Maritime Sediments and Atlantic Geology

Vol. 20 APRIL, 1984 No. 1

Systematic ichnology of the Middle Ordovician Trenton Group. St. Lawrence Lowland, eastern Canada D. TjJULLon and R.H. PlcJayiAM De.pcvitmejtt o-fL Qe.o£ogy, llrviveju>-ULy o/ New Bnuru>uxick, T/iejd£JU-d.on, N.B. £38 5A3 Carbonate sediments of the upper Middle Ordovician Trenton Group between Montreal and Quebec City in the St. Lawrence Lowland, eastern Canada, contain a diverse and abundant trace fossil assemblage consisting of Arenicolites sp., ?Calycraterion sp., Chondrites spp., Circulichnis montanus, Clematis- chnia sp., ?Conostichnus sp., Cruziana problematica, Cruziana sp., cf. Diplichnites sp., Furculosus car- pathicus, Helminthopsis hieroglyphica, Helminthopsis sp., Oichnus paraboloides, Palaeophycus tubularis, Palaeophycus sp., ?Plagiogmus sp., Planolites beverleyensis, P. montanus, Planolites sp., ?Rhizocorallium cf. R. irregulare, ?Rosselia sp., Scalarituba misouriensis, Scolicia sp., Skolithos linearis, Skolithos sp., Teichichnus rectus, Teichichnus sp., Trichichnus sp., Trypanites weisei, Vermiforichnus clarkei and Zoo- phycos sp. as well as informally diagnosed loop, oblique and pronged burrows and bryozoan borings. Of these forms, only Chondrites spp., Palaeophycus tubularis, Palaeophycus sp., Planolites spp., Teichichnus spp. and Trypanites weisei are abundant; the remainder are rare to only moderately common. Neverthe- less, in this paper we describe all the trace fossils in detail and in doing so attempt to resolve several current and controversial problems of nomenclature regarding certain ichnogenera. Sediments of the Trenton Group were deposited initially in lagoons followed in turn by offshore "bar", shallow and, finally, deeper offshore shelf environments. The trace fossils do not exhibit significant variation with respect to these broad depositional regimes and, instead, each environment is character- ized by assemblages typical of the Cruziana ichnofacies as recognized in clastic sequences. The major factor primarily limiting the spatial and temporal trace fossil distribution was substrate and its primary characteristics, in turn a reflectance of food availability and environmental energy levels.

Dans les Basses Terres du St-Laurent de l'Est canadien, les calcaires du Groupe de Trenton (Ordovicien moyen superieur) renferment, entre Montreal et Quebec, un assemblage d'ichnofossiles abondants et varies comprenant: Arenicolites sp., ?Calycraterion sp., Chondrites spp., Circulichnis montanus, Clema- tischnia sp., ?Conostichnus sp., Cruziana problematica, Cruziana sp., cf. Diplichnites sp., Furculosus carpathicus, Helminthopsis hieroglyphica, Helminthopsis sp., Oichnus paraboloides, Palaeophycus tubu- laris, Palaeophycus sp., ?Plagiogmus sp., Planolites beverleyensis, P. montanus, Planolites sp., ?Rhizo- corallium cf. R. irregulare, ?Rossetia sp., Scalarituba missouriensis, Scolicia sp., Skolithos linearis, Sko- lithos sp., Teichichnus rectus, Teichichnus sp., Trichichnus sp., Trypanites weisei, Vermiforichnus clarkei et Zoophycos sp. On y trouve Sgalement des galeries en boucle et fourchues, des terriers obliques ainsi que des perforations de bryozoaires, tous identifies de faipon informelle. De toutes ces formes, seules les suivantes sont presentes en abondance: Chondrites spp., Palaeophycus tubularis, Palaeophycus sp., Planolites spp., Teichichnus spp., et Trypanites weisei; les autres sont rares ou peu communes. N6an- moins, nous decrivons en detail toutes les traces fossiles et tentons par le fait mSme de rSsoudre plusieurs problemes de nomenclature chez certains ichnogenres qui font a l'heure actuelle l'objet de debats. Le dep5t des sediments du Groupe de Trenton s'est initio dans un milieu de lagune puis de barre, de plate-forme de haut-fond et, finalement, de plate-forme d'eau profonde. Les traces fossiles ne varient pas de fagon significative d'un environnement de deposition a l'autre. On remarque plut0t que chaque environnement se caract6rise par des assemblages typiques de l'ichnofacies a Cruziana tel qu'on l'observe dans les sequences detritiques. Le substrat et ses caracteristiques de base, qui refletent la disponibilite de la nourriture et les conditions energetiques du milieu, constituent l'eiement majeur qui a, avant tout, limite la distribution spatiale et temporelle des traces fossiles. [Traduit par le journal] INTRODUCTION Detailed and systematic trace fossil abundant and well-preserved body fossils studies of carbonate rocks, particularly and are generally frustrated by the typi- of Lower Paleozoic age, are far out- cally poor preservation, usually as a re- numbered by comparable studies in their sult of carbonate cementation and re- clastic counterparts. In part this is be- crystallization, of their contained trace cause paleontologists are generally at- fossils. Yet many Lower Paleozoic car- tracted to carbonates by their normally bonate sequences abound in trace fos- sils of various kinds and with careful MARITIME SEDIMENTS AND ATLANTIC GEOLOGY 20, 1-41 (1984) and detailed analysis their systematics

071l-1150/84/0l0001-41$7.50/0 2 FILLION and PICKERILL can be resolved at the ichnogeneric and reviews can be found in the referenced even the ichnospecific level. papers. In this paper we describe the systema- tics of a suite of trace fossils contain- STRATIGRAPHIC, SEDIMENTOLOGIC ed in carbonates of the upper Middle AND ENVIRONMENTAL SETTING Ordovician Trenton Group exposed bet- In the St. Lawrence Lowland between ween Montreal and Quebec City in the Montreal in the southwest and Quebec St. Lawrence Lowland, eastern Canada City in the northeast, a distance of '(Fig. 1). This study is one of the first approximately 250 km (Fig. 1), the up- of its kind in Lower Paleozoic carbon- per Middle Ordovician Trenton Group ate sequences and highlights several con- has been subdivided into many litho- troversial and taxonomical problems re- stratigraphic units several of which, be- garding certain ichnogenera. A com- cause of their overall similarity, are panion paper (Fillion et al. 1984) has candidates for synonymy (Harland and evaluated their spatial and temporal Pickerill 1982). Because such strati- distributional patterns and primary con- graphic revision has not been attempted trolling parameters within the sequence to date, the previously established units but, until now, the systematics remain are retained herein. Figure 2 summarizes unpublished despite the many recordings the Trenton Group stratigraphy and by previous authors of 'burrows', 'tracks' places it in context with underlying and and 'trails' (eg. Flower 1945; Clark et overlying formations. More detailed al. 1972; Clark and Globensky 1975, lithostratigraphic and biostratigraphic 1976a). summaries are available in Clark (1972), As the content of this paper is de- Clark and Globensky (1975, 1976a,b,c) cidedly taxonomic only a brief summary Riva (1968, 1969, 1972). Figure 3 sum- of the stratigraphic, sedimentologic and marizes the essential characteristics environmental setting of the Trenton and interpreted environment of deposi- Group is presented below. More detailed

ng. 1 - Simplified geological map of the St. Lawrence Lowland between Montreal and Quebec City. Numbered localities are: 1. Saint Constant, 2. Tie Bizard, 3. Montreal, 4. Cap Saint Martin, 5. Saint Vincent de Paul, 6. Terrebonne, 7. L'Epiphanie, 8. Saint Roch, 9. Saint Esprit, 10. Saint Jacques, 11. Highway 158/Riviere Ouareau, 12. Highway 158/Riviere Rouge, 13. Pont Corriveau, 14. Barrage savoie, 15. Pont Bournival, 16. Saint Louis de France, 17. Radnor des Forges, 18. Grondines, 19. Saint Casi- mir, 20. Pont Saint Alban, 21. Pont Rouge, 22. Neuvitle. Maritime Sediments and Atlantic Geology 6

SW NE Montreal Joliette Trois-Rivieres St Marc Pont-Rouge Quebec City Utica Group Tetreauviile ;Tetreauvilie ;•-"••; Gtondinti Mimbcf •. .GfondBiwi Member.". Gtondntf Mitnbif.. ••Kieuvilte"-.".."" •Neuville-."..".." '.Neuville .* IH/O* Trenton Mnnrranl -rSatnl Catimir MamtMr-: T^AINT CAIINVR MMIBV Group -£oirit tosW Member - "Saint Commt Member" .... . x _ Deschambault_L _ Deschambaytt •Deschamlpau,J .! J l j ._L _ )eschambaylt _p - Deschambault -L -Deschambaylt i 1 i I , 1 . r~ 1 1 m i.J , t , f -J\^ -Mile End^M -.Ouqreay. . i'. . | \ — ilMTtapp.'.y.'.j^ 1 Black SLeray -_— jj^yT ' t^'V^III'Aigle lili River S Low vi 1 te- - - £ Lowville Group •rPamelia 'I*! ! ! ! -Iftmelia-I-I-I-Z"! "-'^'•K'^'V: Chazy J Laval* 1 7 - T • r Laval \ Group ftPpecambrianiiV-. 1 .(Vecambnan ^ i . Pwoambrian r> mm '•.---•.Mlw'.'n Fig. 2 - Stratigraphic profile of Middle Ordovician lithostratigraphic units between Montreal and Quebec City showing the relationship of the upper Middle Ordovician Trenton Group to underlying and overlying units. Modified after Harland and Pickerill (1982). tion of each formation, generalized de- environment of migrating subtidal shoals, tails of which are given below. bars and large ripples (Harland and The basal Trenton Group formations Pickerill 1982). Additionally, the forma- comprise, from southwest to northeast, tion contains more continuous parallel- the Mile End, Ouareau, Fontaine, Saint laminated or low-angle cross-stratified Alban and Pont Rouge Formations. beds which represent ephemeral beaches These are generally thin (2-6m) and con- developed when shoals and bars were sist of irregular beds of micritic and intermittently built up to sea level. argillaceous calcisiltites which alternate Scouring appears to have been common with thin shales. The shales and most and subtidal channels and runnels cut limestones are thoroughly bioturbated. through the shoals and bars. From Mont- Coquinas, lenses and layers of shell and real northeastwards, these skeletal lime- skeletal hash and muddy skeletal intra- stones developed progressively closer to clastic layers also occur. Each of these shore so that in the Quebec City area formations is similar to the underlying they accumulated along rocky shorelines Lowville and Leray Formations of the and around islands (Harland and Pickerill Black River Group and are interpreted 1984). as having originated in broadly similar Overlying the Deschambault Forma- conditions, that is in a low to moderate tion is the Montreal Formation in the energy outer lagoon or sheltered near- Montreal-Joliette area and the St. Cas4- shore environment which became nar- mir Member of the Neuville Formation rower and more confined from Montreal from Trois Rivieres to Quebec City to Quebec City (Harland and Pickerill (Fig. 2). The former is 110-140m thick 1982). The common occurrence of shell and the latter 40-60m but generally layers indicates a fairly close proximity thins northeastwards. Both consist of to fully marine environments. highly variable limestones interbedded The overlying Deschambault Forma- with dark shales. The limestones are tion, present throughout the St. Law- predominantly thinly and irregularly bed- rence Lowland, is generally 24-28 m ded, bioturbated argillaceous micrites thick, but thins to about 5 m at Mont- but dark micritic rubbly beds or skeletal real, and is characterized by 20cm - lm calcarenites and coquinas (generally 5- thick, fine- to coarse-grained, tabular 10cm but up to 25 cm thick) may also and trough cross-stratified skeletal lime- occur. Their high diversity marine faunas stones with numerous coquinas and and predominantly fine-grained nature layers of shell hash and only minor indicate a shallow subtidal, mainly low shales. These represent high energy de- to moderate energy environment in posits which accumulated in a complex which skeletal layers were produced by 4 FILLION and PICKERILL storm reworking of indigenous shelly SYSTEMATIC PALEONTOLOGY faunas or periodically derived, perhaps As in Hantzschel (1962, 1975), the during storm-ebbs or other unusually trace fossils are described below in strong offshore currents, from shore- alphabetical order. In adopting this ward shoaling environments (Harland scheme, subjective and often overlap- and Pickerill 1982). ping ethological groupings, as utilized The uppermost units of the Trenton by several previous studies (e.g. Osgood Group are the Tetreauville Formation 1970, Chamberlain 1971, Fursich 1974a), (150m thick) in the Montreal- Joliette are avoided and the procedure enables area and the Grondines Member of the ease of reference to a single ichno- Neuville Formation (100-120m thick) from genus. Informally named trace fossils are Trois Rivieres to Quebec City (Fig. 2). listed separately after the formally Both consist of 3-15cm thick argillaceous named ichnogenera. Preservational ter- and dark micritic limestones interbedded minology follows that adopted by with dark shales. Infrequent thin shelly Seilacher (1964), Webby (1969) and Hantz- or skeletal beds, which are commonly schel (1975). For brevity, the geographic graded, are also present. Nodular or and stratigraphic distribution and en- irregularly bedded limestones are com- vironmental occurrence of each ichno- mon and typically progressively increase genus is omitted from the descriptions. in frequency through the sequence. Fully Instead reference should be made to marine faunas dominated by brachiopods Figures 4, 5 and 6 which summarize this and trilobites are sparse but diverse and information. Representative examples of widely distributed whereas bioturbation the majority of collected specimens are is high. The sediments were deposited now housed in the Department of Geo- essentially from suspension in a rela- logy, University of New Brunswick, as tively deep, low energy subtidal environ- Collection S161 - D. Fillion. ment. The infrequent thin skeletal layers are interpreted as distal storm deposits Ichnogenus Arenicolites Salter, 1857. (Harland and Pickerill 1982). Type species - Arenicola carbonaria

FORMATION LITHOLOGY ENVIRONMENT

Interbedded argillaceous and micritic Offshore , deep subtidal , low TETREAUVILLE 150 m limestones and shales. energy shelf subject to periodic NEUVILLE Occasional thin skeletal layers. storm influxes. Grondines Member 100-120m Rubbly or nodular beds increase upwards.

Interbedded fine- coarse limestones Shallow subtidal, low to moderate MONTREAL 110 -140 m with thin shales. energy environment. NEUVILLE Argillaceous micrites predominant Periodic influxes of skeletal Saint Casimir Member 40 - 60 m but also thin skeletal limestones material from shoreward shoaling

GROU P and coquinas. environments.

Cross-stratified skeletal limestones Shallow subtidal shoals, bars , Z DESCHAMBAULT 5-28 m coquinas and shell hash. large-scale migrating ripples and o Minor shales. ephemeral beaches. High energy.

z MILE END Micrites , argillaceous limestones Low energy outer lagoon or LU DC OUAREAU 2-6 m and shale interbeds , with sheltered nearshore environment h- FONTAINE skeletal hash layers and lenses. with periodic influxes of storm and SAINT ALBAN tidal overwashes from offshore PONT ROUGE shoals.

BLACK RIVER GROUP — Mixed tidal flat, lagoonal and nearshore sikicbstics and carbonate muds.

