Storm-induced coral rubble deposition: Pleistocene records of natural reef disturbance and community response.

Shallow-water Pleistocene coral reef facies in Barbados (dominated by Acropora palmata rubble) record evidence of deposition under contrasting non-catastrophic - (fair-weather?) and storm-induced conditions. Depositional styles are interpreted on the basis of coral rubble fabrics and calcareous encruster successions. Terrace exposures on the west of the island comprise uniform (3-4 m thick) depositional sequences. Individual coral samples exhibit similarities in encruster community composition and thickness, and a transition from photophilic to sciaphilic encrusting forms. These are indicative of colonization during gradual burial in an accumulating rubble pile. By contrast, NE coast sites comprise repetitive sequences of discrete (0.4-1 m thick) depositional units. Rubble colonization within each unit is characterized by a vertical succession from thin (1-2 mm), sciaphilic encruster-dominated sequences at the base, to progressively thicker (up to 20 mm), photophilic encruster-dominated sequences at the top. These are interpreted as multiple storm deposits, with upper surfaces colonized by opportunist coral species (primarily Agaricia agaricites). In contrast to many modern hurricane-impacted reef systems, however, there is no evidence of long-term shifts in coral community composition following physical disturbance. Colony sizes of opportunist corals at the tops of storm units are consistent with growth over timescales of <10 years. These are overlain on each occasion by a new A. palmata rubble pile, indicating successful recovery over successive physical disturbance cycles.     

Microfacies and depositional environment of an Upper Triassic intra-platform carbonate basin: the Fatric Unit of the West Carpathians (Slovakia)

Facies associations of the Rhaetian Fatra Formation from the Veká Fatra Mts. (West Carpathians) were deposited in a storm-dominated, shallow, intra-platform basin with dominant carbonate deposition and variable onshore peritidal and subtidal deposits, with 21 microfacies types supported by a cluster analysis. The deposits are formed by bivalves, gastropods, brachiopods, echinoderms, corals, foraminifers and red algae, ooids, intraclasts and peloids. A typical feature is the considerable variation in horizontal direction. The relative abundance and state of preservation of components as well as the fabric and geometric criteria of deposits can be correlated with depth/water energy-related environmental gradients. Four facies associations corresponding to four types of depositional settings were distinguished: a) peritidal, b) shoreface, above fair-weather wave base (FWWB), c) shallow subtidal, above normal storm wave base and d) above maximum storm wave base. The depositional environment can be characterized as a mosaic of low-relief peritidal flats and islands, shoreface banks and bars, and shallow subtidal depressions. The distribution and preservation of components were mainly controlled by the position of base level (FWWB), storm activity and differences in carbonate production between settings. Poorly or moderately diverse level-bottom macrobenthic assemblages are dominated by molluscs and brachiopods. The main site of patch-reef/biostrome carbonate production was located below the fair-weather wave base. Patch-reef/biostrome assemblages are poorly diverse and dominated by the branched scleractinian coral Retiophyllia, forming locally dm-scale autochthonous aggregations or more commonly parautochthonous assemblages with evidence of storm-reworking and substantial bioerosion by microborings and boring bivalves.Facies types and assemblages are comparable in some aspects to those known from the Upper Triassic of the Eastern and Southern Alps (Hochalm member of the Kössen Formation or Calcare di Zu Formation), pointing to similar intra-platform depositional conditions. The absence of large-scale patch-reefs and poor diversity of level-bottom and patch-reef/biostrome assemblages with abundance of eurytopic taxa indicate high-stress/unstable ecological conditions and more restricted position of the Fatric intra-platform setting from the open ocean than the intra-platform habitats in the Eastern or Southern Alps.     

FOSSIL DIAGENESIS IN THE BURGESS SHALE

Abstract:  Current models for the exceptional preservation of Burgess Shale fossils have focused on either the HF-extractable carbonaceous compressions or the mineral films identified by elemental mapping. BSEM, EDX and microprobe analysis of two-dimensionally preserved Marpolia , Wiwaxia and Burgessia identifies the presence of both carbonaceous and aluminosilicate films for most features, irrespective of original lability. In the light of the deep burial and greenschist facies metamorphism documented for the Burgess Shale, the aluminosilicate films are identified as products of late-stage volatilization and coincident mineralization of pre-existing compression fossils, whereas the three-dimensionally preserved gut-caecal system of Burgessia is interpreted as an aluminosilicate replacement of a pre-existing carbonate phase. The case for late diagenetic emplacement of aluminosilicate minerals is supported by the extensive aluminosilicification of trilobite shell and (originally) calcareous veinlets in the Burgess Shale, as well as documentation of other secondarily aluminosilicified compression fossils. By distinguishing late diagenetic alteration from the early diagenetic processes responsible for exceptional preservation, it is possible to reconcile the range of preservational modes currently expressed in the Burgess Shale.     

