u c t u r e o f f o s s i l b i o e l e c t r i c b a c - terialactivityattheoxic–
anoxicinterfaceintheseafloor.Thedisappearanceofbr achiopods(RackiandBalin
´ski1998),a n d b e n t h i c o s t r a c o d s ( O l e m p s k a2 0 0 2 )a b o u t t h e F–
Fboundary,confirmsoxygendeficiency.However,othercause s,s u c h a s a d r a s t i c d e c r e a s e i n t e m p e r a t u r e , w h i c h isevidencedbyδ18O,havebeenalsoinvoked(Joachims kiandBuggisch2007).
Thea b ove-
presentedvariationsi n c o m p o s i t i o n a n d distributiono f t r a c e f o s s i l s , b i o t u r b a t i o n a l s t r u c t u r e s a n d l aminatedintervalscouldbeassociatedwithfluctuationsinoxy genconditionsfromanoxic(laminated)tostronglydys-
oxic( h o r i z o n s w i t h Trichichnus)a n d d y s o x i c ( s p o t t y a n d mottledi c h n o fabrics,w i t h M u l t i n a a n d ?
Planolites).T h e presenceofstromatoporoidsandcoralsp ointtogoodoxy-
genationduringtheFrasnian.Indetail,theinterpretationisas follows:
1. belowtheF–
Fboundary,theKowalasectionshowsV/Crvaluescloset o0reflectingoxicconditionsandthennumerous,minor,ch anges(valuesof0–0.8)associ-
atedwithfluctuationsbetweenoxicandanoxiccondi -
tionsclose,fromaround100cmbel ow,tothebound- ary( R a c k i e t a l .2 0 0 2);t h i s a g r e e s w i t h t h e p r e s e n c e ofs everali n t e r valsw i t h Trichichnusa l t e r n a t i n g w i t h unbioturbated,n o n -
l a m i n a t e d o r o c c a s i o n a l l y l a m i -
natedlimestonebeds.InthePłuckisection, V/Crv al-uesa r e s i m i l a r t o o x i c l evelsb e l owt h e F – F w i t h o u t markedfluctuations(Rackietal.2002).T hiscouldbecorrelatedw i t h t h e g e n e r a l d o m i n a n c e o f b i o t u r b a t e d beds( Trichichnusa n d u n d e t e r m i n e d b i o t u r b a t i o n a l structures)s e p a r a t e d b y a f ewt h i n n e r,u n b i o t u r b a t e d intervals,whichsugg esttheexistenceofminorfluctua-tionsinoxygenation;
2. attheF–
Fboundary,importantchangesinV/Crvaluesarer e c o g n i z e d i n b o t h s e c t i o n s r evealingi m p overish-
menti n o x y g e n c o n d i t i o n s ( R a c k i e t a l .2 0 0 2).
T h eyaree s p e c i a l l y m a r kedi n t h e P ł u c k i s e c t i o n w i t h val-
uesfavouringtheinterpretationofanoxia(Bondetal.2 004),whiletheKowalasectionrepresentsfluctuationsbetwee noxicanddysoxic(Rackietal.2002)oranoxicconditions(Jo achimskietal.2001).Thispatternagreeswitht h e g e n e r a l a b s e n c e o f t r a c e f o s s i l s i n b o t h s e c -
tions,aroundtheF–FboundaryatKowalaandincoin- cidencewi t h t hi s b ou nd ar y a t P ł uc ki . M o r e over,δ34 Sandδ13Cclearlypoint toanoxiaontheseafloorasar esultofanincreaseino rganiccarbondepositionand r eductionofsulphur(Buggisch1991);
3. abovet h e U p p e r Kellwasserl evel,V/Crvaluesr eveal ar e l a t ivelyr a p i d i m p r ovemento f o x i c c o n d i t i o n s i n theKowalas e c t i o n , a n d a p r o l o n g e d d e t e r i o r a t i o n i n oxygenationmaintainedinthePłuc kisection(sub-oxic–
anoxic;R a c k i e t a l .2 0 0 2).These i m p r ovementsin o x i c c o n d i t i o n s c o r r e s p o n d w e l l t o t h e a p p e a r a n c e ofMu l t i na or ?
Planolitesso me c en t i m e t r es ab ovetheF–
FboundaryintheKowalasection.However,thedom i na nceofunbi ot urbat edb eds,wi t honl yl ocal appe aranceofTrichichnus,apossiblesinglespeci- meno f ?
Curvolithuso r u n d e t e r m i n e d b i o t u r b a t i o n a l structuresinthePłuckisectionstillsuggestsprevai linganoxicconditions,aswasalsointerpretedforthewat ercolumnbasedonthepreviousgeochemicalstudies (Joachimskietal.2001).