Fig. 3 - Facies analysis and inferred depositional environments of the Trenton Group between Mont- real and Quebec City (after Harland and Pickerill 1982). Maritime Sediments and Atlantic Geology 5

Binney, 1852, by subsequent designation as paired circular openings on upper of Richter (1924, p. 137). bedding plane surfaces of parallel lami- Diagnosis - Vertical U-tubes without nated calcilutites. Burrow height is up spreite (after Fiirsich 1974a). to 18mm and width ranges from 5.6 - 28mm. The tubes are vertical, parallel Discussion - Arenicolites is generally and of identical diameter, range from considered to be both the dwelling and 0.5 - 3mm and are typically lined. In feeding burrow of suspension-feeding U-plane sections they exhibit an evenly (particularly polychaete) annelids (Hakes rounded base. 1976, Chamberlain 1978a) or possibly crustacean-like organisms (Goldring 1962). Remarks - No single specimen provided Though typically a shallow-water form both vertical U-plane and bedding sur- it has also been reported from deep- face expression thus negating ichnospeci- water deposits (e.g. Crimes 1977, fic identification. In bedding plane ex- Pickerill and Keppie 1981, Hill 1981). It pression the Trenton material differs ranges in age from Cambrian-Recent from A. curvatus Goldring, 1962, A. (?) (Chamberlain 1978b). compressus (Sowerby 1829) and A. sub- compressus (Eichwald 1860) in having Arenicolites sp. burrows with a circular cross-section. (Figure 7d) Additionally, they are too small and too regular to be included within A. varia- Material - 8 specimens. balis Fiirsich 1974a. U-Plane sections Description - Specimens are observed compare favourably with A. carbonaria as either vertical U-plane sections or (Binney 1852) whereas bedding plane ex-

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1 c H B N A clarkei sp . sp . tubularis O hieroglyphica missouriensis sp . sp . sp . sp . montanus sp . typ e typ e boring s T carpathicus problematica sp . burrow s burrow s A paraboloides X burrow s Plagiogmus Vermiforichnus A Zoophyco s Loo p Pronge d Bryozoo n Teichichnus rectus ? Conostichus sp . cf . Diplichnites sp Trypanites weise i Cruziano Scalarituba Teichichnus Trichichnu s ? Planolites montanus Planolites sp . ? Rhizocorallium cf.Rjrregulare Clematischnio sp . Cruziano Helminthopsis . sp Palaeophycus Palaeophycus Rusophycu s Obliqu e Furculosus Planolites beverleyensis ? Rosselia sp . ^ Chondrites spp . Chondrites Circulichnis Helminthopsis Oichnus ? Calyeraterion sp . Scolicia sp . Chondrites Skolithos linearis Skolithos sp . Arenicolites

T.

NfeM) F II I ill O M. ? R L M N(SCM) ? A 1 I T D. ' i 1 O ME. N S o. II I II i F. S.A RR. 1 1 II Fig. 5 - Stratigraphic distribution and abundance of trace fossils in the Trenton Group between Montreal and Quebec City. The relative bar widths represent respectively from thinnest to thickest 1<10 specimens, 10<50 specimens, 50<100 specimens and >100 specimens. Questionable occurrences refer to uncertain identifications. Formational nomenclature is P.R. = Pont Rouge, S.A. = Saint Alban, F = Fontaine, O = Ouareau, M.E. = Mile End, D = Deschambault, N(SCM) = Saint Casimir Member (Neuville), .M = Montreal, N(GM) = Grondines Member (Neuville), and T = Tetreauville For- mations. pressions compare with A. sparsus duced by annelids, the smaller depres- Salter 1857) and A. statheri Bather 1925, sions or knobs representing burrow out- particularly with respect to the presence lets. The ichnogenus has, to date, only in some specimens of a distinct wall been recorded in nearshore (Karaszew- lining. Thus, more than a single ichno- ski 1971), deltaic (Chaplin 1980) and in- species may be present but the paucity tertidal environments (Fillion and of material and the nature of its preser- Pickerill, in prep.) from, respectively, vation precludes more detailed analysis. Jurassic, Carboniferous and Lower Ordo- vician strata. The asymmetry of the Ichnogenus Calyeraterion burrow and the smaller knobs or depres- Karaszewski, 1971 sions differentiate Calyeraterion from Type species - Calyeraterion samsono- the closely related ichnogenera Berg- wiczi Karaszewski 1971, by monotypy. aueria Praritl 1945 and Mammillichnis Chamberlain 1971. Diagnosis - Regular and smooth calyx- shaped depressions (on upper bedding ?Calycraterion sp. surfaces) or knobs (on both upper and (Figure 7e) lower bedding surfaces) possessing up to three smaller depressions or knobs (after Material - 1 specimen. Karaszewski 1971). Description - The trace occurs as a Discussion - Karaszewski (1971) con- slightly elongated and ovoid mound pre- sidered the structures to have been pro- served in convex epirelief on the upper Maritime Sediments and Atlantic Geology 7

nose the Trenton specimen as such. In the Newfoundland examples it is evident ICHNOTAXA u. sf that Calyeraterion can be repeated vertically and its preservation as convex Arenicolites sp. ? Calyeraterion sp. epirelief structures is related to subse- Chondrites type A quent infilling of calyx-shaped burrow Chondrites type B outlets. This style of preservation is Chondrites spp. Circu/ichnis monfonus similar to that of Margaritichnus Bandel C/emotischnio sp. 1973 but the Trenton specimen can be ? Conostichus sp. differentiated from this ichnogenus be- Cruziano problematica Cruziana sp. cause Margaritichnus is usually remark- cf. Diplichnites sp. ably symmetrical and flattened. Furculosus carpathicus Helminthopsis hieroglyphica Helminthopsis sp. Ichnogenus Chondrites Oichnus paraboloides von Sternberg, 1833 Palaeophycus tubularis Palaeophycus sp. Type species - Fucoides antiquus Brong- ? Plagiogmus sp. niart 1828, by subsequent designation of Planolites beverleyensis Planolites montanus Miller (1889, p. 114). Planolites sp. Diagnosis - Regularly branching tunnel ? Rhizocoro//ium cf.R.irregu/ore ? Rosselia sp. systems consisting of a small number Rusophycus sp. of mastershafts open to the surface Sco/orifubo missouriensis Scolicia sp. which ramify at depth to form a den- SJcofithos /ineoris dritic network (after Osgood 1970, Fiir- Skolithos sp. sich 1974a). Teichichnus rectus Teichichnus sp. Discussion - Although the trace is easily Trichichnus sp. recognizable at ichnogeneric level, dis- Trypanites weisei Vermiforichnus c/arfcei tinction between the more than 170 sup- Zoophycos sp. posed ichnospecies (Chamberlain 1977), Loop burrows many of which are candidates for sy= Oblique burrows Pronged burrows nonymy, is at present impossible without Bryozoan borings a thorough and detailed taxonomic study. Many of the ichnospecies proposed Fig. 6 - Environmental distribution of trace fossils in ; in the literature are based on variation the Trenton Group between Montreal and Quebec 'in size, preservation and angle of brach- City. Relative bar widths and questionable occurrences are as indicated in Figure 5. ing and our review of this literature has indicated that the majority of these can surface of a 25mm thick calcilutite probably be placed into synonymy with which immediately overlies a 10cm thick the 11 chondritid ichnospecies (from a skeletal calcarenite. The 'mound' is total of 36 Fucoides species - see filled with this bioclastic material, is Osgood 1970) originally designated by 7mm in height and has a diameter of Brongniart (1823, 1828). Chamberlain 9 - 11mm. A poorly preserved knob is (1971, 1977) has in part resolved some present on the upper extremity of the of the ichnospecific nomenclatural prob- structure. lems but the taxonomic status is still Remarks - Since knowledge of the verti- unsatisfactory. Because of these diffi- cal aspect of the structure is unknown, culties we do not attempt to identify even its ichnogeneric assignment is un- our material beyond the ichnogeneric certain. However, the asymmetry and level but recognize various form types the presence of the problematical knob (cf. Osgood 1970). are closely comparable to Ordovician examples of Calyeraterion we have ob- Chondrites has been interpreted as a served on Bell Island, eastern Newfound- plant (Brongniart 1823), the burrow of land and we therefore tentatively diag- a sipunculoid (Simpson 1957), or poly- 11 FILLION and PICKERILL chaete annelid (Osgood 1975), or the exhibits concentric or meniscate struc- burrow of unknown tentacle-bearing tures (cf. Taylor 1967). The largest ob- organisms (Taylor 1967), anthoptiloid sea- served network covers an area of 120 pens (Bradley 1981) and even small arth- x 150mm . ropods (Ekdale 1977). Simpson (1957) and Remarks - The burrow systems frequent- Osgood (1970) have discussed the manner ly run in close proximity without inter- of production and preservation at some penetration, thus suggesting phobotactic length. It is a marine but eurybathic behaviour of the producer. The internal form, having been reported from mar- meniscate structures present in some of ginal marine (e.g. Hakes 1976, 1977; the specimens suggests, at least for Fisher 1978), shallow subtidal (e.g. Hof- these, an origin as stuffed burrows mann 1979, .Marintsch and Finks 1982), rather than open burrow systems (for deep sea fan (e.g. Crimes et al. 1981, a review on the genesis of Chondrites Hill 1981) and abyssal (e.g. Ekdale 1977, see Osgood 1970, Ekdale 1977). Although 1978; Larson 1982) environments. It Chondrites type A has not been formti- ranges in age from Cambrian-Recent ly speciated it bears strong resemblance (Hantzschel 1975). to Buthotrephis succulens Hall (1847, Finally, it is noteworthy that the pi. 22, fig. 2a). nomenclatural type of Chondrites was designated by Miller (1889, p. 130) as Fucoides antiquus and subsequent desig- Chondrites sp. type B nations (e.g. Bassler 1915, p. 217; (Figures 8a, c) Andrews 1955, p. 130) of Fucoides tar- gionii are invalid by rule of priority. Diagnosis - As for ichnogenus. Additionally, Fucoides lycopodioides as listed in the Treatise by Hantzshel Material - Several hundreds of specimens. (1975) and adopted by some subsequent Description - Oblique networks preser- authors (e.g. Marintsch and Finks 1982) ved in endorelief with burrow diameters is in fact an error. This species is the varying from 0.6 - 2mm and burrow nomenclatural type of Caulerpites von cross-sections circular or elliptical. Bur- Sternberg, 1833. row fill is of the same or coarser grain size as the host rock and is ubiquitously Chondrites sp. type A calcilutite or calcisiltite. The fill is ty- (Figures 8a, b) pically of a different colour than the enclosing sediment and is derived from Diagnosis - As for ichnogenus. immediately overlying strata. The tubes, which are up to 25mm long, are often Material - 64 specimens. lined with micrite. Individual networks Description - Dense, horizontal networks may possess 3 orders of branching and preserved in convex epirelief or hypo- have been observed to penetrate at least relief, composed of straight cylindrical 36mm of host sediment. burrows which exhibit regular symmetri- cal dichotomous branching at angles bet- Remarks - These smaller networks of ween 20 - 65°. Burrow diameter varies Chondrites sp. type B also exhibit good from 1.5 - 7mm but is constant within evidence of phobotactic behaviour. They a single specimen. Branches are up to are frequently observed between Chon- 150mm long and a maximum of 5 orders drites type A networks but never pene- have been observed although there can trate them or, indeed, each other. Type be long segments without bifurcations. B networks, although always initially Burrow cross-sections are ellipsoidal and oblique to stratification, may turn to their contact with host material is sharp become parallel at their distal extremi- and well-defined. Burrow fill is variable ties. The producer(s) of the trace is but is typically finer than the host rock, responsible for much of the bioturbation is usually structureless but occasionally observed in the Trenton Group. Maritime Sediments and Atlantic Geology 9

Chondrites spp. cular (or oval) formed by some cylindri- (Figures 8a, d, e, f) cal object" (Vialov 1971, p. 91, trans, Specimens included here exhibit poor litt.). and variable preservation or are too Discussion - The original diagnosis by irregular to permit assignment to types Vialov (1971) is somewhat confusing in A and B as described above. Three vari- - that it is clear in the remainder of his ants can be recognized:- text that the circular (or oval) trace is (i) Elaborate, moderately dense and in fact a burrow of 0.7 - 1.5mm in dia- now flattened networks preserved at meter. Since the original designation it shale-calcilutite interfaces of the Gron- has only been formally recorded from dines Member of the Neuville Formation the Tremadoc of Nova Scotia (Pickerill (Figs. 8a, d). Individual networks are and Keppie 1981). Similar structures have composed of sinuous tunnels which vary been reported and, or, figured but not in diameter along their length from 2 described by various authors (Ulrich - 8.5mm. Up to 5 orders of branching 1904, pi. 22, fig. 3; Hantzschel 1975, are present and the branching is very fig. 44, 2a; Ksiazkiewicz 1977, fig. 32; irregular. Branching angles are equally Nicholson 1978, pi. 3, fig. 2; Crimes irregular varying from 20 - 90°. Burrow et al. 1981, p. 975). Preferably such fill is of the same grain size as the structures should be included within the enclosing material (micrite), is typically ichnogenus Circulichnis with which they structureless but occasionally exhibits best conform. Accordingly, Circulichnis a crudely developed meniscate appear- would range from Tremadoc-Palaeocene ance. (ii) A loose horizontal network age. Although its producer and environ- preserved at a calcisiltite - calcarenite mental range are uncertain Pickerill and bed in the Deschambault Formation Keppie (1981) suggested a worm-like (Figs. 8a, f). Tunnels vary in diameter originator and a preference for deeper- from 2 - 4mm but the diameter of a water regimes. Crimes (pers. comm., single tunnel remains constant along 1983) has also recorded it from deep- its length. The branching is curved, water turbidites of Carboniferous age asymmetrically dichotomous but never in Spain and England. crossing, occurs at angles of 35 - 60° and up to 5 orders can be observed. Circulichnis montanus Vialov, 1971 Burrow fill is micrite. In many respects, (Figure 7g) this network is very similar to C. fur- catus as figured by Marintsch and Finks Diagnosis - As for ichnogenus. (1982, pi. 2, fig. 4) and C. antiquatus as figured by Hofmann (1979, pi. 15, fig. Material - 1 specimen. F). (iii) A widely spaced regularly Description - Slightly elliptical trace branching network of asymmetrically preserved in convex epirelief on the dichotomous tunnels (Figs. 8a, e). upper surface of a 7cm calcarenite. The Individual tunnels are straight and retain trace measures 115 x 90mm and the bur- a constant diameter of 1 - 3 mm. row forming it is 4mm in diameter and Branching angles vary from 30 - 40° consistent in width throughout the el- and up to 2 orders were observed. Bur- lipse. Evidence of burrow collapse is row fill is calcilutite but a dark shale present locally (Fig. 7g). Burrow fill is lining can be observed in some speci- identical to the host rock. mens. This variety resembles C. tar- Remarks - Since Vialov (1971) only gave gionii sensu Marintsch and Finks (1982). size measurements of his holotype, the full variation in ellipse size and burrow Ichnogenus Circulichnis Vialov, 1971 diameter of the original material is un- Type species - Circulichnis montanus, known. In view of the incomplete des- Vialov, 1971, by original designation. cription and the unavailability of type material for restudy, the Trenton speci- Diagnosis - "Annular trace, almost cir- men can best be regarded as conspeci- 10 FILLION and PICKERILL

fic. Its occurrence in the Trenton ducer and stratigraphic range. Its simi- Group, although rare, suggests that it larity to chondritids and Palaeophycus is possibly more a eurybathic form. suggest, however, that it is the feeding and dwelling burrow of an unknown an- Ichnogenus Clematischnia Wilson, 1948 nelid. Similarly, although to date it has only been reported from Ordovician Type species - Buthotrephis succulens strata, its stratigraphic range is likely Hall, 1847, by original designation of to be considerably more extensive. Wilson (1948, p. 10). Diagnosis - Irregularly branched or un- Clematischnia sp. branched cylindrical burrows with promi- (Figure 7i) nent external ornament of transverse ribs which completely or partially cross Material - 12 specimens. the burrow; in the latter case the bur- row assumes a distinctive mammilated Description - Unbranched cylindrical bur- appearance (modified after Wilson 1948). rows up to 7mm in diameter and 10cm in length preserved on upper surfaces Discussion - Wilson (1948) designated and all parallel to stratification. In all Buthotrephis succulens Hall as her type of the observed specimens the burrows ichnospecies of Clematischnia but it is occur in groups of four to seven, ap- apparent that Hall's (1847) two figured parently radiating from some common specimens are morphologically separate; centre which is not preserved. Burrow his figure 2a, plate XXI is more akin surfaces are mammilated, this being to a chondritid (Chondrites type A as produced by an alternating series of described herein) and his figure 2b, plate transversely oriented interlocking pear XXI falls within Clematischnia as de- or cylindrical shaped protruberances, fined by Wilson (1948). Furthermore, even each separated by a narrow depression. Wilson's (1948) material is taxonomically Burrow fill (siltstone) is coarser than the variable; her plesiotype G.S.C. 9283 surrounding matrix and is structureless. (plate 1, fig. 4) is likewise more akin to Palaeophycus sp. or Chondrites sp. Remarks - Congeneric material from the whereas her plesiotype G.S.C. 13442 Trenton Group immediately around Que- (plate 1, fig. 5) falls within Clematis- bec City was previously identified by chnia and is very similar to Hall's (1847) Pickerill and Forbes (1979) as cf. Bifor- figure 2b. Clearly a detailed systematic mites. However, both their material and review of the ichnogenus is warranted that described herein lacks the initial but until this is undertaken we regard short and longitudinally furrowed section it as a useful form to describe these more typical of Biformites and the distinctive mammilated burrows. Because material is therefore best included with- of the rarity of actual recordings of in Clematischnia. In view of its unresol- Clematischnia (for example the ichno- ved taxonomic status it is herein only genus was only listed as an unrecognized identified at ichnogeneric level. form in Hantzschel (1975) and briefly discussed by Osgood (1970) in his re- Ichnogeneus Conostichus Lesquereux, 1876 description of Buthotrephis succulens) Type species - Conostichus ornatus Les- it is difficult to comment on its pro- quereux, 1876, by original monotypy.

