Lower Cambrian trace fossils from the Holy Cross Mountains, Poland

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Geological Quarterly, 2002, 46 (2): 135–146

Lower Cambrian trace fos sils from the Holy Cross Mountains, Poland

Stanisław ORŁOWSKI and Anna ŻYLIŃSKA

Orłowski S. and Żylińska A. (2002) — Lower Cam brian trace fos sils from the Holy Cross Moun tains, Po land. Geol. Quart., 46 (2):

135–146. Warszawa.

Ad di tional data on the Lower Cam brian ichnotaxa from the Holy Cross Moun tains are pre sented. Nine ichnotaxa are de scribed for the first time from the area, ad di tional de scrip tions of three known ichnotaxa are sup plied, and two ichnotaxa are shown to have ex tended strati graphic ranges. Most Lower Cam brian trace fos sils be long to the Cruziana ichnofacies. The high tax o nomic di ver sity of the Early Cam brian trace fos sil as sem blages from the Holy Cross Moun tains in com par i son to the trace fos sil as sem blages of the Mid- and Late Cam brian of the area is con firmed.

Stanisław Orłowski and Anna Żylińska, In sti tute of Ge ol ogy, Uni ver sity of War saw, Żwirki i Wigury 93, 02-089 Warszawa, Po land;

e-mail: zylinska@geo.uw.edu.pl (re ceived: July 7, 2001; ac cepted: De cem ber 5, 2001).

Key words: Holy Cross Moun tains, Lower Cam brian, trace fos sils.

INTRODUCTION

Trace fossils were first recognised in the Cambrian rocks of the Holy Cross Mountains over eighty years ago (Czarnocki, 1919). Descriptions of particular ichnotaxa appeared successively since the early 1960s (for details concerning bibliography see Orłowski and Żylińska, 1996).

Through out the years the col lec tion of Cam brian trace fos - sils, par tic u larly from Lower Cam brian strata, has con sid er ably in creased. In com par i son with the spec i mens pre sented pre vi - ously from the Lower Cam brian strata of the area (Orłowski, 1989, 1992), the ichnotaxa de scribed in this pa per are rep re - sented by few, in many cases only sin gle spec i mens.

This pa per de scribes ichnotaxa not known pre vi ously from the Cam brian rocks of the Holy Cross Moun tains, gives ad di - tional de scrip tions of known ichnotaxa, and de tails their ex - tended strati graphic ranges (Figs. 1 and 2). Par tic u lar at ten tion is given to trace fos sils com monly known from youn ger deep-wa ter en vi ron ments, those pre sented in this pa per as well as those de scribed ear lier by Orłowski (1989) and oth ers (Kowalski, 1978, 1987; Pacześna, 1985).

In relation to ichnotaxonomy we fol low here the approach outlined in the International Code of Zoological Nomenclature (International Commission …, 1999). Particular ichnotaxa are applied to traces of animal behaviour preserved within strata of different age. Such traces reflect various types of animal

behaviour within the sediment, and therefore their morphology often strongly depends on taphonomic processes (Uchman, 1998). As animals belonging to different biotaxa can produce quite similar traces (Seilacher, 1953; Bromley, 1990), ichnotaxa do not reflect phylogenetic relationships between the particular trace makers.

The spec i mens stud ied are housed in the In sti tute of Ge ol ogy of the Uni ver sity of War saw (ab bre vi ated as IGPUW).

LOCALITIES INVESTIGATED

Ociesęki For ma tion. Lo cal ities with the great est num ber of trace fos sils in clude the Sterczyna, Jaźwina and Igrzyczna hills, form ing the Ociesęki Range, as well as the Leśniakowa Dębina Hill and other out crops oc cur ring in the vi cin ity of Ociesęki in the cen tral part of the Klimontów Anticlinorium. At pres ent, the larg est ac ces si ble out crop in the area is a small quarry on the north ern side of Sterczyna Hill; other out crops of this for ma tion are temporary ones, though these typ i cally show a rich trace fos sil con tent. Fine-grained, yel low sand stones of the Ociesęki For ma tion with many bioturbated ho ri zons yield trilobites in di cat ing the Early Cam brian Holmia-Schmidtiellus Zone (Orłowski, 1985a, c). The trace fos sil as sem blage is equally rich both in non-arthropod and ar thro pod trace fos sils (Orłowski, 1989, 1992), among them Phycodes palmatum,

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Rhizocorallium jenense, Teichichnus rectus, Bergaueria perata, Cruziana rusoformis and Rusophycus crebrus. An other out crop of the same for ma tion, Malkowska Hill near Malkowice, oc curs in the east ern part of the Klimontów Anticlinorium, about 8 km south-westwards from Iwaniska in the Wygiełzów Range. Fine-grained sand stones yield few fos - sils, such as Schmidtiellus nodosus and Holmia sp., indicating the Holmia-Schmidtiellus Zone, as well as frag ments of Velumbrella czarnockii and Rotadiscus sp. (Orłowski, 1985a;

Masiak and Żylińska, 1994). Trace fos sils in clude Halopoa imbricata. A few spec i mens of Diplichnites isp. were also col - lected in this part of the Klimontów Anticlinorium near Rybnica in the Koprzywianka River val ley, about 2 km south of Nawodzice.

