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    Ekdale - Pitfalls of Paleobatimetrci Interpretation Based On Trace Fossils Assamblages

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    464 PALEOBATHYMETRY THEME ISSUE Pitfalls of Paleobathymetric Interpretations Based On Trace Fossil Assemblages A.A. EKDALE. Department of Geology and Geophysics, University of Utah, Salt Lake City, UT 84112 PALAIOS, 1988, V. 3, p. 464-472 Trace fossils have ben used widely as indicators of original water depth in palecenvironmental studies, but most paleobathymetric interpretations in ichnology have been based on only @ small ‘number of standard ichnofacis. In addition to bathymetry, itis important to recognize that salinity, oxygen concentration, and substrate character also play a major role in controling the distribution of trace fosils. Several pitfalls of paleobathymeiric interpretation based on trace fossils can lead to erroneous palecencironmental reconstructions. The most common ofthese ‘pitfalls, of which historical geologists should be wary, include the following: 1) occurrence of a particular ichnogenus does not necessarily indicate the presence of the icknofactes of the same name, and a standard ichnofacies may be identified without the -rosence of the namesake trace fossil; 2) environmental shifs of ‘certain trace fossils have occurred through time, so the paleoen- sironmental significance of a particular ichnotaxon may have ‘hanged during its history: 3) the nine standard ichnofacis that ‘have enjoyed wide use by ichnologists have broad environmental significance that may include, but certainly extends beyond, bathymetry: 4) not all environmental situations are represented fn the nine standard ichnofacies, so many trace fossil assoca- tions in the real world cannot be easily fit into those categories. INTRODUCTION “Trace fosis have been known since the last century to be the preserved evidence of organism behavior, but only forthe past tree or four decades have they been employed widely in Jaleoenvconmentalstuties. The important pioneering work of Adolf Selacher established our modern frameworks for both ethologie and ecologic interpretation of trace fosss, especially invertebrate trace fossils in the marine realm. Seiacher’s approach, published initially in German (Seilacher, 1953, 1955, 1959) and later in English (Seilacher, 1962, 1964, 1967, 1974, 1977, 1978), has been to examine both ndvidal trace oss and entre associations in order to make reasonable pleocnvi- Tonmental interpretations ‘Copyrigt © 1988, The Society of Econ Paleorblgiss and Mineralogie Seilcher (1964, 1967) noted that recurrent assemblages of particular trace fossil taxa must have paleoenvironmental implications, and he established the concept of ichnofacies to ilustrate such implications. An ichnofacies (sometimes also called a trace fossil facies) is a characteristic association of, trace fossils that directly reflects certain environmental conditions, such as bathymetry, salinity, substrate character, and so forth. This is basically diferent from an ichnocoenosis (also called a trace fossil community), which represents a group of traces emplaced more or less simultaneously at various levels within a sedimentary deposit by a single endobenthic community of organisms. The ichnofacies concept js based on the common recurrence of particular types of ichnocoenoses in rocks that were deposited in similar environments Seilacher’s ichnofacies model, at least initially, was based explicitly on water depth (Fig. 1). However, other aspects of benthic habitats, such as sediment grain size, energy of the depositional environment, and availabilty and predictability of food resources, were implicit. Nevertheless, many paleobathy- metric interpretations in ichnology have been based rather strictly on a few standard ichnofacies. Ichnologists now realize that the distribution of trace fossils may be a direct result of water depth in some cases but not in others. In fact, itis safe to say that trace fossils are distributed according to bathymetry only insofar as other ecologic factors governing the distribution of trace-making organisms are cor- related with water depth. These factors of course may include bathymetric gradients in sediment composition and texture in some settings, wave andior current energy in others, of otganic content of the sediment in still others. ‘The distribution of marine organisms themselves certainly cannot be attributed to any single environmental parameter, and the distribution of marine organism traces likewise can- not be explained so simplisticaly. However, general bathy- metric gradients can indeed be observed in ‘the composition cof ‘many organism communities, and thus trace fossil assemblages also may exhibit general bathymetric trends. Seilacher’s main point was not that individual trace fossil taxa can be used as precise indicators of absolute water depth, but ngs. 1551680009-046083 00 PALEOBATHYMETRIC PITFALLS 465 that trace fossil assemblages often occur in sequences that ‘may indicate relative water depths within a particular basin. ‘Therefore, although Seilacher's ichnofacies model can be a valuable too! in palecbathymetrc interpretations, many pitfalls wait the reckless