Stratigraphy, geometry, and facies of a Middle Devonian ramp-to-basin transect (Eastern Anti-Atlas, SE Morocco)

Summary In the castern Anti-Atlas (SE Morocco), a small sedimentary basin (Mader Basin) evolved during the late Palacozoic. The Middle Devonian deposits consist of shales and limestones with a thickness up to 700 m in the depocentre. Sedimentary structures and sole marks of Middle Devonian limestones indicate transport from the northwest and the south towards the basin centre, located in the central Mader area. Lithostratigraphic, biostratigraphic, and dynamic stratigraphic approaches were applied to correlate stratigraphic sections. Five correlatable large-scale base-level hemicycles were recognized in the Middle Devonian succession. Thickness trends of Middle Devonian deposits, regional correlations, and facies-trends reveal the geometry of a carbonate ramp. The carbonate ramp was slightly inclined (<1°) to the NE. A sedimentary wedge, consisting of limestones and limestone/marl alternations, was deposited during the Eifelian and marks the transition from the ramp to the adjacent basin. Middle Devonian water depths are estimated as close to and within the storm wave-base at the southern area of the ramp and far below storm wave-base in the northeastern part of the ramp. Shallowest conditions (inner-ramp environment), close or within the fairweather wave-base, existed during the early Givetian as documented by the abundance of recfal fauna (stromatoporoids, corals) and calcimicrobes (lumps, micritic envelopes) in the eastern and southeastern area of the ramp.     

Taphonomy of Sheridan College Quarry 1, Buffalo, Wyoming: Implications for reconstructing historic dinosaur localities including Utterback's 1902-1910 Morrison dinosaur expeditions

The Upper Jurassic Morrison Formation has yielded some of the most important, voluminous and diverse dinosaur bonebeds in western North America, yet many of its historic sites were excavated during the celebrated period of vertebrate paleontology in western North America referred to as the first and second “Great Dinosaur Rush” (1870-1910s). Because of the large quantity of fossils collected during this era, a considerable amount of data pertaining to patterns of sedimentation, preservation, and paleoecology across broad portions of the Morrison Formation (and indeed many other Mesozoic and Cenozoic units) is still poorly understood. This paper critically re-evaluates the Sheridan College Quarry 1 dinosaur bonebed which lies along the western rim of the Powder River Basin in the region of localities excavated during Utterback's expeditions in the 1900s. Sedimentologically, the bonebed is interpreted as having been formed by episodic flooding events affecting the proximal floodplain depocenter of a meandering river system. Limited evidence of bone abrasion or rounding and progressive upsection changes in bone orientations suggest that minimal transport occurred, but that at least four episodes of overbank flooding resulted in the concentration and burial of attritional, time averaged vertebrate skeletal material that accumulated in topographic lows on the floodplain. Taphonomic analysis indicates that multiple unassociated to partially associated fossil elements excavated represent at least three taxa of sauropod dinosaurs, whereas isolated elements from the site indicate the presence of several other small vertebrate taxa. This work provides significant new information not only about the Sheridan College Quarry 1, but also about local sedimentary and taphonomic conditions that were likely influential to burial and preservation of other nearby Morrison dinosaur localities in the Big Horn Mountains, most notably those excavated during the famous Utterback expeditions. This study highlights the research potential in reconstructing lost data for historic dinosaur localities.     

