Alfred UCHMAN
Institute o f Geological Sciences, Jagiellonian University, Oleandry 2a: 30-063 Kraków, Poland Uchman, A., 1996. Taxonomy and ethology of flysch trace fossils: revision of the Marian Książkiewicz collection and studies of complimentary material. Ann. Soc. Geol. Polon., 68: 105-218.
A bstract: Ichnotaxonomy o f 1,840 of flysch trace fossil specimens from the Książkiewicz collection is revised.
The specimens derive from diverse Tithonian-Miocene flysch deposits of the Polish Carpathians. Their ichno
taxonomy, based on morphology, was published by Książkiewicz in 1997. In the revision presented in this publication, the ichnotaxonomic subdivisions are based on type o f behaviour represented by the considered trace fossil. As a result, several diagnoses are changed and several ichnospecies ascribed to other ichnogenera.
Ichnogenera Hormosiroidea, Saerichnites, IParahaentzschelinia, Halopoa, Nereites, Beaconites, Cladichnus, Protovirgularia, Ubinia and Oscillorhaphe are used the first time in the Polish Carpathians. The ichnogenera Pararusophycus, Rhabdoglyphus, Fucusopsis, Traucumichnis, Sabularia, Granularia, Bostricophyton, Halyme- nidium, Buthotrephis, Tubuliclmium, Tuberculichnus, Helminthoida, Muensteria, Keckia, and Taphrhelminthoida are not recommended for further use. The new ichnogenus Belocosmorhaphe n. igen. and the new ichnospecies name Cosmorhaphe carpathica nom. nov. are proposed. Ichnotaxa Trichichnus linearis Frey, Imponoglyphus torquendus Vialov, Palaeophycus tubularis Hall, Chondrites Irecurvus (Brongniart) and Arenituba isp. are distinguished for the first time in the Carpathian flysch on the base o f Książkiewicz material.
A bstrakt: 1840 okazów fliszowych skamieniałości śladowych z kolekcji Książkiewicza poddano ichnotaksono- micznej rewizji. Okazy te pochodzą z różnych tytońsko-mioceńskich utworów fliszowych Karpat polskich. Ich ichnotaksonomia, oparta na morfologii, była przedstawiona przez Książkiewicza w 1977 roku. W rewizji przedsta
wionej w niniejszej publikacji, wydzielenia ichnotaksonomiczne oparto na typie behawioru reprezentowanego przez daną skamieniałość śladową. W rezultacie, zmieniono szereg diagnoz. Wiele ichnogatunków zaliczono do innych niż dotąd ichnorodzajów. Ichnorodzaje Hormosiroidea, Saerichnites, IParahaentzschelinia, Halopoa, Nereites, Beaconites, Cladichnus, Protovirgularia, Ubinia i Oscillorhaphe użyto po raz pierwszy w Karpatch polskich. Ichnorodzaje Pararusophycus, Rhabdoglyphus, Fucusopsis, Traucumichnis, Sabularia, Granularia, Bostricophyton, Halymenidium, Buthotrephis, Tubulichnium, Tuberculichnus, Helminthoida, Muensteria, Keckia i Taphrhelminthoida nie są rekomendowane do dalszego używania. Zaproponowano nowy ichnorodzaj Belo
cosmorhaphe n. igen. i nową nazwę ichnogatunkową Cosmorhaphe carpathica nom. nov. Ichnotaksony Trichi
chnus linearis Frey, Imponoglyphus torquendus Vialov, Palaeophycus tubularis Hall, Chondrites 1 recurvus (Brongniart), Arenituba isp. są wyróżnione po raz pierwszy we fliszu karpackim, w oparciu o materiał z kolekcji Książkiewicza.
Key words: Ichotaxonomy, trace fossils, flysch, Cretaceous, Paleogene, Carpathians.
Manuscript received 2 April 1996, accepted 27 July 1998.
C ontents
Abstract... 105 Halopoa T o r e ll...
Aim, material, and m e th o d s... 106 Hormosiroidea S c h a ffe r...
Classification and principles of ichnotaxonomy... .. 107 Strobilorhaphe Ksi^zkiewicz...
Systematic p a r t... . . 107 Imponoglyphus Vialov...
Circular and elliptical structures... . . 107 Planolites N icholson...
Mammillichnis C ham berlain... 107 Palaeophvcus H all...
Bergaueria P ra n tl... . . 109 Chondrites Sternberg...
Simple and branched structures... .. 109 Trichichnus F re y ...
Arthrophvcus H a ll... 109 Ophiomorpha Lundgren...
112
117 118 120 120
121
121 125 125
106
A. UCHMANThalassinoides E hrenberg... ...128
Spongeliomorpha de Saporta...129
Phycodes Richter...130
Saerichnites B illin g s... ...130
?Parahaentzschelinia Chamberlain...131
Anemonichnus Chamberlain & C l a r k ...131
Radial structures... ...132
Lorenzinia G abelli... ...132
CapodistriaV i a l o v ... ...139
Glockerichnus P ickerill...141
Arenituba Stanley & P ickerill...145
Fascichnium Książkie w ic z ...146
Gyrophyllites G lo ck er...146
Spreite structures... ...148
Zoophycos M assalongo...148
Phycosiphon Fischer-O oster...148
Lophoctenium R ichter...149
Rhizocorallium Z e n k e r ...149
Winding and meandering stru ctu re s... ...149
Nereites M acL eay... ...149
Scolicia de Q uatrefages...152
Naviculichnium K siążkiew icz...159
Taenidium H eer...160
Beaconites V ialo v ... ...161
Cl'adichnus D'Alessandro & Bromley...162
Protovirgularict M cC oy...162
Oniscoidichnus B ra d y ...168
Gordia Emmons...168
Cosmorhaphe F u c h s ... ...168
Cochlichnus H itchcock...173
Helicorhaphe Książkiewicz... ...173
Helicolitlnis Azpeitia M o ro s ...174
Helminthorhaphe Seilacher... ...175
Helminthopsis H ee r... ...176
Spiral structures... ...179
“Spirophycus” H antzschel... ...179
Spirorhaphe Fuchs... 181
“Rotundusichnium” Plicka... 182
Branched winding and meandering traces... 182
Belorhaphe F u c h s ... 182
Belocosmorhaphe n. igen... 183
Paleomeandron Peruzzi... 184
Acanthorhaphe K si^zkiew icz... 186
Desmograpton Fuchs... 186
Urohelminthoida Sacco... 187
Ubinia G rossgeim ... 189
OsciHorhaphe Seilacher... 189
Protopaleodictyon Ksi^zkiewicz... 190
N etw orks... 192
Megagrapton Ksi^zkiewicz... 192
Paleodictyon M eneghini... 195
Ichno(taxa) not recommended for further u s e ... 195
Pararusophycus Ksi^zkiewicz... 195
Rhabdoglyphus Vassoevich... 195
Fncusopsis Palibin in V assoevich... 196
Traucumichnis Ksiazkiewicz... 196
Sabularia Ksiazkiewicz... 196
Granularia P o m e l... 196
Bostricophyton S quinabol... 196
Halymenites Sternberg and Halymenidium Schimper 196 Buthotrephis Hall... 198
Tubulichnium Ksiazkiewicz... 198
Tuberculichnus K siazkiew icz... 198
Helminthoida Schafhaut!... 199
Muensteria Sternberg... 199
Keckia G lo c k e r... 199
Taphrhelnnnthoida K siazkiew icz... 199
M iscellanea... 200
Ichnotaxonomy of the Ksiazkiewicz collection after the rev isio n ... 202
Acknowledgments... 205
References... 205
AIM, MATERIAL, AND METHODS
Im precisely defined ichnotaxa are still the w eak point o f ichnology. This problem still negatively influences various applications o f trace fossils and is a hinderence in their study. The problem m ay be am eliorated by ichnotaxonom ic revisions using the application o f clearly defined principles.
Such revision o f the K siazkiew icz collection is the main aim o f this paper.
The K siazkiew icz collection, housed in the Institute o f G eological Sciences o f the Jagiellonian U niversity, Krakow (acronym U J TF), is one o f the largest flysch trace fossil col
lections in the world. It contains 1,840 specim ens including 57 ichnogenera and 147 ichnospecies described by Ksi^z- kiew icz (1958, 1960, 1961, 1968, 1970, 1977) from alm ost 220 localities in the Polish C arpathians, from the T ith o n ian - M iocene flysch form ations. A lm ost all the specim ens w ere labelled. Each full label includes num ber o f specim en, nam e o f locality and lithostratigraphic unit, and ichnotaxonom ic detennination. A num ber o f the labels appeared to be not readable, having been dam aged because o f im proper hous
ing. For this reason, a lot o f im portant inform ation from la
bels was lost. As a result o f the inventory w ork connected
w ith the present revision, the first catalogue o f the speci
m ens o f the collection was prepared, w hich com prises the inform ation from the labels and place o f housing. The col
lection contains a few ichnotaxa, w hich w ere neither de
scribed nor labelled, for instance T richichnus and Im pono- glyphus, w hich are included in this study. A bout a dozen specim ens are lost.