geologist who uncrtically applies the Seila- ‘cher bathymetric model too strictly in certain situations. This aper examines some of the most commonly encountered pitfalls, namely ichnofacies that are controlled by environmen tal factors other than water depth, such as salinity, oxygen, and substrate character. Because the potential dangers of paleobathymetric interpre- tation alluded to here do not apply directly to the subaerial realm, only subaqueous environments are considered in this paper; however, itis wise forthe geologist to keep in mind that ‘many’ subaerial facies in the geologic record easily can be Confused with certain subaqueous facies. Also, because inver- tebrate traces predominate in the fossil record, they are emphasized in the following discussions, and trace fossils created by vertebrates and plants are not treated. BATHYMETRY ‘Trace-making organisms in subaqueous environments are distributed according to three primary factors: water depth, water chemistry, and substrate character. Therefore, so are trace fossils. In marine settings (and possibly also in large, deep, fresh-water lakes) as water depth increases, there usually is a concomitant decrease in temperature, light pene- tration, wave or current energy, sedimentation rate, sediment s7ain size, and food supply on the bottom. Thus, bathymetric tends in these critical environmental factors yield a bathymet- ric zonation of the benthic fauna and their traces. ‘Seilacher described the paleoenvironmental, and especially paleobathymetric, implications of trace fossil assemblages in {terms of “universal types of ichnofacies” (1964, p. 307), which he named after particular ichnogenera that seem to be espe- ially characteristic. In the marine realm, from the intertidal zone down to the abyssal plain, four soft-bottom ichnofacies were discerned. In terms of relative bathymetry, the Skolithos Ichnofacies represented intertidal and very shallow subtidal environments, the Cruziana Ichnofacies represented the sub- tidal continental shelf, the Zoophycos Ichnofacies represented ‘outer-shelf and continental-slope (bathyal) depths, and the Nercies Ichnofacies represented the deep (abyssal) sea floor (Fig. 1). He pointed out the tectonic implications of these ichnofacies as wel, indicating for example that the Skolithos Ichnofacies is very prominent in “epicontinental” regimes, and the Nereites Ichnofacies is especially characteristic of “geosyn- inal areas” Geilacher, 1967, p. 414). Although ostensibly’ related to water depth, these four standard ichnofacies have additional environmental significance Geilacher, 1953, 1964, 1967). The Skolithos Ichnofacies typi- fies rather high-energy hydrodynamic conditions in clean, winnowed sediments that are subject to abrupt erosion or deposition. The Cruziana Ichnofacies typically occurs below ‘normal wave base (but not necessarily below storm wave base) in environments of somewhat lower-energy hydrodynamic conditions in muddy and clean sands and silts. The Zooplycos Ichnofacies indicates quiet-water, offshore conditions, probably Decreosng tanpecta sedimentation rate. sediment rin sze, Tood supely IOURE 1—Sellachers basic model of bathymetric zonation of trace {essls, including his fve “universal Ichnofacles” that charecterize marine and non-marine soft-sediment habitats (modified trom Sela. cher, 1967, figure 2). Note that as water depth of the ocean Increases, thee is a decrease in magnitude of several other environ- mental factors below storm wave base, in muds or muddy sands that often are rich in organic matter and sometimes are associated with turbidity current deposits. The Neretes Ichnofacies character- izes fine-grained turbidites in flysch or flysch-lke basinal ‘environments, In addition to the above-mentioned four, several other universal ichnofaces subsequently have been introduced inthe literature. A number of these likewise have been generally accepted by ichnologists as useful in paleoenvironmental recon structions. These incude the folowing: ‘The Scoyenia Ichnofacies represents trace fossil associations in non-marne (Chiefly fluvia-acustrine) sands and. shales Geilacher, 1967; Frey et al, 1984). The Glassifungites Ichno- facies occurs in marine frmgrounds, which typical are mani- fested asst, compacted muds, into which suspension-feeders (cnainly crustaceans and bivalves) have excavated their dwell ing burrows (Sellacher, 1967; Pemberton and Frey, 1985). ‘The Trypanites Ichnofacies encompasses associations of bor- ings in hard, cemented marine substrates (rocky coasts, beachrock, hardgrounds) or shell material (rey and Seilacher, 1980; Ekdale et al, 1984, p. 315). Bromley et al. (1984) distinguished between trace fossil associations in rocky (Clithie") and woody Cyc”) substrates by establishing the Teredoites Ichnotacies for marine woodgrounds. Ekdsle and Berger (1978) described a “Deep-sea Ichnofacies” of infaunal traces that characterize abyssa/hadal sediments deposited by pelagic modes. Because it is mainly the unique ichnofabric rather than a particular ichnotaxonomic composition of that ichnofacies that distinguishes it from Sellacher’s Zaophyos and Nereites Ichnofacies, no ichnogeneric name was applied to it. Stil other universal ichnofacies have been