Opposite page Fig. 7 - a. Cruziana sp. on lower bedding surface in convex hyporelief, Grondines Member, locality 22. b. Cruziana problematica on lower bedding surface in convex hyporelief, Deschambault Formation, locality 12. c. Scolicia sp. and Diplichnites sp. (arrowed) preserved on upper bedding surface of thin chert, Grondines Member, locality 18 (see text for details), d. Arenipolites sp. tubes (arrowed) on upper bedding surface, Saint Casimir Member, locality 16. e. ?Caly- craterion sp. on upper bedding surface in convex epirelief with arrow indicating possible location of burrow outlet, Deschambault Formation, locality 21. Coin diameter 2 cm. f. ?Conostichus sp. actually preserved on lower bedding sur- face in convex hyporelief but here reversed to exhibit faint longitudinal fluting, Grondines Member, locality 18. g. Circulichnis montanus on upper bedding surface in convex epirelief, arrow indicating burrow collapse, Deschambault Formation, locality 21. h. Rusophycus sp. on lower bedding surface in convex hyporelief, Grondines Member, locality 22. i. Clematischnia sp. on upper bedding surface in convex epirelief, Pont Rouge Formation, locality 21. Maritime Sediments and Atlantic Geology 11 12 FILLION and PICKERILL

Fig. 8 - a. Variation in chondritids in the Trenton Group. 1 = Chondrites sp. type A, 2 = Chondrites sp. type B, 3 = Chondrites sp. dense and elaborate networks, 4 = Chondrites sp. loose horizontal networks, and 5 = Chondrites sp. widely spaced horizontal regularly branching networks (see text for details). All redrawn from actual specimens and all x 0.5. b. Chondrites sp. type A on upper bedding surface, Pont Rouge Formation, locality 21. c. Chondrites sp. type B in vertical cross-section, Tetreauville Formation, locality 8. d. Dense horizontal network of Chondrites sp. on lower bedding surface, Grondines Member, locality 18 (cf. Fig. 8a [3]). e. Loose horizontal network of Chondrites sp. on upper bedding surface, Deschambault Formation, locality 21 (cf. Fig. 8a [5]). f. Chondrites sp. on upper bedding surface, Deschambault Formation, locality 16 (cf. Fig. 8a [4]). Maritime Sediments and Atlantic Geology 13

Diagnosis - Variable but mostly small regularity of the structure, its associ- conical to subconical structures preser- ation on the same bedding surface with ved in full relief. Bodies commonly abundant trace fossils, particularly Chon- fluted by transverse constrictions and drites sp. type B and Planolites bever- longitudinal furrows and ridges originat- leyensis, and a complete absence on ing from the apical end. Concentric that bedding surface of additional physi- laminae may be present internally (after caly produced inorganic structures. Hantzschel 1975, Hakes 1976). Discussion - Following its original inter- Ichnogenus Cruziana d'Orbigny, 1842 pretation as an alga (Lesquereux 1876) Type species - Cruziana rugosa d'Orbigny the > structure has been interpreted as 1842, by subsequent designation of Miller a sponge, a scyphomedusoid, a burrow (1889, p. 115). of a filter-feeding organism, a funnel- shaped opening of Arenicola-like bur- Diagnosis - Elongate band-like furows rows and a trace produced by upward covered by herringbone-shaped ridges, migration of actinian-like organisms (see with or without two outer sooth or fine- Pfefferkorn 1971, Chamberlain 1971, Bar- ly longitudinally striated zones outside thel and Barth 1972, Hakes 1976). Its the V-markings, occasionally with lateral origin still remains enigmatic and, in- ridges and, or, wisp-like markings when deed, its producers may be polyphyletic. preserved on bedding soles (after Hantz- Many of its numerous species are prob- schel 1975). ably synonymous and it is a prime can- Discussion - Although Seilacher (1970) didate for serious taxonomic revision united under Cruziana both long furrows (see Branson 1960). It ranges in age from and short coffee bean-like excavations Arenig (pers. observ. - Wabana Group (=Rusophycus) most subsequent authors of eastern Newfoundland) to ?Lower have, as we do, preferred to retain the Cretaceous (Imlay 1961) and, although two as distinctive ichnogenera (see exceptions . do occur (e.g. Chamberlain Crimes et al. 1977). Furthermore, we 1971), is typically associated with near- herein accept the reasoning by Bromley shore environments subject to rapid and Asgaard (1979) to include Isopodich- sedimentation. nus Bornemann, 1899 within Cruziana as the only essential differences between ?Conostichus sp. the two previously separated ichnogenera (Figure 7f) are in accessory features (sensu Fiirsich Material - 1 specimen. 1974b) which should be employed only for classification at the ichnospecific Description - A truncated conical struc- level. It is, nevertheless, clear that ture 15mm in diameter and 13mm in Bromley and Asgaard (1979) 'lumped' height preserved in convex hyporelief heterogeneous material into a single below a 25mm-thick bioclastic calcilu- ichnospecies (C. problematica) and tite. Poorly preserved longitudinal ribs revision of their and other material is (ca. 6-7) are present. still necessary. Remarks - The specimen, although not Undoubtedly, the majority of Cruziana well enough preserved for definite furrows were produced by trilobites identification, may be distinguished from (Seilacher 1970), although the small Tri- the allied ichnogenus Conichnus Myannil, assic forms described by Bromley and 1966 by the presence of longitudinal Asgaard (1970) may have been produced fluting. Additionally, Bergaueria Prantl, by trilobite-like arthropods or notostra- 1945 is more flattened and less conical. can branchiopods and the large Middle A more important difficulty is demon- Ordovician and Triassic forms described strating that the structure is in fact by, respectively, Fisher (1978) and Shone organic and not inorganic in origin. The (1978, 1979) may be the result of aglas- former, however, is suggested by the pids or even vertebrates. Many ichno- 17 FILLION and PICKERILL

species of Cruziana now exist and it is (sensu Trewin 1976) typical of C. strom- perhaps the most useful of all trace fos- nessa and the well-defined scratches, sils in stratigraphic studies (Seilacher often arranged in a V-pattern, of C. 1970, Crimes 1975). The ichnogenus eutendorfensa. Conspecific material was ranges in age from Cambrian - Triassic described by Pickerill and Forbes (1979) and is a typically shallow-water marine from the Trenton Group northeast of form constituting the type ichnogenus Quebec City. of Seilacher's (1967) Cruziana ichno- facies, though it has also allegedly been Cruziana sp. observed in freshwater deposits (e.g. (Figure 7a) Helwig 1972). Material - 8 specimens. Cruziana problematica (Schindewolf, 1921) Description - The specimens consist of (Figure 7b) variably preserved, straight or slightly sinuous bilobate furrows preserved in Diagnosis - Straight or curved, often convex hyporelief. Mean width is 20mm self-crossing, Cruziana up to 5mm wide and length is variable, up to a maximum possessing faint, unevenly spaced trans- observed of 100mm. Endopodal lobes are verse striations which may reach the poorly to moderately well-preserved giv- margins of the trace in low relief speci- ing the trace a depth of l-8mm. V- mens or terminate before reaching the markings, when preserved, are moderate- margins in high relief specimens (adapt- ly deep, closely spaced and V at 160 - ed from Schindewolf 1928, Hantzschel 180°. Genal spine markings are absent. 1975, Trewin 1976, Bromley and Asgaard 1979). Remarks - Though not well enough pre- Material - 2 specimens. served for ichnospecific identification the furrows can be clearly diagnosed as Description - Two poorly preserved Cruziana. They are reminiscent of speci- specimens, one in convex hyporelief at mens described as Cruziana sp. by the base of a 30mm thick calcisiltite, Alpert (1976, pi. 1, fig. 4) from the the second in concave epirelief on an Cambrian of California and also re- 80mm thick calcilutite. The former is semble C. pudica described from Middle a short 8mm long straight band-like Ordovician to Middle Silurian strata of trace, 4mm wide with lobes 1.5mm deep the United States, southern Jordan and and the latter a 120mm long slightly Wales by, respectively, Osgood (1970), sinuous trace, 2mm in width, composed Seilacher (1970) and Pickerill (1977). Con- of two furrows each 0.7mm wide separ- generic material has also been figured ated by a median ridge. Both specimens by Pickerill and Forbes (1979) from the possess transverse but poorly preserved Trenton Group northeast of Quebec and faint scratch markings irregularly City. developed along their exposed length. Remarks - Although poorly preserved Ichnogenus Diplichnites Dawson, 1873 the material lacks the 'Isopodichnus' - Type species - Diplichnites aenigma type bilobed expanded terminations Dawson, 1873, by original montypy.

Opposite page

Fig. 9 - a. Furculosus carpathicus on lower bedding surface in convex hyporelief, Montreal Formation, locality 11. b. Unbranched Palaeophycus tubularis on upper bedding surface in convex epirelief, Deschambault Formation, locality 21. Coin diameter 2 cm. c. ?Plagiogmus sp. on upper bedding surface in convex epirelief, Deschambault Formation, locality II. d. Helminthopsis sp. on upper bedding surface in convex epirelief, Deschambault Formation, locality 21. Coin diameter 2 cm. e. Helminthopsis hieroglyphica on upper bedding surface in convex epirelief, Pont Rouge Forma- tion, locality 21. Pen length 10 cm. f. Lined Planolites sp. on upper bedding surface in concave epirelief, Grondines Member, locality 18. Coin diameter 2 cm. g. Planolites beverleyensis on upper bedding surface in convex epirelief, Pont Rouge Formation, locality 21. h. Planolites montanus on upper bedding surface, Grondines Member, locality 18. i. Helminthopsis sp. (arrowed) and Palaeophycus sp. (to left of coin) on upper bedding surface in convex epirelief, Deschambault Formation, locality 21. Coin diameter 2 cm. Maritime Sediments and Atlantic Geology 16 FILLION and PICKERILL Maritime Sediments and Atlantic Geology 17

Diagnosis - Morphologically simple trails 1mm wide and 1mm deep and set width up to 36cm wide and consisting of two (sensu Osgood 1970, p. 354) is 7-lImm. parallel series of tracks (each up to 9cm Up to 13 sets have been observed in a wide); individual tracks elongate, closely single track, giving a maximum length and regularly spaced up to one per of 45mm. centimeter and roughly normal to trace Remarks - The poor preservation pre- axis (emend. Briggs et al. 1979). cludes positive identification even at Discussion - Although Dawson (1873, p. ichnogeneric level, although of all pub- 7) originally interpreted Diplichnites as lished 'trackways' they compare most the result of 'a large crustacean or favourably to Diplichnites. gigantic annelid or myriapod' subsequent workers, following Seilacher (1955), have Ichnogenus Furculosus frequently utilized the ichnogenus to Roniewicz and Pienkowski, 1977 describe morphologically similar but Type species - Furculosus carpathicus smaller scaled trilobite-produced traces Roniewicz and Pienkowski, 1977, by (e.g. Crimes 1970, Osgood and Drennen original designation. 1975). Although Briggs et al. (1979) questioned the trilobite origins of origi- Diagnosis - Cylindrical burrows forming nally described specimens and suggested tight fork-line loops with endings paral- a more restricted usage to exclude trilo- lel or diverging. bite trails, suggesting that those speci- Discussion - Roniewicz and Pienkowski mens of Diplichnites previously inter- (1977) hypothesized that the loops of F. preted as trilobite trails to be included carpathicus were formed by a deposit- in one of the junior synonyms listed by feeding organism passing through an in- Osgood (1970). the taxonomic confusion terface boundary. Although to date no has still not been resolved. Because of other occurrence of the ichnogenus has the limited material and its poor formally been made, similar forms are preservation and the unresolved taxo- known in the external Carpathians nomic difficulties, we herein retain (Kai^zkiewicz, pers. comm. in Roniewicz Diplichnites even though we assume it and Pienkowski 1977). The occurrence to have been produced by trilobites. of Furculosus in the Trenton Group ex- The ichnogenus has been reported from tends its known stratigraphic range from Cambrian-Triassic strata and though Middle Ordovician - Oligocene and its typically a shallow marine form (e.g. environmental range from shallow to Crimes 1977) has also been recorded deep-water regimes. Gordia arcuata from deep-water sediments (e.g. Ksicjzkiewicz, 1977 is a morphologically Pickerill 1980, 1981). similar form but can be distinguished from Furculosus by its more arcuate cf. Diplichnites sp. shape and smaller burrow diameter. (Figure 7c) Furculosus carpathicus Material - 2 specimens collected and 5 Roniewicz and Pienkowski, 1977 recorded in the field. (Figure 9a) Description - Paired(?) elongated dot- Diagnosis - As for ichnogenus. like impressions preserved in concave epirelief on the upper surface of a chert Material - 1 specimen. bed. Each impression is 2.5mm long, Description - Tight-looped smooth and

