Occurrence of the trace fossil Zoophycos from the Upper Viséan Paprotnia Beds of the Bardo Structural Unit (Sudetes, SW Poland)

Occurrence of the trace fos sil Zoophycos from the Up per Viséan Paprotnia Beds

of the Bardo Struc tural Unit (Sudetes, SW Po land)

Jolanta Muszer & Joanna Haydukiewicz

Uni ver sity of Wroc³aw, In sti tute of Geo log i cal Sci ences, Cybulskiego 30; 50-205 Wroc³aw, Po land, e-mails: jolanta.muszer@ing.uni.wroc.pl; joanna.haydukiewicz@ing.uni.wroc.pl

Key words: Zoophycos, trace fos sils, Up per Viséan, Sudetes, Po land.

Ab stract

Manu script re ceived 16 Au gust 2009, ac cepted 18 De cem ber 2009

IN TRO DUC TION

Zoophycos is a term that de scribes a com mon trace fos - sil formed of a large num ber of three-di men sional, heli - cally coiled spreiten struc tures that can have very vari able de tailed morphologies and that have been de scribed from ma rine sed i ments rang ing in age from Cam brian to Qua - ter nary (see Bradley, 1973; Wetzel & Werner, 1981;

Kotake, 1989, 1997; Miller, 1991; Olivero, 2003; Löwe- mark et al., 2006; Knaust, 2009). Zoophycos have also been rarely de scribed from Pre cam brian rocks (Crimes, 1987;

Uchman, 1998). This enig matic and cos mo pol i tan ichno- fos sil has chal lenged gen er a tions of ge ol o gists and ichno- logists for over a cen tury (for the ear lier pa pers see Oli- vero, 2007; Olivero & Gaillard, 2007; Knaust, 2009). The Zoophycos ichnofacies is one of the four orig i nal ichnofa - cies pi o neered by Adolf Seilacher (Seilacher 1963, 1964;

see Frey et al., 1990; Bromley & Asgaard, 1991). Tra di - tional in ter pre ta tions place Zoophycos within Seilacher’s ethologic cat e gory fodinichnia (Seilacher, 1986), which in - cludes the semi-per ma nent struc tures pro duced by endo- ben thic de posit feed ers (see Miller & D’Alberto, 2001).

The broad col lec tion of re lated forms pres ently grouped as Zoophycos are cov ered by the ichnofamily Alectoruridae (see Bromley & Hanken, 2003). Ac cord ing to Miller (2003) and Olivero (2007), Zoophycos is a very com plex trace fos sil and there is still no real agree ment

con cern ing its tax on omy and sig nif i cance. Thus, Uchman (1995) pro posed the term “Zoophycos group” to in clude all the trace fos sils that share some com mon mor pho log i - cal “Zoophycos” char ac ter is tics.

De spite Zoophycos be ing com mon in both an cient and mod ern sed i ments, the an i mal or an i mals re spon si ble for these traces have yet to be dis cov ered; and a be hav ioural ex pla na tion is still strongly de bated (e.g. Bromley, 1991).

Zoophycos (sensu lato) is gen er ally as sumed to be the trace of an as-yet un dis cov ered de posit feeder (see Uchman, 1998; Rodríguez-Tovar & Uchman, 2006). Zoophycos pro - duc ers have been var i ously hy poth e sized to be si pun cu lids (Wetzel & Werner, 1981), polychaete annelids, ar thro pods (Ekdale & Lewis, 1991) or echiuran worms (Kotake, 1992).

Zoophycos have been de scribed from a di verse range of sed i ments, in clud ing sand stones, mudstones, shales, lime - stones, marly lime stones, marls, chalk (Olivero, 2007) and hemipelagic mud (Leuschner et al., 2002). These rock types rep re sent a very wide range of palaeoenvironments, from the infralittoral to the abys sal (Frey et al., 1990).

Zoophycos have even been de scribed from re stricted la goon en vi ron ments (Osgood & Szmuc, 1972) and from a glacio- ma rine en vi ron ment (Bhattacharya & Bhattacharya, 2007;

Gong et al., 2008).