Kamieniec For ma tion. Trace fos sils within this for ma tion are gen er ally rare and rep re sented mainly by Bergaueria-type traces. These were col lected at Nowa Łagowica, south-east of Łagów and at Kamieniec, north of Konary, both in the east ern part of the Klimontów Anticlinorium, in strata rep re sent ing the Protolenus-Strenuaeva Zone (Orłowski, 1985a). Out crops in Kamieniec were al ready known to Jan Samsonowicz in the early 1930s. They oc cur near the dam on the Koprzywianka River.

OUTLINE OF STRATIGRAPHY

Trace fos sils have been col lected from var i ous lev els of the Lower Cam brian strata of the Holy Cross Moun tains, Po land (Fig. 2). Most of the trace fos sils pre sented in this pa per oc cur in the Lower Cam brian part of the Ociesęki For ma tion. The for ma tion con sists of fine-grained, thin- to me dium-bed ded hard sand stones, with siltstones and rare shale in ter ca la tions. Its

thick ness reaches the great est val ues in the cen tral part of the area (up to 1200 m), where it ranges up to the low er most Mid - dle Cam brian (Orłowski, 1985b, 1988). The age of the strata is de ter mined on the ba sis of trilobites. The lower part of the for - ma tion in cludes Holmia marginata, Kjerulfia orcina, Schmidtiellus panovi and Strenuella polonica, in di cat ing the Holmia-Schmidtiellus Zone, while the up per part yields Ellipsocephalus sanctacrucensis and Strenuaeva orlowinensis, in di cat ing the Protolenus- Strenuaeva Zone (Orłowski, 1985a).

Trace fos sils are much less fre quent in the Lower Cam brian Kamieniec For ma tion. This for ma tion is lim ited to the east ern part of the Holy Cross Moun tains area and is rep re sented by clayey and silty shales with fine-grained sand stone in ter ca la - tions. Its thick ness reaches about 600 m. Biostratigraphically the strata com prise the up per part of the Holmia-Schmidtiellus as well as the Protolenus-Strenuaeva zones, yield ing such trilobites as Holmia marginata, Kjerulfia orcina, Micmacca klimontowi as well as Protolenus (Protolenus) expectans, P.

(Latoucheia) glabellosus, Strenuaeva trifida and Serrodiscus primarius (Bednarczyk et al., 1965; Orłowski, 1985a).

SYSTEMATIC DESCRIPTIONS

CIRCULAR STRUCTURES

Bergaueria Prantl, 1945

R e m a r k s . — Cubichnial or domichnial trace fos sil, pro - duced prob a bly by sus pen sion-feeders (Fürsich, 1975).

Bergaueria Prantl is com monly re ferred to sea-anemones or

Fig. 1. Geological sketch-map of the Holy Cross Mountains (modified after Orłowski, 1992) with localities of the described trace fossils

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sea-pens (Prantl, 1945; Alpert, 1973; Pem ber ton et al., 1988;

Seilacher-Drexler and Seilacher, 1999).

Bergaueria perata Prantl, 1945 (Fig. 3a, b)

M a t e r i a l : Five spec i mens (IGPUW Tf/1/213, 240–243).

D e s c r i p t i o n . — Cy lin dri cal con vex hyporeliefs with rounded base bear ing a shal low cen tral de pres sion. Sin gle spec i mens, di am e ter from 15 to 45 mm. Walls oc ca sion ally with rings par al lel to bed ding planes.

R e m a r k s . — The spec i mens described oc cur in strata older than those de scribed pre vi ously from the area (Orłowski, 1989), thus ex tend ing the strati graphic range of the taxon in the Holy Cross Mts. The par al lel rings pres ent on sides of spec i - mens may re sult from com pac tion of sed i ment.

S t r a t i g r a p h i c d i s t r i b u t i o n . — From the late Pre cam brian (Crimes, 1987) to the Mio cene (Uchman, 1995).

O c c u r r e n c e i n t h e H o l y C r o s s M o u n t a i n s .