Opposite page Fig. 10 - a. Trichichnus sp. (arrowed) in vertical section, Montreal Formation, locality 14. b. Zoophycos sp. on upper bedding surface in convex epirelief, Grondines Member, locality 18. Note the peripheral tube (arrowed), c. Teichichnus rectus in transverse vertical section with the longitudinal burrow arrowed, Grondines Member, locality 21. d. Thickly lined Skolithos sp. burrow (arrowed) in transverse vertical section, Pont Rouge Formation, locality 21. e. Scalarituba missouriensis on upper bedding surface in convex epirelief, Ouareau Formation, locality 11. f. g. ?Rosselia sp., f. on upper bedding plane surface, and g. in transverse vertical section, Saint Casimir Member, locality 16. h. Teichichnus sp. on upper bedding surface, Grondines Member, locality 18. 18 FILLION and PICKERILL cylindrical burrow preserved in convex and is retained herein as the type. hyporelief on the base of a parallel It must be noted that although most laminated calcarenite. Burrow diameter authors have utilized the ichnogenus for is 3.5 - 4mm and the loop is 35mm long winding or loosely meandering, unbranch- and 11mm wide. The burrow terminates ed, generally unornamented 'burrows', as a result of both distal extremities the original diagnosis did not specify turning up into the overlying cal- that the traces had necessarily to be carenite. Burrow fill is calcarenite. of burrow origin. Thus, it seems logical Remarks - Although Roniewicz and Pien- that morphologically and geometrically kowski (1977) discussed the 3-dimensional similar 'trails' (e.g. Hall 1852, pi. 11, fig. aspect of F. carpathicus they omitted 2b; Dorjes 1978, fig. 4 and Swinbanks to include it in their diagnosis, accord- and Murray 1981, fig. 10A) should also ing to which, knowledge of the bedding be included within the ichnogenus. plane expression of the burrow is suffi- Many authors, too numerous to men- cient for positive identification. The tion here, have reported the ichnogenus tight fork-like loop is therefore the in strata of ?Late Precambrian (Webby most characteristic feature of the 1970, fig. 17, H. hieroglyphica) to Oligo- species and the Trenton material is cene (Ksiazkiewicz 1977) or even Recent therefore regarded as conspecific. (Dorjes 1978, Swinbanks and Murray 1981) age. It is a eurybathic form, though Ichnogenus Helminthopsis Heer, 1877 more frequently reported from deep- water flysch successions (Pickerill 1981) Type species - Helminthopsis abeli, by and is generally regarded as having been subsequent designation of Ksiazkiewicz produced by polychaete and possibly (1977, p. 116). priapulid annelids. Diagnosis -' "Unbranched, very long, very much snake-like winding tubes or Helminthopsis hieroglyphica cylinders of very different sizes. Hel- Heer in Maillard, 1887 minthopsis magna was undoubtedly tubu- (Figure 9c) lar but this is uncertain for the two other species .... The outer surface is Diagnosis - Helminthopsis in which the smooth. It is similar to Helminthoida windings are irregular, low and the Schafh. in which, however, the meanders course of the trace is often alternately are closer and the string-like or cylin- winding and straight (after Ksiazkiewicz drical body touches itself" (Heer 1877, 1977). p. 116 - trans, litt.). Material - 7 specimens. Discussion - Heer (1877) originally estab- lished three species of Helminthopsis but Description - The traces are moderately did not designate a type. Subsequently, well-preserved in convex epirelief on the Ulrich (1904, p. 144) designated H. magna upper surface of a massively bedded a somewhat unfortunate choice as this crinoidal calcarenite. They consist of trace is a bilobate structure (see Heer burrows, up to 2cm in diameter and 1877, pi. XLVII, figs. 1,2) resembling 50cm in length which are characterized Taphrhelminthopsis Sacco, 1888 or Scoli- by long straight segments up to 30cm cia de Quatrefages, 1849. In view of this long, but usually smaller, with a low problem Ksiazkiewicz (1977) proposed to turn between individual segments. Bur- utilize a specimen originally figured by row diameter is constant along the Abel (1935, p. 290, fig. 261B) and subse- length, burrow fill is identical to the wuently illustrated by Hantzschel (1962, enclosing calcarenite material and the p. W197, fig. 4a; 1975, p. W71, fig. 2b) burrows are smooth and unornamented. as the type ichnospecies under the name Remarks - As noted by Ksiazkiewicz Helminthopsis abeli. Compared to H. (1977, p. 119), this ichnospecies is not magna this latter ichnospecies conforms well defined and the material originally best to Heer's (1877) original diagnosis figured in Maillard (1887, pi. 2, fig. 4; Maritime Sediments and Atlantic Geology 19

pi. 1, fig. 2) resembles both a Helmin- of biogenic origin bored into hard sub- thoida (pi. 1, fig. 2) and a loose irregu- strates. The hole may pass right through larly winding trace (pi. 2, fig. 4) which the substrate as a penetration, where Ksiazkiewicz argued be adopted as the the substrate is a thin shell; or end nomenclatural type of the species. To within the substrate as a shallow to maintain stability this approach is adopt- deep depression or short, subcylindrical ed herein. pit (Bromley 1981).

Helminthopsis sp. Remarks - The borings are considerably (Figures 9d, i) shorter than Trypanites Magdefrau, 1932, range in age from possibly early Cam- Material - 5 specimens and numerous brian to Recent, and certainly Tertiary occurrences recorded in the field. and younger examples are produced by Description - Variably preserved burrows predatory snails, as more fully discussed preserved in convex and concave epi- in Bromley (1981). relief on upper surfaces of thinly bed- ded calcilutites or bioclastic calcisil- Oichnus paraboloides Bromley, 1981 tites. Burrows are smooth, slightly sinu- (Figure 11a) ous or meandering but meanders are al- Diagnosis - Oichnus having a spherical ways incomplete. Burrow fill is coarser paraboloid form, truncated in those or of the same grain size as underlying cases where the boring penetrates right sediments. They are up to 150cm in through the substrate. Where it does not length and range in diameter from 1.8 so penetrate, the paraboloid may be - 30mm (mean 7mm), the diameter re- deformed by a slightly raised central maining constant in a single specimen. boss (Bromley 1981). Collapse in some specimens has resulted in their preservation as two epichnial Material - 1 specimen. ridges separated by a narrow furrow. Description - A round perforation within Remarks - The most workable scheme the pygidium of the trilobite Isotelus for ichnospecific identification of Hel- gigas, 1.2mm in diameter at the initial minthopsis is that proposed by Ksiazkie- point of penetration of the dorsal sur- wicz (1977), who utilized the nature and face narrowing to 1mm at its ventral type of winding, the burrow diameter exit. Walls of the truncated paraboloid and the presence or absence of surface are smooth. ornamentation. Because of their general- ly poor and incomplete preservation, Remarks - Although we observed no particularly regarding the meander pat- evidence of etching patterns on the bor- terns, the Trenton specimens can there- ing it is identical to and diagnosed as fore best only be identified at ichno- O. paraboloides Bromley, 1981, as O. generic level. Although Helminthopsis simplex Bromley, 1981 possesses a cylin- is generally regarded to be a deposit- drical to subcylindrical form. The pro- feeding burrow (Ksiazkiewicz 1977) the ducer of the Trenton specimen remains presence of collapsed tunnels in some enigmatic. of the Trenton specimens suggests, at least for these specimens, that they re- Ichnogenus Palaeophycus Hall, 1847 mained open after the passage of the Type species - Palaeophycus tubularis producing organism(s). Hall, 1847, by subsequent designation of Miller (1889, p. 130). Ichnogenus Oichnus Bromley, 1981 Diagnosis (emended) - Branched or un- Type species - Oichnus simplex Brom- branched, straight to slightly curved ley, 1981, by original designation. to slightly undulose or flexuous, smooth Diagnosis - Circular to subcircular holes or ornamented, lined essentially cylin- 20 FILLION and PICKERILL

drical, predominantly horizontal burrows Diagnosis - Smooth, unornamented Pala- of variable diameter; infillings typically eophycus of variable diameter, thinly but structureless, of same lithology as host distinctly lined (Pemberton and Frey rock (after Pemberton and Frey 1982). 1982).

Discussion - There is still much con- Material - Several hundreds of speci- fusion regarding the original designation mens. of the type of the ichnogenus Palaeo- phycus. In their thorough and detailed Description - Straight to slightly sinu- review of Palaeophycus and Planolites ous, generally smooth, horizontal to Pemberton and Frey (1982) followed slightly inclined, frequently collapsed Hantzschel (1975) and suggested the type cylindrical burrows, l-24mm in diameter was P. tubularis Hall, 1847, subsequent- and up to 57cm in length. Burrow dia- ly designated by Bassler (1915, p. 939). meter is constant along the length of At an earlier date, however, Miller a single burrow. True branching is rare, (1889, p. 130), in a frequently overlooked but when observed is irregular and oc- and comprehensive text, had also desig- curs at 30-70°; cross-cutting is common. nated P. tubularis Hall, 1847. Unfortun- Burrow walls are thinly lined, burrow ately, both Bassler (1915) and Miller fill is identical to enclosing material (1889) did not design any type material, and rare backfill structures are present. and indeed, the first to designate such The burrows are preserved in convex material was Andrews (1955, p. 202). hyporelief and convex and concave epi- In the absence of any formal and uni- relief typically on thin beds of fine cal- versal code, however, at present we carenite, calcisiltite or calcilutite. regard the subsequent designation of Miller (1889) and not Bassler (1915) as Remarks - Some of Hall's (1847) original the most acceptable. material which he described as P. rugo- sus and P. simplex, but now both includ- Minor emendation of the diagnosis of ed in P. tubularis, was collected from Palaeophycus by Pemberton and Frey the Trenton Group of New York and (1982) is considered necessary to exclude therefore the ichnospecies is apparently systematic and non-systematic winding widespread and abundant throughout the and, or, meandering burrows (e.g. Gordia Group. Although we have chosen to Emmons, 1844, Helminthopsis Heer, 1877, adopt the classification scheme of Pem- Cochlichnus Hitchcock, 1858). Otherwise, berton and Frey (1982), some of the the review of the taxonomy of Palaeo- Trenton P. tubularis specimens illustrate phycus by Pemberton and Frey (1982), some of the difficulties of its universal following several decades of inconsistent application. In particular, several speci- usage and confusion, is considered the mens were observed to pass from cal- most workable nomenclatural system and cilutites and calcisiltites into and even is adopted herein, even if problems may through patchily distributed lenses of arise in the case of concealed bed-junc- calcarenite, though still retaining a cal- tion preservation (Hallam 1975), inter- cisiltite or calcilutite fill throughout. face-crossing burrows (Hofmann 1979) or Had only the portion of the burrows faintly meniscate burrows (Clifton and within the calcarenite been observed Thompson 1978). then they would have had to be refer- Palaeophycus is a eurybathic form red to as Planolites which, of course, ranging in age from Proterozoic to Re- would necessitate, in such cases, er- cent (Hantzschel 1975) and is usually roneous ethological interpretations. attributed to represent the burrows of Nevertheless, in most cases it was pos- predaceous or suspension-feeding organ- sible to observe the transition and such isms, commonly but not universally poly- 'compound' specimens were named chaete annelids. according to their dominant component Palaeophycus tubularis Hall, 1847 (cf. Bromley and Frey 1974, Frey et (Figure 9b) al. 1978). Maritime Sediments and Atlantic Geology 21

Fig. 11 - a. Oichnus paraboloides (upper arrow) and Vermiforichnus clarkei (lower arrow) in the pygidium of the isotele- nid Isotelus gigas, Grondines Member, locality 18. b. Micrite filled borings of Trypanites weisei in a hemispherical colony of the bryozoan Prasopora sp., Pont Rouge Formation, locality 18. c. Longitudinal section through the bryozoan Prasopora sp. illustrating essentially vertical Trypanites weisei borings, Grondines Member, locality 18. d. Numerous small borings of Vermiforichnus clarkei in the cephalon of the isotelenid Isotelus gigas, Grondines Member, locality 18. e. f. Two views of reticulate bryozoan borings on a pygidium of the isotelenid Isotelus gigas, with f. an enlargement of the central area exhibited in e. Grondines Member, locality 18. 22 FILLION and PICKERILL

Palaeophycus sp. (1897) but not formally erected until (Figure 9i) 1929 by Roedel, who defined two species, P. simplex and P. arcuatus, the Material - 67 specimens. latter subsequently being designated as Description - Ellipsoidal, thinly lined bur- the type by Hantzschel (1962). Roedel (1929) believed the structures were trails rows, 3-7mm in diameter and observed produced by polychaetes, but later mainly as full relief structures in trans- Glaessner (1969) and more recently Jae- verse section. Burrow fill is identical ger and Martinsson (1980), who restudied to host rock and is apparently structure- the type material, demonstrated their less. The few examples observed in true burrow nature. Although the nature semirelief are poorly preserved and in- of the producing organism is enigmatic, complete, but essentially straight to molluscs (Glaessner 1969, Crimes et al. slightly sinuous, horizontal to gently 1977) and segmented annelids (Roedel inclined, smooth, unbranched and rarely 1929, Whitaker 1979 - described as branched at up to 25°. Steinsfjordichnus brutoni) have been con- Remarks - Since the specimens are ob- sidered as likely producers. Jaeger and served essentially in transverse section Martinsson (1980) have demonstrated that and their complete 3-dimensional char- irrespective of the geological affinities acter is unknown they are only identi- of the producer, the movement was con- fied at ichnogeneric level. ' Semirelief trolled and locomotory and not peristalic specimens are apparently smooth and in any sense. therefore possibly P. tubularis. This smoothing, however, could be a result The ichnogenus commonly occurs in of weathering as the specimens are bad- strata of early Cambrian age (e.g. Banks ly preserved and weathered throughout. 1970, Alpert 1977, Crimes et al. 1977) P. heberti (de Saporta 1872-73) can be though Hantzschel (1975, p. W95) reports dismissed because of the presence of it from the Upper Precambrian of only thin linings, but whether or not the Russia. If, as we suspect, specimens of material also represents one or several Steinsfjordichnus brutoni described by of the ichnospecies P. striatus Hall, Whitaker (1979, p. 87) are congeneric, 1852, P. sulcatus (Miller and Dyer 1878) its stratigraphic range must be extended or P. alternatus Pemberton and Frey, to the Lower Devonian. It has previously 1982, whose differentiation is based on been recorded from shallow marine external sculpture, is unknown. (Banks 1970) to possibly fluvial and lacustrine environments (Whitaker 1979). Ichnogeneus Plagiogmus Roedel, 1929 ?Plagiogmus sp. Type species - Plagiogmus arcuatus (Figure 9c) Roedel, 1929, by subsequent designation of Hantzschel (1962, p. W210). Material - 1 specimen recorded in the Diagnosis - Horizontal endichnial burrow, field. straight or slightly curved 10 - 28mm Description - Preserved in convex epi- wide, possessing sausage-shaped or fusi- relief on the upper surface of a 15cm form transverse ventral ridges (as viewed thick massive bioclastic calcilutite as on upper surfaces) and corresponding a series of 13 parallel and equally spaced dorsal furrows (as viewed on lower sur- (4mm) ridges, each 1mm high, 2mm wide faces), the spacing of which varies con- and 10.5mm long, oriented transversely siderably. Narrow smooth zones separate to the trace axis. The areas between the ends of the ridges from the margins the ridges are smoother, flatter and of the trace (after Hantzschel 1975, slightly wider. Burrow walls are not pre- Jaeger and Martinsson 1980). served. Discussion - Plagiogmus was originally Remarks - Poor preservation and lack described from Sweden by Nathorst of knowledge of the 3-dimensional struc- Maritime Sediments and Atlantic Geology 23