named in the Feature, but the aforementioned nine are the ones that have been adopted most widely. “Thus, at least nine ichnofacies of seemingly universal envi- ronmental significance have been employed successfully by ge- clogists around the world in reconstructing paleoenvironmental settings within particular basins. However, big problems 466 EKDALE TABLE 4—Most common trace-making taxa characterizing various salinity facies BRACKISH WATER’ FRESHWATER VARIABLE SALINITY MARINE WATER HYPERSALINE Insects Tingullds Echinoderms Insects Crustaceans Grustaceans Crustaceans Crustaceans Oligochaetes Oligochaetes Polychaetes Bivalves Polychaetes Bivalves Pulmonates Bivalves Prosobranchs Vertebrates Prosobranchs Opisthobranchs, (fish, etc.) Vertebrates Sipunculans: (fish, ete.) Echiurans ‘Triobites Vertebrates can arise from applying these standard ichnofacies in paleo- bathymetric reconstructions too strictly. Four major pill, which easily can entrap the unwary geologist, include the following: 1. Occurrence ofa particular ichnogenus does not necessar- iy indicate the presence of the ichnofacies ofthe same name. For example, the trace fossil Skoliias itself has been deserbed in fil, tertidl,bathyal, nd even abyssal deposits; yet the trace fossil association known as the Skoliths Ichnofacies does tot have such broad implications. Simlarly, a standard ichno- facies may be identified without the presence ofthe namesake trace fossil For example, Cruziana is a trace fossil created mainly by tlobites, which have been extinct since the end of the Paleozoic Era; yet the trace fossil association known as the CCruziana Ichnofaces may be recognized in the absence of Cnuziana itself, and indeed it does occur in post-Paleozoic rocks as well as Paleozoic rocks (Bromley and Asgaard, 1979). 2. Ina similar vein, itis important to separate aividual ichnotaxa (e.g., Zoophyces) from their namesake ichnofacies (e.g, Zoophycas Ichnofacies) because of the possible incon- stancy oftheir environmental implications. Environmental (Le, bathymetric) shifts of certain trace fossils have occurred through time, so the paleoenvironmental significance of a particular ichnotaxon may have changed during its history. For example, shallow-water occurrences of Zoophycas are quite ‘common in Late Paleozoic rocks, but shallow-water Zoopycos are uncommon in Mesozoic rocks and are virtually unheard of in Cenozoic rocks, where Zooplycos is known primarily a8 a deep-sea trace fossil 3. All nine ofthe standard ichnofacies have broad environ: mental significance that may include, but certainly extends beyond, bathymetry. Use of characteristic ichnofacies as strict bathymetric indicators without considering other environmen- tal factors, as well as other nonichnologic evidence of paleoen- vironments, can lead an investigator to an inaccurate interpre- tation. 4. Not all environmental situations are represented in the above-mentioned nine ichnofacies. These standard ichnofacies simply represent the most common environmentally inked (ish, cetaceans, ete.) trace fossil associations. Therefore, it should not be surprising to a geologist to learn that many trace fossil assemblages ‘observed in the real world do not easily fit into any of the nine categories. SALINITY Salinity isa critical factor for nearly all types of benthic life that produce trace fossils. Stenohaline marine organisms in- clude virtually all echinoderms, articulate brachiopods, sipun- culans, and echiurans as well as many molluscs (e.g., all cephalopods and opisthobranch gastropods and most proso- branch gastropods), arthropods (e.g., all tiobites and most crustaceans), annelids (e.g., almost all polychaetes), and sponges (e.g., the boring clionids). Euryhaline groups include certain inarticulate brachiopods as well as many crustaceans, bivalves, and prosobranch gastropods. Stil other taxa that are restricted to freshwater environments include aquatic insects, pulmonate gastropods, most oligochaete annelids, and certain bivalves (¢-g., unionids and pisdids) and crustaceans (e.g., some amphipods, decapods and ostracods) Itis quite apparent that salinity isa major controt of organism distribution, and thus the distribution of traces created by those organisms likewise is salinity-controlled (Table 1). Trace fossil assemblages in freshwater environments may be moder- ately diverse; that is, they typically are more diverse than in brackish-water and hypersaline environments but less diverse than in normal marine environments (Fig, 2) Originally, Seilacher (1964, 1967) erected a single non- marine ichnofacies, named the “Scoyenia Ichnofacies” after a ‘common ichnogenus that seems to be restricted to fhuvial- lacustrine facies. This association is supposed to represent the characteristic trace fossils of virtually all non-marine deposits, especially redbeds deposited under subaqueous conditions, and it has no implications at all for water depth, The Scoyenia Tchnofacies probably is an unnatural grouping of different trace fossil associations from sedimentary sequences formed in a broad spectrum of settings, including ephemeral, braided, and meandering streams as well as small ponds and large lakes; PALEOBATHYMETRIC PITFALLS 467 theoretically, it also incudes subseril