The geological effects of hurricanes on coral reefs and the interpretation of storm deposits

Hurricanes occur in belts 7° to 25° north and south of the equator. Reefs growing in these belts suffer periodic damage from hurricane-generated waves and storm surge. Corals down to 20m depth may be broken and removed, branching colonies being much more susceptible to breakage than upright massive forms. Sand cays may be washed away and former storm ridges may migrate to leeward across reef flats to link with islands. Reef crest and reef front coral debris accumulate as talus at the foot of the fore-reef slope, on submarine terraces and grooves, on the intertidal reef flat as storm ridges of shingle or boulders and isolated blocks of reef framework, as accreting beach ridges of leeward migrating shingle, as lobes and wedges of debris in back-reef lagoons, as drapes of carbonate sand and mud in deep off-reef locations in the fore-reef and lagoonal areas. In addition to the coarse debris deposited, other features may aid the recognition of former hurricane events, including the assemblage of reef biota, its species composition and the structure of the skeletons; graded internal sediments in framework cavities; characteristic sequences of encrusting organisms; characteristic shapes of reef flat microatoll corals; and submarine cement crusts over truncated reef surfaces. The abundance of reef flat storm deposits whose ages cluster around 3000–4000 y BP in certain parts of the world most likely relate to a slight fall in relative sea level rather than an increase in storminess during that period. A higher frequency of storms need not result in more reef flat storm deposits. The violence of the storm relative to normal fair-weather conditions influences the extent of damage; the length of time since the previous major storm influences the amount of coral debris created; the length of time after the hurricane, and before a subsequent storm influences the degree of stabilization of reef-top storm deposits and hence their chances of preservation.     

Optimal hurricane overwash thickness for maximizing marsh resilience to sea level rise

The interplay between storms and sea level rise, and between ecology and sediment transport governs the behavior of rapidly evolving coastal ecosystems such as marshes and barrier islands. Sediment deposition during hurricanes is thought to increase the resilience of salt marshes to sea level rise by increasing soil elevation and vegetation productivity. We use mesocosms to simulate burial of Spartina alterniflora during hurricane‐induced overwash events of various thickness (0–60 cm), and find that adventitious root growth within the overwash sediment layer increases total biomass by up to 120%. In contrast to most previous work illustrating a simple positive relationship between burial depth and vegetation productivity, our work reveals an optimum burial depth (5–10 cm) beyond which burial leads to plant mortality. The optimum burial depth increases with flooding frequency, indicating that storm deposition ameliorates flooding stress, and that its impact on productivity will become more important under accelerated sea level rise. Our results suggest that frequent, low magnitude storm events associated with naturally migrating islands may increase the resilience of marshes to sea level rise, and in turn, slow island migration rates. Synthesis: We find that burial deeper than the optimum results in reduced growth or mortality of marsh vegetation, which suggests that future increases in overwash thickness associated with more intense storms and artificial heightening of dunes could lead to less resilient marshes.     

Paleolimulus, an early limuline (Xiphosurida), from Pennsylvanian-Permian Lagerst tten of Kansas and taphonomic comparison with modern Limulus

The Pennsylvanian-Permian horseshoe crab Paleolimulus signatus (Beecher), incorporating as a junior synonym P. avitus Dunbar, is one of the earliest species of the Limulina (Xiphosurida). Some specimens from Kansas, USA, are exceptionally well preserved, retaining intact book gills and appendages. Indistinct, bilobed burrowing traces of variable width occur in association with some examples of P. signatus and may have been produced by that animal. Based on actualistic taphonomic experiments on Limulus polyphemus, ancient horseshoe crabs and other arthropods having non-mineralized exoskeletons are inferred to have become pliable soon after death or moulting, and to have disarticulated slowly prior to burial. Extreme compression, wrinkling, and loose folding of sclerites are attributed to burial of a pliable exoskeleton. Slow preburial disarticulation partly accounts for the exceptional preservation of Paleolimulus remains. Also relevant for the exceptional preservation of these arthropods was burial in estuarine, tidal flat, or lacustrine environments. Because of fluctuating salinity and possibly dessicating conditions, these settings were limiting to scavengers, burrowers, and some microbes that could potentially disarticulate or decompose xiphosurid remains.     

Reassessment of the mode of life of Pagetia Walcott, 1916 (Trilobita: Eodiscidae) based on a cluster of intact exuviae from the Kaili Formation (Cambrian) of Guizhou, China

Pagetia is the most abundant trilobite in the Kaili Formation (lower to middle Cambrian). During the course of studying the museum collections of Kaili trilobites ( n  > 1000), a cluster containing 22 pagetiid individuals at various growth stages is noted. Specimens record the growth range from the degree 0 meraspid to late holaspid phases. Based on the relative completeness of the moults except for missing free cheeks, these specimens are interpreted as intact exuviae that had undergone minimal transport prior to burial. If Kaili pagetiids had pelagic or planktonic living habits as previously suggested, it would be difficult to explain the presence of intact exuviae in clusters. Therefore, Kaili pagetiids are interpreted here as having a benthic mode of life after the onset of the meraspid phase.     