ture precludes positive identification even Planolites beverleyensis Billings, 1862 at ichnogeneric level but the specimen (Figure 9g) does resemble Plagiogmus to which it is tentatively referred. The specimen is Diagnosis - Relatively large, smooth, very similar, both in scale and morpho- straight to gently curved or undulose logy, to Steinsfjordichnus brutoni cylindrical Planolites (after Pemberton Whitaker, 1979, which as noted previous- and Frey 1982). ly, is also best included in Plagiogmus. Material - 21 specimens and several hun- dreds observed in the field. Ichnogenus Planolites Nicholson, 1873 Description - Smooth, straight to slightly Type species - Planolites vulgaris curved, horizontal to slightly inclined Nicholson and Hinde, 1875, by subse- (up to 20°) regularly cylindrical, un- quent designation of Miller (1889, p. branched or branched (at varied angles 520). of 20-115°), isolated or cross-cutting bur- Diagnosis (emended) - Unlined, rarely rows. Burrow fill is typically coarser lined, rarely branched, straight to tortu- than the host rock but may be finer ous, smooth to irregularly walled or an- and of different colour. Burrow fill is nulated burrows, circular to elliptical typically structureless but rare speci- in cross-section, of variable dimensions mens exhibit thin shale or micritic lin- and configurations; infillings essentially ings up to 9mm thick which may be structureless, differing in lithology from composed of a series of concentric host rock (after Pemberton and Frey laminae, giving the burrows an apparent 1982). central core. Specimens are preserved Discussion - Although Alpert (1975, p. in convex or concave epirelief, convex 512) was the first to designate a type hyporelief and endorelief in a wide specimen of Planolites, Miller (1889, p. variety of lithologies and vary from 5- 520) and later Bassler (1915, p. 982) were 20mm in diameter and 2.7-100cm in the first authors to designate the (same) length. type species. Like Palaeophycus, Plano- Remarks - Although the diagnosis pro- lites has been a topic of considerable posed by Pemberton and Frey (1982) did confusion since its original establishment not include lined burrows, those obser- by Nicholson (1873). Detailed summaries ved in the Trenton Group are so rare have been presented by Alpert (1975), and are so well mixed within crowded Benton and Trewin (1978) and more re- P. beverleyensis that the use of another cently Pemberton and Frey (1982). As name would obscure the close similarity with Palaeophycus, we accept the most of the burrows. Arguably, strict adher- useful and workable nomenclatural system ence to the original definition of Cylin- as that of Pemberton and Frey (1982) drichnus Toots in Howard, 1966, might and adopt it herein. Nevertheless, we suggest their inclusion within this ichno- slightly emend their diagnosis to include genus. However, the original definition lined forms, which, although rare in the of Cylindrichnus encompasses a wide Trenton Group are consistent with the variation in morphology, often overlap- overall diagnosis of Planolites as origin- ping into related forms such as Astero- ally described by Nicholson (see Benton soma von Otto, 1854 and Rosselia Dah- and Trewin 1978,pp. 5-8). mer, 1937. Additionally, most authors Planolites is a eurybathic form (Cham- now adopt an undefined but emended berlain 1978b) ranging in age from Late usage of Cylindrichnus as vertical or Precambrian-Recent (Hantzschel 1975) steeply inclined, unbranched concentri- and is commonly attributed to the sedi- cally lined structures (e.g. Fursich 1974b, ment-ingesting activity of vagile deposit 1981; McCarthy, 1979). We are therefore feeding vermiform organisms (Alpert hesitant to adopt Cylindrichnus as a use- 1975, Pemberton and Frey 1982), though ful description of these Trenton speci- several phyla may in fact be responsi- mens and include them within Planolites, ble. to which they best conform. 24 FILLION and PICKERILL

Otherwise, the specimens described in convex and concave epirelief. here exhibit all the attributes of P. Remarks - Their preservation as endo- beverleyensis and are definitively con- relief and only occasionally as semi- specific. relief structures, and hence the lack of knowledge of the wall characteristics Planolites montanus Richter, 1937 and detailed 3-dimensional geometry (Figure 9h) precludes ichnospecific assignment. They Diagnosis - Relatively small, curved to are certainly larger than P. montanus contorted Planolites burrows less than and although extremely similar to P. 5mm in diameter (after Pemberton and beverleyensis differ in that they are oc- Frey 1982). casionally slightly sinuous. Material - 10 specimens and more than a hundred observed in the field. Ichnogenus Rhizocorallim Zenker, 1836 Description - Straight to slightly sinu- ous, smooth, horizontal unbranched and Type species - Rhizocorallium jenense rarely branched (angles of 25-83°), ir- Zenker, 1836,by original monotypy. regularly cylindrical burrows, l-5mm in Diagnosis - U-shaped spreiten burrows, diameter and 14-74mm in length. Cross- parallel or oblique to bedding plane, overs, interpenetration and reburrowed limbs more or less parallel and distinct; segments are common. Burrow fill is ty- tube diameter: diameter of spreite 1.5 pically structureless but rare examples (after Fursich 1974c). preserve thin wall linings ( 0.6mm) and, or, concentric laminae. They occur in Discussion - Two of the three ichno- a wide variety of lithologies and are species of Rhizocorallium, namely R. preserved mainly in endorelief but also irregulare Mayer, 1954 and R. uliarense in convex epirelief and concave hypo- Firtion, 1958, are clearly the result of relief. deposit-feeding organisms, whilst the third, R. jenense Zenker, 1836, is the Remarks - As the various ichnospecies product of a suspension feeder (Fursich of Planolites are based, according to 1974c). In both cases, the most likely Pemberton and Frey (1982), on size, cur- producers have been suggested to be vature and wall sculpture, the Trenton crustaceans or even nymphs of ephe- material described here can be con- merids (Fursich 1974c, Fursich and Mayr fidently assigned to P. montanus. Lined 1981) or even possibly annelids (Basan and and concentrically laminated specimens, Scott 1979). The ichnogenus is cosmopo- otherwise exhibiting the same basic size litan, Cambrian-Tertiary in age and and morphology as P. montanus, are here although commonly reported in marginal included within the ichnospecies based or shallow marine strata (e.g. Hakes on the same reasoning as discussed 1976; Fursich 1974c, 1981) has also been under P. beverleyensis. reported in deep-water (Crimes 1977, Pickerill et al. 1982) and non-marine Planolites sp. (Fursich and Mayr 1981) environments. (Figure 9f) A full review of its nomenclatural and Material - 57 specimens. interpretive history is given in Fursich Description - Horizontal, straight to (1974c). slightly sinuous, unbranched, irregularly cylindrical (where observed) burrows, ?Rhizocorallium cf. R. irregulare 3-20mm in diameter and up to 12cm in Mayer, 1954 length. The burrows are most typically (Fig. 10, Pickerill and Forbes 1979) preserved in endorelief in calcisiltites Diagnosis - Long, sinuous, bifurcating and calcilutites, where their coarser fill or planispiral U-shaped spreiten burrows, and rare concentric laminae can be in the main horizontal (after Fursich readily observed, and only occasionally 1974c). Maritime Sediments and Atlantic Geology 25

Material - 5 specimens. ing its use with respect to the morpho- Description - Straight, parallel or slight- logically related forms Cylindrichnus ly oblique to stratification, isolated, U- Toots in Howard, 1966 and Asterosoma shaped burrows, with tube diameter von Otto, 1854. Asterosoma should only (where discernible) 2-3mm, total width be used for star-shaped, concentrically 12-15mm and length 60-70mm with ir- lined and longitudinally wrinkled burrows regularly shaped and often discontinuous and, for example, the specimens figured poorly preserved protrusive spreiten. by Frey and Howard (1970) are not con- The burrows are smooth and fill is sistent in any sense of the original diag- identical to the enclosing sediment thus nosis (cf. Hantzschel 1975). Many authors making for extremely poor preservation now employ Cylindrichnus for vertical which is in both full relief and concave or steeply inclined, straight to weakly epirelief. curved, unbranched Skolithos-like struc- tures with multiple layered walls (e.g. Remarks - The poor preservation and Fursich 1974a, 1981; McCarthy 1979), that the extremely small size of the burrows is in contradiction to its original and precludes confident assignment to extremely broad diagnosis (see Howard Rhizocorallium. However, slightly larger 1966). Strict adherence to this emended material and with comparable discontinu- usage is to be recommended, because ous or even no spreiten has been recent- once morphologically similar structures ly described by Fiirsich and Mayr (1981) assume a cone or funnel-shape, they from the Miocene of southern Germany. should be included within Rosselia and If, indeed, the specimens are true Rhi- Asterosoma. Although the three ichno- zocorallium, then they are conspecific genera are clearly end members of in- with R. irregulare Mayer, 1954, as R. tergradational burrow systems (e.g. uliarense is a 3-dimensional spiral and Chamberlain 1971, Fursich 1974a, R. jenense is shorter and typically obli- McCarthy 1979), providing caution is ob- que. Several attempts to photograph the served, ichnogeneric assignment should specimens proved futile but a schematic be possible. figure is reproduced in Pickerill and Rosselia ranges in age from Lower Forbes (1979, p. 2032, fig. 10). Cambrian (Seilacher 1955) to Upper Eocene (Ksiazkiewicz 1977) and although Ichnogenus Rosselia Dahmer, 1937 most commonly reported from shallow Type species - Rosselia socialis Dahmer, sublittoral environments (Garcia-Ramos 1937, by original monotypy. 1976, McCarthy 1979, Pemberton et al. 1982) has also been reported in associ- Diagnosis - "Cylindrical pencil-thick bur- ation with deep-water flysch (Ksiazkie- rows, filled with quartzite, dug in the wicz 1970, 1977). It may represent the sediment at various distances and orien- feeding or dwelling burrows of annelids tations. Most are inclined at an angle (Chamberlain 1971), crustaceans (Frey of 30° or more to bedding. The lower 1970) or sea anenomes (Chamberlain and end is not observed; the opening is ex- Clark 1973, Ksiazkiewicz 1977). panded and filled with imbricated con- centric layers of matrix which as a rule are strongly weathered; these structures ?Rosselia sp. are clearly distinctive in their appear- (Figures lOf, g) ance" (Dahmer 1937, pp. 532-533 - trans, Material - 10 specimens. litt.). Description - Vertical to slightly in- Discussion - Dahmer (1937) originally clined, funnel-shaped burrows preserved erected Rosselia for cone or funnel- only in concave epirelief on upper sur- shaped burrows with a concentric back- faces of thinly bedded (20-35mm) cal- fill structure and, as outlined briefly in cilutites. Burrow diameter is 4-9mm; the discussion of P. beverleyensis, there depth is up to 9mm. In most examples has been considerable confusion regard- the filling has been eroded but some 26 FILLION and PICKERILL exhibit a poorly preserved concentric The morphology, interpretation and wall surrounding a central tube up to detailed taxonomy of Rusophycus has 3mm in diameter. been fully discussed by Osgood (1970) and Remarks - The material is so poorly Osgood and Drennen (1975). It ranges preserved even ichnogeneric identifica- in age from Cambrian-Triassic (Crimes tion is uncertain. However, its vertical 1975, Bromley and Asgaard 1979) and, attitude and funnel-like cross-section following Seilacher (1955) and Osgood and its poorly preserved concentric (1970), was produced by trilobites and layering and central tube are sufficient is a trace diagnostic of the Cruziana to tentatively assign it to Rosselia ichnofacies of Seilacher (1967). Dahmer, 1937 . Rusophycus sp. Ichnogenus Rusophycus Hall, 1852 (Figure 7h) Type species - Fucoides biloba Vanuxem, Material - 9 specimens. 1842, first? subsequent designation by Description - Variably and generally Seilacher (1955, p. 110) as R. bilobatus poorly preserved (in convex hyporelief) (=R. bilobatum). posteriorly tapering bilobate traces; Diagnosis - Shallow to deep, short, maximum length 44mm, maximum width posteriorly tapering bilobed resting bur- 26mm and a mean length/width ratio of rows (cubichnia) of trilobites. Lobes may 1.7:1. Depth variable, up to 8mm. Lobes be smooth or exhibit transverse to obli- are smooth and outer margins grade im- que scratch marks in various arrange- perceptibly into surrounding material. ments directed anterolateral^. Coxal, Remarks - Poor preservation and the exopodal, spinal, cephalic and pygidial absence of detailed scratch markings markings may be present (after Osgood precludes ichnospecific assignment. They 1970, Alpert 1976). are certainly extremely similar to speci- Discussion - If the ichnogenera Cruziana ments of Rusophycus described from the and Rusophycus are to be retained as Trenton Group northeast of Quebec City distinctive forms, the designations by by Pickerill and Forbes (1979), who noted Miller (1889, p. 138), Bassler (1915, p. a close similarity to R. pudicum. Sec- 1132), Andrews (1955, p. 230) and Hakes tioning reveals the traces were formed (1976, p. 32) of the type species of at the sediment-water interface. Rusophycus as R. clavatus must be re- jected, as Hall's (1852) original material Ichnogenus Scalarituba Weller, 1899 should more correctly be diagnosed as Type species - Scalarituba missouriensis Cruziana clavata (Hall), as suggested by Weller, 1899, by original montypy. Osgood and Drennen (1975). Other Diagnosis - Mainly horizontal "burrows authors (e.g. Osgood 1970, Hantzschel ...., subcylindrical in form, never 1975, Osgood and Drennen 1975, Alpert straight for more than a few centi- 1976) quote Fucoides biloba (Vanuxem) meters, curving in all directions, and as the type, even though Hall (1852) did marked by transverse ridges situated at not designate R. bilobatus (=R. bilo- batum) as his nomenclatural type. Argu- distances of one or two millimeters, ably, his R. pudicus (=R. pudicum) could filled with material similar to that equally be regarded as the type as it which surrounds them, .... detected by was described prior to R. bilobatus. To their slightly different color ...." (after retain some semblance of nomenclatural Weller 1899, p. 12). stability, however, particularly as R. Discussion - Scalarituba has been the bilobatum is more closely representa- subject of much nomenclatural contro- tive of typical rusophycids, as originally versy since Seilacher and Meischner inferred by Hall himself (1852, p. 23), (1965) first considered it to be a be- we accept F. biloba Vanuxem, 1842 as havioural variant but not strictly synony- the type species. mous with associated Nereites and Neo- Maritime Sediments and Atlantic Geology 27

nereites traces. Chamberlain (1971), with ionally meniscae possess coarser bio- extremely well-preserved material, de- clastic material. Burrow diameter is 2.5- monstrated that for a single specimen, 5mm and is consistent within a single preservation in hyporelief resulted in specimen; length is variable, a maximum biserially arranged pustules characteris- of 18cm having been observed. Meniscae tic of Neonereites biserialis whereas are irregularly distributed but up to 3 preservation in epirelief resulted in a per cm may be observed. median gallery with lateral lobes char- Remarks - The Trenton material can un- acteristic of S. missouriensis, Nereites doubtedly be diagnosed as S. missourien- sp. (=Phyllodocites view) and Neonereites sis and is extremely similar to conspeci- uniserialis. He placed N. uniserialis and fic material described by Pickerill (1981) N. biserialis in synonymy with S. mis- and by Marintsch and Finks (1982) as S. souriensis and expanded Scalarituba to missouriensis var. "meniscate form". include a 'Nereites view' in convex hyporelief and a 'Phyllodocites view' in Ichnogenus Scolicia de Quatrefages, 1849 concave epirelief. Although this approach has subsequently gained a large follow- Type species - Scolicia prisca de ing (e.g. Chamberlain 1977, 1978b; Quatrefages, 1849, by original monotypy. Chaplin 1980; Jordan 1981; Larson 1982) Diagnosis - Concave, top-surface mean- several authors have preferred to retain dering trails broadly U-shaped in section the forms as distinctive ichnogenera, with floor and lateral walls. Most of the particularly because poorer preserved floor shows a single or biserially alter- material than that described by nating series of laminae, but the edges Chamberlain (1971) does not illustrate the of the floor are smooth tracks, some- morphological overlap and, instead, con- times consisting of discrete sediment forms more closely to individual ichno- strings. Lateral walls with distinct cur- genera as originally defined (e.g. Hantz- ved laminae, concave surface anterior schel 1975; Hakes 1976; Brasier and and steep (after Smith and Crimes 1983). Hewitt 1979; Pickerill 1980, 1981; Marintsch and Finks 1982). We adopt the Discussion - Scolicia was erected from latter approach herein. the Paleocene flysch of Spain but its Scalarituba represents the deposit- original description was inadequate and feeding activity of a worm-like organism had no accompanying figure (Smith and burrowing in the sediment and periodi- Crimes 1983). This led to utter confusion cally backfilling its burrow (Weller 1899, in the literature and the "Scolicia group" Conkin and Conkin 1968). It is a marine now comprises extremely variable traces but eurybathic form (Crimes 1977) hav- interpreted as the trails of annelids, ing been reported from a wide spectrum gastropods, crustaceans, polychaetes, of environments ranging from tidal flat echinoids, holothuroids, Pennatulacea (as (Conkin and Conkin 1968) to deep-water reviewed by Bradley 1980) and more re- flysch (Chamberlain 1971, Pickerill 1980). cently by Smith and Crimes (1983) as It ranges in age from Ordovician-Per- heart urchins. Following Ksiazkiewicz mian (Hantzschel 1975). (1977), Crimes et al. (1981) and Smith and Crimes (1983) used Scolicia for either shallow burrows or furrows and Subphyl- Scalarituba missouriensis Weller, 1899 lochorda for cylindrical backfilled bur- (Figure 10c) rows. Although this approach is followed Diagnosis - As for ichnogenus herein we do not agree with the sug- gestion of Smith and Crimes (1983) that Material - 5 specimens. all Palaeozoic forms referred to Scolicia Description - Horizontal, sinuous menis- should be transferred to other ichno- cate burrows, that straighten over short genera because they cannot possibly be distances, preserved in endorelief on up- related to heart urchins. Knowledge of per surfaces of calcilutites. Burrow fill the producer of a trace fossil is not is identical to host material but occas- taxonomically significant in ichnology 28 FILLION and PICKERILL