Zoophycos have been re corded from subtidal to basinal en vi ron ments: yet there is ev i dence that the bathymetric range of Zoophycos has in creased over time (see Kotake, 1997). Most Palaeozoic Zoophycos were rec og nized from shal low-wa ter de pos its, whereas Me so zoic–Ce no zoic spe- cimens seem re stricted pre dom i nantly to deep-wa ter sed i - ments (e.g. Ekdale & Bromley, 1984; Seilacher, 1986;

Kotake, 1989, 1997; Miller, 1991; Bromley, 1996; Leusch- ner et al., 2002; Olivero, 2003; Wetzel et al., 2007; Knaust, 2009). And in mod ern seas, Zoophycos is only known from mid dle bathyal to abys sal sed i ments (Wetzel & Werner, 1981; Kotake, 1989).

The real sedimentological value of the Zoophycos ich- nofacies is still enig matic (Ekdale, 1992; Goldring, 1993).

Ac cord ing to Frey & Pem ber ton (1985) and Frey et al.

(1990), the Zoophycos pro ducer could live in circalittoral to bathyal en vi ron ments, in quiet wa ters, and be ac tive in rel - a tively ox y gen-de fi cient con di tions.

Spe cif i cally with re spect to Lower Car bon if er ous de - pos its, Zoophycos is com mon in South Wales (Wu, 1982), Bel gium (Gaillard et al., 1999), the Moravian–Silesian Zone (Zapletal & Pek, 1999; Mikuláš et al., 2004), Texas (Yurewicz, 1977), New Mex ico (Coo per & Dutro, 1982), West Vir ginia (Bjerstedt, 1988), north west Al berta (Sir- man & Pem ber ton, 2003) and Ja pan (Kotake, 1997). This ichnogenus oc curs ei ther in low-en ergy car bon ate fa cies

(Yurewicz, 1977; Gaillard et al., 1999), or in the Culm fa - cies, de pos ited from in ner- to the mid dle-deep-sea fans (Zapletal & Pek, 1999; Mikuláš et al., 2004), or in shales, which rep re sent a fully ma rine en vi ron ment (Sirman &

Pem ber ton, 2003). Zoophycos trace fos sils have also been rec og nized from Lower Car bon if er ous deltaic fa cies (Bjer- stedt, 1988).

In Po land, Zoophycos ichnospecies have been descri- bed mainly from the Tithonian–Mio cene flysch for ma - tions of the Carpathians (e.g. Ksi¹¿kiewicz, 1977, Wetzel

& Uchman, 1998; Uchman, 1998) and from the Ju ras sic car bon ates of the Pieniny Klippen Belt (Tyszka, 1994).

Rotnicka (2005) cited pos si ble Zoophycos from the fine- grained Up per Cre ta ceous rocks of the Sto³owe Moun - tains (Sudetes). Paszkowski & Uchman (1992) and recen- tly Hoffmann et al. (2009) rec og nized Zoophycos from the car bon ate de pos its of the Rudawa Group (Mid dle-Up per Tournaisian) of the Kraków Block.

To date, ichnogenera have not been de scribed from the Palaeozoic de pos its of Sudetes. The first dis cov ery of Zoophycos from the Palaeozoic strata of the Pol ish Sudetes has been shortly repoprted by Muszer & Haydukiewicz (2010). This paper spe cif i cally de scribes Zoophycos that oc - cur rence from the Up per Viséan Paprotnia Beds of the Bardo Struc tural Unit.

GEO LOG I CAL SET TING

The Bardo Unit is a sep a rate com plex struc ture within the cen tral Pol ish Sudetes (Fig. 1). Ac cord ing to the tec- tono-strati graphic model pro posed by Wajsprych (1995), this unit is built up of both allochthonous and autochtho- nous/parautochthonous suc ces sions. The allochthonous rocks are as signed to the Up per Or do vi cian–De vo nian and oc cur as large olistoliths in the up per most part of the autochthonous se quence. The autochthonous/parauto- chthonous suc ces sion com prises Up per De vo nian–Lower Car bon if er ous rocks. The lower part of Car bon if er ous suc ces sion is rep re sented by sev eral dif fer ent fa cies, whereas its up per part con sists of flysch and wildflysch sed i ments with olistoliths (see Wajsprych, 1995).

