<strong>Continental</strong> environments <strong>and</strong> their <strong>trace</strong>-fossilassociations can be classified as the behaviouralproxies of biological community assemblages orichnocoenoses (Figure 4), rather than archetypalichnofacies. The redefined Scoyenia ichnofacies(Frey et al. 1984) <strong>and</strong> the purported Mermia,Coprinisphaera <strong>and</strong> Termitichnus ichnofacies (e.g.,Genise et al. 2000, 2004, Buatois <strong>and</strong> Mángano2004), are merely large lists of <strong>trace</strong> <strong>fossils</strong> that occurin a substratum that is largely ignored, not integratedinto the facies scheme <strong>and</strong> too broad to be of any use(e.g., Hasiotis, 2004). Construction of ichnocoenosesproperly incorporates patterns in bioturbation withthe biophysicochemical controls <strong>and</strong> processes thatoperate in the continental realm, <strong>and</strong> correspond tocharacteristic environmental conditions. Localizedremnants of above- <strong>and</strong> below-ground, <strong>trace</strong>-making,ecological communities are preserved as <strong>trace</strong>-fossilassociations or ichnocoenoses. An ichnocoenosescan contain tiered <strong>trace</strong>s of arboreal, epigeal <strong>and</strong>fossorial organisms that lived together, <strong>and</strong> hadtransient, temporary, periodic or permanentrelationships with the substratum (geophiles <strong>and</strong>geobionts).An ichnocoenoses would be named for the mostabundant or significant pedological <strong>and</strong> ecologicalmodifyingbehaviour in that ichnocoenosis <strong>and</strong>subenvironment. For example, if crayfish burrowsare the dominant <strong>trace</strong> <strong>fossils</strong> of an ichnocoenosis<strong>and</strong> environment, then the <strong>trace</strong>-fossil association istermed the Camborygma ichnocoenosis. If sphericaltermite nests are the dominant <strong>trace</strong> <strong>fossils</strong> of anichnocoenosis <strong>and</strong> environment, then the <strong>trace</strong>-fossilassociation is termed the Termitichnus ichnocoenosis.All things considered, continental <strong>trace</strong> <strong>fossils</strong> providea vast amount of information when studied carefullywith respect to the deposits in which they are found.Traces preserve evidence of differing amounts of soilmoisture <strong>and</strong> the position of the palaeo-water table,<strong>and</strong> their fluctuations through time. Trace <strong>fossils</strong>indicate the presence of large numbers of plants,invertebrates <strong>and</strong> vertebrates, <strong>and</strong> record biodiversity<strong>and</strong> palaeoecologic relations that otherwise areoverlooked when body <strong>fossils</strong> are absent orunderrepresented in sedimentary deposits. Inferencescan be made about the palaeoclimatic setting of anarea in terms of precipitation <strong>and</strong> its seasonalitywhen ichnologic data <strong>and</strong> interpretations are combinedwith other palaeontologic, sedimentologic <strong>and</strong>geochemical field <strong>and</strong> laboratory data. All of theseaspects of continental <strong>trace</strong> <strong>fossils</strong> <strong>and</strong> the study ofichnology can be related to the public using <strong>museum</strong><strong>exhibits</strong>.Trace <strong>fossils</strong> as <strong>museum</strong> <strong>exhibits</strong>The best way to teach the public about the importance<strong>and</strong> utility of continental <strong>trace</strong> <strong>fossils</strong> is to displaythem alongside casts of modern <strong>trace</strong>s that representhomologs or analogues to those ancient behaviours.Some <strong>museum</strong>s already have <strong>exhibits</strong> of dinosaur orreptilian trackways displayed with the trackmaker’sskeleton or its restoration (Hannibal <strong>and</strong> Lucas 2006),however, there is much more to continental <strong>trace</strong><strong>fossils</strong> than