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.     

Late Triassic Tetrapod-Dominated Ichnoassemblages from the Argana Basin (Western High Atlas,Morocco)

Diverse tetrapod track assemblages with Scoyenia invertebrate traces were discovered in the Triassic Timezgadiouine and Bigoudine formations of the Argana Basin (Western High Atlas, Morocco). The ichnofossils occur in alluvial plain sandstones and mudstones of the Irohalène Member (T5) and Tadart Ouadou Member (T6) considered Carnian-Norian in age by vertebrate remains and palynomorphs. Tetrapod footprints are assigned to Apatopus, Atreipus-Grallator, Eubrontes isp., Parachirotherium, cf. Parachirotherium postchirotherioides, Rhynchosauroides ispp., and Synaptichnium isp. They can be referred to lepidosauromorph/ archosauromorph, basal archosaur, and dinosauromorph trackmakers. Apatopus, represented by 11 tracks of a more than 4 m long trackway, is recorded for the first time outside of North America and Europe. The assemblage concurs with the proposed Late Triassic age of the track-bearing beds by the occurrence of Apatopus, Atreipus-Grallator, and Eubrontes. If this is accepted, the stratigraphic range of Synaptichnium and Parachirotherium, hitherto known only from Early or Middle Triassic deposits, has to be extended to the Carnian-Norian. The occurrence of Eubrontes in the Irohalene Member (T5) provides further evidence for large theropods in pre-Jurassic strata. All assemblages are referred to the Scoyenia ichnofacies indicating continental environments with alternating wet and dry conditions.     

Early Triassic Archosaur-Dominated Footprint Assemblage from the Argana Basin (Western High Atlas,Morocco)

An assemblage of abundant and well-preserved tetrapod footprints has been discovered in the Tanamert Member (T3) of the Triassic Timezgadiouine Formation (Argana basin, western High Atlas, Morocco). It is the first fossil record from T3. Surfaces from different localities show a uniform tetrapod ichnofauna that consists of chirotherian and small lacertoid forms. The chirotherians are assigned to the plexus Protochirotherium—Synaptichnium, their trackmakers interpreted as basal archosaurs. The lacertoid imprints show close affinities with Rhynchosauroides and may reflect archosauromorphs or lepidosauromorphs. Protochirotherium—Synaptichnium assemblages are characteristic of the Early Triassic and were known previously only from units of this age in central Europe. Biostratigraphically, the European record implies a wide-spread pre-Anisian Protochirotherium—Synaptichnium dominated assemblage preceding the first appearance of Chirotherium barthii near the Olenekian-Anisian boundary. The stratigraphic position of T3 between Late Permian (uppermost T2) and Middle Triassic (T4) and the European correlatives suggest an Early Triassic age of this unit. It is the first record of Early Triassic continental deposits in Morocco. The surfaces from T3 open up perspectives for further contributions to ecology, biogeography and locomotion of early archosaurs. Furthermore, excellent outcrops and quality of footprint preservation in the Argana basin offer a potential for clarification of ichnotaxonomic and biostratigraphic issues.     

Middle–Late Permian tetrapods from the Rio do Rasto Formation,Southern Brazil: a biostratigraphic reassessment

Dias‐da‐Silva, S. 2011: Middle–Late Permian tetrapods from the Rio do Rasto Formation, Southern Brazil: a biostratigraphic reassessment. Lethaia, Vol. 45, pp. 109–120. The Rio do Rasto Formation (Permian of Southern Brazil) was previously regarded as Guadalupian–early Lopingian age. Three tetrapod‐based localities are known: the Serra do Cadeado area, Aceguá and Posto Queimado. The latest tetrapod‐based biostratigraphic contribution considers that the Posto Queimado and Aceguá faunas are coeval and Wordian (middle Guadalupian) in age, correlated to the Isheevo faunas from Eastern Europe and to the Tapinocephalus Assemblage Zone of South Africa; whereas the Serra do Cadeado fauna is Capitanian (late Guadalupian), correlated to the Kotelnich fauna of Eastern Europe and, from bottom to top, to upper Pristerognathus, Tropidostoma and lower Cistecephalus assemblage zones of South Africa. A re‐evaluation of the tetrapods from the Rio do Rasto Formation and new fossil discoveries in the localities of Posto Queimado and Serra do Cadeado area (melosaurine and platyoposaurine temnospondyls, a basal anomodont, a dinocephalian and a basal dicynodont) supports a new tetrapod‐based biostratigraphic scheme for the Rio do Rasto Formation. Accordingly, the age of the fauna at Aceguá is late Roadian‐early Wordian, whereas the locality of Posto Queimado is late Wordian‐Capitanian. The Serra do Cadeado Area is correlated with both southernmost ones (Guadalupian) but also Wuchiapinghian (early Lopingian). □Paraná Basin, Passa Dois Group, tetrapod biostratigraphy, Western Gondwana.     