The de pos its con tain ing the trace fos sil Zoophycos be - long to the Paprotnia Beds, which oc cur only in the west - ern part of the Bardo Unit and prob a bly un der lie the flysch strata. The Paprotnia Beds (for merly termed the Paprotnia Se ries – see Haydukiewicz & Muszer, 2002) are an in for mal unit be long ing to an autochthonous/parauto- chthonous plat form-to-fore land suc ces sion. The out crop of these beds is sit u ated about 500 m SE of Paprotnia Hill.

The ex posed sec tion is 13.7 m thick, con tains clay- stones and mudstones in its lower part (Fig. 2) and a few in ter ca la tions of thin dark-grey micritic lime stone beds.

These de pos its are over lain by dark-grey and dark-ol ive mudstones that are suc ces sively in ter ca lated up wards with thin (2–4 cm thick) lay ers of grey claystones and grey- wackes. These mudstones also con tain sev eral ben ton ite lay ers and ir reg u larly dis trib uted small (centi metre-scale)

mudstone nod ules (intraformational sub-rounded con cre - tion frag ments) that can be found within the 5.3 m thick shale pack age (Fig. 2). The mid dle part of the sec tion com - prises sandy mudstones and greywackes with lenses and nod ules of dark-grey organodetrital lime stones, which usu ally form dis tinct ho ri zons. The sed i ments are over lain by greywackes con tain ing rare sandy and mudstone in ter - ca la tions. These greywackes form the high est parts of the Paprotnia Beds and pass grad u ally into the polymict Wilcza Con glom er ates (Fig. 2).

The Paprotnia Beds con tain a very rich palaeontolog i - cal re cord that has been stud ied by ge ol o gists and palaeon- tologists for more than 170 years (e.g. von Buch, 1839;

Schmidt, 1925; Haydukiewicz & Muszer, 2002). Brachi o - pods and cor als dom i nate the ben thic fauna; goniatids and nautiloids rep re sent the less com mon nektonic fauna; and ma rine microfossils in clude ostracods, foraminifers, nu - mer ous frag ments of cal car e ous al gae, and var i ous kinds of cal car e ous tubes and spines. Within this ma rine as sem - blage are ter res trial plant de bris that are dis persed throug- hout the de pos its. Haydukiewicz & Muszer (2002) sug - gested that the Paprotnia Beds be long to the Goniatites cre- nistria Zone, which cor re sponds to the Up per Asbian (re - gional substage) of the Up per Viséan (V3b). Haydukie- wicz & Muszer (2002) noted ichnofossils in the Paprotnia Beds, but they did not de scribe any ichnogen era.

Both the lithological and palaeontological fea tures of the Paprotnia Beds re flect a grad ual en vi ron men tal change from off shore to on shore con di tions (Haydukiewicz &

Muszer, 2002). These plat form de pos its are in ter preted as the shal lower-wa ter equiv a lent of the pe lagic crenistria lime stone, which is wide spread in the Culm fa cies of Vari- scan Eu rope.

A re cent U–Pb SHRIMP geo chron ol ogi cal study of

vol ca nic zir cons from a ben ton ite within the Paprotnia Beds gave an age of 334 ± 3 Ma (Kryza et al., 2007; Kryza et al., 2008; Kryza et al., 2010 ). This date is in per fect agreement with cur rent age of the (lower) Asbian (see Menning et al., 2006).

Fig. 1. A – Lo ca tion of the Bardo Unit in the Mid-Eu ro pean Varsicides. Ab bre vi a tions are as fol lows: EFZ (Elbe Fault Zone), H (Harz), O (Odenwald), OFZ (Odra Fault Zone), S (Spessart), Sd (Schwarzwald), TTL (Teisseyre–Tornquist Line), V (Vosges). B – Gen - er al ized geo log i cal map of the Bardo Unit (mod i fied af ter Oberc, 1957 and Haydukiewicz, 2002) in clud ing the lo ca tion of the Paprotnia Beds.

