A diverse deep‐marine Ichnofauna from the Eocene Tarcau sandstone of the Eastern Carpathians,Romania

The Paleocene to Middle Eocene Tarcau Sandstone at Buzau Valley, eastern Carpathians, Romania, records sedimentation in a turbidite system. These strata contain a diverse and abundant pre‐ and postdepositional ichnofauna consisting of 35 ichnogenera and 54 ichnospecies. The predepositional assemblage is rich in graphoglyptids and ornate grazing trails; simple grazing trails, resting traces, and feeding structures also occur. The predepositional assemblage includes Acan‐thorhaphe, Belorhaphe, Cardioichnus, Circulichnus, Coch‐lichnus, Cosmorhaphe, Desmograpton, Fustiglyphus, Gordia, Helicolithus, Helminthopsis, Helminthorhaphe, Lorenzinia, Megagrapton, Paleodictyon, Paleomeandron, Protopaleodictyon, Scolicia (S. strozzii), Spirorhaphe, Spirophycus, Treptichnus, and Urohelminthoida. The ich‐nodiversity, composition, ethology, and morphologic complexity of the predepositional association are indicative of the Nereites ichnofacies. The postdepositional association essentially consists of dwelling, feeding, and grazing traces, and is represented by Chondrites, Glockerichnus, Halopoa, Nereites, Ophiomorpha, Phycodes, Planolites, Polykampton, Scolicia(S. prisca. S. striata), Taenidium, Thalassinoides, and Zoophycos. Palaeophycus occurs in both assemblages. Allochthonous Teredolites is present in wood fragments, The postdepositional association includes elements of the Skolithos ichnofacies and facies‐crossing forms that are commonly present in deep‐marine deposits, Elements of the Skolithos ichnofacies are present not only in the most proximal parts of the turbidite system, but also in distal parts. The number of predepositional forms greatly exceeds postdepositional ones, reflecting a dominance of K‐selected over r‐selected population strategies in a stable environment. High levels of ichnodiversity in the Tarcau Sandstone are comparable with deep‐sea ichnofaunas from the Polish Carpathians and with other flysch trace‐fossil assemblages of similar age. This abundant and diverse Eocene ichnofauna supports the idea of extremely rich deep‐sea ichnofaunas in the Cenozoic.     

Triassic trace fossils from lacustrine shoreline deposits of the Passaic formation,douglassville, pennsylvania

Lake‐margin deposits of the Late Triassic Passaic Formation, Douglassville, Pennsylvania, have yielded a moderate variety of trace fossils. The greatest diversity and abundance of trace fossils occurs on the sole of a thin gray claystone overlain and underlain by gray siltstones. Specimens of Cochlichnus anguineus, Helminthoidichnites tenuis, Helminthopsis hieroglyphica, Treptichnus pollardi, and paired trails reflect simple, unspecialized, horizontal grazing as well as feeding traces under very shallow water lacustrine conditions. The lack of commonly associated Scoyenia burrows in these deposits may possibly be related to the degree of maturation of the organic debris available. The traces, as well as a lack of meniscate burrows, compare favorably to the Mermia ichnofacies, except Passaic deposits exhibit evidence of subaerial exposure. This unit most likely records a lacustrine expansion whereby grazing trails were emplaced under fully subaqueous conditions. Subsequent shallowing and desiccation, followed by sediment influx during rainstorms, favored preservation of these traces.

In contrast, the Scoyenia ichnofacies consists of feeding burrows of Scoyenia gracilis and Spongeliomorpha milfordensis within reddish brown siltstones and mud‐stones (redbed sequence). The Scoyenia ichnofacies records limited exploitation, by opportunistic infaunal deposit feeders, of lake‐margin nutrients carried in during occasional rainstorms that punctuated otherwise extended periods of aridity.     