Fossil analogues of present-day Cladocora caespitosa coral banks: Sedimentary setting, dwelling community, and taphonomy (Late Pliocene, W Mediterranean)

 Cladocora caespitosa is a common zooxanthellate, ahermatypic, constructional scleractinian coral in shallow waters of the present-day Mediterranean. Extensive coral banks in Upper Pliocene shallow marine deposits of the Almería-Níjar Basin (SE Spain) contain the same species. These banks occur on debris-flow conglomerates deposited in a fan delta, or on bioclastic accumulations interpreted as storm deposits. Direct relationships of coral beds with coastal facies indicate that C. caespitosa colonized shallow settings near the paleocoast, probably not deeper than 20–30 m. Low turbulence allowed corals to colonize substrates, which remained stable for long periods. Activity of organisms in the coral community, storms, and detritic discharges from the fan delta were the most significant mechanisms disturbing the coral development. The hard substrata provided by coral banks promoted colonization by cemented and epibyssate organisms. Coral banks marked maximum flooding surfaces at the end of transgressive systems tracks. They were suddenly buried by sediment input into the basin. Taphonomic signatures measured on components of the coral bank communities indicate a low turbulence environment, probably a bay. The low hydraulic energy further inhibited post-burial reworking, thus promoting the in situ preservation of a great part of the organisms inhabiting the bioconstructions. Accepted: 2 December 1997     

The Montceau-les-Mines Lagerstätte (Late Carboniferous,France)

The Montceau-les-Mines Lagerstätte (Late Stephanian, Late Carboniferous) is located northeast of the French Massif Central. Situated at equatorial latitudes during the Pennsylvanian, this Lagerstätte, probably a freshwater environment, preserves a rich and diverse flora (lycopsids, sphenopsids, ferns, pteridosperms, and cordaites) and fauna (bivalves, annelids, crustaceans, myriapods, insects, chelicerates, myxinoids, actinopterygians, sarcopterygians and tetrapods). These exceptionally preserved fossils can be found either flattened in shales or three-dimensionally preserved in sideritic nodules. The fossils from the nodules are exceptional for at least two reasons: the absence of major disarticulation of their body structure and the preservation of soft parts and extremely fragile cuticular structures. Such preservation was made possible by the combination of several factors: rapid burial in fine anoxic mud, early siderite precipitation (inducing the nodule formation) and phosphatization of cuticles and soft-bodied features.     

Exceptional occurrence of fossil baleen in shallow marine sediments of the Neogene Pisco Formation,Southern Peru

Fossil baleen is rare in the sedimentary record. This paper documents the exceptional occurrence of thirty seven fossil whale specimens with preserved baleen in the Neogene Pisco Formation during a transect survey in a limited area west of the Ica River Valley near the town of Ocucaje in southern Peru. The sedimentary layers consist of tuffaceous and diatomaceous sandstones, diatomaceous mudstones, and dolomites, deposited in a shallow marine embayment. Observations of modern whale carcasses on the seafloor and stranded individuals indicate that baleen detaches from the mouth of the whales very rapidly after death, and that bones deteriorate very rapidly as a result of scavenging activity and abrasion. In contrast, the bones of the Pisco Formation whales are exceptionally well preserved, and their baleen is often found in life position suspended from the rostrum. Sedimentary structures found associated with some skeletons indicate tidal and storm processes, suggesting that the environment was not anoxic. This exceptional occurrence of fossil baleen suggests early mineralization of the baleen attachment to the rostrum or rapid burial of the skeletons before any detachment or loss could occur.     

Gyrophyllites cristinae isp. nov. from Lower Ordovician Shallow-Marine Deposits of Northwest Argentina

Abstract

The lower Paleozoic marine siliciclastic succession of the Central Andean Basin, northwestern Argentina, provides a valuable record of the onset of the Great Ordovician Biodiversification Event in western Gondwana. A new ichnospecies of rosette trace fossil, Gyrophyllites cristinae, is documented from lower and upper Tremadocian (Tr1 and Tr2) deposits of this basin. It is characterized by five to six non-overlapping petaloid lobes and can be easily differentiated from the other four formally defined ichnospecies. Gyrophyllites cristinae occurs at the top of hummocky cross-stratified sandstone regularly interbedded with mudstone. These deposits are interpreted as reflecting the alternation of high-energy storm events and low-energy fair-weather conditions immediately below the fair-weather wave base, representing deposition in offshore transition environments. Gyrophyllites has been traditionally interpreted as the product of worms of uncertain taxonomic affinity that mined the sediment in search for food (fodinichnia). The occurrence of Gyrophyllites cristinae in these Ordovician deposits records post-storm colonization. Storms may have increased oxygenation and supplied fresh organic detritus that were exploited by worm-like, surface detritus- or shallow deposit-feeders exploring the uppermost silt-rich fine-grained sediments.     