In order to resolve certain ichnotaxonom ic problem s, com plem entary m aterial was studied. This derives from the author’s collection, the collections in the N aturhistorisches H ofm useum in V ienna, the Sternberg collection in the N a tional M useum in Prague, the H eer collection in the G eo
logical Institute o f the ETH, Zurich, and the F ischer-O oster collection in the N atural H istory M useum in Bern.
Som e specim ens from the K siazkiew icz collection un
derw ent additional preparation, and/or w ere cut and studied in polished slabs after treatm ent w ith the “ B ushinsky oil technique” (B rom ley, 1981).
The first problem is th e definition o f trace fossils. I fol
low the idea that trace fossils are the sedim entary expres
sions o f fossil behaviour (Seilacher, 1986). Thus, the m ost im portant questions o f ichnotaxonom y are w hat behaviour is represented by an exam in ed trace fossil and how to clearly define it, not only th e exact description o f the ap
pearance o f the exam ined trace fossil. M orphology o f trace fossils can be strongly d ep en d an t on taphonom ic processes.
Especially, trace fossils preserv ed as sem i-reliefs are incom plete bedding-plane expressions o f com m only m uch m ore com plicated three-dim ensional burrow s. The num erous dif
ficulties encountered in reconstruction o f com plete trace fossils and in understanding o f the bahaviour they represent cannot be argum ents th at influence principles o f ichnotax
onom y.
The other problem is th e rank o f ichnotaxonom ic divi
sions, especially the hierarchy o f features used for diagnoses o f ichnogenera and ichnospecies. I tried to follow the idea o f Fiirsich (1974a), according to w hich im portant m orphologi
cal features (so-called significant features) diagnostic o f ich
nogenera are these related to distinct behaviour are diagnos
tic o f ichnogenera. O ther features related to m inor variation o f tracem aker behaviour and to som e preservational features are regarded as diagnostic o f ichnospecies (so-called acces
sory features). O ther features o f third-order rank, reflecting m ainly preservational aspects, m ay be used inform ally at the subichnospecies level as variations (U chm an, 1995). D ivi
sion o f this level is not forbidden by the 1985 International Code o f Z oological N o m en clatu re (R indsberg, 1990). Ex
am ples o f their use are given by Ekdale & Lew is (1991b) for Diplocraterion. The m ain prin cip le in the presented ichno
taxonom y is th e application o f ichnogeneric nam es only for trace fossils representing d istinctly different fossil behav
iour. O nly ichnogenera d iagnosed in this w ay m ay be used for further palaeoecological interpretations and com pared w ith other ichnocom m unities. A pplication o f accessory or low er-rank features at the ichnogeneric level w ould give a false picture o f behavioural diversity.
In order to recognize the fossil behaviour represented by som e trace fossils, their careful ethological interpretation is necessary. Such interpretation w as applied for instance to passively filled Palaeophycus versus actively filled Plano- lites (Pem berton & Frey, 1982). P urely m orphological clas
sification, w ithout references to any ethological m odel and w ithout taphonom ic interpretation, seem s to be one o f the dead-ends o f ichnotaxonom y.
A nother problem o f ichnotaxonom y is classification o f trace fossils above the ichnogeneric rank (cf. Brom ley, 1996). K si^zkiew icz (1977) introduced classification o f flysch trace fossils based solely on m orphological criteria.
He distinguished ten m orphological groups, nam ely: (1) cir
cular and elliptical, (2) sim ple, (3) branched, (4) rosetted, (5) spreite, (6) w inding, (7) spiral, (8), m eandering, (9)
(e.g., Arthrophycus, Halymenidium=Spongeliomorphd). For this reason the sim ple and branched structures are u nited in one group in this paper. In the K siqzkiew icz, classification genetically close ichnotaxa, such as Scolicia and Taphr- helminthoida (=Scolicia in this paper), belong to different groups. Thus, a classification based on rigid criteria is not satisfactory.
SYSTEMATIC PART
In order to shorten the system atic part, a num ber o f ich
notaxa descriptions provided by K siazkiew icz (1977) that have not been changed or have been supplem ented only by new diagnoses, are om itted.
CIRCULAR AND ELLIPTICAL STRUCTURES
Mammillichnis C ham berlain 1971 a
Emended diagnosis: H ypichnial m ound w ith a convex or concave apex and a teat-like tu b ercu le at the centre o f the apex.
Mammillichnis aggeris C ham berlain 1971a Figs. 1-2
* 1971a Mammillichnis aggeris n. ichnog. and sp..- Chamberlain, 238, pi. 30, figs. 6-7.
1977 Mammillichnis aggeris Chamberlain - Chamberlain, 14, figs. 2b’, 7G.
1977 Mammillichnis aggeris Chamberlain - Stanley el al., 267, fig. 18.
v 1977 Mammillichnis aggeris Chamberlain - Ksiazkiewicz, 53, text-fig. 5a-b, pi. 1, figs. 1-2.
1979 Mammillichnis aggeris Chamberlain - Chamberlain, 17, fig. 3.
1981 Mammillichnis aggeris Chamberlain - Crimes el al., 975, pi. 4, figs. 9-10 [specimen from fig. 10 also in Pemberton el al., 1988. fig. 11С].
1985 Mammillichnis Chamberlain - Eager el al., 140, pi. ID.
?non 1986 Mammillichnis ichnosp. - Paczesna, 32, pi. 5, fig. 2.
1993 Mammillichnis aggeris Chamberlain - Alexandrescu el al.; 10, figs. 2-5.
?non 1996 Mammillichnis ichnosp. - Paczesna, 57, pi. 7, figs. 1-2.
? 1997 1 Mammillichnis ichnosp. Zagora, 355, fig. 5.3 Diagnosis: As for ichnogenus by its monotypy.
Material: 9 specimens (UJ TF 117-118, 122, 1464, 1470, 1762- 1764, 2501).
Description: Small, circular or elliptical hypichnial mounds with apical depression and tubercule at the centre o f the depression.
Only one specimen does not display the apical depression and the tubercule is located at the convex top o f the mound. The mounds are 8-10 mm in diameter and 3-4 mm in height.
Remarks: The specim ens w ith th e apical depressions re
sem ble Laevicyclus Q uenstedt. S om e o f them (UJ TF 118, 1470) had been labelled by K siazkiew icz as Laevicyclus
108
A. UCHMANFig. 1. Mammillichnis aggeris Chamberlain, soles o f turbiditic sandstone beds. A. UJ TF 1649, Krosno beds (Oligocene), Stonne Gory range near Zahiz. B. UJ TF 2501, Krosno beds (Oligocene), Wujskie. Scales in mm
ph ilip p i (W urm ), but then th e labels w ere changed to M. a g geris. The trace fossils d escribed by W urm (1911) are dis
tinctly larger and display a double ring.
O rigin o f M. aggeris is not clear. A ccording to C ham berlain (1971a), this is a resting or hiding trace, a fossil egg case (a body fossil), or th e u pper term ination o f a burrow.
Later, C ham berlain (1977) suggested that M. aggeris is a resting or dw elling trace pro d u ced by a ?"w orm " and th at it is probably a preservational v ariant o f Alcyonidiopsis p h a r- m aceus R ichter & Richter. H ow ever, it is difficult to find re
lationships betw een these tw o ichnotaxa. O nly the granular surface in som e specim ens o f M. aggeris (especially in the C ham berlain’s m aterial) m ay be related to the pelletal struc
ture o f Alcyonidiopsis. H ow ever, the m orphology o f the apex in M. aggeris has no relationship to hom ogenously fae
cal-stuffed trace fossil o f A lcyonidiopsis. K si^zkiew icz (1977) suggested that M am m illichnis is an actinian burrow
ORIGIN OF MAMMILICHNIS
IMI 1 ! I i Mil MM
r
IIII Mill II111 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 [ 1 1 1 1 1 i 1 1
MM MM IMI 1 M i
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1111 IMI 1111111111111111111111111 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1
DEPRESSION IN THE SEA-FLOOR IN CROSS-SECTION
RESULTED TRACE FOSSILS AFTER CASTING IN CROSS-SECTION
Fig. 2. General model (top) and morphological variations (be
low) of Mammillichnis. The morphological variants depend on the shape o f the depression in the sea floor
because o f supposed radial sym m etry in C h am b erlain ’s (1971a) fig. 7J, w hich is, how ever, a draw ing o f the trace fossil m odel. Sim ilarly, C rim es e t al. (1981) referred to sup
posed radial sym m etry o f M. aggeris in their pi. 4, fig. 9.