Middle Pennsylvanian Ichnofauna from Eastern Oklahoma,USA

The Keota site, in the Middle Pennsylvanian (Desmoinesian) Keota Sandstone Member, McAlester Formation (Krebs Group) of Haskell County, Oklahoma, yields an extensive ichnofossil assemblage of arthropod trackways, insect resting traces, arthropod feeding and grazing traces, fish swimming traces, and tetrapod footprints. This ichnofossil assemblage occurs in a 0.8 to 1.5-m-thick unit of thinly laminated and ripple-laminated sandstone at the base of the Keota Sandstone Member that is interpreted as a tidal flat sandstone. The arthropod traces are assigned to the ichnotaxa Diplichnites gouldi Gevers types A and B, Diplopodichnus biformis Brady, Paleohelcura tridactyla Gilmore, Tonganoxichnus buildexensis Mángano, Buatois, Maples and Lanier, Gordia marina Emmons, cf. Cochlichnus sp. and Treptichnus bifurcus Miller. The fish swimming traces are assigned to Undichna britannica Higgs, and the tetrapod tracks to Notalacerta ichnosp. and Pseudobradypus ichnosp. The Keota ichnofossil assemblage thus documents the first North American Pennsylvanian record of Paleohelcura tridactyla and the first record of Pseudobradypus from the western United States. This ichnofossil assemblage also provides compelling evidence of a freshwater habitat in a tidal flat setting during deposition of the fossil-bearing sandstone at the Keota site. Indeed, the Keota ichnofossil assemblage is characteristic of a Carboniferous-Permian Tonganoxichnus assemblage that identifies nonmarine tidal flat settings.     

Scolicia and its producer in shallow-marine deposits of the Miocene Chenque Formation (Patagonia,Argentina): functional morphology and implications for understanding burrowing behavior

Abstract

Scolicia is one of the most conspicuous trace fossils in lower shoreface deposits of the Miocene Chenque Formation of Patagonia, Argentina. This ichnotaxon consists of horizontal, sinuous or meandering trace fossils with a laminated backfill and two parallel strings located at the base. Abundant body fossils attributed to Brisaster iheringi occur in close association to these trace fossils. The echinoids are very well preserved, and most of the specimens have some areas with their spines attached in life position. In particular, preservation of the subanal tufts of spines, associated to isopores (which are interpreted as being associated to funnel building tube feet), supports the interpretation that these organisms are the producers of Scolicia isp. A shaft connecting the burrow with the sediment-water interface was not observed in vertical sections of the excavations, although on bedding plane surfaces, some small circular tube outlets are recognized. However, the great diversity and abundance of ichnofossils co-occurring in these deposits may preclude adequate identification of vertical structures. These deposits provide an excellent opportunity to integrate trace fossil data, body fossil information and observations from modern analogues in order to perform a morpho-functional analysis of Scolicia.     

Short-term colonization trace-fossil assemblages in a carbonate lacustrineKonservat-Lagerstätte (Las Hoyas fossil site, Lower Cretaceous, Cuenca, Central Spain)

Summary Rhythmically laminated fine-grained lacustrine carbonatcs of the Cretaceous La Huérguina Limestone Formation at Las Hoyas fossil site, central Spain, contain not only a spectacular body fossil fauna, but also vertebrate and invertebrate trace fossils. The study of the Las Hoyas invertebrate ichnofauna provides valuable taphonomic, paleoecological and paleoenvironmental information and represents one of the first attempts to study in detail a lacustrine trace-fossil assemblage in carbonate rocks. The Las Hoyas ichnofauna includesCruziana problematica, Helminthoidichnites tenuis, Lockeia isp.,Palaeophycus tubularis, andTreptichnus pollardi. This assemblage is characterized by low ichnodiversity and small burrow size, and it is dominated by surface trails and extremely shallow burrows produced mostly by detritus feeders. The Las Hoyas ichnofauna indicates low to moderately low energy, permanently submerged, shallow-lacustrine areas. The ichnofauna is suggestive of environmental stress, most likely oxygen-depleted conditions in interstitial waters. Occurrence of traces in event beds indicates opportunistic colonization by epifauna and very shallow infauna during brief periods of improved oxygenation related to the passage of density underflows and dilute turbidity currents. The Las Hoyas ichnofauna resembles other shallow lacustrine trace-fossil assemblages of the archetypalMermia ichnofacies and differs significantly from the archetypalScoyenia ichnofacies. It is therefore best regarded as a proximal, depauperate example of theMermia ichnofacies. The Las Hoyas assemblage is similar to the slightly older lacustrine Montsec ichnofauna, but it is remarkably different from marine ichnocoenoses of other Mesozoickonservat-lagerst?tten in lithographic limestones such as those of Cerin, Solnhofen and Nusplingen.     