SYS TEM ATIC ICHNOLOGY

Ichnogenus Zoophycos Massalongo, 1855 Zoophycos ichnospecies

Figs 3–5

Ma te rial: Sev eral dozen in com plete spec i mens, cat a logue num bers ING/P-1 to ING/P-35, housed at the In sti tute of Geo log i cal Sci ences, Uni ver sity of Wroc³aw.

De scrip tion: Large spreiten bur row struc tures with ei ther sim ple flat forms or unilobate he li cal forms pre served par - al lel to bed ding. Most spec i mens are parts of larger struc - tures whose ar chi tec ture is un known. Some he li cal forms have down ward growth. The spreiten have a sim ple mor - phol ogy com pa ra ble to the morphotype A as de scribed by Fig. 2. Lithological col umn of the Paprotnia sec tion (based on Haydukiewicz & Muszer, 2002) show ing strati graphic po si tion of the Zoophycos oc cur rence.

Olivero (2003) from Me so zoic de pos its in France. The spe- cimens from the Paprotnia Beds rep re sent two types that can be dif fer en ti ated by the pres ence of sec ond ary lamellae and size. The first type, termed here as morphotype A1, com prises the larger forms, which range in size be tween 110 mm and 160 mm. The sec ond type, termed here as morphotype A2, in cludes the smaller forms, which have a length of up to 70 mm. Mea sure ment of these trace fos sils is of ten dif fi cult be cause many spec i mens cross cut one an - other. Some slabs of Zoophycos are en cir cled by cy lin dri cal mar ginal tubes, pre served only as frag ments. This tubes are 1–2 mm (morphotype A2) to 3–5 mm (morphotype A1) wide. The min i mal heights of the in di vid ual he li cal struc - tures range from about 14 mm (the smaller forms) to about 20–35 mm (the larger forms). The lobes are filled with ar - cu ate, well-marked pri mary lamellae, with the dis tance be - tween two sub se quent lamellae rang ing from 1 to 2 mm (morphotype A2) and from 3 to 7 mm (morphotype A1).

The sec ond ary lamellae oc cur of ten only in the larger forms (morphotype A1) and the dis tance be tween each lamellae are in the range 0.7–1 mm. The acute an gle be - tween pri mary lamellae and sec ond ary lamellae var ies from 10° to nearly 30°, which is sim i lar to Zoophycos de - scribed from the Kitakami Moun tains by Kotake (1997).

The laminae are spi rally coiled around a ver ti cal axis that is ap prox i mately per pen dic u lar to bed ding. The ax ial tun nel is oval in out line and its di am e ter ranges from 12–24 mm (morphotype A2) to 19–30 mm (morphotype A1). The an - gle be tween the bed ding plane and the up per part of the lamina var ies from 20° to 55°. The Paprotnia Beds spec i - mens lack pel lets in the spreite, fea tures that are, by con - trast, of ten pre served in Ce no zoic Zoophycos (see Wetzel &

Werner, 1981; Kotake, 1997).

Oc cur rence and ver ti cal dis tri bu tion: All the Zoophycos spec i mens de scribed here are from the Paprotnia Beds of the Bardo Unit (Sudetes). These trace fos sils are rel a tively com mon.

The ichnogenus Zoophycos is lo cally abun dant in the lower part of the Paprotnia sec tion. Dis tinct con cen tra - tions of this trace fos sil oc cur within the 2.0 m-thick pack - age of dark-grey and dark-ol ive mudstones that im me di - ately underly the first ho ri zon of nod ules of organodetri- tal lime stones. Zoophycos also oc curs lo cally in mudstones be tween the lime stones (Figs 2, 5B), but it has not been pos si ble to dis cern any three-di men sional struc tures from these de pos its. Lo cally, neigh bour ing bur row sys tems inter pen etrate one an other. We ob served that morpho- types A1 (the larger forms) and A2 (the smaller forms) oc - cur in the same ho ri zon.

