Dec 17, 2008 - figs. 2,4. 1937 Archaeonassa fossulata Fenton and Fenton, p. 455 and pi. 1, figs. 1,2. 1971 Scolicia vada Chamberlain, fig 4i, and pi.29, fig. 8.
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Archaeonassa Fenton and Fenton 1937 reviewed James O. Buckman
a b
a
Division of Geology, School of Sciences , Staffordshire University , College Road, Stoke‐on‐Trent, Staffordshire, ST4 2DE, United Kingdom b
Department of Civil Engineering , Paisley University , High Street, Paisley, PA1 2BE, Scotland Published online: 17 Dec 2008.
To cite this article: James O. Buckman (1994) Archaeonassa Fenton and Fenton 1937 reviewed, Ichnos: An International Journal for Plant and Animal Traces, 3:3, 185-192, DOI: 10.1080/10420949409386387 To link to this article: http://dx.doi.org/10.1080/10420949409386387
Ichnos, v. 3, pp. 185-192, 1994 An International Journal for Plant and Animal Traces
Archaeonassa Fenton and Fenton 1937 reviewed James O. Buckman† Division of Geology, School of Sciences, Staffordshire University, College Road, Stoke-on-Trent, Staffordshire, ST4 2DE, United Kingdom
Archaeonassa is a poorly known ichnogenus, originally described from the Cambrian of North America, which is a member of the Scolicia 'group', but can be regarded as distinct from Scolicia. Archaeonassa was originally based on modern material, and therefore falls into a taxonomic grey zone concerning its availability, but as A. fossulata was erected on fossil material the ichnogenus is here retained. Although exhibiting a degree of morphological plasticity in its style of ornament, Archaeonassa is considered to be monospecific, with only the ichnospecies A. fossulata recognised. The ichnogenus appears to be restricted to the Paleozoic, although similar material is well documented from modern environments. Thus the ichnogenus is expected to be geographically and stratigraphically more widespread than it otherwise appears. Archaeonassa is a valuable tool for environmental reconstruction within the Paleozoic as it occurs within intertidal deposits. The ichnogenus is interpreted chiefly as the work of gastropods, although in many cases may have been produced by trilobites or even echinoids, and represents the exogenic expression of either surface or shallow subsurface locomotion. Key Words: Archaeonassa, Palaeobultia, Scolicia, gastropod, trilobite, Carboniferous, Paleozoic, intertidal, northwest Ireland.
Archaeonassa are discussed, based on well developed Irish material and other examples in the literature.
GEOLOGIC SETTING OF THE IRISH MATERIAL Archaeonassa has been recorded from the Lower Carboniferous of northwest Ireland, within the Donegal Bay Basin, which contains a mixed carbonate to clastic sequence of sediments, with environments of deposition ranging from open shelf, prodelta, deltaic, marginal marine and nonmarine (Buckman, 1992a). Archaeonassa is restricted to the Mullaghmore Sandstone Formation (Buck-
Mountcharles Sandstone
N INTRODUCTION The ichnogenus Archaeonassa Fenton and Fenton 1937 was erected for Cambrian trace fossils believed to be the product of gastropods. Since its erection the ichnogenus has been infrequently referred to (Häntzschel, 1962,1965, 1975; Alpert, 1975,1977; Crimes, 1987; Buckman, 1992a, b; Parnell et al. 1992), with Alpert (1975), Buckman (1992a, b) and Parnell et al. (1992) apparently the only new records of the ichnogenus; but with material only discussed or illustrated in Buckman (1992a, b). In this paper examples of Archaeonassa are fully described from the Lower Carboniferous of northwest Ireland (Fig. 1). The taxonomic position of Archaeonassa is discussed in detail, with similarities and differences noted to Palaeobullia Götzinger and Becker 1932 and Scolicia de Quatrefages 1849, as well as other trace fossils (including modern lebensspuren). The toponomy, ethology, environmental significance, age range and likely producer of †
Current address: Department of Civil Engineering, Paisley University, High Street, Paisley, PA1 2BE, Scotland. Photocopying permitted by license only Reprints available directly from the publisher
Fig. 1. Locality map. (A) Ireland, indicating the position of the detailed map. (B) The position of Mullaghmore Sandstone Formation outcrop within the Donegal Bay area, is shown in stipple, based on George and Oswald (1957) and Oswald (1955). (C) Enlargement of the Mullaghmore Head area. All major locations of Archaeonassa are indicated: localities 1-6 on (B) and (C). Key: 1 = Pollayarry bay [G699571], 2 = N.W. Mullaghmore Head [G698580] (see Buckman 1992a for details), 3 = E. Mullaghmore Head [G710579], 4 = Blackrocks [G725564], 5 = Skerrydoo [G740573], and 6 = Illanmore [G845664]. 'K' numbers = specimen numbers of material within the collections of the Ulster Museum, Belfast, Northern Ireland. © 1994 Harwood Academic Publishers GmbH Printed in the United States of America
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man, 1992a, b), which represents a prograding deltaic complex, with delta advancement from the north and northwest (Sheridan, 1972). The deltaic complex represents a major siliciclastic wedge, with a maximum recorded thickness in the study area of 213 metres (Oswald, 1955; George and Oswald, 1957; Sheridan, 1972). The formation contains a rich and diverse ichnofauna, with at least fifty recorded ichnogenera (Buckman, 1992a, b; Parnell et al., 1992), of which Archaeonassa is a common component, particularly from sections interpreted as tidal in origin.