H ow ever, neither the K si^zkiew icz m aterial nor the speci
m en in the photograph o f C rim es et al. (1981) display the radial patterns. The photographs and descriptions published by C ham berlain (1971a) and part o f the photographs o f C rim es e t al. (1981) show a granular surface o f the trace fos
sils, and therefore their slightly lobate outline, w hich cannot be treated as the radial pattern. The K si^zkiew icz specim ens are m ostly sm ooth and display convex or concave apex.
Som e o f them , sim ilarly to the m aterial o f C rim es e t al.
(1981), are slightly fluted. This reveals the pre-depositional origin o f the trace fossil flange. H ow ever, it cannot be ex
cluded that the apical knob is a term ination o f a vertical bur
row that m ay be also active after deposition o f turbidite.
N evertheless, there is no vertical burrow structure in the pol
ished slab o f one cross-sectioned K si^zkiew icz specim en (U J TF 1762).
In tw o cases (UJ TF 1763, 2501), the trace fossils dis
play a bulb-like m orphology in cross-section, i.e. their di
am eter is sm aller at the base than at the half-w ay poin t o f their height.
M ost probably, M. aggeris represents the upper part o f a thin vertical or subvertical cylindrical burrow , w hich d is
plays a hem ispherical funnel-like u pper term ination (Fig. 2).
hesive sedim ent. In contrast, the funnel o f M. aggerisw as m ost probably produced actively by the tracem aker in cohe
sive m ud and possibly it w o rk e d as a trap. Thus, the funnel is a record o f a unique b ehaviour and is a basis for separa
tion o f M. aggerisfrom o ther ichnotaxa. H ow ever, it is not excluded that Mammillichnisand LaevicyclusQ uenstedt are related. N evertheless, there is so far no satisfactory explana
tion o f Laevicyclusand th erefo re such com parison cannot be drawn. Probably, the flatter specim ens w ith deeper apical depressions, w hich resem ble Laevicyclus, are only preserva
tional variants o f the sam e ichnotaxon. T hese specim ens can occur if the term ination o f the thin burrow in the funnel w as rim m ed by a sm all m ound (Fig. 2).
Palaeoenvironm ent: D eep-sea flysch deposits, except one occurrence in the C arboniferous deltaic deposits (Eager et al., 1985) and on eocurrence (?) in teh C am brian shallow - w ater deposits (Paczesna, 1986, 1996).
Stratigraphic range: (?)C am brian (Paczesna, 1986, 1996), O rdovician (C ham berlain, 1977), O ligocene (Ksi^z- kiewicz, 1977; A lexandrescu et al., 1993).
Bergaueria Prantl 1945
Diagnosis: C ylindrical or h e m isp h erical, vertical structures h a v ing sm ooth, u n o m am en ted w a lls, c irc u la r to ellip tical in cro ss-sec
tion; fill essen tially stru ctu reless; ro u n d ed base, w ith o r w ith o u t shallow , central depression an d radial ridges (after P em berton et al., 1988).
Remarks: Bergaueria is pro b ab ly a cubichnial o r dom ich- nial trace fossil produced by suspension-feeders (Ftirsich,
1975). These w ere pro b ab ly coelenterates, chiefly sea anem ones (e.g. Prantl, 1945, A lpert, 1973; P em berton et al., 1988) sim ilar to living Cericmthusor Edwardsia(Pem berton et a l, 1988). The present-day sea anem ones are very w idely distributed from estuaries to abyssal plains (e.g. C arney, 1981). H ow ever, in the case o f non-radiated IB. prantli K si^zkiew icz (1977) and B. hemispherica C rim es et al.
(1977) that additionally has no apical depression, the com parison w ith sea-anem one burrow s should be treated with caution.
Bergaueriaoccurs in diverse facies from shallow -w ater deposits (N arbonne, 1984; C rim es & A nderson, 1985) to flysch deposits (Prantl, 1945; K si^zkiew icz, 1977; C rim es
& C rossley, 1991) from the late Precam brian (e.g. Crim es, 1987) to (?) the Eocene (K si^zkiew icz, 1977) and M iocene (U chm an, 1995).
?Bergaueria prantliK si^zkiew icz 1977 Fig. 3
partim 1977 Bergaueria prantli n. ichnosp. - Ksi^zkiewicz, 53, pi. 1, figs. 3-5, text-fig. 5c-e.
? 1996 Bergaueria prantli Ksi^zkievvicz - Paczesna, 56, pi. 1, figs. 8-9.
Fig. 3. ?B erg a u eria p ra n tli K si^ zk iew icz, sole o f tu rbiditic sandstone bed. U J T F 907, R opianka beds (S e n o n ia n -P ale o c e n e), K ^clow a. S cale b ar = 1 cm
Emended diagnosis: B ergauerians that display an irregular shape and apical, m ore or less distinct, irregular depression.
M aterial: 3 specimens (U J TF 115 (holotype), 967, 1178).
Remarks: In confrontation w ith the revised Bergaueria (Pem berton et al., 1988), the original K si^zkiew icz (1977) diagnosis o f IB. prantli is too b road and does not suffi
ciently express differences betw een this taxon and B. perata Prantl. P em berton et al. (1988) reservedly included IB.
prantliin B. perata. This suggestion cannot be accepted. B.
perata is m ore regular, w ith steeper sides, m ore or less cir
cular apical depression, and faint radial ridges at the apex.
T he differences have been noticed b y K si^zkiew icz (1977, p. 53). In contrast, IB. prantliis ch aracterized by an irregu
lar outline and irregular apical depression. T herefore, it should be treated as a separate ichnospecies. H ow ever, it is not clear if this trace fossil is a real Bergaueria, w hich is typified by B. perataPrantl. It cannot be excluded th at som e specim ens are casts o f the upper term inations o f som e verti
cal or subvertical partially filled burrow s. T herefore, the question-m ark is placed in front o f the ichnogeneric nam e.
Som e specim ens labelled by K si^zkiew icz as B. prantli are undeterm inable trace fossils (U J TF 1471).
SIMPLE STRUCTURES
ArthrophycusH all 1852 Ich n o sp ecies in cluded in A rth ro p h ycu s H all:
1831 Fucoides alleghaniensis - Harlan, 393-398.
1838 Fucoides Harlani - Conrad, 113.
1843 Fucoides Harlani - Hall, 46, figs. 1 -2.
1852 Arthrophycus harlani - Hall, 5, pi. 1 ,p i.2 , figs. la-c.
1852 Harlania Hallii Gopp. -G oppert, 98, pi. 41, fig. 4.
110
A. UCHMAN1856 Harlania Hallii Goepp. - Bronn & Roemer, pi. 4, fig. 1.
1869 Harlania Goepp. — Schimper, 195.
1874 Harlania Hallii Goepp, - Schimper, pi. 2, fig. 6.
1881 Harlania Hallii Goepp. - Saporta, fig. 1.2.
1882 Arthrophycus Harlani Hall - Saporta, 49, pi. 7, fig. 2.
1883 Arthrophycus Harlani Hall - Saporta & Marion, fig. 21.
1886 Arthrophycus cfr. Harlani Hall - Delgado, 75, pis. 23, 25-26.
1887 Arthrophycus cfr. Harlani Hall - Delgado, 66, pis. 9-10.
1890 Arthrophycus Harlani Hall - Schimper & Schenk, fig. 41.
1895 Arthrophycus Harlani Gopp. - Fuchs, pi. 9, fig. 2.
1901 Arthrophycus harlani Conrad - Grabau, 132, pi. 16.
1908 Arthrophicus Harlani Hall - Hernandez Pacheco, 84 (lap
sus calami).
1918 Problematicum - Zuber, fig. 123.
1940 Harlania harlani (Conrad) - Desio, 68, fig. 7, pi. 6, figs.
1-2, pi. 7, pi. 13, fig. 6.
non 1942 Harlania G oeppert-P icard, 14, pi. 2, figs. 6-8.
1948 Arthrophycus alleghaniensis (Harlan) - Shinier & Shrock, 719, pi. 303, figs. 26-27.
' ? 1949 Fraena (?) - Gómez de Llarena, fig. 1.
1952 Arthrophycus - Becker & Donn, 214, figs. 1-2.
1955 Harlania harlani Conrad - Lessertiseur, pi. 74, fig. 11.
? 1960 Arthrophycus — Henbest, fig. 177.IF.
1961 Arthrophycus alleghaniensis (Harlan) - Wolfart, 81, pi. 7, fig. 2.
1963 Harlania alleghaniensis (Harlan) - Bender.pl. 13, fig. 3.
1969 Arthrophycus — Seilacher, 120, pi. I.
non 1970 Arthrophycus-Wke burrows - Frey, 23, fig. 4E.
non 1970 Arthrophycus (?) sp. - Frey & Howard, 159, fig. 3e.
1970 Arthrophycus sp. - Książkiewicz, 285, fig. la.
1970 Harlania - Selley, pi. 2, fig. b.
non 1971 Arthrophycus — Maberry, 11, fig. 8.
non 1972 Arthrophycus-Wke burrows - Frey, fig. 9A.