The origin of echinoid shell beds in siliciclastic shelf environments: three examples from the Miocene of Sardinia,Italy

Three mass accumulations of sea urchins from the Miocene of Sardinia show a number of taphonomic features which set them apart from previously described echinoid assemblages from the Cenozoic in which they represent: (1) monotypic assemblages; (2) include very well‐preserved remains of either regular or spatangoid echinoids; and (3) originate in deeper water environments. These accumulations are compared using a detailed sedimentological and taphonomic analysis including preservational fabrics, taphonomic signatures, size frequency distributions, density of occurrences and preferred orientations. The possible role of gregarious behaviour contributing to mass occurrences and the specific sedimentary events leading to the excellent preservation are discussed. The interpreted depositional environment of all three deposits is that of a storm‐dominated, siliciclastic shelf environment. A phymosomatid assemblage represents rapid burial through obrution of a highly dense, freshly dead community. A Brissopsis‐dominated spatangoid assemblage represents a mixed accumulation of parautochthonous and transported skeletons. The third assemblage consisting of regular echinoid spines and rare tests represents a composite tempestite. Differences in the depositional environments are related to their position along onshore–offshore gradient with the first two beds originated in a deeper setting than that of the spine accumulation. This study shows that the preservation of assemblages containing complete regular echinoids and spatangoids is higher in deeper water settings than in shallow water environments.     

Modern and fossil traces in terrestrial lithic substrates

A number of biogenic processes leads to the formation of distinctive traces in terrestrial lithic substrates. These include: burrowing by vertebrates in moderately lithified rocks; scraping by mammals; smoothing and polishing of limestone surfaces by the locomotion of mammals; excavation by bees, wasps, and ants producing nesting and dwelling tunnels; dissolution of limestone surfaces by terrestrial snails; endolithic activity of fungi, algae, and lichens on subaerial rock surfaces; root corrosion; etc. Processes of biochemical weathering, biophysical erosion, and enlargement of cracks and fissures by the pressure of plant roots do not leave distinctive traces and therefore lie outside the ichnological realm. The fossil preservation of terrestrial bioerosional traces is expected to be uncommon. Nevertheless, various possible means of preservation must be considered, such as by rapid burial by volcanic material, by fluvial sediments, by travertine or tufa, by loess, “conservation”; in caves, case hardening of surfaces of porous rocks, and preservation of subsoil traces below fossil soils.     

Exceptionally well‐preserved vegetal remains from the Upper Cretaceous of ‘Lo Hueco’, Cuenca,Spain

Vegetal remains are considered labile structures that quickly become decayed in ecosystems. However, certain lignified tissues (woody plants) can largely resist decomposition, becoming sometimes exceptionally well preserved. At the Upper Cretaceous site of ‘Lo Hueco’ (Cuenca, Spain), those woody remains (trunks and branches) with resinous material in the inner tracheids and parenchyma cells that were buried rapidly under anoxic conditions experienced a low degree of maturation, becoming exceptionally well preserved. Those woody remains deposited under oxic conditions (sub‐aerial or sub‐aquatic exposure) were more intensely biodegraded and subsequently carbonified, partially or completely mineralized in gypsum and covered by a ferruginous crust. These two modes of preservation are scarce, with silicification or carbonification processes much more common, and both can be considered as ‘exceptional preservation’. Other vegetal remains, such as carbonified leaves, stems and roots, were collected in the site. The different modes of preservation depend directly on: depositional micro‐environment (sandy distributary channel, muddy flood plain); and type (trunk, branch, stem, leave, root) and state (presence or absence of resinous material) of the material. The great abundance and diversity of fossils in ‘Lo Hueco’ identify it as Konzentrat‐Lagerstätten, sequentially formed by alternated events of flooding and drying depositional events, but the exceptional quality and rarity of determinate vegetal macroremains preservation suggest that certain deposits of this site can be considered as conservation deposits.     