— Lower Cam brian of the Kamieniec For ma tion at Nowa Łagowica and Kamieniec (Orłowski, 1989); Lower Cam brian part of the Ociesęki For ma tion at Sterczyna Hill and Ociesęki;

Mid dle Cam brian Góry Pieprzowe For ma tion at Kobyla Hill (Orłowski and Żylińska, 1996); Up per Cam brian Wiśniówka and Klonówka for ma tions at the Wiśniówka Duża and

Wiśniówka Mała quar ries and Krzemionka Hill (Orłowski and Żylińska, 1996).

Bergaueria baltica Pacześna, 1996 (Fig. 3c, f)

M a t e r i a l : Three spec i mens (IGPUW Tf/1/244, 245, 275).

D e s c r i p t i o n . — Cy lin dri cal con vex hyporeliefs with con vex lower ter mi na tion and with reg u lar, broad, slightly con - vex, well marked mar gin. The bur rows are 10 to 20 mm long and their di am e ter var ies from 20 to 30 mm. The “lobes” on spec i men IGPUW Tf/1/275 (Fig. 3c) are ap par ent.

R e m a r k s . — This ichnospecies dif fers from B. perata in pos sess ing a broad, well vis i ble mar gin around the cen tral con - vex part. In this re spect the trace is iden ti cal to Bergaueria baltica Pacześna from north ern Po land and the Lublin re gion (Pacześna, 1996, pl. 1, figs. 2–4). The trace re sem bles spec i - mens of B. hemispherica fig ured by Crimes et al. (1977), which are, however, generally with out any or na men ta tion. Al though Bergaueria-type trace fos sils are typ i cally con sid ered as of sea-anemone or i gin, those with out a cen tral de pres sion (B.

hemispherica or B. baltica) might pos si bly be of other or i gin.

S t r a t i g r a p h i c d i s t r i b u t i o n . — Early and Mid - dle Cam brian (Pacześna, 1996).

Lower Cam brian trace fos sils from the Holy Cross Moun tains, Po land 137

Fig. 2. Stratigraphic ranges of the trace fossils in the Lower Cambrian strata of the Holy Cross Mountains, after Kowalski (1978, 1987), Orłowski (1985b, 1989, 1992) and Pacześna (1985)

Trace fossils discussed in this paper are marked with an asterisk; OSF — Osiek Sandstone Formation

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Fig. 3. a, b — Bergaueria perata Prantl, Lower Cam brian Kamieniec For ma tion: a — IGPUW Tf/1/213, Kamieniec, the ra dial struc tures around the trace fos sil are prep a ra tion marks; b — IGPUW Tf/1/240, Nowa Łagowica. c, f — Bergaueria baltica Pacześna: c — IGPUW Tf/1/275, Lower Cam brian part of the Ociesęki For ma tion, Sterczyna Hill; f — IGPUW Tf/1/244, Lower Cam brian Kamieniec For ma tion, Nowa Łagowica. d — ?Bifungites isp., IGPUW Tf/1/259, Lower Cam brian part of the Ociesęki For ma tion, Sterczyna Hill. e — Protovirgularia isp., IGPUW Tf/1/257, Lower Cam brian part of the Ociesęki For ma tion, Igrzyczna Hill. g — Treptichnus rectangularis Orłowski and Żylińska, IGPUW Tf/1/255, Lower Cam brian part of the Ociesęki For - ma tion, Sterczyna Hill. Scale bar 1 cm

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Fig. 4. a, b — Halopoa imbricata Torell, Lower Cam brian part of the Ociesęki For ma tion: a — IGPUW Tf/1/269, Igrzyczna Hill; b — IGPUW Tf/1/271, Ociesęki. c, d — Gordia arcuata Książkiewicz, Lower Cam brian part of the Ociesęki For ma tion: c — IGPUW Tf/1/262, Igrzyczna Hill; d — IGPUW Tf/1/261, Sterczyna Hill. Scale bar 1 cm

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Fig. 5. a — Helminthopsis abeli Książkiewicz, IGPUW Tf/1/77, Lower Cam brian part of the Ociesęki For ma tion, Leśniakowa Dębina Hill. b — Nereites missouriensis (Weller), IGPUW Tf/1/256, Lower Cam brian part of the Ociesęki For ma tion, Igrzyczna Hill. c, e — Helminthopsis hieroglyphica Wetzel and Bromley, Lower Cam brian part of the Ociesęki For ma tion, Sterczyna Hill: c — IGPUW Tf/1/248; e — IGPUW Tf/1/267. d — ?Asterophycus isp., IGPUW Tf/1/258, Lower Cam brian part of the Ociesęki For ma tion, Sterczyna Hill. f, g — ?Crossopodia isp., Lower Cam brian part of the Ociesęki For -

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Fig. 6. a — Rusophycus carbonarius (Dawson), IGPUW Tf/1/250 with five in di vid u als (ar rowed) and Gordia arcuata Książkiewicz, Lower Cam brian part of the Ociesęki For ma tion, Sterczyna Hill. b–d — Diplichnites isp., Lower Cam brian part of the Ociesęki For ma tion: b — IGPUW Tf/1/252, Sterczyna Hill; c — IGPUW Tf//253, Rybnica Quarry; d — IGPUW/Tf/1/254, Rybnica Quarry. Scale bar 1 cm

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— Lower Cam brian part of the Ociesęki For ma tion at Sterczyna Hill; Lower Cam brian of the Kamieniec For ma tion at Nowa Łagowica.