1972 ?Arthrophycus - Frey & Chows, 37, fig. 5C.
non 1975 Arthrophycus sp. - Chiplonkar & Ghare, 72, fig. IB.
1977 Arthrophycus alleghaniensis (Harlan) - Baldwin, 26, pi.
2a.
? 1977 1 Arthrophycus Hall - Kern, 30.
1977 Arthrophycus strictus n. ichnosp. - Książkiewicz, 57, pi.
1, figs. 11-12.
non 1977 Arthrophycus annulatus n. ichnosp. - Książkiewicz, 56, pi. 1, figs. 8-10 [=Ophiomorpha am ulata (Książkiewicz)]
non 1977 Arthrophycus^?) dzulynskii n. ichnosp. - Książkiewicz, 58. pi. 1, figs. 13-14 [=Protovirgularia dzulynskii (Książkiewicz],
partim 1977 Buthotrephis spec, indet. - Książkiewicz, 76, text-fig. 10b [non text-fig. lOd, f-g, i-j, m, o-s, u-y = Thalassinoides suevicus (Rieth)].
1977 Sabularia tenuis n. ichnosp. - Książkiewicz, 71, pi. 2, fig.
3.
1978 Arthrophycus alleghanensis - Acenolaza, 40, fig. 25.
? 1978 1 Arthrophycus Hall - Kern, 251.
? 1981 Lorenzinia ci. m onzaeRenz-Crim es era/., 972, pi. 3,figs.
4-5.
? 1982 Arthrophycus cf. strictus K siążkiewicz- Alexandrescu &
Brustur, 36, pi. 3, fig. 4.
1983 Arthrophycus alleghaniensis (Harlan) - Turner & Benton, 451, fig. 4A.
1984 Arthrophycus alleghaniensis (Harlan) - Lilian, 57, pi. 2, fig. 5.
1984 Arthrophycus alleghaniensis (Harlan) - Pickerill, Ro
mano & Melendez, 251, fig. 2a.
1985 Arthrophycus alleghaniensis (Harlan) - Durand, 41, pi. 4, figs. 3-4.
1985 cf. Arthrophycus-E a g e r et al., 137, fig. 10A.
non 1986 Arthrophycus - Ghare & Kulkami, 45, pi. 1, fig. 1.
1987 Arthrophycus Hall - Bjerstedt, 875, fig. 8.1.
1987 Arthrophycus alleghaniensis (Harlan) - Valle, 26, fig. 5.
1988 Arthrophycus - Seilacher & Alidou, fig. ld-f, fig. 2.
?non 1991 Arthrophycus isp. — Pickerill, Fill ion & Branchley, 73, fig.
2
.
?non 1992b "Arthrophycus" corrugatus (fr\lsch )-M \ku \is, 27, pi. 14, fig. I.
? 1993a Arthrophycus sp. A - Li, 94, pi. I, fig. 5.
?non 1993a Arthrophycus sp. A - Li, 94, pi. 2, fig. 7.
? 1994 Arthrophycus minoricensis Bourrouilh - Orr, 209, figs. 7, 13, 15 (nomen nudum).
?non 1994 Arthrophycus qiongzhusiensis ichnosp. nov. - Luo et al., 33, pi. 1, fig. 4., pi. 2, fig. 3 [=ITorrowangea isp.].
1996 Arthrophycus strictus K si^zkiewicz- Paczesna, 59, pi. 10, fig. 3.
1996 Arthrophycus hunanensis ichnosp. nov. - Zhang & Wang, 484, pi. 3. fig. I.
Emended diagnosis: O blique to horizontal, cylindrical or subcylindrical structures w ith regular, perpendicular fine ribs and a tendency to plunging into bed surfaces. C om m only, the trace fossils are grouped in bundles.
Remarks: Probably, the above diagnosed Arthrophycus em braces only three ichnotaxa, i.e. A. alleghanienis (H ar
lan) and tw o ichnotaxa lum ped so far in A. strictus Ksiqz- kiew icz, w hich are separated as A. strictusK si^zkiew icz and A. tenuis(K si^zkiew icz) in this paper. O rr (1994) illustrated Arthrophycus minoricensis B ourrouilh, w hich w as origi
nally described in an unpublished thesis. The description and illustration o f this trace fossil is not sufficient for assess
m ent o f its diagnostic features. Arthrophycus qiongzhusien
sis described b y Luo et al. (1994) from the C am brian o f C hina, is a hypichnial, w inding trace fossil w ith fine trans
verse striae. A photograph o f this trace fossil suggests that the striae can be an external expression o f internal sedim ent pads typical o f Torrowangea W ebby. R elationships be
tw een these two ichnotaxa should be explained. Z hang &
W ang (1996) described Arthrophycus hunanensisfrom Silu- rian -D ev o n ian deposits o f C hina. It is not excluded that it is a ju n io r objective synonym o f A. alleghanienis.
Arthrophycus annulatus K si^zkiew icz and Arthrophy- cus(T) dzulynskii K si^zkiew icz (K si^zkiew icz, 1977), and several other ichnotaxa are excluded from Arthrophycusas indicated in the above list.
A. alleghanienis (H arlan) is com m on in neritic silici- clastic facies o f the M iddle P alaeozoic and regarded as rep
resenting a burrow (Sarle, 1906) o f feeding arthropods (e.g., T urner & B enton, 1983). H ow ever, Schiller (1930) tried to explain it as a tectonic structure, and B ecker & D onn (1952) regarded this ichnotaxon as an “ algal structure” . A. al
leghanienism ay occur as linear, protrusive, flat palm ate, or deep palm ate retrusive trace fossil (S eilacher & A lidou, 1988). The palm ate specim ens have been included in Phy- codespalmatus(H all) (Seilacher, 1955, p. 386) [incorrectly spelled P. palmatum;Fillion & P ickerill, 1990]. Sim ilarities o f this ichnotaxon to Phycodes w ere p ointed out by Beck (1916). A. strictusK si^zkiew icz displays sim ilar tendencies.
Thus, grouping in bundles can be regarded as the com m on feature o f Arthrophycus, w hich is included in th e diagnosis.
H ow ever, this feature does not occu r constantly, and there
fore Arthrophycuscannot be included in PhycodesRichter.
Stratigraphic range: Low er C am brian (Linan, 1984) - L ow er M iocene (A lexandrescu & B rustur, 1984).
Arthrophycus strictusK siazkiew icz 1977 Figs. 4, 5A, 6
1970 Arthrophycus sp. - Ksiazkiewicz, 285, fig. 1 a.
*v 1977 Arthrophycus strictus n. ichnosp. - Ksiazkiewicz, 57, pi.
1, figs. 11-12.
£ # i
m
Fig. 4. Arthrophycus strictus Książkicwicz (arrowed) and Arlhrophycus tenuis (Książkiewicz). Sole of a turbiditic sandstone bed. UJ TF 131, Lgota beds (Albian), Rzyki near Andrychów.
More details are shown in Fig. 5. Scale in mm
partim 1977 Bulhnirepius spec. mdel. - Ksiazkiewicz. 76, text-tig. 10b [non text-fig. lOd, f-g, i-j, tn, o-s, u-y = Thalassinoides suevicus (Rieth)].
? 1981 Lorenzinia cf. moreae R enz-C rim es et al., 972, pi. 3, figs.
4-5.
? 1982 Arlhrophycus cf. strictus Ksiazkiewicz - Alexandrescu &
Brustur, 36, pi. 3, fig. 4.
non 1996 Arthrophvciis strictus Ksiazkiewicz- Paczesna, 59, pi. 10,
% 3.
E m e n d e d d ia g n o sis: Sm all A rthrophycus, approxim ately circular in cross-section, w ith very fine, delicate perpen
dicular ribs. Structures are distinctly arcuate in vertical plane and plunge into sole o f beds.
M aterial: 5 slabs (U JTF 124, 131 (holotype), 1318, 1508, 1523).
R e m a rk s: The holotype o f A rthrophycus strictus K si^zkie- w icz (U J T F 131, Fig. 4) contains tw o different trace fossils treated by K siazkiew icz (1977) as variations o f the sam e ichnospecies. H ow ever, these trace fossils display enough different features for th eir separation at the ichnospecies level. The first trace fossil is represented by short hypich- nial, perpendicularly striated, sim ple ridges, w hich dip into the bed at both ends (Fig. 5). T hey are elevated at the m iddle part and form w ide, variably oriented arches, w hich are 4 -6 m m in diam eter. Ridges o f the sam e m orphology are better and m ore com pletely preserv ed in other specim ens (e.g. UJ TF 124), w hich w ere determ ined by K siazkiew icz as A.
strictus. In som e more com plete specim ens, the ridges tend to form bunches, w hich plunge into the sole o f beds (Fig. 6).