environments, such as paleosos and eolanits, although traces in these environments generally are not attributed to the Scoyenia (or any other named) Ichnofacies. Clearly, itis salinity rather than water depth that separates this particular ichnofacies fom the other, more familar Ichnofaces of the marine realm (e.g., the ‘Skolithos, Crusiana, Zoophyeos and NerctesIchnofaces) The Scoyenia Ichnofacies surely needs to be divided into separate trace fossil associations characterizing the various alluvial, fluvial, and lacustrine (not to mention subaerial) env- ronments, if not broken down further into the numerous assorted facies within those environments, suchas point bars, chanel bars and beaches. Frey et al. (1984) examined the ‘environmental sigicance of common ichnotaxa within this particular ichnofacis, and they sought to restrict the applca- tion of the name Scoenia Ichnofaces to situations. where Scoyenia gracis itslt (andlor the very simlar Ancorichnus coronus) predominates. Frey and Sellacher (1980) described the Scoyenia Iehnofaces a a low-diversity association of traces produced mainly by burrowing arthropods (insects and crusta- ceans). They down-played the contribution of pulmonate gas- tropods, nematodes and oligochaete anneliés to this ichnofa- cies, although traces produced by these groups (not to mention wating birds as well) are not uncommon in fluvablacustrine trace fossil assemblages. Th a thick and varied sequence of Triassic freshwater deposits (Fleming Fjord Formation) in East Greentand, Brom- ley and Asgaard (1979) reported considerable variation in the composition of trace fossi assemblages that could be related to different sedimentary facies. They concluded tat at least some of the trace fossil assemblages were bathymetrcally zoned ‘within an ancient lake environment. Distinct diferences could be observed between trace fossis in shallow lacustrine and deep lacustrine facies. Theit “Scoyenia Assemblage” is inter~ preted as a lake-margin setting subject to occasional subaerial exposure; their “Avenicolites Assemblage” represents a shal Jw lake bottom; and their “Fuersicinus Assemblage” corre- sponds to a deeper environment within the same lake. Studies such as this one by Bromley and Asgzard (1979) promise to elucate the bathymetric relations of freshwater ichnofaces, bt too few ichnologc investigations of this nature in fluviaHlacustrine depositional environments have been ac- complished to alow meaningful generalizations a this time, In certain marginal marine environments, such as estuaries and coastal lagoons, salinity gradients correspond to salnity- controled benthic communities. Lateral changes inthe nature and composition of trace fossil assemblages in such situations may be confused with a simple bathymetnic zonation, when in fact it is salinity rather than water depth that controls the ichnofacies transitions. In marginal marine settings, one usualy can discern an obvious decrease inthe diversity of benthic ite corresponding to a decrease in salinity (Dorjes and Howard, 1975; Frey and Howard, 1980; Miler, 1984), Tnestuaries that are characterized by variable sanites, both the abundance and diversity of benthic organisms tend to be lowest where salinity fuctuations are greatest Miller, 1984) Preservation of organism traces may be less likely in estuarine situations than in other environments, oving to the common dominance of primary bedforms and physical sedimentary aivens Bianes EstuagiEs Diversity SLACOONS o 0 2% 3 40 9 Solinity (Be) —> IOURE 2—Non-quantitative plot of trace fossil versity in suboque us environments relative to various salinity conditions. Diversity of Preserved trace fossils (if present at al) typically is moderete in freshwater environments, low In brackish-water environments, high to very high in normal marine environments, and very low In hyper saline environments. The salintybased trend of trace diversity feflects a similar salinty-based trend of species diversity of organ- structures over biogenic sedimentary structures in river mouth environments. Nevertheless, several trends may allow the geologist to recognize safinity-controlled trace fossil ass0- Giations in the ancient record. These include the following: 1) identification of traces created by strictly non-marine aquatic organisms, such as insects, tubificid oligochaetes, pulmonate sastropods, or unionid bivalves (e.g., see Chamberlain, 1975; Ratcliffe and Fagerstrom, 1980; McCall and Tevesz, 1982), a5, ‘opposed to those created by strictly marine organisms, such as ‘most polychaetes and bivalves and all anthozoans and echino- derms; 2) recognition of ichnotaxa that are known only from non-marine aquatic environments, such as Scoyenia gracitis, Ancorichnus coronus, and Fuersichnus communis, as opposed to the great host of ichnotaxa that have been described only in ‘marine environments; and 3) observation ofa general increase in trace fossil diversity that parallels a transitional fluvialto- ‘marine sequence exhibited by other (non-ichnologc) sedimen- tary criteria, such as bed geometries and primary sedimentary structures OXYGEN Oxygen concentrations of bottom water and interstitial water directly influence the faunal composition of benthic communities, and vertical