SIMPLE AND BRANCHED STRUCTURES

Protovirgularia Mc Coy, 1850

R e m a r k s . — The trace fos sil is re garded as of mol lus - can or i gin, and neoichnological ex per i ments have shown that it re sults from the ac tion of the cleft-foot of protobranchs and sca - pho pods (Seilacher and Seilacher-Drexler, 1994).

Protovirgularia isp.

(Fig. 3e)

M a t e r i a l : One spec i men (IGPUW Tf/1/257).

D e s c r i p t i o n . — Straight cy lin dri cal con vex hyporelief, nar row ing slightly at one end, with di ag o nal chev - ron ribs. The spec i men is 20 mm long, 3 mm wide and about 1.5 mm high.

R e m a r k s . — The trace fos sil is rather small and in com - pletely pre served, how ever, it re sem bles Protovirgularia-type trace fos sils with wide chev ron marks. It might there fore indicate the early ad ap ta tion of molluscs to cleft-foot sed i ment pen e tra tion. Trace fos sils rep re sent ing this taxon dis play strong mor pho log i cal vari a tion (Uchman, 1998).

S t r a t i g r a p h i c d i s t r i b u t i o n . — From the Early Or do vi cian (Fillion and Pickerill, 1990) to the Miocene (D’Alessandro, 1982).

— Lower Cam brian part of the Ociesęki For ma tion at Igrzyczna Hill.

Bifungites Desio, 1940

?Bifungites isp.

(Fig. 3d)

D e s c r i p t i o n . — Dumb bell-shaped con vex hyporelief in the form of a straight oval tube with a rounded and con vex struc ture at its end. The struc ture seems to extend into the sed i - ment. Spec i men partly de stroyed, about 40 mm long.

R e m a r k s . — Ac cord ing to Häntzschel (1975), the trace fos sil can be in ter preted as a cast of a pro tru sive, U-shaped, spreite-bearing bur row. How ever, closer ex am i na tions have shown that Bifungites lacks spreite struc tures (Gutschick and Lamborn, 1975; Miller, 1979). The com plete struc ture has a shape of an in verted pi (π), which is also partly shown by this spec i men. Its in com plete ness does not, how ever, al low a def i - nite as sign ment.

S t r a t i g r a p h i c d i s t r i b u t i o n . — From the Early Cam brian (Häntzschel, 1975) to the Early Car bon if er ous (Gutschick and Lamborn, 1975).

— Lower Cam brian part of the Ociesęki For ma tion at Sterczyna Hill.

Treptichnus Miller, 1889

R e m a r k s . — The trace rep re sents a 3-D bur row sys tem com posed of sim ple or zig zag, straight or curved seg ments as - so ci ated with ver ti cal or oblique tubes (Buatois and Mángano, 1993). It ranges from the Early Cam brian (Pacześna, 1989; this pa per) to the Eocene (Crimes et al., 1981).

Treptichnus rectangularis Orłowski and Żylińska, 1996 (Fig. 3g)

M a t e r i a l : Two spec i mens (IGPUW Tf/1/255, 256).

D e s c r i p t i o n . — Hor i zon tal sys tems of cy lin dri cal elon gated units, oval in cross-section, pre served as con vex hyporeliefs. Mi nor units formed as a rule on one side of sys tem.

An gle of branch ing vari able, close, how ever, to right an gles.

One of the sys tems con tains six units. Dis tal ends of units slightly thicker and di rected to ward the sed i ment sur face.

R e m a r k s . — In com par i son with the rich and well pre - served forms of this ichnospecies from the Up per Cam brian of the Holy Cross Moun tains (Orłowski and Żylińska, 1996), the cy lin dri cal units de scribed herein are lon ger and of smaller di - am e ter. No lon gi tu di nal ridges or traces of fae cal pel lets are pres ent. Priapulid worms, com mon Cam brian infauna, were pos si bly re spon si ble for the pro duc tion of these traces (Orłowski and Żylińska, 1996; Sören Jensen, pers. comm., 1999).