The lo cal range of Zoophycos from the Paprotnia Beds cor re sponds to the up per most part of taphocoenosis (II), an as sem blage char ac ter ized by a dom i nance of epifaunal sus pen sion feed ers and the pres ence of bioturbational structures (see Haydukiewicz & Muszer, 2002). Other ichnofos sils that oc cur in this taphocoenosis are rare, poorly pre served frag ments that are dif fi cult to iden tify:

they are partly filled with a po rous ferruginous sub stance

(see Figs 10G and 10H in Haydukiewicz & Muszer, 2002).

The fos sils of this com mu nity are usu ally ir reg u larly dis - persed in mudstone that con tain greywacke in ter ca la tions and thin ben ton ite lay ers (Fig. 2). Un for tu nately, Zoo- phycos struc tures and other ichnofossils have not been ob - served within these bentonites.

This taphocoenosis (II) as sem blage in di cates os cil la - tions in en vi ron men tal con di tions from storm wave base to the fair-weather wave base, but al ways in ox y gen ated wa ter (Haydukiewicz & Muszer, 2002). The rel a tive fre - quency of oc cur rence of par tic u lar tax o nom i cal groups in the Paprotnia Beds var ies de pend ing on li thol ogy. A dis - tinct de crease in the amount of ben thic fauna is no tice able in the greywacke in ter ca la tions, in the in ter val of mud- stones within the thin ben ton ite lay ers, and just be low the thick est ben ton ite layer in the se quence. Above this thick - est ben ton ite layer, a pro gres sive in crease in ben thic fauna is ob served. The ben thic or gan isms of this taphocoenosis could have ex isted in an en vi ron ment whereby there was a grad ual in crease in the in flux of terrigenous ma te rial and bot tom wa ter tur bu lence. Any rapid in flux of terrigenous ma te rial, com bined with in creas ing amounts of vol ca nic ash, would cause a short-term re gress of ben thic de vel op - ment (Haydukiewicz & Muszer, 2002).

Dis cus sion: Spec i mens of Zoophycos ichnospecies in the Papronia Beds are well pre served but un for tu nately in com - plete. The host rock type it self (mudstone) is very frag ile, and is a rea son why pre cise de scrip tion of the Zoophycos bur row sys tem is im pos si ble. Nev er the less, these Zoophy- cos clearly cor re spond to the con struc tional model of Gail- lard & Olivero (1993) and are very sim i lar to Tournaisian forms from Bel gium (Gaillard et al., 1999), de spite the sec - ond ary lamellae of the Tournaisian spec i mens not be ing vis i ble.

In com par i son to the Lower De vo nian spec i mens from Bolivia (Gaillard & Racheboeuf, 2006) the Paprotnia Zoophycos are sim i lar, though much smaller. The acute an - gle be tween the ma jor and mi nor lamella of the Paprotnia Bed spec i mens are sim i lar to those of Zoophycos from the Lower Car bon if er ous of Ja pan (Kotake, 1997). Fur ther - more, the dis tance be tween two sub se quent pri mary lamellae and the cen tral axis of the Paprotnia Zoophycos are very sim i lar to those from Zoophycos from the Mid dle Penn syl va nian of Nova Sco tia (McIlroy & Fal con-Lang, 2006). In our spec i mens, how ever, the cen tral axes are much smaller, pos si bly due to frag men ta tion. Pel lets within the spreite are not pres ent.

In con trast to the forms from the glaciomarine Tal- chir For ma tion in In dia (Bhattacharya & Bhattacharya, 2007) Zoophycos from the Paprotnia Beds are much larger, and this may be due to dif fer ences in cli mate and tem per a - ture of the lo cal sea wa ter.

It seems that the larger forms (our morphotype A1) are close to the ichnospecies Zoophycos villae Massalongo, 1855, and the smaller forms (our morphotype A2) are very sim i lar to ichnospecies Zoophycos brianteus Massalongo, 1855 (see Olivero, 2007).