changes in trace fossil associations actually may reflect fluctuations in the oxygen available to benthic organisms. Oxygenation of the water is important to all animals, but some taxa can tolerate lower oxygen levels than others (Table 2). Thus, certain types of trace fossils can indicate original oxygen conditions at the sea floor, and transitions from one type of ichnocoenosis to another may reflect changes in the amount of oxygen at the sea floor (Bromley and Ekdale, 1984; D'Alessandro et al, 1986; Savrda EKDALE TABLE 2—Most common trace-making taxa character- izing various oxygen facies in the marine realm AEROBIC DYSAEROBIC ANAEROBIC Crustaceans Crustaceans (Crustaceans Nematodes Nematodes Nematodes Polychaetes Polychaetes Bivalves Bivalves Echiurans Echiurans ‘Sipunculans Priapulids Gastropods Echinoderms Nore: Insfcent published data precide generalizations about charac teristic faunas of nonmarine oxygen faces and Bottjer, 1986). In general, trace fossil diversity correlates positively with oxygen concentrations of bottom water (Fig. 3). Although they are sometimes confused in paleoenviron- mental reconstructions, it is very important to. distinguish between the oxygen content ofthe bottom water (immediately above the seafloor) and that of the interstitial water (within the sediment). An animal may be able to lve in anoxic sediment if it maintains an open connection to the sea floor through which to circulate oxygenated bottom water. In oxyges-poor environments, the sediment commonly con- tains abundant unoxidized organic matter for depositfeeding animals to feed upon. Such environments usually lack sufficient current activity to bring a constant stream of suspended food particles into the water column for fiter feeders to consume. For this reason, deposit-feeding burrow systems (fodinichnia and pascichnia) are common, and permanent dwelling struc- tures of fter-feeders (domichnia) are rare in sediments depos- ited under reducing conditions. Endostratal pascichnia (horizontal deposit-feeding trails within the sediment) usually lack an open connection to the sediment surface. This suggests that the sediment may be corganic-rich, but the interstitial water may not be totally devoid of oxygen. On the other hand, complex fodinichnia (deposit- feeding burrow systems constructed for systematically mining the sediment) usually have a connection to the sediment surface. Such fodinichnia may be formed in anoxic, organic-rich sediment at some distance below the water-sediment inter- face, where the organisms can feed upon unoxidized carbona- ‘ceous detritus and at the same time breathe oxygenated water brought down from above. ‘Organisms that can tolerate low-oxygen (dysaerobic) condi- tions tend to be feeding generalists with broad environmental tolerances. Thus, “r-selected” opportunists, as opposed to "K-selected” specialists, usually dominate in oxygen-poor en- vironments (Ekdale, 1985). In fact, the ichnologic signature of coxygen-depleted deposits is a very high-density, very low- diversity assemblage of “r-selected,” deposit-feeding burrows (e-g., Chondrites and/or Zoophycos). in’ summary, different animal taxa have different oxygen Troce Fossil Diversity |AMA#FOR ° os 10 15 Bottom - Water Oxygen (lO HO) [FGURE 3—Non-quanttative plot of trace fossil diversity in subaque- ‘us environments relative to dissolved oxygen concentrations of the bottom water. Diversity of preserved trace fossils typically i ze in completely anaerobic bottom waters, low to moderate in dysaerobic bottom waters, and high to very high in aerabic bottom waters. It the ‘bottom water is low in oxygen (aysserobic) and the sediment s nearly inmpermeable, endostratal mining traces (fodinichnia) dominate: if the bottom water is fow in oxygen but the sediment is somewhat permeable, endostratal grazing Laces (pascichnia) dominate: ifthe Bottom water is rch in oxygen (oerebic), dwelling structures (dom lernia) and locomotion wails (repichnia) are common, but virally ll ‘ypes of traces may occur tolerances, ad itis deposit-feeders that tend to dominate the infauna in oxygen-poor environments (Bromley and Ekdale, 1984; Ekdale and Mason, in press) Animals may inhabit totally anoxic sediments as long as they maintain a connection to at least partially oxygenated bottom water. Endostratal pascch- ni, such as Helminthoida and Spirophycus, dominate in per- meable sediments that are organicrich but aerobic (ora least dysaerobic). Complex fodinichnia, such as Chondrites and Zoopiycos, dominate in impermeable sediments that are ongani-rich and anaerobic, as long as the bottom water is aerobic (or at least dysaerobic) Oxygen-controlled ichnocoenoses might be confused with bathymetrically zoned ichnofacies in some geologic settings ‘where transitions between certain kinds of ichnofacies occur. General relationships between trace fossis and oxygen cond tions are 2s follows: 1. Most environments where oxygen isnot a iting factor contain a moderate to high diversity of trace fossil, including domichna of filter feeding organisms as well as various repich- nia, cubichnia, pascichnia, and so on '2. Oxygen-poor environments generally are dominated by “pselected” Opportunistic ichnotaxa, such as Chondrites, Zoophycos, oF ‘ther