S t r a t i g r a p h i c r a n g e . — Early to Late Cam brian (Orłowski and Żylińska, 1996; this pa per).

— Lower Cam brian part of the Ociesęki For ma tion at Sterczyna Hill; Mid dle Cam brian Słowiec For ma tion at Konary (Orłowski and Żylińska, 1996); Upper Cam brian Wiśniówka and Klonówka for ma tions at the Wiśniówka Duża, Wiśniówka Mała and Wąworków quar ries and at Chabowe Doły (Orłowski and Żylińska, 1996).

Halopoa Torell, 1870

R e m a r k s . — Uchman (1998) showed that the trace is formed as a Teichichnus-type pat tern, how ever the par tic u lar probes were not ex actly ver ti cal and the spreite struc ture is thus not that reg u larly de vel oped as in Teichichnus.

Halopoa imbricata Torell, 1870 (Fig. 4a, b)

M a t e r i a l : Six spec i mens (IGPUW Tf/1/267–272).

D e s c r i p t i o n . — Cylindrical, full re lief con vex hyporeliefs, straight, un branched, cov ered by ir reg u lar, short wrin kles or lon ger striae form ing a plait struc ture. Largest spec - i men is 80 mm long and 10 mm wide.

R e m a r k s . — Ac cord ing to Martinsson (1965), the trace mak ers of this ichnospecies were mainly epipsammonts dig - ging long trails along a silt sur face. How ever, the trace fos sils were rather pro duced by de posit-feeding or gan isms, by prob - ing the sed i ment and push ing out the for merly re worked sed i - ment in older probes (Uchman, 1998), as is tes ti fied by de for - ma tion of laminae above the trace fos sils in cross-section

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views. The spec i mens pre sented here are rather reg u lar, with a dis tinct plait struc ture.

S t r a t i g r a p h i c d i s t r i b u t i o n . — From the Early Cam brian (Jensen, 1997) to the Mio cene (Crimes and Mc Call, 1995).

— Lower Cam brian part of the Ociesęki For ma tion at the Sterczyna, Igrzyczna and Malkowska hills, and at Ociesęki.

WINDING AND MEANDERING STRUCTURES

Nereites Mac Leay, 1839

R e m a r k s . — Ac cord ing to Rindsberg (1994) and Uchman (1995), Neonereites Seilacher, along with Scalarituba Weller, Palaeohelminthoida Ruchholz, and Helminthoida Schafhäutl are con sid ered as syn onyms of Nereites Mac Leay.

Nereites missouriensis (Weller, 1899) (Fig. 5b)

D e s c r i p t i o n . — Ir reg u lar, slightly curved chain of oval con cave marks, about 40 mm long and 3 mm wide. Con - cave epirelief.

R e m a r k s . — This ichnospecies is in ter preted as a postdepositional, in ter nal bur row de vel oped in the course of graz ing within the sed i ment. Neonereites isp. from the Lower Cam brian part of the Ociesęki Sand stone For ma tion near Chęciny (Kowalski, 1987) is most prob a bly syn on y mous.

S t r a t i g r a p h i c d i s t r i b u t i o n . — From the late Pre cam brian (Crimes, 1987) to the Mio cene (Uchman, 1995).

— Lower Cam brian part of the Ociesęki For ma tion at Igrzyczna Hill.

Gordia Emmons, 1844

R e m a r k s . — Fillion and Pickerill (1990) and Pickerill and Peel (1991) have re cently dis cussed the ichnotaxonomy of Gordia Emmons. The trace fos sil is typ i cally un branched, wind ing or ir reg u larly me an der ing, hor i zon tal.

Gordia arcuata Książkiewicz, 1977 (Figs. 4c, d, 6a)

M a t e r i a l : Four spec i mens (IGPUW Tf/1/250, 261–263).

D e s c r i p t i o n . — Slen der, smooth, of even di am e ter, curved, thread-like trace fos sils, less than 1 mm thick and up to 50 mm long. Sin gle or cov er ing sur face with ir reg u lar me an - ders. Con vex hyporeliefs.

S t r a t i g r a p h i c d i s t r i b u t i o n . — From the lat est Pro tero zoic to the Oligocene (Wal ter et al., 1989).

— Lower Cam brian part of the Ociesęki For ma tion of the Sterczyna and Igrzyczna hills; as Gordia isp. in the Upper Cam brian Wiśniówka and Klonówka for ma tions at the Wiśniówka Duża and Wiśniówka Mała quar ries and at Chabowe Doły (Orłowski and Żylińska, 1996).