commonly on rippled muddy sandstone beds, at junctions with shales and siltstones. Description: As for ichnogenus diagnosis, with the addition: total width 2-25 mm (Fig. 4), by up to 900 mm or more in length, with marginal levées of low relief and 0.5-7.0 mm wide. Variation in ornament illustrated in Figures 2a, b, d and 5. Trails parallel to ripple crests typically only have a levée on the ripple trough side (Figs. 2b, c, 6). Bedding-parallel, pattern straight, gently curved, sinuous or occasionally looped (Fig. 2b, e), with crosscutting common. May pass into simple V-shaped plough trails (= Unisulcus) (Fig. 2e).
SYSTEMATIC ICHNOLOGY Ichnogenus Archaeonassa Fenton and Fenton 1937 v 1852 v 1852 1937 1971 1985 1990 1992a 1992b non 1849 non 1932 non 1971
Annelid trails Hall, pi. 13, figs, lb-d, and pi. 14, figs. l , 2 ( c - c ' ) , 3 (d-d', e-e')Gastropod trails Hall, p l . l l , fig. 4, and pi. 12, figs. 2,4. Archaeonassa fossulata Fenton and Fenton, p. 455 and pi. 1, figs. 1,2. Scolicia vada Chamberlain, fig 4i, and pi.29, fig. 8. Palaeobullia Götzinger and Becker; Knox and Miller, pi. 2, fig. f (only). cf.Palaeobullia Götzinger and Becker; Bryant and Pickerill, figs. 8 and 9. Archaeonassa fossulata Fenton and Fenton; Buckman, p. 221, and fig. 8a, b. Archaeonassa fossulata Fenton and Fenton; Buckman, p. 191-201, and pi. 15b-g. Scolicia de Quatrefages, p. 265. Palaeobullia Götzinger and Becker, ?Scolicia prisca de Quatrefages; Chamberlain, p. 225.