The second type is represented by short, horizontal and
Fig. 5. Arthrophycus strictus Ksiazkiewicz (arrowed) and Arthrophycus tenuis (Ksiazkiewicz). A-C. Details o f Fig. 4. Scales in mm
112
A. UCHMANFig. 6. Arthrophycus strictus Książkiewicz. Sole of a turbiditic sandstone bed. UJ TF 124, Ropianka beds (Senonian-Paleocene), Mszana Dolna. Scale in mm
com m only branched ridges, w hich do not form bunches and are not bent in arches (Fig. 5 B -C ). The ridges are covered w ith very fine p erpendicular striae and are distinctly thinner than the ridges o f the first type. T heir diam eter ranges from 0.4 to 1.2 m m . A bout h a lf o f th e ridges are branched, alw ays at a sharp angle. Ridges o f th e second type are alm ost iden
tical to Sabularia tenuis K si^zkiew icz. T hese trace fossils differ only in the lack o f the p erpendicular striation in m ost specim ens o f S. tenuis. H ow ever, som e ridges o f the latter ichnotaxon, including the holotype (U J TF 1686), display faint perpendicular striation, w hich w as noted also by K si^zkiew icz (1977, p. 71). P reservation o f th e fine striation depends strongly on cohesion o f substrate and w eathering.
Thus, degree o f p reservation o f the striation is not a suffi
cient criterion for the ichnotaxonom ic separation o f these trace fossils and therefore S', tenuis is included in A rthrophy
cus. The ichnospecies nam e o f S. tenuis is retained, as A.
tenuis, for th e second type o f ridges. For the first type o f ridges the nam e A. strictus is retained.
The specim en U J TF 1318 indicated by K si^zkiew icz (1977, text-fig. 10b) as B uthotrephis spec, indet. displays distinct features o f A. strictus.
A. strictus w as regarded by K si^zkiew icz as repre
senting a feeding burrow o f polychaetes. It occurs in the Al- bia n -S en o n ian flysch deposits o f the C arpathians (K si^zkiew icz, 1977). A n o t w ell preserved occurrence, treated therefore reservedly, is n oted in the Eocene flysch o f R om ania (A lexandrescu & B rustur, 1982). B rustur & Stoica (1993) reported A. cf. strictus from U pper Eocene flysch.
Paczesna (1996) d escribed A. strictus from the shallow - w ater C am brian deposits, b u t it displays a m edian furrow transecting the p erpendicular ribs. T herefore, it is excluded from this ichnospecies.
A rthrophycus tenuis (K si^zkiew icz 1977) Figs. 4 - 5 , 7
1918 [...] drobne hieroglify - Zuber, fig. 3.
*v 1977 Sabularia tenuis n. ichnosp. - Ksi^zkiewicz, 7 1, pi. 2, fig.
3.
partim 1977 Arthrophycus strictus n. ichnosp. - Ksi^zkiewicz, 57, pi.
1, figs. 11-12.
1984 Sabularia tenuis Książkiewicz - Alexandrescu & Brustur, 20, pi. 1, figs. I -2, pi. 2, fig. 1.
1986 Sabularia tenuis Książkiewicz - Alexandrescu, 6, pi. 1, figs. 1-2, pi. 2, figs. 1-2.
1993 Sabularia ichnosp. - Alexandrescu et al., pi., figs. 1-3.
?non 1995 Sabularia tenuis Książkiewicz - Bąk, fig. 10.
?non 1995 ISabularia tenuis Książkiewicz - Bąk, fig. 21.
Emended diagnosis: Sm all, hypichnial, short, straight, ra
rely branched ridges, covered w ith very fine perpendicular striae. T he striation com m only is not preserved.
M aterial: 9 slabs (UJ TF 131, 373-374, 1493, 1579, 1583, 1614, 1687 (holotye), 2680.
R e m a rk s : A. tenuis is noted only in the flysch deposits o f the C arpathians from the V alanginian (K siążkiew icz, 1977) to the (?) L ow er M iocene (A lexandrescu & Brustur, 1981).
H alopoa Torell 1870 Ichnospecies included in Halopoa Torell:
1870 Halopoa imbricata n. sp. - Torell, 7 [no illustration]
1878b Trichophycus sulcatum n. sp. - Miller & Dyer, 4, pi. 4, fig.
5. [also James, 1884, pi, 6, fig, 5].
1932 Fucusopsis angulatus Palib. gen. et sp. n. - Vassoevich, 51, pi. 2, figs. 2, 6.
1946 Fucopsis angulatus - Grossheim, fig. 4b (lapsus calami).
? 1949 Pistas de arthropodos - Gómez de Llarena, fig. 8.
?non 1959 Fucusopsis angulatus Palibin - Birkenmajer, 229, pi. 22 [non text-fig. 1].
1959 Fucusopsis angulatus Palibin-Seilacher, 1070, tab. 2, fig.
30.
1962 Fucusopsis — Seilacher, fig. 1, pi. 1, fig. 2.
1964 Gyrochorda fraeniformis nov. iscp. - Farres Malian, 92, pi. 5, fig. 2.
1965 Halopoa imbricata Torell - Martinsson, 219, fig. 29.
1965 Halopoans -- Martinsson, 219, figs. 30-32.
v 1970 Fucusopsis angulata Palibin - Książkiewicz, 286, fig. Is.
v 1970 Fucusopsis annulata ichnosp. n. - Książkiewicz, 286, fig.
lr.
1970 Fucusopsis sulcatum (Miller and Dyer) - Osgood, 380, pi.
64, fig. 1, pi. 70, fig. 1, pi. 71, fig. 5.
?non 1970 Fucusopsis cf. angulatus Vassoevich - Chiplonkar &
Badve, 9, pi. 3, fig. 6.
? 1973 Feddenichnus feddeni sp. n. - Chiplonkar & Borkar, 572, figs. 1-2.
1976 Fucusopsis - Hakes, 27, pi. 8, fig. 2a-b.
v 1977 Fucusopsis angulata Palibin - Książkiewicz, 59, pi. 2, fig.
5 [also Leszczyński, 1992, pi. 15, fig. 1].
v 1977 Fucusopsis annulata Książkiewicz, 60, pi. 2, figs. 6-7.
v 1977 Fucusopsis striata (Hall) - Książkiewicz, 61. pi. 2, fig. 8.
? 1977 Fucusopsis Vassoevich - Kumar et al., 421. pi. 9, fig. 6.
1978 Fucusopsis angulata Palibin - Radwański, 51, pi. 1, fig.
2.
?non 1980 Fucusopsis - Soudant, pl„ figs. C-D.
? 1978 Halopoa indica ichnosp. n. - Badve & Ghare, 132, pi. 4, fig. 5.
non 1981 ?Fucusopsis sp. - Bradshaw, 639, figs. 38-39 [=Palaeo- phycus isp.].
partim 1981 Fucusopsis angulata Palibin - Crimes et a l, 968, pi. 2, fig.
1 [?non pi. 2, fig. 2].
1981 Fucusopsis sp. - Pickerill, 44, fig. 5b.
non 1982 Radionereites annulata (Książkiewicz) - D’Alessandro, 535, pi. 38, fig. 2, pi. 39, fig. 4, pi. 40, fig. 1, pi. 42, fig. 4.
? 1983 Halopoa sp. - Singh & Rai, 77, pi. 7, figs. 74-76.
1983 Halopoa sp. - Palij et al.. pi 67, figs. 1-2.
? 1986 Fucusopsis angulatus Palibin in Vassoevich - Ghare &
Kulkami, 47, pi. 1, fig. 2, pi. 5, fig. 2.
? 1988 Fucusopsis? isp. - Yang, 322, pi. 1, fig. 3a.
? 1990 Palaeophycus striatus! Hall - Fillion & Pickerill, pi. 11, fig. 1.
1990b Fucusopsis - Seilacher, pi. 32.2f.
Fig. 7. Arthrophycus tenuis (Książkiewicz). Soles of a turbiditic sandstone beds. A. UJ TF 1687, holotype. Krosno beds (Oligocene), Wujskie. B. UJ TF 2680, label lost. C. UJ TF 374, Kąclowa, Ropianka beds (Senonian-Paleocene). Coll. W. Szajnocha. D. UJ TF 373, Krosno beds (Oligocene), Slonne near Zaluż. Scale bars = 1 cm, scale in C in mm
1993 Bioglife (“Pietrificarea de la Pinsdorf’) Contescu ct at., pi. 8, fig. 3.
? 1994 Fucusopsis isp. - Gong, 487, pi. 6, fig. 4.
? 1995 Palaeophycus (Fucusopsis) angulci/a Palibin in Vasso- evicli - Crimes & McCall, 239, fig. 4a.
? 1995 Palaeophvcus striatus Hall - Crimes & McCall, 239, fig.
4b.