Insights from functional morphology and neoichnology for determining tracemakers: a case study of the reconstruction of an ancient glacial arthropod‐dominated fauna

Apportioning trace fossils to producers is significant for palaeoecological and palaeoenvironmental studies. A tracemaker is usually attributed through the comparison of the morphology of the trace to the body plans of modern and extinct animals. However, hypotheses of tracemaker identifications may be tested with the use of further methods. This contribution aims to evaluate the potential tracemakers of a trackway‐dominated ichnofauna preserved in the siltstone–claystone rhythmites of the Pennsylvanian Rio do Sul Formation in the southern Brazil. These deposits are typical of the deglaciation phase of the Late Palaeozoic Ice Age of Gondwana in the Paraná Basin. Thus, the parameters applied in this work are as follows: (1) functional analyses of the traces to infer the biomechanics of the locomotion of the producer; and (2) neoichnology, whereby modern traces whose producers are known are compared to those of the ichnofauna in question. The conclusions obtained with these methods strongly support millipedes as the producers of Diplichnites gouldi and Diplopodichnus biformis. The malacostracan crustacean‐related ichnogenera are Umfolozia sinuosa and Gluckstadtella elongata, produced by syncarids or peracarids, while isopods are suggested as the tracemakers of Glaciichnium isp. and Protovirgularia dichotoma. In terms of branchiopod crustaceans, Cruziana problematica is attributed to notostracans. Helminthoidichnites tenuis, Mermia carickensis and Treptichnus pollardi are considered traces produced by insect larvae. The palaeobiology of the tracemakers that produced the ichnofauna preserved in the rhythmites of the Rio do Sul Formation permits inferences that terrestrial (millipedes) and aquatic (crustaceans and insect larvae) animals colonized the sediments. This conclusion points to a dynamic depositional environment, with changing physical and chemical parameters conditioning the biota.     

Lunulichnus Tuberosus Ichnogen. and Ichnosp. Nov. from the Early Eocene Wasatch Formation,Fossil Butte National Monument,Wyoming: An Arthropod-Constructed Trace Fossil Associated with Alluvial Firmgrounds

A new trace fossil, Lunulichnus tuberosus, is described from fluvial deposits of the Wasatch Formation (early Eocene) at Fossil Butte National Monument, southwestern Wyoming, USA. L. tuberosus are straight, vertical to obliquely oriented, unlined cylindrical burrows with pronounced crescent-shaped wall sculptings. In situ examples of these trace fossils are most commonly preserved as sand-filled casts emanating from the erosional bases of fluvial channel sandstone bodies into underlying floodplain mudstone/siltstone beds. L. tuberosus is interpreted as the dwelling trace of a stream-dwelling decapod crustacean. Excellent preservation of fine detail, particularly their diagnostic crescent-shaped wall sculptings, support the hypothesis that L. tuberosus were excavated in firm substrata subjacent to fluvial erosional surfaces. As such, they are interpreted as constituents of alluvial Glossifungites trace fossil assemblages.     

Trace fossils in middle to late triassic fluvial redbeds,pranhita‐godavari valley,south India

Several morphological varieties of trace fossils abound in Middle and Late Triassic fluvial redbeds in the Pranhita‐Godavari Valley, south India, including Skolithos, Palaeophycus, Taenidium, escape burrows, and a type of trace very similar to ‘small stuffed burrows’ from the Triassic of Greenland. Burrow morphology was influenced by local hydrodynamic conditions. The distribution of burrows was facies controlled; some forms are restricted to channel deposits whereas others occur only in floodplains. Vertical dwelling burrows (Skolithos) occur in both channel and floodplain deposits. Horizontal structures representing deposit feeding (Taenidium) are confined to nondepositional surfaces within parallel‐laminated sandstones having parting lineations that represent catastrophically emplaced sand‐sheets in channels and proximal floodplains. Vertical escape burrows are confined to what were slowly but continually accreting parallel‐laminated sands of channel bars. Horizontal dwelling burrows (Palaeophycus) and ‘small stuffed burrows’ are virtually restricted to the smaller sandsheets of floodplain drainage systems.