Type ichnospecies: Archaeonassa fossulata Fenton and Fenton 1937; by monotypy. Diagnosis: Epireliefs composed of two convex parallel lateral levées separated by a flat, convex, or concave central zone, which is greater or less commonly equal in width to that of a single levée. Levées and central zone either smooth or variably ornamented (typically by oblique or transverse elements). Not representing the lower surface of a three-dimensional back filled burrow such as Palaeobullia or Scolicia. Archaeonassa fossulata Fenton and Fenton 1937 Figs. 2, 3 Occurrence: Common within the Mullaghmore Sandstone Formation, of Co. Sligo and Donegal, from at least six localities (Fig. 1). Diagnosis: As for the ichnogenus. Preservation: Occurring as positive epireliefs, most
TAXONOMIC DISCUSSION The original description of Archaeonassa is clearly based upon modern lebensspuren, and therefore is not covered under zoological nomenclature; article la and lb (7) of the 1985 ICZN code (see Rindsberg, 1990, p. 60); nevertheless, as A. fossulata was erected on fossil material, the ichnogenus is here retained. Modern traces similar to Archaeonassa (Abel, 1935, p. 209-213; Knox and Miller, 1985, pi. la-c, e) are not included within this ichnogenus as they have yet to pass the fossilization barrier, and are therefore not trace fossils; albeit representing excellent analogues of the fossilized material. According to Häntzschel (1975, p. W37-W38), Archaeonassa is probably referable to the Scolicia 'group', although Archaeonassa cannot be considered as synonymous with Scolicia, given that the latter is a much more complex back-filled burrow (see D'Alessandro and Bromley, 1987, p. 749), and the differences in toponomy between Archaeonassa and Scolicia (see Fig. 7). Archaeonassa may therefore be broadly considered as belonging to the Scolicia 'group', as indicated by Häntzschel (1975), although the taxonomic position of the Scolicia 'group' requires further investigation. Material that can be considered as representing Archaeonassa is indicated in the synonymy list (see systematics), which includes material previously described as Palaeobullia, Scolicia, or referred to informally as annelid and gastropod trails. Material identified as Palaeobullia by Knox and Miller (1985) can in part be referred to Archaeonassa (see systematics). Certainly the modern trails recorded by Knox and Miller (1985, pi. la-c, e) are very like Archaeonassa, to which Knox and Miller compared Pennsylvanian material (e.g. Knox and Miller, 1985, pi. 2, fig. f)- Other Palaeobullia in Knox and Miller's paper are here not considered synonymous with Archaeonassa. The hyporeliefs in particular (Knox and Miller, 1985, pi. 2, fig. a, d), having greater affinities to the Cruziana of Osgood and Drennen (1975) rather than Archaeonassa. Palaeobullia illustrated by Bryant and Pickerill (1990) can be clearly
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n=25 6U
1
420
(a)
. I ..
TOTAL WIDTH (MM) Fig. 4. Graphs of Archaeonassa fossulata from the Mullaghmore Sandstone Formation. (A) Histogram of trail width, with (a) range of material recorded in Häntzschel (1975, p. W37). (B) Scatter diagram of trail width against levée width.
Fig. 5. Sketches demonstrating the variation in observed ornament from the Mullaghmore Sandstone Formation Archaeonassa fossulata. (A) Smooth. (B) Standard forms, exhibiting variable ornament definition, with broad levées (1), or narrow levées (2). (C) An example with axial ornament divided in a tripartite fashion.
Fig. 6. Sketch of part of block BELUM K21873, from Skerrydoo, indicating that where trails follow parallel to ripple crests (RC), that a lateral levée is only developed on the ripple trough side. Therefore, apparently appearing to represent a trace fossil comprising a single positive ridge. Arrows indicate cross-over points, at ripple crests, where the true morphology of the trail can be ascertained; cross-hatch indicates wash-outs. Insert (top right): profile of trail cross-sectional shape, normal to trail length, at point a-b.
movement, and therefore is ethologically distinct from both Palaeobullia and Scolicia. Some of the material described by Hall as gastropod and annelid trails is here also considered to represent Archaeonassa (see systematics), although it should be noted, that even given visual inspection of the original material, it is not clear whether Hall's material is preserved in epirelief or hyporelief. Hall's material (Hall, 1852, pi. 13, figs, lb-d; pi. 14, figs. 1, 2 (c-c'), 3 (d-d', e-e'); pi. 11, fig. 4; pi. 12, fig. 2, 4) is assumed to be preserved in epirelief. In the unlikely outcome that the latter are hyporeliefs, they would be more appropriately applied to an ichnogenus such as Cruziana or Didymaulichnus. Only one ichnospecies of Archaeonassa has been erected, namely A. fossulata Fenton and Fenton 1937. However, a number of apparently morphologically distinct types of axial ornament can be differentiated: simple (smooth or transversely ornamented) [standard A. fossulata]; with ring like structures (Hall, 1852, pi. 14, figs. 2, 3; Fig. 8 herein), or swellings (Bryant and Pickerill, 1990, fig. 8); or with a tripartite axial ornament (Figs. 2d, 5c). Variability in the prominence of transverse ornament has been interpreted as a factor of the degree of water
Archaeonassa: taxonomy
FEATURE
189
PALAEOBULLIA Fenton and Fenton 1937
Gotzinger and Becker 1932
SCOLICIA de Quatrefages 1849
A
g 3 z o
ag-a g
As ECHINOIDS (JURASSIC-PRESENT)
MAINLY POSTPALEOZOIC
INTERTIDAL
DEEP (FLYSCH)OFFSHORE SHELF
1, 2, 3, 5
1, 2, 3, 4, 5
> Ü
o _l
o
cu
OS
o
Fig. 7. Comparison between Archaeonassa and ScolicialPalaeobullia. (A) Archaeonassa produced by surface movement, (B) Archaeonassa produced by subsurface movement, but without producing an obvious subsurface burrow; both forming positive epireliefs at the sediment-water/air interface. (C) Palaeobullia produced at the surface (negative epirelief), or an eroded endogenic burrow ofScolicia; in both cases the constructor of the trace fossil occurring above the observed negative relief. (D) Scolicia produced by subsurface movement (endogenic), and comprising a complex backfill (illustration much simplified). Key to morphology: 1 straight, 2 curved, 3 sinuous, 4 meandering, 5 looped.