1995 Palaeophycus sulcal u\ (Miller & Dyer) - Crimes &
McCall, 2 41, fig. 4c.
? 1995 Fucusopsis ichnosp.-B ą k , fig. 19.
1997 Palaeophycus imbricatuas (Torell) - Jensen, 69, figs. 8D, 45-46, 47B. D, 48B.
E m e n d e d d ia g n o sis: Long, generally horiżontal trace fos-
114
A. UCHMANFig. 8. Halopoa imbricata Torell in turbiditic sandstone bed.
UJ TF 87, Beloveza Formation (Eocene), Zubrzyca Górna. A.
Bedding-plane view. B-C. Cross-section views. Polished and oiled surfaces. Scale bars = 1 cm
sils covered w ith longitudinal irregular ridges or w rinkles, w hich are com posed o f several im perfectly overlapping cy
lindrical probes.
R e m a rk s : H alopoa w as described, but not illustrated by T orell (1870). A ndrew s (1955) designated H. im bricata Torell as the type ichnospecies o f this ichnogenus. M artins- son (1965) synonym yzed H alopoa com posita Torell (1870),
Scotolithus m irabilis Linnarson (1871) and H alopa im bri
cata, and illustrated the lectotype o f the latter ichnotaxon.
He tried to com pare H. im bricata to G yrochorte H eer. H ow ever, Jensen (1997) show ed that H. com posita and S. miri- abilis are different ichnospecies and that the com parison to G yrochorte cannot be accepted. H akes (1976) tu rn ed his at
tention to the strong sim ilarities betw een H alopoa Torell and Fucusopsis Palibin in V assoevich (1932). A ccording to the description by Jensen (1977), H alopoa displays the sam e significant features as F ucusopsis. H ow ever, Jensen included this tw o ichnogenera in P alaeophycus. A lso P em berton & Frey (1982) included F ucusopsis in P ala eo p h ycu s, w ho related P alaeophycus to passively fdled, open, w alled burrow s, including specim ens th a t display longitudinal striation. H ow ever, the origin o f the longitudinal striation m ay be diverse and related to diverse tracem aker behaviour or taphonom ic processes. The longitudinal striation m ay be produced actively by locom otory organs and/or passively by body appendages o f tracem aker. Its preservation strongly depends on cohesion o f the substrate. O ther types o f longi
tudinal striation m ay be produced by m icrofaulting con
nected w ith collapse o f burrow s or w ith com pressional faulting due to tension caused by tracem aker from interior o f burrow . This type o f striation is a p ro d u ct o f a unique behav
iour, w hich can be interpretet as diagnostic feature at the ichnogeneric level. The latter idea w as proposed for F u cusopsis (O sgood, 1970, p. 380; Seilacher, 1990b). In the case o f the K si^zkiew icz m aterial, there is no wall and no evidence that the burrow s w ere open, and therefore it does not conform w ith the idea o f P alaeophycus sensu P em ber
ton & Frey (1982). Trace fossils o f this type, especially H.
im bricata (=F. angulata) from the Ksic(zkiewicz m aterial are com m only densely crow ded and they rew ork the low er
m ost p art o f turbiditic beds. T hey are rather produced by de
posit feeding organism s. T heir outline is not sharp and di
am eter is not constant.
The cross-sections o f the H alopoa specim ens display several overlapping probes, w hich are com m only stacked in vertical plane (Fig. 8 B -C ). The trace fossils form a Teichi- chnus-like structure, how ever m uch less vertically and regu
larly developed than in Teichichnus Seilacher. Som e probes o f H alopoa diverge aw ay from som e objects, for instance other fills o f burrow s o f the sam e ichnotaxon, and jo in once again behind the object (Fig. 9B). F orm ation o f each new com ponent probe is connected w ith pushing out o f the for
m erly rew orked sedim ent in the older probe w hich results in the form ation o f an irregular longitudinal striation. D efor
m ation o f the lam inae above the trace fossls (Fig. 8) indi
cates that the sedim ent w as pushed out o f the burrow inte
rior. The burrow s w ere presum ably form ed by intruding in the sedim ent by the tracem aker.
Teichichnus is never so long, lacks external sculpture, and is m uch m ore extended in the vertical plane. Therefore, H alopoa and Teichichnus should rem ain separate. O ne can conclude th at H alopoa represents sufficently unique behav
iour for its retention as ichnogenus.
K si^zkiew icz (1977) regarded th e striation o f F ucusop
sis (= H alopoa) as resulting from the sculpture o f the trace
m aker, w hich probably belonged to priapulid w orm s. Such explanation does not conform w ith the trace fossil m orphol
ganic-rich lam inae can be observed. H alopoa exploits rarely occupy tiers in the sandstone part o f turbiditic beds.
H alopoa im bricata T orell 1870 Figs. 8 -9
* 1870 Halopoa imbricata n. sp. - Torell, 7 [no illustration]
1878b Trichophycus sulcatum n. sp. - Miller & Dyer, pi. 4, fig.
5 [also James, 1884: pi. 6, fig. 5]
1932 Fuciisopsis angulatus Palib. gen. et sp. n. - Vassoevich, 51, pi. 2, figs. 2, 6.
1946 Fucopsis angulatus - Grossheim, fig. 4b (lapsus calami).
? 1949 Pistas de arthropodos - Gómez de Llarena, fig. 8.
?non 1959 Fuciisopsis angulatus Palibin - Birkenmajer, 229, fig 1, pi. 12.
1959 Fuciisopsis angulatus Palibin - Seilacher, 1070, tab. 2, fig.
30.
1962 Fuciisopsis - Seilacher, fig. 1, pi. 1, fig. 2.
1962 Fucusopsisangulatus Palib. - Dimitrievae/ al., pi. 75, fig.
1.
1964 Gvrochorda fraenifonnis nov. iscp. - Farres Malian, 92, pi. 5, fig. 2.
1965 Halopoa imbricata Torell - Martinsson, 219, fig. 29.
1965 Halopoans - Martinsson, 219, figs. 30-32.
1970 Fucusopsis angulata Palibin - Książkiewicz, 286, fig. Is.
1970 Fucusopsis sulcatum (Miller and D yer)-O sgood, 380, pi.
64, fig. 1, pi. 70, fig. 1, pi. 71, fig. 5.
1976 Fucusopsis - Hakes, 27, pi. 8, fig. 2a-b.
v 1977 Fucusopsis angulata Palibin - Książkiewicz, 59, pi. 2, fig.
5 [also Leszczyński, 1992, pi. 15, fig. 1].
v 1977 Fucusopsis striata (Hall) - Książkiewicz. 61, pi. 2, fig. 8.
1978 Fucusopsis angulata Palibin - Radwański, 51, pi. 1, fig.
2.
partim 1981 Fucusopsis angulata P alibin- Crimes et al., 968, pi. 2, fig:
1 [?non pi. 2, fig. 2].
? 1990 Palaeophvcus striatus? Hall - Pillion & Pickerill, pi. 11, fig. 1.
1990b Fucusopsis - Seilacher, pi. 32.2f.
1993 Bioglife ("Pietrificarea de la Pinsdorf’) - Contescu et al..
pi. 8, fig. 3.
1995 Palaeophvcus (Fucusopsis) angulata Palibin in Vasso
evich - Crimes & McCall, 239. fig. 4a.
■ ? 1995 Palaeophvcus sinol/is Hall - Crimes & McC all. 239, fig.
4b.
1995 Palaeophvcus sulcatus (Miller & Dyer) - Crimes &
McCall, 241, fig. 4c.
E m e n d ed d ia g n o sis: U nbranched H alopoa w ith horizontal, relatively long and continuous furrow s and w rinkles.
M aterial: 9 specimens (UJ TF 69-70, 87, 93, 588, 1158, 1179, (?) 1570, 2507).
Description: As in Książkiewicz (1977) description o f Fucusopsis angulata Palibin.
R e m a rk s: The C arpathian m aterial displays com parable features to H alopoa im bricata Torell (1870) and Trichophy- cus sulcatum M iller & D yer (1877). P em berton & Frey (1982) included T richophycus sulcatum M iller & D yer, the type m aterial o f F. sulcatum (O sgood, 1970), in P alaeophy- cus sulcatum (M iller & D yer). T hey also excluded F. a n g u lata described by K siążkiew icz (1970, 1977) from this ich- nospecies; how ever, these authors included the type mate-
Fig. 9. Halopoa imbricata Torell. Soles o f turbiditic sandstone beds. A. UJ TF 93, Jarmuta Formation (Senonian), Jaworki. B. UJ TF 1158, Szydłowiec beds (Senonian), Kobielnik. Scales in mm rial o f F. angulatus in P. sulcatum . T hey included also, re
servedly, the K siążkiew icz m aterial in P alaeophvcus stria tus. N evertheless, the m entioned ichnotaxa display features o f H alopoa, and are excluded herein from P alaeophvcus, and described under H. im bricata.