(see Fillion and Pickerill 1984). Clearly, Ichnogenus Skolithos Haldeman, 1840 a full and detailed taxonomic revision of the ichnogenus is still required and Type species - Fucoides? linearis Halde- until such a study is undertaken we re- man, 1840, by original montypy. gard Scolicia as a variable and facies Diagnosis - Unbranched, vertical or transgressive form which ranges in age steeply inclined, cylindrical or subcylin- from Lower Cambrian to Recent (Hantz- drical, lined or ur.lined burrows, l-15mm schel 1975). in diameter and a few centimetres to up to a meter in length. Burrow walls Scolicia sp. distinct or indistinct, smooth to rough, (Figure 7c) possibly annulated; burrow diameter may Material - 3 specimens and several oc- vary slightly along its length. Burrow currences on a single bedding plane. fill structureless (after Alpert 1974). Description - All specimens are preser- Discussion - As with several other com- ved on the upper surface of a 5mm monly occurring ichnogenera, Skolithos thick chert bed in concave 'epirelief and has received considerable discussion and consists of two parallel furrows, each systematic treatment. Alpert (1974) re- 4mm wide, separated by a slightly raised cently published a systematic review, central lobe 4-4.5mm wide. The latter conducted solely on literature research, is smooth and rounded whereas the fur- and suggested that many of the numer- rows exhibit crude transverse striations ous previously published ichnospecies and oriented approximately normal to the even ichnogenera such as Tigillites, central lobe, usually less than 2mm Stipsellus and Foralites could confidently apart. Total depth of the furrows is 1.5 be placed within five ichnospecies of mm. The traces are sinuous, often tens Skolithos. He later proposed a sixth, of centimetres in length and never self- S. bulbus (Alpert 1975), and more recent- crossing. ly Hofmann (1979) erected a new ichno- Remarks - Because of the generally poor species, S. gyratus. James (1891), Richter preservation of the traces and the un- (1920, 1921), Howell (1943) and Osgood satisfactory taxonomic status of the (1970) have also extensively discussed the ichnogenus, the Trenton material is only ichnogenus. Undoubtedly, a thorough identified at the ichnogeneric level. systematic review still seems necessary, Although we know of no identical analo- but until this is undertaken we regard gues, they do resemble specimens of Alpert's (1974, 1975) scheme as the most Scolicia figured by both Garcia-Ramos satisfactory. (1976) from the Devonian of Spain and Chamberlain (19?1) from the Carboni- Skolithos has been reported from a ferous of Oklahoma. variety of marine environments and is typically found in association with 'high energy' conditions, particularly in near- .4 I ASYMMETRICAL O cm O O ^ ° shore shallow-water conditions but also O INTACT in deep-water submarine channels and canyons (see Seilacher 1967, Alpert 1974, •3 • BORED . . g*£S ° Crimes 1977, Pickerill 1981 etc.) or even in floodplain and dune environments (e.g. Ratcliffe and Fagerstrom 1980). It con- o ao° stitutes the type ichnogenus of the Sko- lithos ichnofacies of Seilacher (1967) and • O DIAMETER is generally regarded as the dwelling and 1 21 3i 4i 5i 6i cm feeding burrow of annelids or phoronids (Alpert 1974) and ranges in age from Fig. 12 - Height: Diameter plot of intact and bored, symmetric and asymmetric colonies of the bryozoan Late Precambrian-Recent (Crimes and Prasopora spp. Germs 1982). Maritime Sediments and Atlantic Geology 29

Skolithos linearis Haldeman, 1840 structures consisting of a pile of gutter (Figure 13a) shaped laminae (Seilacher 1955, p. 378 -trans. litt.). Diagnosis - Burrows cylindrical to subcy- lindrical, perfectly straight to slightly Discussion - Teichichnus was introduced curved and inclined, up to a meter in by Seilacher (1955) to describe a type length. Diameter 3-7mm, occasionally up of spreiten trace fossil formed by a to 12mm. Burrow wall distinct to indis- series of long horizontal burrows stacked tinct, may be annulated (after Alpert vertical to bedding. He interpreted the 1974, p. 663). burrow as resulting from continual up- Material - 13 specimens. ward-shift by the producing deposit-feed- ing organism. Since its original descrip- Description - Straight, vertical, un- tion, it has been shown to intergrade branched burrows 3-5mm in diameter and with Phycodes (Hantzschel and Reineck up to 66mm in length. Burrow fill is 1968, Hofmann 1979), Thalassinoides, typically coarser than host material, Ophiomorpha and Gyrolithes (Hester and rarely finer. Burrow walls are distinct, Pryor 1972, Frey et al. 1978, Frey and parallel, apparently smooth and unlined. Seilacher 1980), Rhizocorallium (Chisholm Preservation is in endorelief. 1970, Sellwood 1970) and even Cruziana (Garcia-Ramos 1976). As such, its pro- Remarks - Of the seven existing ichno- ducers are probably polyphyletic, though species of Skolithos, the Trenton mater- annelids and arthropods are usually most ial most closely resembles S. linearis favoured (Martinsson 1965, Chisholm Haldeman, 1840 and is diagnosed as such 1970, Chamberlain 1977). Although they superficially resemble S. verticalis (Hall), this ichnospecies is It is a strictly marine but eurybathic generally shorter and smaller and more form, most commonly occurring in shal- commonly inclined and curved (Alpert low subtidal sediments (e.g. Fursich 1981) 1974). but also in association with submarine fans and abyssal environments (Ekdale 1977, Wetzel 1981). It ranges in age from Skolithos sp. Lower Cambrian-Recent. (Figure lOd) Material - 4 specimens and several oc- Teichichnus rectus Seilacher, 1955 currences in the field. (Figure 10c) Description - Straight, vertical to steep- Diagnosis - Straight, unbranched Teichi- ly inclined (up to 60°) burrows 4-10mm chnus with an exclusively retrusive in diameter and up to 35mm in length. spreite (after Seilacher 1955). Burrow fill is finer or identical to the Material - 138 specimens. host rock (calcisiltite). Burrow walls are distinct and iregular; some speci- Description - Straight, smooth, un- mens possess a thick calcilutite wall branched burrows, parallel or slightly (2-3mm) around an inner tube of 4mm oblique to stratification possessing re- diameter. Preservation is in endorelief. trusive spreiten. Length 8.5-125mm, Remarks - Though clearly lacking the width 4-35mm and height 14-25mm; indi- characteristics of S. linearis, the speci- vidual specimens are consistently deeper mens are not well enough preserved for than wide, are preserved in full relief ichnospecific identification. Similar and possess a calcilutite or calcisiltite structures have been figured by Pickerill fill of identical grain size to host rock. and Forbes (1979, p. 2033, fig. 116). Remarks - T. rectus is the product of a shallow burrower usually associated Ichnogenus Teichichnus Seilacher, 1955 with low sedimentation rates (Fursich 1975, Hakes 1976) but occasionally obser- Type species - Teichichnus rectus, Seil- ved in environments subject to rapid de- acher 1955, by original monotypy. position (Chisholm 1970). Specimens from Diagnosis - Long wall-shaped septate the Trenton Group within Quebec City 30 FILLION and PICKERILL

Fig. 13 - a. Skolithos linearis in vertical section, Grondines Member, locality 18. b. 'Oblique burrow1 in transverse sec- tion, Grondines Member, locality 18. c. 'Loop burrows' on lower bedding surface in convex hyporelief, Grondines Mem- ber, locality 21. d. 'Pronged burrows' with typical prongs arrowed, on upper bedding surface, Grondines Member, locality 21.

have also been figured in Pickerill and spreite are concave-up and retrusive. Forbes (1979, p. 2031, figs. 9c, 9d). Remarks - The total character of this form is not fully preserved and not Teichichnus sp. clearly understood, but the absence of (Figure lOh) a tube clearly distinguishes them from Material - 12 specimens. the similar Polarichnus Narbonne (in Description - All specimens preserved Narbonne et al. 1979). They most closely in full relief on upper bedding plane sur- accord to Teichichnus Seilacher, 1955 and faces as elongate, unbranched, straight are diagnosed as such but only at ichno- endoreliefs with parallel sides and round- generic level. ed terminations. Length 50-105mm, width ll-20mm and depth, where observed, 16- Ichnogenus Trichichnus Frey, 1970 21mm. Plan views exhibit bidirectional spreite concave towards the centre of Type species - Trichichnus linearis Frey, the burrow; transverse sections indicate 1970, by original designation. Maritime Sediments and Atlantic Geology 31

Diagnosis - Branched or unbranched, surface with a single opening to the hairlike, cylindrical, straight to sinuous surface (after Bromley 1972). burrows distinctly 1mm in diameter, Discussion - As recently noted by oriented at various angles with respect Bradshaw (1980), Magdefrau (1932) to bedding. Burrow walls distinct, com- named Trypanites weisei, the type monly lined with diagenetic minerals species, for slender more or less (after Frey 1970). straight-sided and long (up to 4cm) Discussion - Frey (1970) interpreted but very narrow (1mm) vertical borings Trichichnus as the feeding and dwelling in the Middle Triassic Muschelkalk burrow of a minute deposit-feeding hardgrounds. At a later date, however, vermiform or crustacean-like organism, Bromley (1972) expanded Trypanites to and since its original formulation re- include all single-entrance pouch-shaped mains a monospecific form, even though borings (e.g. Teredolites, Gastrochaeno- it has been reported from strata of lites) resulting from the activities of Tremadoc-Recent age (Wetzel 1981, a variety of organisms but excluding Fillion and Pickerill 1982). It is strictly acrothoracican cirripeds. The expanded marine but eurybathic, having been re- definition has generally been accepted ported from shallow (Frey 1970, 1975; apart from the inclusion by Bromley Pickerill and Forbes 1979) and less com- (1972) of Vermiforichnus Cameron, 1969, monly deep-water (Kennedy 1975) envir- which is still adopted by many authors onments. (e.g. Pickerill 1976, Elias 1976, Hofmann 1979) as a useful descriptor of un- Trichichnus sp. branched horizontal or subhorizontal (Figure 10a) cylindrical borings. In this paper we Material - 15 specimens and several also exclude borings more closely akin occurrences recorded in the field. to Vermiforichnus and adopt Trypanites as originally defined by Magdefrau Description - Unbranched, straight to (1932). slightly cu.rved, thin cylindrical bur- rows less than 1mm in diameter and Trypanites ranges in age from Lower up to 30mm long. Commonly vertical Cambrian (James et al. 1977) to Recent but may be inclined and rarely horizon- (Kobluk et al. 1978) and is commonly tal. Burrow walls distinct and fill is attributed to the activities of poly- typically coarser than enclosing mater- chaetes, sipunculids, phoronids and ial but may be finer. Preservation is thoracican cirripeds of which the sipun- in full relief. culids (Pemberton et al. 1980) or poly- chaetes (Kobluk and Nemcsok 1982) are Remarks - The Trenton material is only most favoured. It constitutes the type identified at ichnogeneric level as of the recently defined Trypanites specimens lack the diagenetic wall lin- ichnofacies of Frey and Seilacher ings diagnostic of T. linearis (see Frey (1980). 1970, p. 22), thus suggesting that the producing organism was not required to reinforce its burrow. Pickerill and Trypanites weisei Magdefrau, 1932 Forbes (1979, p. 2031, fig. 9e) have also (Figures lib, c) figured identical material from the Diagnosis - More or less straight and Trenton Group at Pont Rouge (see Fig. vertical Trypanites, diameter ca. 1mm 1). and reaching over 4cm in length (after Bromley 1972). Ichnogenus Trypanites Magdefrau, 1932 Material - 97 specimens and several Type species - Trypanites weisei Mag- hundred recordings in the field. defrau, 1932, by original monotypy. Description - All borings are preserved Diagnosis - Simple, unbranched borings in colonies of the calcareous red alga in hard substrates, non-parallel to the Solenopora compacta or the bryozoans 32 FILLION and PICKERILL