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TOPONOMY & ETHOLOGY
10 mm Fig. 8. Sketch oiArchaeonassafossulata (AMNH 30975, previously described as "trails of Annelida?"), from the Clinton Group of North America. Note the ring-shaped additions along the trails length. Based on Hall (1852, plate 14, figure 3).
saturation of sediments (Knox and Miller, 1985), and therefore to be environmentally/facies controlled. If other observed variations in ornamentation (Figs. 2d, 5c, 8) are also dependent on environmental factors such as sediment consistency, then such features would be of little value as ichnospecific criteria. If, however, such variations represent behavioural differences, they might be considered as of ichnospecies significance (see Bromley, 1990, p. 157). Such material (Figs. 2d, 5c, 8) is for the meantime considered as representing within ichnospecies variation of A. fossulata, given that the significance of the axial structures is at present unknown, their small number of known occurrences, and their axial continuity with 'normal' A. fossulata. Therefore, at present, Archaeonassa is considered to be monospecific. Archaeonassa can pass into Unisulcus (simple V-shaped surface plough trails), as observed from one example from the study area (Fig. 2e). Whilst Archaeonassa and Unisulcus can appear similar, the two are usually distinguishable. The taxonomic, ethological, and environmental implications of this relationship is, however, worthy of future study.
The ichnogenus Archaeonassa occurs as epireliefs on sandstones. From examination of the overall trail morphology (Fig. 7) and comparison to modern analogues of Archaeonassa (Fenton and Fenton, 1937; Abel, 1935; Frey, 1975; Knox and Miller, 1985) this ichnogenus can be interpreted as locomotory or possibly predatory trails. Modern analogues of Archaeonassa are produced as epireliefs at the sediment-water/air interface (= exogenic structures), either by surface movement of epifauna (Fenton and Fenton 1937, fig. 1) or by shallow subsurface movement of infauna (Knox and Miller 1985, pi. 1, fig. c, e), whereas members of the Scolicia 'group' are typically assumed to represent subsurface endogenic or intergenic structures without exogenic expression. A number of features can be observed from the Irish Archaeonassa which indicate that they represent exogenic trails: these include cross cutting by the funnel-shaped openings of Arenicolites carbonarius (and fluidized sand ejected from the funnels of A. carbonarius); cross cutting by small Asteriacites lumbricalis (c. 10 mm) which represent exogenic resting traces (Cubichnia) or shallow (in the region of 5 mm deep) hiding burrows (Domichnia); the washing out of Archaeonassa on ripple crests (Fig. 6); and the occurrence of ripples slumping over ripple parallel sections. Archaeonassa from the Mullaghmore Sandstone Formation can, at least in part, be interpreted as the product of infaunal movement (Fig. 3), in which no distinct burrow is produced below the exogenic trace of the Archaeonassa.