116
A. UCHMANFig. 10. Halopoa anindata (Ksiqzkie-wicz), the holotype. Sole of a turbiditic sandstone bed. A. U.F TF 1263, Hieroglyphic beds (Eocene), Kamionka Wielka. A. General view. B. Detail o f A. Scales in mm
K siazkicw icz (1977) distinguished F ucusopsis striata (Hall) (labelled also as F ucusopsis longistriata) and com pared it to P alaeophycus stria tu s described by H all (1852, p.
22, pi. 10, fig. la). P em berton & Frey (1982) included re
servedly the K siqzkiew icz F ucusopsis striata in P alaeophy
cus striatus Hall. O ne o f the Ksicjzkiwicz specim ens (U J TF 70) is a hypichnial striated trac e fossil on the sole o f biotur- bated turbiditic sandstone, lacking wall. It displays features o f H. im bricata and is included in this ichnospecies.
B irkenm ajer (1959) considered th at F. angulata (= //.
im bricata) w as produced by deposit-feeding annelids.
Stratigraphic range: L ow er C am brian (Seilacher, 1990b; Jensen, 1997) - M iddle M iocene (C rim es & M cC all, 1995).
H alopoa annulata (K si^zkiew icz 1977) Figs. 10, 15
*v partim 1970 Fucusopsis annulatei ichnosp. n. - Ksi^zkiewicz, 286, fig.
Ir2 (non fig. lrl).
v 1977 Fucusopsis annulata Ksi^zkiewicz, 60, pi. 2, figs. 6-7.
Diagnosis: Commonly branched Halopoa with perpendicular con
strictions.
M aterial: 10 specimens (UJ TF 71, 587, 767, 1263 (holotype), 1525, 1696, 2503-2504, 2506, 2509).
R e m a rk s : The discussed ichnotaxon, described by Ksi%z- kiew icz as Fucusopsis a n n u la ta , is included in H alopoa Torell in this paper. F. annu la ta w as reservedly included in Palaeophycus: alternus Pem berton & Frey (1982). In the
sam e year, D ’A lessandro included F. annulata K si^zkie- w icz (1970, non 1977) in R adionereites G regory. F ucusop
sis annulata described by K si^zkiew icz in 1970 w ithout designation o f the holotype, com prises tw o different trace fossils. In 1977, K siazkicw icz designated the holotype, w hich corresponds only to the specim en illustrated in 1970 in fig. Ir2. The specim en illustrated in fig. l r l is related to Im ponoglyphus V ialov in this paper. Irrespective o f these facts, D ’A lessandro (1982) included both these speciem ns o f F. annulata K siazkicw icz 1970 non 1977 in R adionere
ites G regory. H ow ever, D ’A lessandro e t al. (1987) then ex
cluded the m aterial o f D ’A lessandro (1982) from R adione
reites and erected for it a new ichnogenus, R utichnus, w hich lum ps certain m eniscate trace fossils. They reservedly in
cluded F. annulata Ksiq.zkiewicz (1977, non 1970) in R u tichnus and suggested the presence o f a m eniscate internal structure in the K siazkiew icz specim ens.
The K siazkiew icz m aterial, including the holotype, does not display any evidences o f internal m eniscate struc
ture, including in specim ens sectioned along the trace fos
sils. It displays, how ever, features o f H alopoa (see the dis
cussion o f this ichnogenus). The m orphology o f H. annulata is due to a special behaviour o f tracem aker. A part from re
peatedly rew orking o f sedim ent along the sam e general course and the outw ard tension o f the sedim ent typical o f H alopoa, the tracem aker m oved by steps searching obli
quely in sedim ent, retreated, and rep eated its action along wide, shallow , vertical or oblique, and asym m etrical U- shaped sub-courses. This resulted in the occurrence o f p er
L ow er O ligocene (personal observations, M alcov F orm a
tion, Sam orody n e a rN o w y T arg in the Polish Carpathians).
H orm osiroidea S chaffer 1928 Ichnospecies included in Hormosiroidea Schaffer:
? 1856 Corallinites rosarium Massal. -M assalongo, 41, pi. 6, fig.
4.
partim 1877 Fucoides Moeschii Hr. - Heer, 113, pi. 39, fig. 15b, pi. 40, 5a. 6a, 11 [non pi. 40, fig. 7a, ?non pi. 40 fig. 12a],
?non 1888 Taenidium carboniferum Sacc. - Sacco, 14, pi. 2, fig. 1.
1897 Halimeda Fuggeri Lor. - Lorenz von Liburnau, 177, pi. 1.
1902 Halimedaites Fuggeri Lor. - Lorenz von Liburnau, 710, pis. 1-2
1908 Spungolithus annulalus Fr. -- Fritsch, 16, pi. 12, fig. 17.
1928 Hormosiroidea florentina n. een. n. sp. - Schaffer, 215, fig. 3.
1959 Hormosiroidea - Seilacher, tab. 2, fig. 20.
1962 Rhabdoglyplmsgrossheimi Vassoevich - Boucek& Elias, 146, pi. 8, fig. 3 (also Vialov, 1971, 91, fig. 3).
1962 Hormosiroidea florentina - Hantzschel, W 241.
1965 Hormosiroidea florentina - Hantzschel, 47.
1965 Halimedides fuggeri - Hantzschel, 42.
1970 Hormosiroidea florentina Schaffer - Osgood, pi. 78, fig.
7.
? 1970 Ichnofossil gen. et sp. indet. - Meiburg & Speetzen, 12, figs. 1-3.
partim 1970 Rhabdoglyplms ichnosp. a - Ksiqzkiewicz, 285, fig. 1 h, k (non fig. li-j).
1971 Fustiglyphus anirulatiis gen. et sp. n. - Vialov, 91, fig. 3.
? 1972 1Hormosiroidea - Frey & Chows, 37, pi. 5G-H.
1975 Hormosiroidea florentina - Hantzschel, 70, fig. 43.3.
1975 Halimedides fuggeri - Hantzschel, 65, fig. 42.1.
v 1977 Rhabdoglyplms caliciformis n. ichnosp. - Ksi^zkiewicz, 66, pi. 3, figs. 6, 11, text-fig. 6c-d.
v ?non 1977 Rhabdoglyplms sulcatus n. ichnosp. - Ksi^zkiewicz, 67, pi. 3, fig. 9, text-fig. 6e.
v 1977 Rhabdoglyplms spinosus n. ichnosp. - Ksi^zkiewicz, 66, text-fig. 6b, pi. 3, figs. 7-8.
v ?non 1977 Rhabdoglyplms compos it us n. ichnosp. - Ksi^zkiewicz, 67, pi. 3, fig. 9, text-fig. 6.
1977a Hormosiroidea florentina Schaffer - Seilacher, 309, fig.
6n.
non 1977a Hormosiroidea beskidensis - Seilacher, 309, fig. 6k-l.
1977 Hormosiroidea florentina Schaffer - Chamberlain, 14, figs. 2q, 5G.
1979 Hormosiroidea florentina Schaffer - Chamberlain, 17, part o f fig. 3.
1979 Fustiglyphus roselandensis new species - Boyer, 75, figs.
2-3.
non 1985 Hormosiroidea canadensis n . ichnosp. — Crimes & Ander
son, 325, fig. 8.1-9
? 1979 H orm osiroidea- Powichrowski, fig. 3.4.
?non 1989 Hormosiroidea arumbera sp. nov. - Walter et al„ 244, figs. I4D-E, I5B, D.
non 1990 Hormosiroidea isp. - Bryant & Pickerill, 49, fig. 4.
non 1991 Hormosiroidea (Saerichnites) cf. beskidensis (Plicka) - Crimes & Crossley, 32, fig. 3d-e (=Saerichnites abniptus Billings).
1993 Fustiglyphus annulalus Vialov - Stanley & Pickerill, 58, figs. 2-3A-D.
partim 1993 Rhabdoglyplms grossheimi Vassoevich - Stanley & Pick
erill, 62, fig. 51 (copy from Ksi^zkiewicz, 1977, fig. 6b).
v 1995 Hormosiroidea florentina Schaffer - Uchman, fig. 15A.
?non 1997 Fustiglyphus isp. - Jensen, 50, fig. 34B.
der Hormosiroidea S chaffer at the ichnogeneric level is rather the problem o f tradition and m inor m orphological features than use o f significant diagnostic features.
S eilacher (1977a) regarded the spherical “cham bers” o f Hormosiroidea as sectioned vertical burrow s th at protrude from the horizontal burrow tow ard the bottom . H e included Saerichnites Billings and Rhabdoglyphus V assoevich in Hormosiroidea Schaffer. This ichnotaxon w as tentatively included in the graphoglyptids (Seilacher, 1977a). Such a concept o f Hormosiroidea cannot be accepted.
T he type ichnospecies o f Hormosiroidea, //. florentina S chaffer (1928), consists o f a row o f sm all bulbs connected by a string (Schaffer, 1928; H antzschel, 1975, p. W 70, fig.