The burrow assemblages do not occur as recurrent associations throughout the redbed sequence, and variations in different stratigraphic levels seem to be controlled by minor differences within a broadly similar environment. The entire assemblage has components of both the Scoyenia and Rusophycus ichnocoenoses reported from East Greenland but may be considered as the Scoyenia ichnofacies characteristic of redbeds deposited in extensive floodplains dissected by small streams, even though no Scoyenia individuals are present.     

Bowl-shaped structures in a Pleistocene clastic carbonate wedge on the Island of Rhodes,Greece

Abstract

Bowl-shaped structures are for the first time reported from the Greek Island of Rhodes. They occur in Pleistocene deposits of the Cape Arkhangelos Formation in the Rhodes Synthem. The regularity of their three-dimensional appearance is the argument for a biological origin. This gives the reason to assign the structures to Piscichnus waitemata for which we issue a formal diagnosis. They were formed on a clastic carbonate wedge that built into a coastal, steep-sided basin. Associate trace fossils are Thalassinoides suevicus and Bichordites monastiriensis.     

The insect trace fossil Tonganoxichnus from the middle Pennsylvanian of Indiana: Paleobiologic and paleoenvironmental implications

The ichnogenus Tonganoxichnus, produced by one or more monuran insect taxa, is now recorded from the Middle Pennsylvanian Mansfield Formation of Indiana. Tonganoxichnus is a resting trace that has three important implications. First, it represents a recurrent behavioral pattern in Upper Carboniferous to Lower Permian marginal marine environments of North America. Second, it provides finely resolved anatomical information for axial and appendicular body structures and behaviors that are difficult to determine from body‐fossil material alone. Third, integrated sedimentologic and ichnologic observations indicate that the Tonganoxichnus assemblage, inclusive of other ichnotaxa, is common in tidal rhythmites that were developed under freshwater conditions, probably in the innermost part of estuarine systems, close to or at the fluvioestuarine transition.     

The origin and paleoecologic significance of the trace fossil Asteriacites in the Pennsylvanian of Kansas and Missouri

The trace fossil Asteriacites, recorded in Cambrian to Recent shallow- and deep-marine facies, is traditionally interpreted as the resting trace of asterozoans. Well-preserved specimens of A. lumbricalis are abundant in Pennsylvanian (Upper Carboniferous) shallow- and marginal-marine siliciclastic deposits of eastern Kansas and western Missouri. Detailed morphologic analysis of these specimens suggests that they record the activities of mobile epifaunal ophiuroids. Evidence of a brittle star (ophiuroid) producer rather than sea star (asteroid) is provided by (1) trace-fossil morphologic features reflecting the anatomy of the producer (e.g., well-differentiated central structure, slender vermiform arms) and ophiuroid burrowing technique (e.g., proximal arm expansion, arm branching), and (2) mode of occurrence (e.g., gregarious behavior, horizontal and vertical repetition). Vertical and horizontal repetition produces complex aggregates of A. lumbricalis that are interpreted either as escape structures (fugichnia) or as feeding structures, respectively. Ophiura texturata is proposed as a modern analogue for the A. lumbricalis producer, based on inferred life habit and feeding behavior. Asteriacites lumbricalis is present in two different intertidal trace-fossil assemblages. The first assemblage is characterized by high diversity and records tidal flats developed outside of embayments under normal marine conditions. The second assemblage consists of A. lumbricalis together with a few other ichnotaxa and represents a depauperate association that developed in restricted tidal flats within an embayment or estuarine setting. This challenges the conventional view of Asteriacites as a normal-marine salinity indicator. Some echinoderms, and particularly asterozoans, penetrate and inhabit modern environments of depressed salinity. The presence of Asteriacites in Pennsylvanian marginal-marine facies of Kansas and Missouri provides evidence that ophiuroids had adapted to brackish-water conditions by the late Paleozoic.     