ENVIRONMENTAL SIGNIFICANCE Archaeonassa is a useful environmental indicator, being typical of intertidal regimes (see examples in Hall, 1852; Fenton and Fenton, 1937; Miller and Knox, 1985; Knox and Miller, 1985; Bryant and Pickerill, 1990; Buckman, 1992a, b). Additionally, material recorded by Alpert (1975, 1977) comes from an environment regarded as shallow water in character (Alpert, 1975, p. 509), presumably marine (containing amongst others Bergaueria, Cruziana and Zoophy cos), and Scolicia vada (= Archaeonassa) recorded by Chamberlain (1971,1978) occurs in a Skolithos-Cruziana ichnofacies (Chamberlain, 1978, tab. 1). Within the Mullaghmore Sandstone Formation Archaeonassa is found in coarsening-upward sequences capped by thick channel-sandstone deposits, is closely associated with Olivellites and Diplocraterion and is commonly underlain by Arenicolites, Skolithos and Teichichnus. Two different environmental associations are observed: Archaeonassa can occur sparsely and sporadically on large-scale rippled surfaces, commonly in close association with V-shaped surface plough trails (association 1); or more
Archaeonassa: taxonomy
typically occur within thinly-bedded, wave-rippled, muddy sandstones, interbedded with shales, where it is commonly prolific in occurrence (association 2). Both types of association represent shallow-water environments. The Archaeonassa of association 1 represent the colonization of shallow-water sandflats within an interdistributary bay type environment, comprising relatively clean sands. The more common association (association 2) represents the preferred conditions for the occurrence and preservation of the Irish Archaeonassa, within muddy sandstones, of an intertidal flat environment of variable salinity. As such, Archaeonassa appears to have important environmental implications and may be of use in environmental reconstruction, particularly in Paleozoic deposits. It should be noted, however, that modern trails similar to Archaeonassa have been illustrated by Hollister et al. (1975) from shelf environments (146 m depth) and also from abyssal environments (4, 694 m depth); although such material may be more appropriately assigned to Palaeobullia. It has been suggested, by the study of modern analogues, that features such as the definition of transverse ornament may be of environmental significance, indicating the relative position on a sandflat (Knox and Miller, 1985). However, the Irish material can vary in ornament definition over very short distances, indicating that in some cases the small scale local environment, depth of burrowing, speed of locomotion, or other factors may also be of importance in determining the nature of ornamentation. Nevertheless, the presence of strong well marked transverse ornament would tend to suggest a firmer more cohesive substrate that was subaerially exposed (Knox and Miller, 1985).
LIKELY PRODUCER From comparison with similar modern gastropod trails (Abel, 1935; Fenton and Fenton, 1937; Knox and Miller, 1985), the most likely producer would appear to be a gastropod, although no taxa can be identified as the producer for any particular occurrence. The possibility that Archaeonassa were produced by organisms other than gastropods cannot be completely excluded. Hyporeliefs associated with Hall's material have been interpreted as Cruziana (Osgood and Drennen, 1975). As these hyporeliefs are commonly associated with Archaeonassa, from elsewhere other than the Irish study area, it could be that trilobites were responsible for some Archaeonassa. Feeding traces made by sea-birds illustrated by Schäfer (1972) and Cadée (1990) are morphologically similar to Archaeonassa, but can be excluded, as they are typically of much greater width, shorter, with smaller marginal levées, and of less complex pattern. In addition no avian bioturbation is known in the Paleozoic, as birds had yet to evolve. An
191
example of a modern irregular echinoid surface trail illustrated by Hollister et al. (1975, fig. 21.10) appears similar to Archaeonassa, although the details of the central section differ. The latter example illustrates the difficulty that can occur in trying to differentiate Palaeobullia forms of 'Scolicia' from Archaeonassa. It is also of note that modern trails, on beach sand, produced by a struggling moth on its back, can also appear broadly similar to Archaeonassa (Frey, 1975, fig. 2.7c, d), which as noted by Frey (1975, p. 27) is comparable to a snail-like crawling trace. Although such traces can be considered as 'accidents', with little chance of preservation (Frey, 1975, p. 27), their occurrence is of importance, further indicating that not all material that appears to correspond to Archaeonassa was necessarily the product of gastropods. Although gastropods are the most likely constructor of these trails, the use of Archaeonassa should not be restricted to trails produced by gastropods (as per Scolicia and echinoids), as no single organism, family or grouping of organisms, can be held as solely responsible for producing Archaeonassa.