Helminthopsis Heer, 1887

R e m a r k s . — The ichnogenus Helminthopsis Heer was crit i cally ex am ined by Han and Pickerill (1995), who used the sta tis ti cal method of “Fou rier Trans fer Anal y sis” to dis tin guish the var i ous ichnospecies. An in de pend ent eval u a tion of the ichnogenus based on Heer’s type ma te rial (Wetzel and Bromley, 1996) re vealed the va lid ity of three ichnospecies out of the ear lier de scribed ones. Re sults ob tained from FT Anal y - sis (Han and Pickerill, 1995) were criti cised as be ing based on wrongly pro duced func tions by Wetzel and Bromley (1996, p.

18), who pointed out that Heer’s ma te rial can be in cluded in Spirocosmoraphe Seilacher and Taphrhelminthopsis Sacco [=

Scolicia strozzii (Savi and Meneghini) — see Uchman, 1995, 1998]. How ever, in or der to re tain Helminthopsis as an ichnogenus, they se lected an unillustrated spec i men from Heer’s col lec tion, de ter mined by him as Helminthopsis hieroglyphica, as the holotype of the type species.

Helminthopsis is a fa cies-crossing trace fos sil, quite com mon in flysch de pos its. It can be pre served in dif fer ent modes, as a con - vex hyporelief, semirelief or con cave epirelief (Wetzel and Bromley, 1996). Polychaete annelids were prob a bly re spon si - ble for the pro duc tion of such trace fos sils, how ever, priapulids are more likely to have been pro duc ers in Cam brian sed i ments (Książkiewicz, 1977).

S t r a t i g r a p h i c d i s t r i b u t i o n . — Cam brian (Crimes, 1987) to Re cent (Wetzel, 1983).

Helminthopsis abeli Książkiewicz, 1977 (Fig. 5a)

M a t e r i a l : Three spec i mens (IGPUW Tf/1/77, 246, 249).

D e s c r i p t i o n . — Trace fos sils in the form of cy lin dri - cal, partly empty hypichnial semireliefs, with some parts col - lapsed, with smooth walls, with me an ders ir reg u lar and vari - able in shape, typ i cally with horse shoe-like turns, oc ca sion ally with some parts of the trace fos sil straight within the wide open me an ders. The di am e ter of the trace fos sil is 2 to 3 mm, length ex ceeds 100 mm. In col lapsed seg ments a tube wall, about 0.2 mm thick, is vis i ble.

R e m a r k s . — Wetzel and Bromley (1996) have shown the va lid ity of this ichnotaxon. The tube wall is most prob a bly the outer zone of a two-zoned bur row fill, and can be con sid - ered as a man tle (sensu Bromley, 1990, and Keighley and Pickerill, 1994), thus not a true bur row wall. It was prob a bly made by a de posit-feeding or gan ism dur ing its pas sage through the sub strate.

— Lower Cam brian part of the Ociesęki For ma tion at Leśniakowa Dębina Hill and at Ociesęki.

Helminthopsis hieroglyphica Wetzel and Bromley, 1996 (Fig. 5c, e)

M a t e r i a l : Three rock slabs with about twenty spec i mens (IGPUW Tf/1/248, 249, 267).

D e s c r i p t i o n . — Sim ple, elon gate, non-branched semireliefs with ir reg u lar low-amplitude wind ings, of small di - am e ter. Wind ings are com posed of low-angle kinks and

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straight sec tions giv ing the trace an ap pear ance of a box-shaped fold. Trace fos sils 2–3 mm in di am e ter and up to 30 mm long.

BILOBATE TRACES

Crossopodia M’Coy, 1851

?Crossopodia isp.

(Fig. 5f, g)

M a t e r i a l : Two spec i mens (IGPUW Tf/1/265, 266).

D e s c r i p t i o n . — Straight con vex endoreliefs con sist - ing of two rows of branches of dense ribs, di rected di ag o nally to a broad me dian fur row sep a rat ing the rows. The spec i mens are about 40 mm long and 15 mm broad.

R e m a r k s . — The ichnotaxonomic re la tion ship of this trace fos sil is rather un clear (Ma ples and Suttner, 1990), how - ever, the most char ac ter is tic fea ture, which dif fers this ma te rial from other ichnogenera, are the ribs sep a rated from each other by a broad me dian fur row. The ribs orig i nated prob a bly as a re - sult of back fill ac tiv i ties, thus re sem bling Psammichnites Torell.

S t r a t i g r a p h i c d i s t r i b u t i o n . — Early Cam brian (Crimes, 1992) to Car bon if er ous (Häntzschel, 1975; Ma ples and Suttner, 1990).

— Lower Cam brian part of the Ociesęki For ma tion of the Sterczyna and Leśniakowa Dębina hills.

RADIAL STRUCTURES

Asterophycus Lesqueraux, 1876

?Asterophycus isp.