complex fodnichnial burrows, in low- diversity Gf not monospectic) assemblages ‘3. Oxygen-depleted sediments, which were deposited in abiotic environments where even the base ofthe water column ‘was anoxic, usually contain po biogenic structures at all and are PALEOBATHYMETRIC PITFALLS 469 ‘TABLE 3—Most common trace-making taxa characterizing various marine substrate types SOUP/SOFTGROUND FIRMGROUND HARDGROUND ‘WOODGROUND Bivalves Bivalves Bivalves Bivalves Crustaceans Crustaceans Crustaceans (Crustaceans. Echinoderms Echinoids Echinoids Polychaetes Polychaetes Polychaetes Prosobranchs Sipunculans Opisthobranchs Demosponges Echiurans ‘Trilobites Vertebrates sh, etc.) Nore: Insufficient pushed dea preciade generalizations about characteristic faunas of non-marineimyrounds,bardgrounds, aed woodgrounds finely laminated. Exceptions occur in situations where lami- nated deposits have been burrowed downward from the top following recolonization of the sea floor ater a return of oxic conditions. Where transitions from highliversty, domichnia-dominated ‘chnocoenoses to low-diversity, fodinichnia-dominated ichno- ‘coenoses to laminated or structureless sediment can be ob- served, sea-floor oxygen conditions (rather than bathymetry) ‘may be the controling factor. SUBSTRATE Composition, texture, stability, and hardness of the sub- strate are critical factors influencing the types of organisms that inhabit the sea floor as well as the types of behavior patterns that are preserved (Table 3). In fact, virtually all trace fossils are controlled in one way ot another by the original substrate character, and the diversity of trace fossil assem- blages likewise may be influenced by the nature of the ‘substrate (Fig. 4). Most interpretations of ichnofacies transi- tions that are based on variations in bathymetry, salinity, or ‘oxygen must assume that substrate character is a constant rather than a variable; for this reason, variations in the ‘substrate can significantly complicate reconstructions of paleo- bathymetry, paleosalinity, or paleo-oxygenation of the deposi- tonal environment, ‘The substrates occupied by benthic organisms may be described by ecologists and ichnologists as various types of “grounds,” such as soupground, softground, firmground, hard- sgfound, or woodground, depending upon their composition and consistency (Fursich, 1978; Ekdale et al., 1984, p. 73-74; Ekdale, 1985, figure 4). For example, ‘Seilacher’s (1967) ‘Skolithos, Cruziana, Zoophycos, and Nercites Ichnofacies usu- ally reflect relative bathymetry, but all represent marine softgrounds. His Scoyenia Ichnofacies represents. non-marine softgrounds andor firmgrounds, regardless of water depth. ‘The other named ichnofacies,'which were mentioned earlier in this paper, represent other types of substrates in marine environments. The Glossifingites Ichnofacies represents trace Trace Fossil Diversity Substrate Consistency —> FIGURE 4—Hon quantitative plot of trace fossil diversity in subaque- ‘us emvronments relative to substrate consistency. Diversity of reserved trace fossils typically is low In soupgrounds, high to very igh In sofgrounds, low to moderate i rmgrounds, and moderate to high in hardgrounds. Locomotion tals (repichnia) and grazing traces {pascichnia} dominate in soupgrounds: itualy all types of wails and burrows occur in softgrounds; dwelling traces (domihnia) dominate in frgrounds and nardgrounds. fossil associations in firmgrounds, where special types of, burrows have been excavated in stiff, compacted but uncement- ced sediment (Pemberton and Frey, 1985). The Trpanites Tchnofacies represents associations of bioerosion traces, espe- cially borings, produced in hardgrounds and shell material (Bromley, 1972; Frey and Seilacher, 1980). The Teredoites Tchnofacies represents associations of borings made in wood Bromley et a., 1984), ‘Thus, a transition from the Skolithas to Cruziana to Zoophycos Ichnofacies commonly indicates a shift from shallow 470 EKDALE to deep water, but a transition from the Skolithas to Glassi- ungites to Trpanites Ichnofacies definitely indicates a shift from soft to frm to indurated substrates, regardless of water depth. In general, the animals that inhabit firmgrounds and hardgrounds ae tiited by water depth only insofar asthe frm and hard substrates themselves are bathymetrically restricted Although there are few hardground surfaces avaiable for colonization by borers in the deep sea, bioerosion traces have been observed in shells and rocks from deep-sea environments (Zeft and Perkins, 1979; Ekdale et al., 1984, figure 10-11; Golubic et al, 1984). Although bored logs that typify the Teredolites Ichnofacies are most characteristic of shalow- Iharine environments, the ichnogenus Teredoites is known to be produced by wood-boring bivalves inthe deep sea (Turner, 1973) and has been reported in wood deposited in fluvial environments (Plint and Pickeril, 1985), In addition to composition, texture, and hardness of the substrate a5 controling factors of benthic organism distribu tion, the stability of the substrate likewise can be a eritical influence. All other aspects of the sediment being equal, a shifting versus stable condition of the substrate may profoundly limit the occupation of the