AGE RANGE The ichnogenus Archaeonassa is known over a wide span of the Paleozoic; Cambrian (Fenton and Fenton, 1937; Alpert, 1975, 1977; Bryant and Pickerill, 1990), Silurian (Hall, 1852) and Carboniferous (Chamberlain, 1971, 1978; Miller and Knox, 1985; Knox and Miller, 1985; Buckman, 1992a, b). Similar traces are also known from the present day (Abel, 1935; Fenton and Fenton, 1937; Knox and Miller, 1985). Therefore, Archaeonassa has a range of Cambrian-Carboniferous, which may by extended to the present day by comparison to modern trails. The apparent lack of post-Paleozoic fossil examples may be because of identification into the Scolicia 'group'. It may also reflect a post-Paleozoic increase in shallow tier bioturbation, leading to a lower preservation potential of surface trace fossils such as Archaeonassa. Alternatively, it may simply represent a gap in the study of suitable postCarboniferous shallow-water intertidal clastic deposits. Finally, if these trace fossils were produced by trilobites, the lack of post-Paleozoic examples would then correlate with the extinction of the trilobites, the ecological niche represented by this ichnogenus being later taken by gastropods as observed from present day trails.
CONCLUSIONS Material that can be considered as synonymous with Archaeonassa has previously been assigned to Scolicia (Chamberlain, 1971), Palaeobullia (Knox and Miller,
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1985; Knox and Miller, 1985; Bryant and Pickerill, 1990), Crimes, T.P. 1987. Trace fossils and correlation of late Precambrian and early Cambrian strata. Geological Magazine, 124:97-119. or simply been referred to as annelid or gastropod trails (Hall, 1852). Archaeonassa is probably widespread in D'Alessandro, A., and Bromley, R.G. 1987. Meniscate trace fossils and the Muensteria-Taenidium problem. Palaeontology, 30:743-763. occurrence, but assigned to either Palaeobullia or to the Fenton, C.L., and Fenton, M.A. 1937. Archaeonassa, Cambrian snail broad-based Scolicia ' group' (including Palaeobullia). trails and burrows. American Midland Naturalist, 18:454-456. This trace fossil, in general, appears to be the product of Frey, R.W. 1975. T he realm of ichnology, its strengths and limitations. In Frey, R.W. (ed.), T he study of trace fossils. Springer-Verlag, New gastropods, either as a true surface trail, or as the surface York, p. 13-38. expression of shallow subsurface movement. They are George, T.N., and Oswald, D.H. 1957. T he Carboniferous rocks of the interpreted as either locomotory or possibly predatory Donegal Sy ncline. Journal of the Geological Society of London, trails. Some Paleozoic Archaeonassa may have been pro 113:137-183. duced by trilobites, and the ichnogenus should therefore Giitzinger, G., and Becker, H. 1932. Zur geologischen Gliederung des Wienerwaldflysches (Neue Fossilfunde). Geologische Bundesanstalt not be restricted to the presumed work of gastropods. Wien, Abhandlungen; Jahrbuch; Verhandlungen, 82:343-396. Archaeonassa appears to be most common within inter Hall, I. 1852. Palaeontology of New York, V.2. State of New York, tidal environments, and as such is of value for reconstruct Albany, 362 p. ing Paleozoic paleoenvironments. Hantzschel, W. 1962. Trace fossils and problematica. In Moore, R.C.
ACKNOWLEDGMENTS
T his paper represents part of work carried out under the tenure of a research grant from the Department of Education for Northern Ireland, which is gratefully acknowledged. M.J. Benton and A.D. Wright are thanked for their comments on various stages of the manuscript, as are also P.E Carey and R.J. McCaffrey. Bristol University and Queen's University of Belfast are acknowledged for the use of facilities. ET. Fiirsich and one anonymous reviewer are thanked for their useful com ments.