An Early Cenomanian (Late Cretaceous) maximum flooding bioevent in NW Europe: Correlation,sedimentology and biofacies

The Early Cenomanian (mid-Mantelliceras dixoni Zone) Schloenbachia/Inoceramus virgatus Event of NW Europe is recorded in five calcareous ‘marl’-limestone couplets rich in inoceramid bivalves of the Inoceramus virgatus plexus. The 1.3–6.8m thick beds represent a bundle of precession couplets inferred to have been deposited during a short-term cycle of sea-level change triggered by the 100kyr short eccentricity cycle within the Milankovitch periodicities. A burrowed omission surface terminates the Schloenbachia/Inoceramus virgatus Event in most places. Rock accumulation rates vary in different sections between 13 and 68Bubnoff units (mm/kyr) for the complete 100kyr cycle with extreme values ranging from 10 to 100Bubnoffs at shorter (i.e., 20kyr precession couplet) scales. Taphonomic and microfacies trends suggest deposition below fair-weather wave base with frequent to sporadic reworking of the sea-floor by storm waves for proximal and medial sections. The distal sections were close to, or even slightly below, maximum storm wave base. The biofacies of the Schloenbachia/Inoceramus virgatus Event is strongly dominated by inoceramid bivalves (65–85%). The two representatives of Inoceramus virgatus, I. virgatus virgatus Schlüter and I. virgatus scalprum Böhm, are not geographic subspecies as they co-occur in the same beds. However, proximal morphs tend to be more strongly ribbed and flatter whereas distal ones are more inflated and smoother, representing ecophenotypes of one polymorphic species. Taphonomic observations and shell morphology suggest that the inoceramids lived as epibyssate recliners. The low species richness and fossil content of the event strata (without the inoceramid bivalves) are attributed to low fluxes of organic matter to the sea-bed during the maximum flooding interval of Cenomanian depositional sequence Ce III. The inoceramids of the virgatus plexus are inferred to have been very effective filter feeders flourishing under nutrient-depleted, low-energy conditions. The Schloenbachia/Inoceramus virgatus Event is an example of a maximum flooding bioevent, forming a widespread marker dominated largely by autochthonous biota.     

A New Decapod Trackway from the Upper Cretaceous, James Ross Island, Antarctica

A new trace fossil, Foersterichnus rossensis igen. nov. and isp. nov., is described from the Coniacian Hidden Lake Formation, James Ross Island, Antarctica. The trace fossil is a trackway comprising straight, or slightly curving, paired rows of elongate to tear-shaped impressions, parallel or slightly inclined to the long axis of the trackway. Foersterichnus is interpreted to be the trackway of a brachyuran decapod crustacean. It occurs in transgressive shallow marine deposits formed above the storm wave base in a shelf setting. Preservation of the trackway may have been linked to a rapid deoxygenation event during drowning that led to cessation of bioturbation allowing preservation of mixed layer trackways.     

Facies control on lower Cambrian wrinkle structure development and paleoenvironmental distribution, southern Great Basin, United States

Assessing the role that physical processes play in restricting microbial mat distribution has been difficult due to the primary control of bioturbation in the modern ocean. To isolate and determine the physical controls on microbial mat distribution and preservation, a time in Earth’s history must be examined when bioturbation was not the primary control. This restricts the window of observation primarily to the Precambrian and Cambrian, which precede the development of typical Phanerozoic and modern levels of bioturbation. Lower Cambrian strata of the southern Great Basin, United States, record the widespread development of seafloor microbial mats in shallow shelf and nearshore settings. These microbial mats are recorded by wrinkle structures, which consist of millimeter-scale ridges and sinuous troughs that represent the former presence of a surface microbial mat. Wrinkle structures within these strata occur exclusively within heterolithic deposits of the offshore transition, i.e., between fair-weather wave base and storm wave base, and within heterolithic tidal-flat deposits. Wrinkle structures are not preserved in siltstone-dominated offshore deposits or amalgamated shoreface sandstones. The preservation of wrinkle structures within these environments is due to: (1) the development of microbial mats atop clean quartz-rich sands for growth and casting of the structures; and (2) the draping of the microbial mat by finer-grained sediment to inhibit erosion. The exclusion from offshore deposits may be due to a lack of sufficient sunlight, whereas the restriction from the shoreface is likely due to the amalgamation of proximal tempestites, resulting in the erosion of any incipient microbial mat development.