sediment by particular types of burrowers. For example, in a high-energy, wel-oxygenated, shallow-marine environment, such as a beach, a large number of burrowing taxa may be precluded purely’ because of the shifting nature of the sandy substrate, which can cause fre- quent collapse and destruction of uniined burrows. However, some infaunal animals, such a$ certain thalassinidean crusta- ceans (€-g., Calianassa major) and onuphid polychaetes (e.8., Diopatra cuprea), have adapted to such unstable substrate concitions by builing burrows with reinforced walls (e.g., See Weimer and Hoyt, 1968; Myers, 1972). Pellet-walled burrows, lke those made by the shrimp Cal- Fianassa today, are known inthe trace fossil record 2s Ophio- morpha. Such distinctive burrows are quite prevalent in se mentary rocks from the Upper Paleozoic to the present, and they occur in a wide variety of paleoenvironmental situations. Occurrences of Ophiomorpha have been described in fhvil deposits (Stewart, 1978; Bown, 1982; Merril, 1984), beacl’ intertidal sandflat sequences (Hoyt and Weimer, 1963; Frey and Mayou, 1971; Frey et al, 1978; Howard, 1978), shallow subtidal sediments. (Howard, 1972; Curran,’ 1984; ‘Kamola, 1984; Pemberton and Frey, 1984) and turbidites (Kern and Warme, 1974; Bottjer, 1982; Link and Bottjer, 1982). The common denominator of these occurrences iS neither ater depth nor salinity nor oxygen concitions nor even sediment ‘composition (Ophiomorpha occurs in both siliciclastic and car- bbonate deposits). The common denominator of virtually all Opkiomorpha occurrences is texture and stablity of the Sub- strate. Wherever Opkiomorpha is found, it nearly always ‘occurs in clean sand (usually fine- to medium-grained) depos- ited under high-energy conditions. Although most such envi- ronments are in intertidal or shallow subtidal marine settings, the occurrence of Ophiomorpha clearly is linked tothe nature of the substrate and not to the depth of the water. ‘Another example ofthe direct tie between Opiriomorpha and substrate character was described by Kern and Warme (1974) in Upper Cretaceous turbidites (Point Loma Formation) in southern California, In this sequence of alternating sandstone and mudstone, pelleted Ophiomorpha and non-pelleted Thalas- Sinoides burrow systems actualy are interconnected and this are genetically associated with each other. The very same animal, probably a decapod crustacean, constructed knobbly- waled Ophiomorpha burrows in the turbidite sand, which was deposited under high-energy conditions, and. smooth-walled Thalassinoides burrows in the interbedded mud, which repre- sented a more stable substrate Some sedimentary sequences, such as tubidite or storm influenced shelf sequences, attest to frequent shits between high-energy and low-eneray depositional regimes. In such cases, the alternation between two different ichnocoenoses, which represent the highenergy and low-energy conditions respectively, may be interpreted mistakenly as resulting from large-scale fivtuations in water depth. Significant differences between pre-depositional and post-deposiional trace fossil as- sociations in turbidite sequences are wellknown (Seiacher, 1962; Kern, 1980; Ekdale et al, 1984, p. 93-96). Pre- depositional traces, typitied by diverse assemblages of agrch- nial burrows (e.g, Paleoicton, Casmorhaphe, and Desmograp- ton) and pascichnial trails (e.g, Helminthoida, Spirophycus, and Scolca), were produced by sediment-eeders inthe muddy sea floor during the long periods between turbidite events. In con- trast, post depositional traces (e.g, Granular, Opkiomorbha, and Phycosiphon) were dug nthe turbiite sand soon after it was deposited and before hemipelagic sedimentation once again cov- ered the sea floor with mud (e.g. see Seiacher, 1964, figure 4 Bkdale et al, 1964, figure 8-2; Bromley and Ekeale, 1986, figure 7 ‘Similarly, storm-generated sedimentary sequences may ex- hibit ‘alternations between different ichnocoenoses that were produced at approximately the same water depth but under ernating high-energy and low-energy conditions. An espe ially cogent example ofthis phenomenon has been described in Upper Cretaceous shelf deposits (Cardium Formation) of ‘Alberta by Pemberton and Frey (1984), who recognized two sdstint, interbedded trace foss associations. One ichnocoen- sis, exemplified by Skolhas, Diplocratenon, and Ophiomor- ‘Pha, occurs in clean, hummocky cross-stratifed sand beds that ‘obviously were deposited by major storm events. The other ichnocoenosis,typiied by a more diverse assemblage contain- ing Chondrits, Coclichnus, Clindrichnus, Gyrochort, Muen- sera, Palacophycus, ?Phoebicinus, Planoltes, Rhizocoralliom, Rosselia, Thalassinoides, and Zoophycos, occurs in mud layers that were deposited during quiet (non-storm) interval. ‘The interesting point about this occurrence in Alberta is that the two associations represent approximately the same water depth under diferent sets of depositional conditions. However, the former ichnocoenosis in the sandstone beds is stereotypical ‘of Selacher’s Skolithos Ichnofacies, which is supposed to represent intertidal environments, and the latter ichnocoenosis in the mudstone units is stereotypical of Sellacher’s Cruziana Tehnofacies, which