Small Amphibian and Reptile Footprints from the Permian Carapacha Basin,Argentina

This paper contains a taxonomic study of the Permian tetrapod ichnofauna from the Carapacha Basin. Tetrapod traces are analyzed in their environmental context and compared with similar faunas from Europe and North America. This ichnofauna is particularly relevant because of the scarcity of Permian tetrapod tracks from South America and also of Permian tetrapod fossils from Argentina. Ephemeral fluvial and shallow lacustrine deposits compose the sedimentary succession of the basin, which is represented by the Carapacha Formation. Most of the tracks have been collected from the upper member of the formation (Urre-Lauquen Member), mainly from freshwater ephemeral lake deposits as well as from playa-lake mudflats. The deposits of this member have been attributed to the early Late Permian on the basis of a Glossopteris fossil flora. Ichnotaxonomic designations of tetrapod traces are made on the basis of morphologic features that reflect the anatomy of the producer and special attention has been paid to extramorphologic deformations observed in the track assemblage. A total of four footprint ichnotaxa have been recognized, namely Batrachichnus salamandroides (Geinitz, 1861 Geinitz, H. B. 1861. Dyas oder die Zechsteinformation und das Rhotliegende—Die animalischen Überreste der Dyas, 130 ppLeipzig: Wilhelm Engelmann.  [Google Scholar]), Hyloidichnus bifurcatus Gilmore, 1927 Gilmore, C. W. 1927. Fossil footprints from the Grand Canyon: second contribution. Smithsonian Miscellaneous Collection, 80(3): 1–78.  [Google Scholar], cf. Amphisauropus isp. and cf. Varanopus isp. These track taxa are associated with two forms of vertebrate swimming traces (Characichnos isp. and type A swimming trace) and a possible fish trail. Invertebrate trace fossils include abundant arthropod locomotion traces and Scoyenia isp. The ichnofauna is composed of six tetrapod ichnocoenoses that are dominated by tiny amphibian tracks attributed to Temnospondyli (Batrachichnus and type A swimming trace) and Seymouriamorpha (Amphisauropus), and also contain the footprints of small reptiles, mostly Captorhinomorpha and possibly Pelycosauria (Hyloidichnus and Varanopus). Even if the ichnofauna of the Carapacha Basin is slightly younger than typical examples from the literature of the Early Permian “red bed ichnofacies” (Hunt et al., 1995b Hunt, A. P., Lucas, S. G., Lockley, M. G., Haubold, H. and Braddy, S. 1995b. Tetrapod ichnofacies in Early Permian red beds of the American Southwest. Bulletin New Mexico Museum of Natural History and Science, 6: 295–301. Early Permian footprint facies [Google Scholar]), a comparison is made. However, further detailed case studies are needed to formally define this “red bed ichnofacies” and its prospective subdivisions.     

The first record of a nyctiphruretid parareptile from the Early Permian of North America,with a discussion of parareptilian temporal fenestration

The Richards Spur Locality of Oklahoma, USA, long known for its highly diverse Early Permian terrestrial tetrapod assemblage, is particularly interesting for the presence of many endemic taxa. The parareptilian component of the assemblage, rare members of other Early Permian communities, is especially diverse at Richards Spur, consisting of six species. The newest parareptile, A byssomedon williamsi gen. et sp. nov. , consists of an articulated left jaw and various disarticulated cranial and postcranial elements. A new phylogenetic analysis of parareptiles, based on an updated modified data matrix revealed that Ab . williamsi is a member of the small clade Nyctiphruretidae. This makes Ab . williamsi the first and oldest nyctiphruretid, a clade of parareptiles otherwise known from the Middle and Late Permian of Russia, extending the age of the clade back into the Early Permian. This discovery also raises the possibility that nyctiphruretids may have dispersed from western Laurasia to eastern Laurasia. The characteristic jugal morphology of Ab . williamsi shows that it would have possessed a slender, deep, temporal emargination. The current topology of Parareptilia indicates that there was considerable variability in the patterns of lateral temporal openings amongst the various members of this clade, suggesting that there may have been multiple, independent modifications of this region of the skull. © 2014 The Linnean Society of London     

Small Theropod Track Assemblages from Middle Jurassic Eolianites of Eastern Utah: Paleoecological Insights from Dune Ichnofacies in a Transgressive Sequence

New discoveries show that very small theropod tracks (cf. Wildeichnus) are abundant in the upper part of the Moab Member, recently assigned to the Curtis Formation (formerly considered part of the Entrada Formation) in the Mid-?Late Jurassic of eastern Utah. The tracks represent a distinct small-theropod ichnofacies associated with eolian dune deposits that is easily differentiated from the water-lain beds of the overlying Megalosauripus-Therangospodus ichnofacies, which comprises the single-surface Moab megatracksite. Pterosaur track assemblages, representing the Pteriachnus ichnofacies, are found a few meters above the megatracksite surface in the upper tongue of the Summerville Formation.