REFERENCES Abel, 0. 1935. Vorzeitliche Lebensspuren. Gustav Fischer, Jena, 644 p. Alpert, S.P. 1975. Planolites and Skolithos from the Upper Precambrian Lower Cambrian, White-Inyo Mountains, California. Journal-of Paleontology, 49:508-521. Alpert, S.P. 1977. Trace fossils and the basal Cambrian boundary. In Crimes, T.P., and Harper, J.C. (eds.), Trace fossils 2. Geological Journal, Special Issue, 9:1-8. Bromley, R.G. 1990. Trace Fossils: Biology and Taphonomy. Unwin Hy man, London, 280 p. Bryant, I.D., and Pickerill, R.K. 1990. Lower Cambrian trace fossils from the Buen Formation of central North Greenland: preliminary observations. Gr0nlands Geologiske Unders0gelse, Rapport 147: 44-62. Buckman, 1.0. 1992a. Palaeoenvironment of a Lower Carboniferous sandstone succession northwest Ireland: ichnological and sedimen to!ogical studies. In Parnell, J.(ed.), Basins on the Atlantic Seaboard: Petroleum Sedimentology and Basin Evolution. Geological Society Special Publication, 62:217-241. Buckman, 1.0. 1992b. Lower Carboniferous Trace Fossils from north west Ireland. Ph.D. T hesis, Queen's University, Belfast, 356 p. Catlee, G.C. 1990. Feeding traces and bioturbation by birds on a tidal flat, Dutch Wadden Sea. lchnos, 1:23-30. Chamberlain, C.K. 1971. Morphology and ethology of trace fossils from the Ouachita Mountains, southeast Oklahoma. Journal of Paleontol ogy, 45:212-246. Chamberlain, C.K. 1978. A Guidebook to the Trace Fossils and Paleo ecology of the Ouachita Geosy ncline. Tulsa, Society of Economic Paleontologists and Mineralogists, 68 p.
(ed.), Treatise on Invertebrate Paleontology, part W. New York & Lawrence, Kansas, Geological Society of America and University of Kansas Press, W l77-W245. Hantzschel, W. 1965. Vestigia invertebratorum et problematica. Fos silium Catalogus. I: Animalia, 108. s'-Gravenhage, W. Junk, 142 p. Hantzschel, W. 1975. Trace fossils and problematica. In Teichert, C. (ed.), Treatise on Invertebrate Paleontology, part W. Boulder, Colo rado and Lawrence, Kansas, Geological Society of America and University of Kansas Press, 269 p. Hollister, C.D., Heezen, B.C., and Nafe, K.E. 1975. Animal traces on the deep-sea floor. In Frey, R. W. (ed.), T he study of trace fossils. Springer-Verlag, New York, p. 493-510. Knox, L.W., and Miller, M.F. 1985. Environmental control of trace fossil morphology. In Curran, H.A. (ed.), Biogenic structures: their use in interpreting depositional environments. Society of Economic Pale ontologists and Mineralogists, Special Publications, 35:167-176. Miller, M.F., and Knox, L.W. 1985. Biogenic structures and depositional environments of a Lower Pennsylvanian coal-bearing sequence, northern Cumberland Plateau, Tennessee, U.S.A. In Curran, H.A. (ed.), Biogenic structures: their use in interpreting depositional envi ronments. Society of Economic Paleontologists and Mineralogists, Special Publications, 35:67-97. Osgood, R.G., and Drennen, W.T. 1975. Trilobite trace fossils from the Clinton Group (Silurian) of east-central New York State. Bulletins of American Paleontology, 287:299-348. Oswald, D.H. 1955. T he Carboniferous rocks between the Ox Mountains and Donegal Bay. Journal of the Geological Society of London, lll: 167-186. Parnell, J., Monson, B., and Buckman, J.O. 1992. Excursion guide: basins and petroleum geology in the north of Ireland. In Parnell, J.(ed.), Basins on the Atlantic Seaboard: Petroleum Sedirnentology and Basin Evolution. Geological Society, Special Publications, 62: 449-464. Quatrefages, M.A. de 1849. Note sur la Scolicia prisca (A. De Q.), annelide fossile de la craie. Annales des Sciences Naturelles, Zoo logie, series 3, 12:265-266. Rindsberg, A.K. 1990. Ichnological consequences of the 1985 Interna tional Code of Zoological Nomenclature. lchnos, 1:59-63. Schafer, W. 1972. Ecology and palaeoecology of marine environments. Oliver and Boyd, Edinburgh, 568 p. Sheridan, D.J.R. 1972. Upper Old Red Sandstone and Lower Carbo niferous of the Slieve Beagh sy ncline and its setting in the northwest Carboniferous basin. Geological Survey of Ireland, Special Paper, 2:129 p.