trackways. Side-by-side displays ofmodern <strong>and</strong> ancient behaviour allow the public to seehow scientists study <strong>and</strong> interpret the significance of<strong>trace</strong> <strong>fossils</strong>. This kind of exhibit demonstrates thatmodern organisms <strong>and</strong> their behaviours have anevolutionary history through deep geologic time asrecorded by the record of body <strong>and</strong> <strong>trace</strong> <strong>fossils</strong>.Casts of modern <strong>trace</strong>s can be made by pouringfibreglass, epoxy, concrete or dental plaster downinto the burrow, nest or track-bearing surface (e.g.,Shinn 1968, Farrow 1975, Hasiotis <strong>and</strong> Mitchell1993). The burrows <strong>and</strong> nests should be cast so thatthe casting material forms a horizontal plane at theentrance-ground surface interface so that the cast canbe mounted properly with respect to its originalorientation in the subsurface. The constructors of themodern <strong>trace</strong>s can be removed prior to casting or theycan be entombed within the casting medium itself.Specimens of the <strong>trace</strong>maker can also be retrievedfrom similar <strong>trace</strong>s via excavation <strong>and</strong> capture, <strong>and</strong>displayed alongside the cast or actual <strong>trace</strong> examples.Genuine pieces of such nests as those constructed bydung beetles, termites, wasps, bees <strong>and</strong> ants can alsobe displayed alongside their constructors.Photographs of the burrow entrances in which thecasting material was poured or the nest itself can bedisplayed with the <strong>trace</strong> <strong>and</strong> its constructor. Mostpeople have no idea what kind of three-dimensionalstructure lies below a burrow entrance, seen only asan open hole in the ground. Such displays link theopening to the burrow <strong>and</strong> its constructor(s).The exhibit is complete when the modern organism(s)<strong>and</strong> their biogenic structure are displayed togetherwith an ancient continental <strong>trace</strong> fossil that representstheir homologous or analogous behaviour. In thecase of ancient trackways, a display is completedwith the trackmaker’s skeleton or its restoration, <strong>and</strong>complemented with the trackway <strong>and</strong> body of anextant organism with analogous behaviour. Theancient <strong>trace</strong> <strong>fossils</strong> can be displayed in or out of theirsurrounding matrix, depending on how complicatedthe three-dimensional structure is <strong>and</strong> the ease withwhich they can be removed from the matrix. In rarecases, the constructor of the ancient <strong>trace</strong> fossil is-217-
Figure 4. <strong>Continental</strong> ichnocoenoses. Examples of ichnocoenoses found in subenvironments of alluvial,lacustrine, <strong>and</strong> aeolian environments <strong>and</strong> deposits. Tiering <strong>and</strong> distribution in each subenvironment iscontrolled by the behaviour (Fig. 1), groundwater profile (see Fig. 2) <strong>and</strong> depositional processes (Fig. 3 <strong>and</strong>this figure). Abbreviations: AMB-adhesive meniscate burrows, An-Ancorichnus, At-ant nests, Bv-bivalve<strong>trace</strong>s, Ca-Camborygma, Ce-Celliforma, Ck-Cochlichnus, Cl-Cylindrichum, Co-Conichnus, Cp-Coprinisphaera, F-Fuersichnus, G-gastropod trail, Hb-horizontal burrows, Hu-horizontal U-shaped burrow,O-ornithopod <strong>and</strong> theropod tracks, P-Planolites, Rh-rhizoliths, Pt-Pteraichnus, Pts-pterosaur scratchmarks, Sa-sauropod tracks, So-Scoyenia, St-Steinichnus, Tm-termite nest, T/Rh-termite nests in rhizoliths,Ut-ghost U-shaped tubes, Uts-shallow U-shaped tubes, Vb-quasivertical burrows, Vtb-vertebrate burrows,Wp-wasp nest/cocoons, Yt-Y-shaped vertical burrow. Trace fossil illustrations <strong>and</strong> box diagrams are not toscale.-218-
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Figure 8. This Early Cretaceous tra
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