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.     

Tetrapod Ichnofacies: A New Paradigm

We recognize three fundamental terms in ichnology: (1) ichnoassemblage, which is an assemblage of ichnofossils conceptually equivalent to an assemblage of body fossils; (2) ichnocoenosis, which is a trace fossil assemblage produced by a biological community that can be characterized by morphological criteria; and (3) ichnofacies, which refers to recurrent ichnocoenoses that represent a significant portion of Phanerozoic time. There are two different kinds of ichnofacies, ethoichnofacies (mostly invertebrate ichnofacies) and biotaxonichnofacies (mostly tetrapod ichnofacies). Nonmarine invertebrate ichnologists now recognize five archetypal ichnofacies (Mermia, Skolithos, Scoyenia, Coprinisphaera, Psilonichnus) to which we add the Octopodichnus ichnofacies. We propose a coherent and consistent classification and nomenclature for tetrapod ichnofacies. We name five archetypal vertebrate ichnofacies for nonmarine environments: Chelichnus, Grallator, Brontopodus, Batrachichnus and Characichnos ichnofacies.     

The Fish Swimming Trace Undichna from the Mississippian Mauch Chunk Formation,Eastern Pennsylvania

Undichna, the swimming trace of fish, was recovered from the middle member of the Mauch Chunk Formation in eastern Pennsylvania in strata of Late Mississippian (Visean) age. These traces represent the only evidence of fish known from the Mauch Chunk Formation and the Carboniferous of Eastern Pennsylvania. The Mauch Chunk Formation in the study area is characterized by an ephemeral fluvial depositional environment that yields a diverse invertebrate ichnoassemblage of the Scoyenia ichnofacies and a tetrapod footprint assemblage dominated by the tracks of temnospondyl amphibians. The Undichna specimens are preserved on fine-grained, mudstone-draped, rippled sandstone. Two specimens, assigned to U. Britannica, consist of pairs of well-defined, narrowly incised, sinusoidal wave traces that are out-of-phase. Two specimens, assigned to U. quina, consist of two pairs of in-phase sinusoidal waves intersected by an additional single wave with a greater amplitude. Two specimens, each composed of a single wave, are assigned to U. unisulca.

The slabs of rock with the Undichna traces also preserve tetrapod undertracks assigned to Batrachichnus, Matthewichnus, and Hylopus. This trace fossil association indicates the presence of small fish, some with caudal and/or anal fins and some with pectoral, pelvic, and anal and/or caudal fins, that touched the sediment during periods of intermittent flooding of subaerial surfaces walked upon by early tetrapods. Examples of Undichna from Mississippian or older deposits, especially from a fluvial paleoenvironment, are rare; few have been reported from the Devonian and the Lower Carboniferous.     

Scolicia shirahamensis isp. nov.: a triple-corded scolicia and its ichnological implications

Abstract

Peculiar meniscate burrows with three sediment cords occur in early to middle Miocene tidal-flat deposits of southwestern Japan. Two of the cords are situated at the bottom and the other is at its center. Detailed observations of the burrow structures and comparative neoichnological studies of modern spatangoid burrows in a tidal flat revealed that the former two were true drainage tubes and the latter was fecal in origin. The trace fossil was thus assigned to the ichnogenus Scolicia. Based on these findings, a new ichnospecies Scolicia shirahamensis isp. nov. has been described here. The central sediment cord is seemingly identical to the drainage tube of the ichnogenus Bichordites, another ichnogenus that has been commonly ascribed to a fossil spatangoid burrow, similar to Scolicia. Careless ichnogeneric identification of a spatangoid burrow, based only on the central sediment cord, therefore, may produce an incorrect identification.     