Thus,ichnological dataconfirmthefluctuations ontheseaflooroxygenationandtheirinfluenceonthemacro ben-
thict r a c e m a kerc o m m u n i t y d u r i n g t h e U p p e r K ellwasserevent.F l uct uat i onsi noxygenat i onbefor et heF –F bound-
aryandduringtheUpper Kellwassereventwerepossibl e.Thisi s s u g g e s t e d b y t h e p r e s e n c e o f u n b i o t u r b a t e d h o r i -
zonsb e t w e e n b i o t u r b a t e d b e d s . H owever,a b e d o f b i t u -
minous,cephalopodlimestonei n thePłuckiarea,inter- preteda s t h e L owerKellwasser,h a s b e e n r e c o g n i z e d c a . 60mbelow(Rackietal.2002;SzrekandGinter2007)th eUpperKellwasser.Similarly,bedswithTOCmaximu mintheKowalasection,correspondingtotheLowerKellwas serevent,arerecognizedabout15mbelowtheF–
Fboundary(Joachimskietal.2001;DeVleeschouweretal.20 13).OurinterpretationofoxygenationchangesclosetoF–
Fbound-
arycorrespondsrathertotheearlyUpperKellwasserevent.H owever,t heabsenc eofahi gh-
resol ut i onbi ost rat i grap hyprecludesamorepreci se i nterpretati on. Togetherwit hthe generallyoxygen- deficientlowerwatercolumnduringlateFrasnianandearly Famennian,TOCanalysisintheKowalasectionshowstwoma ximaacrosstheFrasnian–Famen-
nianboundarytransitioncorrelatedwiththeLowerandUpper Kellwasserevents( J o a c h i m s k i e t a l .2 0 0 1).T h e s e e nhancedTOCconcentrationswere explainedbyahighe rprimaryproductivity(higherphotoautotrophproduction) thatcouldbeassociatedwithahighersupplyofcontinent -
derivedn u t r i e n t s ( J o a c h i m s k i e t a l .2 0 0 1;F i l i p i a k2 0 0 2;Kazmierczake t a l .2 0 1 2).H owever,R i q u i e r e t a l .
(2006)proposedtwodifferentmechanismsresponsibleforox ygendeficiencyd u r i n g s e d i m e n t a t i o n o f t h e L owera n d U p p e r Kellwasser:t hef i r st on ec an b ea resul t o fa ni n cr ea s ei n productivityinshallow-
waterenvironments,whilethesec-
ondiscausedbyadevelopmentofeutrophicconditionsint hebasin.
Variationsinproductivitycouldbealsoenvisagedbasedon theobtainedichnologicaldata.Low-
oxygenatedbottomwaterandporewatercombinedwithorganic -richsediments
probablyinducepotentiallyidealconditionsforchemo- symbiotica c t ivity,favouringt h e p r e s e n c e o f Trichichn ustracemaker(e.g.Löwemarketal.2006).Accordingtopre -viouspapers(e.g.Rackietal.2002;Pujoletal.2006;Win- ter2015),organicmatterproductionaswellasoxygenationwasc ontrolledbytectono-
magmaticprocesses(volcanism,hydrothermalphenomena )inenrichmentinmetalsandnutrientsandtheloweringofth eoxygenationduringdepo-sitionoftheKellwasserhorizons.
TracefossilsandtheUpperKellwassereventworldwide Previousi c h n o l o g i c a l a n a l y s e s f o c u s i n g o n t h e F r a s n i a n –
Famenniant r a n s i t i o n a r e c o m p a r a t ivelys c a r c e ( S t a c h a c z andUchman2012;Haddadetal.2013;Boyeret al.2014).Boyere t a l .
(2014),b a s e d o n a d e t a i l e d i c h n o l o g i c a l a n d geoche micalanalysisinthewesternNewYorkState(USA)interpreted l o n g -
t e r m e nvironmentalp e r t u r b a t i o n s , r a t h e r thana g e o l o g i c a l l y i n s t a n t a n e o u s o n s e t o f a n o x i a a s t h e causalmechanismoftheextinctionduringtheUpper Kell-
wasserevent.Ichnologicaldatasupportarapiddeoxygena- tioneventbelowthehorizontraditionallyinterpretedastheUp perKellwasseranoxi ci nt er val,int heso-call edprecu r- sorb l a c k s h a l e b e d ( P B S B ) , t h e n a n i n c r e a s e i n o x y g e n levels( n o t f u l l y o x y g e n a t e d ) , c h a r a c t e r i z e d b y a s h a l l owChondrites-
dominateda s s e m b l a g e , f o l l owedb y a g r a d u a l decreas ei n t h e o x y g e n l evelsi n t o t h e U p p e r Kellwasserblac kshaleinterval.Theseconditionsaremaintainedforarela tivelyl o n g p e r i o d o f p e r s i s t e n t o x y g e n s t r e s s ( butn o t persistenta n o x i a , a s r e f l e c t e d b y t h e p r e s e n c e o f s m a l l e r tracemakersbutexcludinglargerburrowingorganis ms)throughthisinterval,andthenrapidlyshifttooxygen- atedenvironmentabovetheUpperKellwasserinterval,asrev ealedbyt heappearanceofthel a rgeThalassi noi des (B oyeretal.2014).