Prasopora spp. and Dlplotrypa spp. Dia- strated that at least some Vermifori- meter is 0.1-3mm and length up to chnus were produced by spionid poly- 17mm; they are straight to gently cur- chaetes and Pickerill (1976) has demon- ved, parallel sided with clearly defined strated at least a single example of unlined walls and typically possess a its host and size specificity. micrite fill. In Prasopora, walls of the borings are smooth provided they paral- Vermiforichnus clarkei Cameron, 1969 lel zoecia but as soon as they turn (Figures 11a, d) they become ragged. Diagnosis - As for ichnogenus. Remarks - Restriction of the borings to Solenopora, Prasopora and Diplotrypa Material - 3 specimens. suggests host specificity as a result of Description - (i) A single boring pre- larval selectivity of the producing served in endorelief in a pygidium of sipunculids or polychaetes probably as the isotelinid Isotelus gigas, 4.5mm a result of the availability of the large long and l-1.3mm wide with slightly surface area substrates afforded by expanded terminations. The boring is these genera. As all the examined smooth, slightly sinuous and is filled colonies do not show any reaction to with calcisiltite (Fig. 11a). (ii) A single the borings we also conclude that indi- boring preserved in endorelief on an vidual colonies were dead prior to bor- undetermined branching bryozoan, 11mm ing activity. In a recent study of com- long and 2mm wide. The boring is parable material from the Middle Ordo- smooth, straight with rounded termina- vician of Ontario, Kobluk and Nemcsok tions and is filled with partially dolo- (1982) suggested that the producing mitized sparite. (iii) Numerous smaller ?polychaetes preferred large and asym- unbranched borings preserved in endo- metrical Prasopora colonies. However, relief surrounding the right eye of Iso- our data (Fig. 12) suggest that popula- telus gigas, 0.09-0.16mm wide and 0.6- tions of bored and intact colonies lmm long. The borings are straight to thoroughly overlap and symmetrical and slightly sinuous, crowded but non-inter- asymmetrical colonies were bored in- secting (Fig. lid). discriminately. Remarks - Despite the small number of specimens, the variability exhibited Ichnogenus Vermiforichnus by the borings of V. clarkei in the Cameron, 1969 Trenton Group suggests that different Type species - Vermiforichnus clarkei groups of organisms were responsible Cameron, 1969, by monotypy. for its production. Type (iii) borings differ from those of Talpina pungens Diagnosis - Unbranched, horizontal or Quenstedt, 1849 in being parallel and subhorizontal, straight to slightly cur- not oblique to the surface (see Voigt ved, rarely irregular hooked or coiled, and Soule 1973). non-intersecting borings (after Cameron 1969a, b). Ichnogenus Zoophycos Massalongo, 1855 Discussion - Although Bromley (1972) included Vermiforichnus within Try- Type species - Zoophycos caputmedusae panites, we regard it as a useful des- Massalongo, 1855, by subsequent design- criptor of horizontal or subhorizontal ation of Andrews (1955, p. 262). unbranched borings, as discussed under Diagnosis - Variably shaped spreiten the description of Trypanites in this structures comprised of numerous small paper. The borings have been reported protrusive more or less U- or J-shaped in a wide variety of hosts (Cameron burrows of variable length and orienta- 1969a, b) and range in age from Middle tion. Spreite tabular or arranged in Ordovician-Permian (Kobluk et al. helicoid spirals giving an overall out- 1978). Cameron (1969a, b) has demon- line of circular, elliptical or lobate Maritime Sediments and Atlantic Geology 33

shape; central vertical tunnel or mar- Szmuc 1972, Bradley 1973, Hantzschel ginal tube may be present (after Frey 1975, Ekdale 1977, Ksiazkiewicz 1977, 1970, Hantzschel 1975, Wetzel and Wetzel and Werner 1981, Marintsch and Werner 1981). Finks 1982). Discussion - The ichnogenus Zoophycos has been plagued with excessive taxo- Zoophycos sp. nomic splitting, different species, many (Figure 10b) of which are probably synonymous, be- Material - 1 specimen. ing created merely on the whorl shape and the width and manner of branching Description - Horizontal arcuate bur- of the rays (Frey, 1970, Hantzschel row 7.5cm in width and preserved in 1975, Ksiazkiewicz 1977). Furthermore, epirelief, possessing disturbed back- there is still disagreement on the type filled coarse spreiten which are species, because following the designa- straight to moderately curved, com- tion of the type as Z. camputmedusae posed of bioclastic material, ca. 3-6mm by Andrews (1955, p. 262), Hantzschel wide and separated by furrows up to (1962, p. 220), Taylor (1967, p. 4) and 10mm wide. A J-shaped tube, 6.5mm Plicka (1968, p. 840) respectively desig- wide and filled with bioclastic material nated Fucoides brianteus, Zoophycos bounds the burrow and gives it its parti- laminatus and Fucoides circinatus. cular outline. Hantzschel (1962), however, noted his Remarks - Because of the poor preser- designation only by an asterisk, and vation identification at the ichnospeci- although still adopted as the type by fic level is uncertain, but the structure several recent authors (e.g. Marintsch clearly has a J-form (sensu Wetzel and and Finks 1982) we regard such a desig- Werner 1981) in which only one side of nation as invalid by rule of priority. the spreite is bounded by a tube, and The original illustration of Fucoides is therefore regarded as Zoophycos. It circinatus by Brongniart (1828, p. 83) resembles parts of Zoophycos insignis is too schematic to ascertain its true as illustrated by Ksiazkiewicz (1977, pi. nature and Zoophycos laminatus is a 10, figs. 1, 7, text-fig. 16) but other- nomen nudum. Until resolved, there- wise we are unaware of any comparable fore, we accept Andrews' (1955) desig- form. nation of the nomenclatural type as Z. caputmedusae by rule of priority. MISCELLANEOUS TRACES Although there is still no general In addition to the morphologically agreement on its origin, Zoophycos has distinctive forms recognized from the been commonly attributed to the de- Trenton Group several forms are pre- posit-feeding activity of polychaetes sent which, because of their poor pres- (Bischoff 1968) and sipunculids (Wetzel ervation, morphological variability and and Werner 1981), to the imprints of sufficient differences in configuration prostomia of sabellids (Plicka 1970) and to already established ichnogenera, or even the feeding activity of umbellu- combinations thereof, are not formally lids (Pennatulaceids) (Bradley 1973). diagnosed. Four broad categories may Though considerable environmental sig- be recognized: nificance has been attached to the (i) 'Loop burrows' (Fig. 13c) - ichnogenus, particularly regarding Irregularly meandering and winding, bathymetry, it is now realized to be subcylindrical, unlined horseshoe or a eurybathic form ranging in age from more commonly loop-shaped burrows, Ordovician-Quaternary and is typical 4-llmm wide and up to 28cm long, pre- of fine-grained substrates characterized served in convex hyporelief on the by low sedimentation rates (for reviews soles of calcilutites. Burrow fill is see Frey 1970, Kennedy 1970, Simpson coarser than enclosing sediment. Al- 1970, Chamberlain 1971, Osgood and though looped, the burrows never inter- 34 FILLION and PICKERILL sect; rather they go under or over pre- wide preserved in two pygidia of Iso- vious pathways. They are obviously the telus gigas. Similar borings or etching product of an unknown deposit feeder traces have been figured by Bromley and resemble Lumbricaria as figured and Surlyk (1973, p. 362, fig. 14c), by Gutschick and Rodriguez (1977, p. Warme (1975, p. 201, fig. 11-14), 206, pi. la-c). Pickerill and Forbes (1979, p. 2033, fig. llg) and Hofmann (1979, p. 57, pi. 22) (ii) 'Oblique burrows' (Fig. 13b) and all have been interpreted as bryo- - Straight, unbranched, inclined (30-50°) zoan in origin. cylindrical burrows preserved in endo- relief, 4-9mm in diameter and down- CONCLUSIONS ward tapering to a rounded or pointed termination. Length 20-35mm, walls As outlined previously, sediments of distinct and regular, lined or more the Trenton Group were deposited commonly unlined, with a structureless initially in lagoons followed in turn by burrow fill of identical grain size to offshore "bar", shallow and, finally, enclosing material. We were initially deeper offshore shelf environments. tempted to include these structures Trace fossils in each of these environ- within Obliquus Marintsch and Finks, ments may be summarized as follows 1982; however, this ichnogenus is con- (see also Fig. 6): sistently lined, inclined at 45° or less Lagoon - Arenicolites sp., Chondrites and upper portions may approach the type A, Chondrites type B, Clematis- horizontal and lower portions the verti- chnia sp., Helminthopsis hieroglyphica, cal. Furthermore, there are difficulties Helminthopsis sp., Palaeophycus tubu- in distinguishing this recently formed laris, Palaeophycus sp., Planolites ichnogenus from Cylindrichnus as origi- beverleyensis, Planolites montanus, nally defined (Toots in Howard 1966) Planolites sp., Scalarituba missouriensis, and inclined specimens Of Palaeophycus. Skolithos linearis, Skolithos sp., Teichi- (iii) 'Pronged burrows' (Fig. 13d) chnus rectus, ?Teichichnus sp., Trichi- - Straight, unbranched, elongate bur- chnus sp., Trypanites weisei and oblique rows preserved in epirelief on the up- burrows. per surfaces of calcilutites, 4-8mm "Bar" - ?Calyeraterion sp., Chondrites wide, up to 9cm long. Burrow fill is type A, Chondrites type B, Chondrites typically structureless calcilutite but spp., Circulichnis montanus, Cruziana occasionally poorly preserved spreite problematica, Helminthopsis sp., Palaeo- can be discerned. Irregular prongs up phycus tubularis, ?Plagiogmus, Plano- to 4mm in width and length project lites beverleyensis, Skolithos linearis, from the burrows (Fig. 13d). Similar ?Skolithos sp. and Trypanites weisei. unnamed material has been figured by Ward and Lewis (1975, p. 884, fig. 2) Shallow shelf - Arenicolites sp., Chon- and Pickerill and Forbes (1979, p. 2033, drites Type A, Chondrites type B, fig. 11a). The presence of a spreite in Chondrites spp., Cruziana problematica, some specimens suggests the burrows Furculosus carpathicus, Helminthopsis were produced by deposit-feeding hieroglyphica, Helminthopsis sp., organisms and their overall morphology Palaeophycus tubularis, Planolites suggests they are akin to teichichnids. beverleyensis, P. montanus, Planolites sp., ?Rhizocorallium cf. R. irregulare, (iv) 'Bryozoan borings' (Figs, lie, ?Rosselia, Rusophycus sp., ?Scalarituba f) - Delicate pinnate or net-like sys- missouriensis, Skolithos linearis, Sko- tems of tear-drop shaped zooecial pits, lithos sp., Teichichnus sp., Trichichnus each 0.3-0.5mm in length, 0.07-0.1mm sp., Trypanites weisei and Zoophycos in width, rarely adjoined and more sp. commonly separated by stolomate im- Offshore shelf - Arenicolites sp., Chon- pressions up to 1mm long and 0.02mm drites type B, Chondrites spp., Clem- Maritime Sediments and Atlantic Geology 35

atischnia sp., ?Conostichnus, Cruziana ACKNOWLEDGEMENTS sp. cf. Diplichnites sp., Helminthopsis We wish to thank Dr. Yvon Globensky sp., Oichnus paraboloides, Palaeophycus and Dr. John Riva for their continual tubularis, Palaeophycus sp., Planolites help and encouragement in our research beverleyensis, P. montanus, Planolites in the Trenton Group over the past sp., ?Rosselia, Rusophycus sp., Scalari- several years and Dr. Guy Narbonne tuba missouriensis, Scolicia sp., Sko- and Dr. Hans Hofmann for providing lithos linearis, Skolithos sp., Teichichnus us with certain relevant literature and rectus, Teichichnus sp., Trypanites wei- advice on terminology. The manuscript sei, Vermiforichnus clarkei and infor- was reviewed by Drs. T.P. Crimes and mally diagnosed loop, oblique and T.L. Harland and for their constructive pronged burrows and bryozoan borings. coments we are extremely grateful. Although there is some obvious vari- One of us (R.K.P.) is grateful to Dr. ation in the abundance of traces in dif- Crimes for permitting access to his ferent environments (Fig. 6), this is be- abundant trace fossil collections and lieved to reflect preservational and for permission to quote unpublished other hazards such as nature and qua- data. Dr. Harland accompanied the lity of outcrop, burrow density, topo- authors in the field on several oc- nomic or diagenetic preservation, grain casions and for his astute observations size and sediment stability as discus- and keen eye we are equally grateful. sed more fully by Fillion et al. (1984). The work was undertaken during the The entire assemblage, irespective of tenure by R.K. Pickerill of Natural environment of deposition, can best be Sciences and Engineering Research equated with the Cruziana ichnofacies Council of Canada Grant A3857 and of Seilacher (1967), which is character- the tenure of D. Fillion of scholarships ized by a mixed association of verti- from Bourse de l'Enseignement cal, inclined and particularly horizontal sup6rieur du ministere de l'Education deposit^feeding but also filter feeding du Quebec and Bourse du Fonds traces. The abundant borings, more F.C.A.C. pour l'aide et le soutien a la characteristic of the recently formu- recherche, all of which are gratefully lated Trypanites ichnofacies of Frey acknowledged. The manuscript was pre- and Seilacher (1980), are no cause for pared by J. Ley and technical assist- concern as their presence merely re- ance provided by R. McCulloch, S. flects the availability of abundant hard Aitken and J. Lynch. substrates, in this case essentially in the form of the bryozoans Prasopora ABEL, O. 1935. Vorzeitliche Lebensspuren. Gustav spp. and Diplotrypa spp., the red alga Fisher, Jena, 644p. Solenopora compacta and the isotelenid ALPERT, S.P. 1974. Systematic review of the genus trilobite Isotelus gigas. Skolithos. Journal of Paleontology 48, pp. 661-669. j 1975. Planolites and Skolithos from the Upper Precambrian-Lower Cambrian White-Inyo Finally, and as discussed more fully Mountains, California. Journal of Paleontology 49, pp. 509-521. by Fillion et al. (1984), the most signi- , 1976. Trilobite and star-like trace fossils ficant environmental parameter govern- from the White-Inyo Mountains, California. Journal ing the production, nature and preser- of Paleontology 50, pp. 226-240. , 1977. Trace fossils and the basal Cam- vation of the trace fossils was sub- brian boundary. In Trace fossils 2. Edited by T.P. strate and its overall characteristics, Crimes and J.C. Harper. Geological Journal Special which in turn reflected the prevailing Issue 9, Seel House Press, Liverpool, pp. 1-8. hydrodynamic regime and depositional ANDREWS, H.N. Jr. 1955. Index of generic names of fossil plants, 1820-1950. United States Geological rate. Substrate not only provided a pri- Survey, Bulletin 1013, 262p. mary control but also directly influ- BANDEL, K. 1973. A new name for the ichnogenus enced diagenetic affects which both en- Cylindrichnus Bandel, 1967. Journal of Paleoento- logy 47, p. 1002. hanced and masked specific traces de- BARTHEL, K.W. and BARTH, W. 1972. Palecologic pending on their original character. specimens from the Devonian of Bolivia. Neues 36 FILLION and PICKERILL