(Fig. 5d)

D e s c r i p t i o n . — Ra dial con vex epirelief, about 30 mm in di am e ter, oval, con sist ing of seven ribs ra di at ing reg u larly from cen tral area, be ing the ter mi na tion of a ver ti cal shaft. Each rib is from 10 to 15 mm long, about 4 mm broad, reg u lar, semioval in cross-section, outer ter mi na tions plunged into the sed i ment.

R e m a r k s . — The sys tem atic po si tion of this trace fos sil is not clear. Ichnogenera in ter preted as ribs ra di at ing around a ver ti cal shaft in clude Asterichnus Bandel, Asterophycus Lesqueraux, Asterosoma von Otto, Stelloglyphus Vialov and Volkichnium Pfeif fer (Häntzschel, 1975). Asterichnus sp. and Volkichnium volki Pfeif fer were de scribed from the Upper Cam brian of the area (Orłowski and Żylińska, 1996). In com - par i son to the ana lysed spec i men the cen tral area in those taxa is less pro nounced and the ribs ra di ate in a more ran dom pat - tern. Stelloglyphus Vialov is char ac ter ised by densely spaced and over lap ping ribs, petaloid in out line (Häntzschel, 1975).

The ana lysed spec i men com prises seven reg u lar, evenly broad smooth ribs ra di at ing from a ver ti cal shaft, ter mi nat ing in the cen tral area. It is thus clos est to Asterophycus Lesqueraux, the

num ber of ribs, how ever, fol lowed by the lack of lon gi tu di nal wrin kles within them and the rather small di am e ter of ribs in cross-section, dis tin guish the spec i men stud ied from the known spec i mens of Asterophycus, e.g. A. coxii (Häntzschel, 1975) or Asterophycus sp. (Bandel, 1967). Asterophycus was prob a bly made by worm-like an i mals or crus ta ceans (Bandel, 1967).

S t r a t i g r a p h i c d i s t r i b u t i o n . — ?Early Cam - brian (this pa per) and Early Car bon if er ous (Häntzschel, 1975).

TRACES OF ARTHROPOD ORIGIN

Rusophycus Hall, 1852

R e m a r k s . — This bilobed trace is typ i cally in ter preted as a rest ing trace of trilobites and re lated ar thro pods.

Rusophycus carbonarius (Dawson, 1864) (Fig. 6a)

M a t e r i a l : One slab with five in di vid u als (IGPUW Tf/1/250).

D e s c r i p t i o n . — Small, oval cof fee-bean-shaped con - vex hyporeliefs, con sist ing of two slightly con vex lobes, sep a - rated by a straight, nar row fur row vis i ble on the whole length of the trace. Very fine striations, not ex tend ing be yond lobes, are vis i ble only on one spec i men. Length of traces 3 to 5 mm, width 2 to 3 mm.

R e m a r k s . — Small cof fee-bean-shaped trace fos sils, of ar thro pod or i gin, are widely known from Cam brian and youn - ger strata. Ac cord ing to Keighley and Pickerill (1996), cof - fee-bean-shaped trace fos sils with out trans verse striations can be con sid ered as taphonomic vari ants of stri ate spec i mens (typ - i cally R. carbonarius) and are here re ferred to as such.

S t r a t i g r a p h i c d i s t r i b u t i o n . — From the Early Cam brian to the Tri as sic (Keighley and Pickerill, 1996).

Diplichnites Dawson, 1873 Diplichnites isp.

(Fig. 6b–d)

M a t e r i a l : Three spec i mens (IGPUW Tf/1/252–254).

D e s c r i p t i o n . — Con cave epireliefs, about 100 mm long, con sist ing of two par al lel rows of nar row de pres sions slightly curved in the same di rec tion. Each row gen er ally con - tains 10 de pres sions, about 10 mm long. In one case (IGPUW Tf/1/254, Fig. 6d) the de pres sions are filled with finer sed i - ment. The dis tance be tween the two rows is about 15 mm.

R e m a r k s . — The trace fos sils are in ter preted as marks of ar thro pod limbs cut ting through the sub strate along with its de for ma tions caused by the an i mal. The most char ac ter is tic fea ture of the trace fos sils is a short dis tance be tween the rows.

This sug gests that the de pres sions could be pro duced by the basal parts of limbs of a tri lo bite or an other ar thro pod. Sim i lar

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trace fos sils were re cently de scribed from the Yu’anshan For - ma tion in Chengjiang, China (Zhu, 1997).

S t r a t i g r a p h i c d i s t r i b u t i o n . — Com mon in Palaeozoic strata, but in a broad con cept of the ichnogenus re - cog nised also in the Tri as sic (Machalski and Machalska, 1994 and ref er ences therein).