Ordovician Bathyal Trace Fossils From Metasiliciclastics in Central Norway and Their Sedimentological and Paleogeographical Implications

Numerous trace fossils are recognized in metasiliciclastic rocks of the Lower-Middle Ordovician Lower Hovin Group, Upper Ordovician Ekne Group and on roofing slates from an unknown locality (probably Ordovician) from the Trondheim region, central Norway. The trace fossil assemblages are dominated by meandering pascichnial forms, i.e., Helminthoidichnites in the Lower Hovin Group and Helminthoidichnites and Nereites in the slates. Protovirgularia, Dictyodora, ?Planolites and ?Palaeophycus are less common. Alcyonidiopsis, ?Trichophycus, Chondrites, cf. Chondrites, Gordia, ?Phycodes, ?Helminthopsis, cf. Naviculichnium, Treptichnus, Saerichnites, Megagrapton and ?Paleodictyon (Squamodictyon) are rare. The trace fossils were collected or observed at 17 different localities and represent assemblages belonging to the Nereites ichnofacies (deep-sea flysch deposits with thinly bedded turbidites). They reflect a distal Nereites subfacies (fan-fringe or basin-plain deposits) transitional to a Nereites/Paleodictyon subfacies (proximal fan facies). The trace fossil assemblages indicate an opportunistic style of colonization related to fluctuations in food supply introduced by turbidity currents.     

Composite Trace Fossil Assemblage in a Distal Carbonate Setting from the Tethys (Middle Jurassic,Betic Cordillera,Southern Spain)

Ichnological analysis of a Middle Jurassic carbonate surface from the Betic Cordillera (southern Spain) reveals a complex trace fossil assemblage, including softground Ophiomorpha, firmground Arenicolites, Thalassinoides and Gastrochaenolites, and hardground Trypanites as well as possible Gastrochaenolites. The ecological replacement in the macrobenthic community is interpreted according to successive suites that are controlled mainly by substrate consistency. Variations in composition and abundance of trace fossils between suites can be ecologically and/or taphonomically determined.     

Facies and palaeoecology of Upper Jurassic (Middle Oxfordian) coral reefs in England

Summary This study documents the facies and fauna of Late Jurassic (Middle Oxfordian) coral reefs in England. Sedimentological and palaeoecological analysis of these reefs distinguishes three generic reef types: (1) small reef patches and thickets associated with siliciclastic deposits; (2) small reef patches and thickets associated with siliciclastic-free bioclastic grainstones and packstones; and (3) biostromal units associated with deep water facies. The depositional environments of these reef types are discussed. Two coral assemblages are identified: (1) the microsolenid assemblage; and (2) theThamnasteria, Isastraea, Fungiastraea andThecosmilia assemblage (Thamnasteria assemblage). TheThamnasteria assemblage developed in all shallow water environments in the study area, regardless of local environmental conditions. The fauna is very eurytopic,r-selected and can tolerate significant environmental fluctuations on short temporal scales (sub-seasonal). The main control on the development of the microsolenid assemblage was low light intensity, low background sedimentation rates and low hydrodynamic energy levels.     

Ichnology of Lower Jurassic beach deposits in the Shemshak Formation at Shahmirzad,southeastern Alborz Mountains,Iran

A 19 m thick package of well-sorted lowermost Jurassic (Hettangian-Lower Sinemurian?) sandstones within the Shemshak Formation of the southeastern Alborz Mountains displays features characteristic of foreshore to upper shoreface environments such as tabular bedding, low-angle lamination, trough cross-stratification, parting lineation, and oscillation ripples. In contrast to most other beach successions recorded in the literature the sandstones contain a trace fossil assemblage characterised by low abundance but comparatively high diversity. The assemblage, comprising 14 ichnotaxa, is dominated by Palaeophycus heberti, Rhizocorallium irregulare, Gyrochorte comosa, and Parahaentzschelinia surlyki. Contrary to predictions, which assume a dominance of suspension-feeders in such high-energy environments, the trace fossil assemblage represents a variety of ethological groups ranging from suspension-feeders to deposit-feeders, detritus-feeders, scavengers, and a possible trap constructor (Ctenopholeus), whereby deposit-feeders predominate. This anomaly is explained by a high amount of organic detritus in the sediment, indicated by abundant plant material, and a position of the beach in the vicinity of a river mouth.