ComparingthestudiedsectionsformCentralPolandandt hesefromN ewYorkState,inbothcases,scarceand lo w-
diversityt r a c e f o s s i l a s s e m b l a g e s a r e p r e s e n t i n t h e Frasnian–
Famenniant r a n s i t i o n i n t e r val,w h i c h a r e d o m i - natedbysm al l burrows,i ncl udi ng C hondrit esi n t heN e wYorkState,andTrichichnusinPoland;inbothcasesreflect- ingloweredoxygenconditions.PriortotheF–
Ftransitioninterval( t h e U p p e r Kellwasserevent),t h e P B S B r e c o g -
nizedinNewYorkStatecouldbecorrelatedwiththerecordofunb ioturbated,laminatedorblack,bituminouscephalo- podl i m e s t o n e b e d s i n C e n t r a l P o l a n d . L a rgers t r
u c t u r e s , suchasThalassinoidesorPlanolites /?
Chondrites,revealareturntobetteroxygenatedconditionsaftert heUpperKell-
wasserevent.InNewYorkState,neitherarapiddeoxygen- ationattheUpperKellwassereventnorapersistentanoxiaisint erpreted,butagradualdecreaseinoxygenationtodys-
oxic/suboxiclevels.Thispatternissimilartothatoccurring
intheKowalasection,withlow-
oxygenconditionspriortotheUpperKellwasserevent,the nmaintenanceofdysoxia, followedbytheimprovementi nbottomwateroxygencon-
ditionsinacomparativelyshorttimeaftertheevent.
Inshort,thiscomparisonconfirmsthegeneralizedi nci-
denceoffluctuatingoxygenconditionsassociatedwithth eUpperKellwasserevent,affectingthemacrobenthictra ce-
makercom munit yinaglobalscale.H owever,it isnec es-
sarytoem phasizet hatt hemacrobenthictracem a ker com -
munityshowsnotauniquepattern,butavariableresponse,pri or,d u r i n g a n d a f t e r t h e event.I n t h e c a s e s t u d y f r o m Poland,t h e p r e s e n c e o f n o n -
b i o t u r b a t e d , l a m i n a t e d i n t e r-
vals,allowinginterpretationofanoxicconditionsisa sig-
nificantfeature.Thisminordifferenceincomparisonw ithNewYorkStatecouldbeassociatedwithlocalsignals,cau sed,fori nstance,by variablet opograp hyand/ orocea ndynamics,whichmaycontroldurationandintensityoft heoxygendeficiency.Itmightbeworthnotingthatwhiletherear ep r e d i c t a b l e d i fferencesi n t h e s i g n a l s , overallt h eya r e quitesi m i l a r,w hi ch sp ea ks t o ag l o ba l s c al ed r iver r at he r thansomethinglocal.Thisseemsanimportantpointto pulloutofthesecomparisons.
Conclusio ns
1. Thelaminat ed,non-
biot urbatedbedsofm arl st onesor shales,usually withpyriteaboveFrasnian–
Famennianboundary,indicateanoxiaontheseafloorjus taftertheF–
Fevent.H owever,t h e a n o x i c evento c c u r s e a r l i e r thantheF–
F,asrecordedbyabedoflaminatedlime- stone.
2. Theincreaseina bundanceoftracefossilsandbiot ur-bationalstructuresinsomebedsabovetheF–
Fbound-
aryisinterpretedasanincreaseinbottom waterox y-genlevels.
3. Lowichnodiversitysuggestsanunfavourableenviro n-
mentforthemostburrowingorganisms.Inthisgeneralco ntext,t h e p r e s e n c e o f l a m i n a t e d i n t e r vals,h o r i z o n s withopportunisticTrichichnus,andonlyspottyan dmottledichnofabrics,withthelargertracefossilsMu l-tinaor?Planolites,allowstheinterpretationoffluctua-
tionsinoxygenationfromanoxic,stronglydysoxicanddys oxicconditions.
AcknowledgmentsResearchbyM.S.andA.U.wassupportedbyJagie llonianUniversity(ProjectDS/MND/WBiNoZ/ING/2/2011and DSfund s,respect ively).M.S.wishestothank toDy c kerhofPolsk aCompany( S i t k ó w k a -
N owiny)f o r t h e f i e l d workp e r m i s s i o n i n t h e Kowalaquarry.
FinancialsupportbyR.- T.wasprovidedbythePro- jectC G L 2 0 1 2 -
3 3 2 8 1 ( S e c r e t a r í a d e E s t a d o d e I +D+I,E s p a ñ a ) a n d therese archgroupRNM-178(JuntadeAndalucía).G.Racki(Univer-
sityofSilesia),D.B oyer(ShinemanScienceCenter)andanano n y-mousreviewerprovidedconstructivereviews.
OpenA c c e s s T h i s articleisdistributedunderthetermsoftheCre ativeCommonsAttribution4.0InternationalLicense(http://crea- tivecommons.org/licenses/by/4.0/),whichpermitsunrestricteduse,d istribution,andreproductioninanymedium,providedyougiveappropria tecredittotheoriginalauthor(s)andthesource,pr ovidealinktot h eCreat iveCo mmon slicen se,and in dicateifch ang eswere ma de.
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