— Lower Cam brian part of the Ociesęki For ma tion at Sterczyna Hill and Rybnica Quarry; Upper Cam brian Wiśniówka For ma tion at the Wiśniówka Duża Quarry (Orłowski, 1992).

DISCUSSION

Early Cam brian trace fos sil as sem blages from the Holy Cross Moun tains show a con sid er ably high tax o nomic di ver - sity (Kowalski, 1978, 1987; Pacześna, 1985; Orłowski, 1989, 1992) in com par i son to those char ac ter is ing the Mid- and Upper Cam brian of the area (Orłowski, 1992; Orłowski and Żylińska, 1996). They in clude sim ple trace fos sils, e.g. sim ple and branched struc tures (Planolites, Gordia, Treptichnus) and cy lin dri cal struc tures (Bergaueria), as well as much com plex forms, e.g. me an der ing traces (Helminthopsis, Cochlichnus), ver ti cal spreite forms (Syringomorpha), ra dial struc tures (Asterophycus) and net works (Paleodictyon), re veal ing di verse sed i ment pen e tra tion pat terns (Kowalski, 1978, 1987;

Pacześna, 1985; Orłowski, 1989, 1992). Most of the traces rep - re sent the Cruziana ichnofacies (Seilacher, 1967; Frey and Seilacher, 1980), how ever there are also individual rep re sen ta - tives of the Nereites ichnofacies, e.g. Helminthopsis or Paleodictyon. Protovirgularia oc curs both in the Cruziana and Nereites ichnofacies. The Ociesęki For ma tion, which yields nu mer ous rep re sen ta tives of the two ichnofacies, com prises fine-grained, strongly bioturbated in ner shelf sand stones. The Nereites ichnofacies in turn is char ac ter is tic of en vi ron ments with sed i men ta tion mostly from sus pen sion (Seilacher, 1967) on an outer shelf and slope. The co-occurrence of trace fos sils of these two ichnofacies within de pos its of the in ner-shelf con - firms that post-Cambrian deeper wa ter sed i ment pen e tra tion pat terns ap peared in shal low ma rine en vi ron ments of the Early Cam brian (Hofmann and Patel, 1989).

Rep re sen ta tives of the gen era Helminthopsis, Paleodictyon, Gordia and Protovirgularia have also been re -

cog nised in the Ter tiary Carpathian flysch (Książkiewicz, 1977; Uchman, 1998). Ob vi ously, the pro duc ers of Cam brian and Ter tiary traces did not rep re sent the same ge nus or spe cies, but also most prob a bly might have be longed to dif fer ent or ders.

How ever, they had sim i lar be hav ioural pat terns, which de vel - oped in Early Cam brian or even Pre cam brian shal low-water en vi ron ments (Crimes and An der son, 1985; Hofmann and Patel, 1989; Crimes and Fedonkin, 1994; Orłowski and Żylińska, 1996). The ap pear ance of such pat terns is ex pressed by a high tax o nomic di ver sity of trace fos sils, to be fol lowed by its de crease at the be gin ning of the Mid dle Cam brian (Crimes, 1992). The colo nis ation of deep-sea en vi ron ments by trace-making or gan isms was de layed un til the Early Or do vi - cian (Crimes et al., 1992; Crimes and Fedonkin, 1994), re sult - ing like wise as an in crease in ichnogeneric di ver sity.

The ex tremely low tax o nomic di ver sity and con tent of trace fos sils within the Czarna Shale For ma tion (Orłowski, 1989, fig.

2) might ei ther be a re sult of sed i men ta tion in ox y gen-depleted en vi ron ments or a con se quence of poor fos sili sa tion po ten tial of trace fos sils in shaly fa cies.

CONCLUSIONS

1. Nine ichnotaxa sup ple ment the list of Lower Cam brian trace fos sils from the Holy Cross Moun tains. These in clude rep re sen ta tives of the gen era ?Asterophycus, Bergaueria,

?Bifungites, ?Crossopodia, Halopoa, Helminthopsis, Protovir - gularia and Rusophycus.

2. Most Lower Cam brian trace fos sils from the Holy Cross Moun tains rep re sent the Cruziana ichnofacies; they are, how - ever, ac com pa nied by sin gu lar spec i mens of ichnotaxa from the Nereites ichnofacies.

3. In com par i son with the Mid- and Late Cam brian trace fos sil as sem blages from the Holy Cross Moun tains, those from the Lower Cam brian of the area are char ac ter ised by a higher ichnotaxonomic di ver sity.

Ac knowl edge ments. The au thors wish to thank Al fred Uchman and an anon y mous re viewer for their re marks and com ments on the pa per. Sören Jensen gave help ful hints to an ear lier draft of the pa per.

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