Jahrbuch fiir Geologie und Palaontologie, Monatshe- Magdefrau. Palaontologische Zeitschrift 46, pp. fte 1972, pp. 573-581. 93-98. BASAN, P.B. and SCOTT, R.W. 1979. Morphology of 1981. Concepts in ichnotaxonomy illus- Rhizocorallium and associated traces from the trated by small round holes in shells. Acta Geo- Lower Cretaceous Purgatoire Formation, Colorado. logica HispSnica, Concept and Method in Paleon- Palaeogeography, Palaeoclimatology, Palaeoecology tology 16(1-2), pp. 55-64. 28, pp. 5-23. BROMLEY, R.G. and ASGAARD, U. 1979. Triassic BASSLER, R.S. 1915. Bibliographic index of American freshwater ichnocoenoses from Carlsberg Fjord, Ordovician and Silurian fossils. Bulletin of the east Greenland. Palaeogeography, Palaeoclimato- • United States National Museum, 92, 718p. logy, Palaeoecology 28, pp. 39-80. BANKS, N.L. 1970. Trace fossils from the late Pre- BROMLEY, R.G. and FREY, R.W. 1974. Redescrip- cambrian and Lower Cambrian of Finnmark, Nor- tion of the trace fossil Gyrolithes, and taxonomic way. In Trace fossils. Edited by T.P. Crimes and evaluation of Thalassinoides, Ophiomorpha and J.C. Harper. Geological Journal Special Issue 3. Spongeliomorpha. Bulletin of the Geological Society Seel House Press, Liverpool, pp. 19-34. of Denmark 23, pp. 311-335. BATHER, F.A. 1925. U-shaped burrows near Blea BROMLEY, R.G. and SURLYK, F. 1973. Borings pro- Wyke. Proceedings of the Yorkshire Geological So- duced by brachiopod pedicles, fossil and Recent. ciety 20, pp. 185-199. Lethaia 6, pp. 349-365. BENTON, M.J. and TREWIN, N.H. 1978. Discussion BRONGNIART, A.T. 1823. Observations sur les and comments on Nicholson's 1872 manuscript "Con- Fucoides. Societg d'Histoire Naturelle de Paris, tributions to the study of errant annelides of the M6moires 1, pp. 301-320. older Palaeozoic rocks". Publication of the Depart- 1828. Histoire des vSgStaux fossiles ment of Geology and Mineralogy, University of ou recherches botaniques et g6ologiques sur les Aberdeen, 1, pp. 1-16. v6g6taux renfermSs dans les diverses couches du BERGSTROM, J. 1976. Lower Palaeozoic trace fos- globe, volume 1. G. Dufour and E. d'Ocagne, Paris, sils from eastern Newfoundland. Canadian Journal 136p. of Earth Sciences 13, pp. 1613-1633. CAMERON, B. 1969a. New name for Palaeosabella BILLINGS, E. 1862. New species of fossils from dif- prisca (McCoy), a Devonian worm boring, and its ferent parts of the lower, Middle and Upper Silur- preserved probable borer. Journal of Paleontology ian rocks of Canada. In Palaeozoic Fossils, Volume 43, pp. 189-192. 1 (1861-1865). Geological Survey of Canada, pp. 96- 1969b. Paleozoic shell-boring annelids and 168. their trace fossils. American Zoologist 9, pp. 689- BINNEY, E.W. 1852. On some trails and holes found 703. in rocks of the Carboniferous strata, with remarks CHAMBERLAIN, C.K. 1971. Morphology and ethology on the Microconchus carbonarius. Manchester of trace fossils from the Ouachita Mountains, Literary and Philosophical Society, Memoirs and southeast Oklahoma. Journal of Paleontology 45, Proceedings, series 2, 10, pp. 181-201. pp. 212-246. BISCHOFF, B. 1968. Zoophycos, a polychaete annelid, 1977. Ordovician and Devonian trace Eocene of Greece. Journal of Paleontology 42, pp. fossils from Nevada. Bulletin of the Nevada Bur- 1439-1443. eau of Mines and Geology 90, pp. 1-24. BORNEMANN, J.G. 1889. Ober den Buntsandstein in 1978a. A guidebook to the trace fos- Deutschland und seine Bedeutung fiir die Trias. sils and paleoecology of the Ouachita Geosyncline. Beitrage Geologie, Palaontologie 1, 61p. Society of Economic Paleontologists and Mineralo- BRADLEY, J. 1973. Zoophycos and Umbellula (Pen- gists, Tulsa (Oklahoma), 68p. natulacea): Their synthesis and identity. Palaeogeo- 1978b. Recognition of trace fossils graphy, Palaeoclimatology, Palaeoecology 13, pp. in cores. In Trace fossil concepts. Edited by P.B. 103-128. Basan. Society of Economic Paleontologists and 1980. Scolicia and Phycodes, trace fossils Mineralogists, Short Course 5, pp. 119-166. of Renilla (Pennatulacea). Pacific Geology 14, pp. CHAMBERLAIN, C.K. and CLARK, D.L. 1973. Trace 73-86. fossils and conodonts as evidence for deep-water deposits in the Oquirrh Basin of Central Utah. 1981. Radionereites, Chondrites and Phy- Journal of Paleontology 47, pp. 663-682. codes; trace fossils of anthoptiloid sea pens. Paci- CHAPLIN, J.R. 1980. Stratigraphy, trace fossil asso- fic Geology 15, pp. 1-16. ciations and depositional environments in the Bor- BRADSHAW, M.A. 1980. Boring bivalves in the New den Formation (Mississippian), northeastern Ken- Zealand Cenozoic with a redefinition of Teredo- tucky. Kentucky Geological Society, Annual Fall lites. Records of the Canterbury Museum 9(5), Field Trip Guidebook. Kentucky Geological Survey, pp. 289-294. Lexington, Kentucky, 114p. BRANSON, C.C. 1960. Conostichus. Oklahoma Geo- CHISHOLM, J.L. 1970. Teichichnus and related trace- logical Notes 20, pp. 195-207. fossils in the Lower Carboniferous of St. Monace, BRASIER, M.D. and HEWITT, R.A. 1979. Environ- Scotland. Bulletin of the Great Britain Geological mental setting of fossiliferous rocks from the Survey 31, pp. 19-35. uppermost Proterozoic-Lower Cambrian of cen- CLARK, T.H. 1972. Montreal area. Quebec Depart- tral England. Palaeogeography, Palaeoclimatology, ment of Natural Resources, Geological Report Palaeoecology 27, pp. 35-57. 152, 244p. BRIGGS, D.E.G., ROLFE, W.D.I, and BRANNAN, CLARK, T.H. and GLOBENSKY, Y. 1975. Grondines J. 1979. A giant myriapod trail from the Namu- area. Quebec Department of Natural Resources, rian of Arran, Scotland. Palaeontology 22, pp. Geological Report 154, 159p. 273-291. 1976a. Sorel area. Quebec BROMLEY, R.G. 1972. On some ichnotaxa in hard Department of Natural Resources, Geological substrates, with a redefinition of Trypanites Report 155, 151p. Maritime Sediments and Atlantic Geology 37

1976b. Laurentides area. ELIAS, R.J. 1976. Solitary rugose corals of the Quebec Department of Natural Resources, Geo- Selkirk Member, Red River Formation (Late Middle logical Reprt 157, U2p. or Upper Ordovician), southern Manitoba. Unpub- 1976c. Trois RiviSres area. lished M.Sc. thesis, University of Cincinnati, Cin- Quebec Department of Natural Resources, Geo- cinnati, Ohio, 232p. logical Report 164, 87p. EfvlMONS, E. 1844. The Taconic System; based on CLARK, T.H., GLOBENSKY, Y., RIVA, J. and observations in New York, Massachusetts, Maine, HOFMANN, H.J. 1972. Stratigraphy and structure Vermont and Rhode Island. Caroll and Cook, of the St. Lawrence Lowland of Quebec. 24th Albany, 68p. International Geological Congress, Guidebook for FILLION, D. and PICKERILL, R.K. 1982. Cambro- Excursion C52, 83p. Ordovician trace fossils from Bell Island, eastern CLIFTON, H.E. and THOMPSON, J.K. 1978. Macaro- Newfoundland (abstract). The Atlantic Geoscience nichnus segregatis, a feeding structure of shallow Society Colloquium on Current Research in the marine polychaetes. Journal of Sedimentary Petro- Atlantic Provinces. Maritime Sediments and Atlan- logy 48, pp. 1293-1302. tic Geology, 18, p. 43. CONKIN, T.E. and CONKIN, B.M. 1968. Scalarituba — 1984. On Arthraria anti- missouriensis and its stratigraphic distribution. quata Billings, 1872 and its relationship to Diplo- The University of Kansas Paleontological Contri- craterion Torell, 1870 and Bifungites Desio, 1940. butions 31, pp. 1-7. Journal of Paleontology 58, (in press). CRIMES, T.P. 1970. Trilobite tracks and other trace (in preparation. Trace fossils from the Upper Cambrian of North Wales. fossils from the Lower Ordovician (Tremadoc- Geological Journal 7, pp. 47-68. Arenig) of Bell Island, eastern Newfoundland. 1975. The stratigraphical significance FIRTION, F. 1958. Sur la prgsence d'ichnites dans of trace fossils. In The Study of Trace Fossils. le Portlandien de l'lle d'016ron (Charente mari- Edited by R.W. Frey. Springer-Verlag, New York, time). Annales Universitatis Saraviensis, (Natur- pp. 109-130. wissenschaftliche) 7, pp. 107-112. 1977. Trace fossils of an Eocene deep- FISHER, W.A. 1978. Trace fossils from the lower sea fan, northern Spain. In Trace fossils 2. Edited Harding Formation (Middle Ordovician), Colorado. by T.P. Crimes and J.C. Harper, Geological In Energy resources of the Denver Basin. Edited Journal Special Issue 9. Seel House Press, Liver- by J.D. Pruit et al. Rocky Mountain Association pool, pp. 71-90. of Geologists, Field Conference, Guidebook for CRIMES, T.P. and GERMS, G.J.B. 1982. Trace fos- 1978, pp. 191-197. sils from the Nama Group (Precambrian-Cambrian) FLOWER, R.H. 1945. Breviconic cephalopods from of southwest Africa (Namibia). Journal of Paleon- Pont Rouge, Quebec. Canadian Field Naturalist tology 56, pp. 890-907. 59, pp. 74-82. CRIMES, T.P., GOLDRING, R., HOMEWOOD, P., FREY, R.W. 1970. Trace fossils of Fort Hays Lime- STUIVENBERG, J. VAN and WINKLER, W. 1981. stone Member, Niobrara Chalk (Upper Cretaceous), Trace fossil assemblages of deep-sea fan deposits, west-central Kansas. The University of Kansas Gurnigel and Schlieren flysch (Cretaceous-Eocene), Paleontological Contributions 53, 41p. Switzerland. Ecologae Geologicae Helvetiae 74, 1975. The realm of ichnology, its strength pp. 953-995. and limitations. In The Study of Trace Fossils. CRIMES, T.P., LEGG, I., MARCOS, A. and ARBO- Edited by R.W. Frey, Springer-Verlag, New York, LEYA, M. 1977. ?Late Precambrlan-low Lower Cam- pp. 13-36. brian trace fossils from Spain. In Trace fossils FREY, R.W. and HOWARD, J.D. 1970. Comparison 2. Edited by T.P. Crimes and J.C. Harper. Geo- of Upper Cretaceous ichnofaunas from siliceous logical journal Special Issue 9. Seel House Press, sandstones and chalk, Western Interior Region, 'Liverpool, pp. 91-138. U.S.A. 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Stutt- bergiana Maritima 10, pp. 117-143. garter Beitrage zur Naturkunde, Serie B (Geologie und Palaontologie) 13, pp. 1-52. EICHWALD, E. 1860. Lethaea Rossica ou palgonto- logie de la Russie I. Stuttgart, E. Schweizerbart, — 1974b. On Diplocraterion Torell 1870 1657p. and the significance of morphological features in EKDALE, A.A. 1977. Abyssal trace fossils in world- vertical, spreiten-bearing, U-shaped trace fossils. wide Deep Sea Drilling Project cores. In Trace Journal of Paleontology 48, pp. 952-962. fossils 2. Edited by T.P. Crimes and J.C. Harper. 1974c. Ichnogenus Rhizocorallium. Palaon- Geological Journal Special Issue 9. Seel House Press, Liverpool, pp. 163-182. tologische Zeitschrift 48, pp. 16-28. 1978. Trace fossils in Leg 42A cores. 1975. Trace fossils as environmental Initial Reports of the Deep Sea Drilling Project indicators in the Corallian of England and Nor- 42, pp. 821-827. mandy. Lethaia 8, pp. 151-172. 38 FILLION and PICKERILL

1981. Invertebrate trace fossils from shoreline deposits - the basal Trenton Group around the Upper Jurassic of Portugal. Communicates Quebec City, Canada. Geological Journal 19, (in dos Servipos Geologicos de Portugal 67, pp. 153- press). 168. HEER, O. 1877. Flora Fossilis Helvetiae. Die vor- FORSICH, F.T. and MAYR, H. 1981. Non-marine weltliche Flora der Schweiz. J. Wurster & Com- Rhizocorallium (trace fossil) from the Upper Fresh- pany, Zurich, 182p. water Molasse (Upper Miocene) of southern HELWIG, J. 1972. Stratigraphy, sedimentation, paleo- Germany. Neues Jahrbuch fur Geologie und geography and paleoclimates of Carboniferous Palaontologie Monatshefte 6, pp. 321-333. ("Gondwana") and Permian of Bolivia. Bulletin of GARCIA-RAMOS, J.C.M. 1976. Morfologia de trazas the American Association of Petroleum Geologists fosiles en dos afloramientos de Arenisca de 56, pp. 1008-1033. Naranco (Devonico medio) de Asturias (NW. de HESTER, N.C. and PRYOR, W.A. 1972. Blade-shaped Espana). 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Geological Journal Valley, especially its footprints. W. White, Boston, Special Issue 9. Seel House Press, Liverpool, pp. 220p. 195-208. HOFMANN, H.J. 1979. Chazy (Middle Ordovician) HAKES, W.G. 1976. Trace fossils and depositional trace fossils in the Ottawa-St. Lawrence Lowlands. environment of four clastic units, Upper Pennsyl- Bulletin of the Geological Survey of Canada 321, vanian megacyclothems, northeast Kansas, The pp. 27-59. University of Kansas Paleontological Contributions HOWARD, J.D. 1966. Characteristic trace fossils 63, 46p. in Upper Cretaceous sandstones of the Book Cliffs 1977. Trace fossils in Late Pennsylvanian and Wasatch Plateau. Utah Geological and Minera- cyclothems, Kansas. In Trace fossils 2. Edited logical Survey, Central Utah Coal Bulletin 80, by T.P. Crimes and J.C. Harper. Geological Jour- pp. 35-53. nal Special Issue 9. Seel House Press, Liverpool, HOWELL, B.F. 1943. Burrows of Skolithos and Plano- pp. 209-226. lites in the Cambrian Hardyston Sandstone at HALDEMAN, S.S. 1840. Supplement to number one Reading, Pennsylvania. Bulletin of the Wagner of "A monograph of the Limniades, and other Free Institute of Science of Philadelphia 3, pp. freshwater uivalve shells of North America", con- 1-33. taining descriptions of apparently new animals IMLAY, R.M. 1961. Characteristic Lower Cretaceous in different classes, and the names and characters megafossils from northern Alaska. United States of the subgenera in Paludina and Anculosa. Geological Survey, Professional Paper 335, 74p. Philadelphia, 3p. JAEGER, H. and MARTINSSON, A. 1980. The Early HALL, J. 1847. Palaeontology of New York, Vol. Cambrian trace fossil Plagiogmus in its type area. 1. C. Van Benthuysen, Albany, 338p. Geologiska Foreningens i Stockholm Forhandlingar 1852. Palaeontology of New York. Vol. 2. 102, pp. 117-126. C. Van Benthuysen, Albany, 362p. JAMES, J.F. 1891. Studies in problematic organisms HALLAM, A. 1975. Preservation of trace fossils. - the genus Scolithus. Bulletin of the Geological In The Study of Trace Fossils. Edited by R.W. Society of America 3, pp. 32-44. (Part of this Frey. Springer-Verlag, New York, pp. 56-63. paper also appears in American Naturalist 26, HANTZSCHEL, W. 1962. Trace fossils and Proble- pp. 240-242, 1892). matica. In Treatise on Invertebrate Paleontology, JAMES, N.P., KOBLUK, D.R. and PEMBERTON, Part W. Edited by R.C. Moore. Geological Society S.G. 1977. The oldest macroborers: Lower Cambrian of America and University of Kansas Press, New of Labrador. Science 197, pp. 980-983. York and Lawrence, Kansas, pp. W177-W245. JORDAN, T.E. 1981. Enigmatic deep-water mechan- 1975. Trace fossils and Problematica. isms, upper part of the Oquirrh Group, Utah. In Treatise on Invertebrate Paleontology, Part Journal of Sedimentary Petrology 51, pp. 879-894. W, Miscellanea, Supplement I. Edited by C. KARASZEWSKI, W. 1971. Some fossil traces from Teichert. Geological Society of America and The the lower Liassic of the Holy Cross Mts., Central University of Kansas Press, Boulder, Colorado, Poland. Bulletin de I'AcadSmie Polonaise des Lawrence, Kansas, 269p. Sciences, S§rie des Sciences de la Terre 19, pp. HANTZSCHEL, W. and REINECK, H.-E. 1968. Fazies- 101-105. Untersuchungen im Hettangium von Helmstedt KENNEDY, W.J. 1970. Trace fossils in the Chalk (Niedersachsen). Geologische Staatsinstitut in Ham- environment. In Trace fossils. Edited by T.P. burg, Mitteilungen 37, pp. 5-39. Crimes and J.C. Harper. Geological Journal Special Issue 3. Seel House Press, Liverpool, pp. 263-282. HARLAND, T.L. and PICKERILL, R.K. 1982. A review of Middle Ordovician sedimentation in the St. 1975. Trace fosils in carbonate rocks. Lawrence Lowland, eastern Canada. Geological In The Study of Trace Fossils. Edited by R.W. Journal 17, pp. 135-156. Frey. Springer-Verlag, New York, pp. 377-398. 1984. 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Reviewers: T.P. Crimes T.L. Harland