40.3) (Fig. 11B). This ichnotaxon w as ten tatively included in the graphoglyptids (Seilacher, 1977a). Saerichnites con
sists o f one or tw o parallel row s o f casts o f vertical shafts (U chm an, 1995). R elation o f Saerichnites to H. florentina is problem atic. P robably, the bulbs o f H. florentina, w hich are m uch greater than the string diam eter and w hich tend to be a little lobate in outline, do not represent vertical burrow s, but m ore likely breeding (?) structures. Thus, single or double row s o f vertical o r subvertical burrow s or single or double row s o f casts o f their outlets represent a separate ichnogenus Saerichnites B illings (U chm an, 1995).
Hormosiroidea S chaffer is proposed as the p ro p e r nam e for the discussed trace fossils. O lder nam es (see the list o f ichnotaxa included in Hormosiroidea) are either applied for trace fossils recently described under o ther ichnogenera or can be treated as forgotten (nom en oblitum ) (see S tanley &
P ickerill (1993).
S tanley & Pickerill (1993) discussed the taxonom y o f Rhabdoglyphus V assoevich (=Protovirgularia M cC oy in this paper) and Fustiglyphus V ialov. T hey kep t these ichno
tax a separate, as w ell as Hormosiroidea Schaffer. N ev e rth e
less, sim ilarities betw een Fustiglyphus and Hormosiroidea are clearly distinct (Fig. 11), as n oticed by O sgood (1970) and H antzschel (1975), and th ese ich n o tax a are com parable at the ichnogeneric level an d the d ifferences betw een them can be regarded as these at the ichnospecies level. T here
fore, Fustiglyphus V ialov is a ju n io r objective synonym o f Hormosiroidea Schaffer. The cham bers o f Fustiglyphus w ere interpreted also as possible bro o d structures, sim ilarly to interpretation o f Hormosiroidea florentina Schaffer (U chm an, 1995).
C ham berlain (1977) included w ith hesitation, expressed by “ cf.” , a few taxa in Hormosiroidea. T hese are: Halimeda saporta Fuchs (1894a), Hormosira moniliformis H eer (1877), and Arthrodendron difussum (U lrich, 1904). H ow ever, these form s represent body fossils o f large foram ini- fers and cannot be included in Hormosiroidea.
The groove-m ound traces, d escribed from deep-sea floor o f the A rctic C anada B asin w ere com pared w ith Hor-
118
A. UCHMANm osiroidea and R h abdoglyphus (K itchell et al., 1978).
H orm osiroidea annu la ta (V ialov 1971) Figs. 11-13
* 1962 Rhabdoglyphusgrossheimi V assoevich-Boucek & Elias, 146, pi. 7, fig. 1, pi. 8, figs. 2 [copied by Osgood, 1970, pi.
78, figs. 8-9).
partim 1970 Rhabdoglyphus ichnosp. a - Ksi^zkiewicz, 285, fig. lh, к (non fig. 1 i-j).
* 1971 Fustiglyphus annulatus gen. et sp. n. - Vialov, 91, fig. 3.
v 1977 Rhabdoglyphus caliciformis n. ichnosp. - Ksi^zkiewicz, 66, pi. 3, figs. 6, 11, text-fig. 6c.
v 1977 Rhabdoglyphus aff. caliciformis n. ichnosp. - Ksi^zkie- wicz, 66, text-fig. 6d.
v 1977 Rhabdoglyphus sulcatus n. ichnosp. - Ksi^zkiewicz, 67, pi. 3, fig. 9, text-fig. 6e.
v? 1977 Rhabdoglyphus sulcatus n. ichnosp. - Ksi^zkiewicz, 67, pi. 3, fig. 9, text-fig. 6e.
v Rhabdoglyphus spinosus n. ichnosp. - Ksi^zkiewicz, 66, text-fig. 6b, pi. 3, figs. 7-8.
1979 Fustiglyphus roselandensis new species - Boyer, 75, figs.
2-3.
? 1993a Fustiglyphus roselandensis Boyer - Metz, 170, fig. 2.
1993 Fustiglyphus annulatus Vialov — Stanley & Pickerill, 58, figs. 2-3A-D.
partim 1993 Rhabdoglyphus grossheimi Vassoevich - Stanley & Pick- erilI, 62, fig. 51 (copied from Ksi^zkiewicz, 1977, fig. 6b).
For additional synonymy o f Fustiglyphus annulatus Vialov see Stanley & Pickerill (1993).
E m e n d e d d ia g n o sis: Straight or rarely w inding H orm osi
roidea w ith angular, trapezoid, oval, sem ispherical or arcu
ate outline o f cham ber-like bodies, w hich are regularly or irregularly distributed along the string. Thin strings, locally
branched, m ay em erge from the cham ber-like bodies.
M aterial: 10 specimens (UJ TF 181-182, 710, 726, 1175, 1505, 1728, 1780, 2018, one specimen in the Naturhistorisches Hof- museum in Vienna).
Description: As in the diagnosis and in the descriptions o f the Ksi^zkiewicz (1977) ichnotaxa included in H. annulata.
Remarks: Stanley & Pickerill (1993) only partially in
cluded the m aterial o f B oucek & Elias (1962) in F usti
glyphus annulatus (V ialov). T hey retained the B oucek &
E lias’ (1962; pi. 8, fig. 2) specim en in R habdoglyphus grossheim i V assoevich. H ow ever, this cannot be accepted.
A lso in B oucek & Elias (1962, pi. 7, fig. 1 and pi. 8, fig. 2), are illustrated trace fossils w ith cham bers, how ever not so w ell preserved as in pi. 8, fig. 3. T he draw ing o f B oucek &
E lias’ specim en in pi. 8, fig. 2 in S tanley & P ickerill (1993, fig. 5A) is sim plified and does not show im portant details.
All these trace fossils are included in H orm osiroidea annu- lata (V ialov) in this paper.
The specim en U J TF 994 (Fig. 13) probably belongs to H. annulata.
Stanley & P ickerill (1993) regarded the cham bers o f H.
annulata as brood structures, w hich is probable. It is diffi
cult to explain the m orphology o f this trace fossil as a de
posit feeding structure as w as previously suggested (B oucek
& Elias, 1962; C ham berlain, 1977). The thin strings that com e out from cham bers (e.g. S tanley & Pickerill, 1993) and cross bedding, i.e. in the direction o f the sea-floor, can be produced by ju v e n ile tracem akers leaving the cham bers or by predators (Fig. 11).
Fig. 11. Hormosiroidea. A. General model, with and without side branchings. B. Ichnospecies of Hormosiroidea
Strobilorhaphe K si^zkiew icz 1968 Ichnospecies included in Strobilorhaphe Ksi^zkiewicz:
v 1968 Strobilorhaphe clavata n. “sp." - Ksi^zkiewicz, 8, pi. 1 figs. 4-5.
v 1968 Strobilorhaphe pusilla n. “sp." - Ksi^zkiewicz, 8, pi. 1 fig. 6.
v 1970 Strobilorhaphe clavata Ksi^zkiewicz - Ksicizkiewicz 288, fig. 1 d, e.
v 1970 Strobilorhaphe pusilla Ksi^zkiewicz- Ksi^zkiewicz, 288 fig. 1c.
v 1977 Strobilorhaphe clavata K si^zkiew icz- Ksi^zkiewicz, 82 pi. 5, figs. 10-11, text-fig. 1 la-r.
v 1977 Strobilorhaphe pusilla Ksi^zkiewicz - Ksi^zkiewicz, 84 pi. 5, fig. 12.
v partim 1977 Strobilorhaphe glandifer n. ichnosp. - Ksi<vzkiewicz, 84 pi. 11, fig. 16, text-fig. 1 Is-z.
1975 Strobilorhaphe clavata - Hantzschel, W 112, fig. 67, 4a.
1977 Strobilorhaphe clavata Ksi^zkiewicz - Chamberlain, 18 fig. 2z(copied in Stanley etal., 1977; 268, fig. 18, and in) 7D.
1979 Strobilorhaphe clavata Ksi^zkiewicz - Chamberlain, 17 fig. 3.
? 1987a Strobilorhaphe - Pickerill, 129, fig. 3 (copy in: Pickerill 1987b; 387, fig. 3s).
? 1990 Strobilorhaphe ichnosp. - Mikulas, 327, pi. 8, fig. 2.
1990 Strobilorhaphe pussila - Uchman, pi. 1, fig. 5 [lapsus calami].
? 1993 Strobilorhaphe ci. c / a v o / o Ksictzkiewicz-Miller, 23, fig.
6E.
E m e n d e d d ia g n o sis: H orizontal trace fossils consisting o f central stem and num erous lateral short, blunt, clavate branches.
R e m a rk s : The sm ooth version o f O phiom orpha annulata