is. supposed to represent subtidal shelf environments. The critical environmental factor controling the ‘composition and distribution of the two trace fossil assem- biages inthis situation, however, is not variation in water depth but rather a difference in sediment texture and stability, which in turn are related to the energy of deposition of each Ithofaces. PALEOBATHYMETRIC PITFALLS 474 A somewhat different example ofthe influence of substrate character on ichnocoenoses can be seen i the tered structure of many infaunal communities. Various infaunal taxa may ive at different depths within the sediment as a result of thei unique Preferences of interstitial oxygen levels and/or substrate frm ness. In most siuations, there is a downward increase in substrate consistency with burial from soupy to soft to fim to hard (ce, indurated) that corresponds toa downward decrease of interstitial oxygen inthe sediment. Because the production af most trace fosss is substrate-controlled, and because the substrate exhibits a distinct vertical variation, burrows tend to be tiered within the substrate (Eka, 1985, fi. 7). Bromley and Ekeale (1986) described several examples of tiered ichnocoenoses in a variety of geologic settings, and they demonstrated that cross-cutting relationships of diferent kinds of burrows can allow ichnologsts to reconstruct the tiered structure of original infaunal communities. Several tiers of different trace fossils may be superimposed upon one another ina composite ichnofabric. The juxtaposed burrows in such an assemblage were produced at diferent horizons below the sea floor, and thus they actually represent diferent substrate conditions (Le., in terms of oxygenation and firmness). Inthe Upper Cretaceous shel-sea chalk of northern Burope, substrate control of infaunal ichnocoenoses is quite apparent (kale et al, 1984, figs. 9-1, 9-2, 17-1, 17-2). The fne- gained calcarcous ooze was totaly bioturbated, and atleast five tiers of burrows can be deciphered from composite ichnofabris inthe chalk (Bromley and Ekdale, 1986, figs. 4 to 6). The shallow tiers are dominated by Planolites, and the deeper tiers are dominated by Thalassinoide, Zooplyes, and Chondrites, usually in that order. ‘The infaunaapparentiy ‘cupid different tiers within the same sediment atthe same time. Planolites-producers preferred the soupy oF soft sedi- rent near the water-sediment interface, whereas Zoophycos- producers and Chondnites-producers preferred the much sifer (ie., compacted and dewatered) sediment farther below the sediment surface. ‘Composite ichnofabrics that reflect tiered ichnocoenoses can greatly confuse attempts at paleobathymetric interpretation of ichnofaces. In the majority of cases, the best preserved and most vivid trace fossils are those that were produced last in the Sequence of bioturbation events and therefore inthe deepest tier Bkdale, 1985; Bromley and Ekdale, 1986). These latest- stage traces (e.g, Zoophycos and Chondvites in the European chal) are usually the ones ichnologists emphasize in theit descriptions, yet these traces were farther removed from the sea floor and actully less indicative of orignal sea-loor conditions than the more poorly preserved traces of the shallower tiers. Thus, in analyzing the paleoenvironmental significance of trace fossis assemblages, itis important. to recognize that substrate-controled tiring of ichnocoenoses is 4 very common phenomenon in the trace fossil record. CONCLUSIONS Paleobathymetric reconstructions often can be aided by careful analysis of ichnofacies. However, in addition to bathy- is absolutely essential for geologists to recognize the vital role of salinity, oxygen concentration, and substrate character in controling the distribution of particular types of trace fossils. Many common trace fossils are useful as paleo- bathymetric indicators in marine rocks (e.g., Diplocraterion in intertidal and shallow subtidal environments; Paleadictyon and ‘other agrichnia in deep-sea settings), but many others are not. For example, rather than original water depth, some common ichnogenera instead may reflect original salinity conditions (e.g., Scoyenia, Fuersichnus, and tubficid worm burrows in fluviuaVlacustrine environments), oxygen conditions (e-g., Chondrites and Zoophycos in oxygen-depleted sediments), of substrate conditions (e.g. Ophiomorpha in shifting sand sub- strates within fluvial, shallow-marine, and deep-marine set- tings). To historical geologists, this presents both bad news and good news. The bad news is that extensive trace fossil research in recent years has burst our bubble of ichnologic imnocence. No longer can we assume that simple, linear variations in the composition of trace fossil assemblages are correlated directly with water depth in all (or perhaps even most) cases. The good news, of course, is that trace fossils have proved to be excellent paleoenvirontmental indicators that tellus a great deal more about ancient depositional systems and animal habitats than just water depth. It is possible that bathymetric zonations of trace fossil assemblages can be established within the other environmental regimes (salinity, oxygen, substrate, etc.) but much additional

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