Fig. 3. Frag ments of Zoophycos ichnospecies (morphotype A1) from the Paprotnia Beds show ing pri mary lamellae and sec ond ary lamellae. Ab bre vi a tions used in this fig ure are as fol lows: a (axil tun nel), mt (mar ginal tube), pl (pri mary lamellae), sl (secondry lamellae).

A, B – Pla nar sprite with high length-to-width ra tio (sam ple ING/P-1; A is lower bed ding sur face, B is up per bed ding sur face). C – Gently curved pla nar spreite with low length-to-width ra tio (sam ple ING/P-9). D – Pla nar spreite with mar ginal tubes (sam ple ING/P-8). E – Spi rally coiled struc ture with ax ial tun nel (sam ple ING/P-3). F – Pri mary and sec ond ary lamellae (sam ple ING/P-3).

CON CLU SIONS

The trace fos sil Zoophycos from the Paprotnia Beds are sim ple in form and cor re spond to the morphotype A de - scribed by Olivero (2003). Our spec i mens closely re sem - ble other Car bon if er ous forms of this ichnogenus (see Kotake, 1997; Gaillard et al., 1999; McIlroy & Fal con- Lang, 2006). We have iden ti fied two morphotypes that can be dif fer en ti ated by the pres ence of sec ond ary lamellae and size: these we term A1 for the larger type with pri - mary and sec ond ary lamellae, which re sem ble Zoophycos villae Massalongo, 1855; and A2 for the smaller type with only pri mary lamellae, which re sem ble Zoophycos brian- teus Massalongo, 1855. Both morphotypes oc cur in the same pack age of de pos its.

Both the lithological and palaeontological fea tures of the Paprotnia Beds in di cate grad ual en vi ron men tal chan- ges from off shore to on shore con di tions. Con cen tra tions of Zoophycos oc cur in the mudstones of the lower part of the sec tion, just be low the ho ri zon of organode tri tal lime - stones. Such a suc ces sion of sed i ments in di cates dis tinct shallowing of the wa ter body, from storm wave base to fair-weather wave base and to more ox y gen ated wa ters.

The pres ence of Zoophycos in such de pos its sup ports pre vi - ous sug ges tions from the ichnological lit er a ture that the trace-mak ing an i mal pre fers shal low-wa ter en vi ron ments

in the Early Car bon if er ous (e.g. Osgood & Szmuc, 1972;

Yurewicz, 1977; Miller & John son, 1981; Wu, 1982;

Bjerstedt, 1988; Ekdale & Ma son, 1988; Gaillard et al., 1999).

We also note that an cient and pres ent-day Zoophycos- group trace fos sils that are sim i lar to the Zoophycos trace fos sils from the Paprotnia Beds of ten seem to oc cur in sed - i ments that con tain peb ble lags, tuffs or vol ca nic ash (e.g.

Cham ber lain, 1975; Kotake, 1989, 1997; Löwemark et al., 2004; Löwemark et al., 2006; Gong et al., 2008; Hoffmann et al., 2009).

Thus, this re port on the oc cur rence of Zoophycos trace fos sils from the Paprotnia Beds of the Bardo Struc tural Unit, SW Sudetes, en riches our knowl edge of the Carbo- niferous of the Sudetes and opens the way for ge ol o gists to study these sed i ments from an ad di tional, and fresh, pers- pective.

Ac knowl edg ments

We thank Al fred Uchman and Pat rick Roycroft for crit i cal re views, for many valu able sug ges tions and for con struc tive re - marks which im proved the manu script. The study was sup ported by the in ter nal grants of Wroc³aw Uni ver sity, 2022/W/ING and 1017/S/ING.

Fig. 4. Frag ments of Zoophycos ichnospecies (morphotype A2) from the Paprotnia Beds show ing pri mary lamellae. Ab bre vi a tions used in this fig ure are as fol lows: a (axil tun nel), mt (mar ginal tube), pl (pri mary lamellae). A – Sam ple ING/P-28. B – Sam ple ING/P-25. C – Sam ple ING/P-23. D – Sam ple ING/P-24.

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