The small theropod ichnofacies is reminiscent of other early Mesozoic dune facies ichnofaunas from the Wingate and Navajo formations (Late Triassic and Early Jurassic) where small theropod tracks occur in association with other small tetrapod footprints. All such examples evidently represent a recurrent dune facies ecosystem dominated by diminutive vertebrates. Because the small theropod ichnofacies is one of three ichnofacies found in a thin stratigraphic sequence (<20 m) that contains no body fossils, it is clear that vertebrate tracks play an important role in providing insight into the paleoecology of units previously considered devoid of any useful fossil evidence. The three successive ichnofacies represent a transgressive transition from sand dunes, through sandy shoreline to shallow marine environments, each with its quite distinct vertebrate fauna.     

Vertebrate assemblages from the north‐central Main Karoo Basin,South Africa,and their implications for mid‐Permian biogeography

The rich fossil vertebrate record from the Beaufort Group, Main Karoo Basin, provides a global standard for mid‐Permian to Mid‐Triassic continental faunas. However, recent studies have demonstrated variability in the composition of contemporaneous faunas across Gondwana. This raises the question of how much the vertebrate faunas differ within the Karoo, where the taxonomic composition of vertebrate assemblage zones (AZs) is mostly considered to be uniform. Although fossil material is known from across the outcrop of the Beaufort Group, the lowest Beaufort strata have received little attention, particularly north of S31°10′. Here, we report two fossil tetrapod assemblages from the lowest Beaufort Group in the southern Free State Province, which represent the northernmost point at which the lowest Beaufort has been targeted for collecting. The lower assemblage is characterized by an abundance of the small dicynodont Eosimops and can thus be attributed to the Tapinocephalus AZ (Guadalupian), but the absence of dinocephalian or pareiasaurian material is unlike contemporaneous assemblages found further south. This suggests that the Tapinocephalus AZ was not uniform across the entire basin and highlights that the abundance, distribution and taxonomic composition of Karoo biozones may vary more than currently appreciated. The upper assemblage, characterized by the dicynodonts Oudenodon, Aulacephalodon and Dinanomodon, is attributable to the upper Cistecephalus AZ to lower Daptocephalus AZ. The juxtaposition of the lower Tapinocephalus AZ and upper Cistecephalus\lower Daptocephalus AZ in the southern Free State implies a stratigraphic gap from the Middle to Late Permian of up to 6 million years.     

Tetrapod Footprint Biostratigraphy and Biochronology

Tetrapod footprints have a fossil record in rocks of Devonian-Neogene age. Three principal factors limit their use in biostratigraphy and biochronology (palichnostratigraphy): invalid ichnotaxa based on extramorphological variants, slow apparent evolutionary turnover rates and facies restrictions. The ichnotaxonomy of tetrapod footprints has generally been oversplit, largely due to a failure to appreciate extramorphological variation. Thus, many tetrapod footprint ichnogenera and most ichnospecies are useless phantom taxa that confound biostratigraphic correlation and biochronological subdivision. Tracks rarely allow identification of a genus or species known from the body fossil record. Indeed, almost all tetrapod footprint ichnogenera are equivalent to a family or a higher taxon (order, superorder, etc.) based on body fossils. This means that ichnogenera necessarily have much longer temporal ranges and therefore slower apparent evolutionary turnover rates than do body fossil genera. Because of this, footprints cannot provide as refined a subdivision of geological time as do body fossils. The tetrapod footprint record is much more facies controlled than the tetrapod body fossil record. The relatively narrow facies window for track preservation, and the fact that tracks are almost never transported, redeposited or reworked, limits the facies that can be correlated with any track-based biostratigraphy.

A Devonian-Neogene global biochronology based on tetrapod footprints generally resolves geologic time about 20 to 50 percent as well as does the tetrapod body fossil record. The following globally recognizable time intervals can be based on the track record: (1) Late Devonian; (2) Mississippian; (3) Early-Middle Pennsylvanian; (4) Late Pennsylvanian; (5) Early Permian; (6) Late Permian; (7) Early-Middle Triassic; (8) late Middle Triassic; (9) Late Triassic; (10) Early Jurassic; (11) Middle-Late Jurassic; (12) Early Cretaceous; (13) Late Cretaceous; (14) Paleogene; (15) Neogene. Tetrapod footprints are most valuable in establishing biostratigraphic datum points, and this is their primary value to understanding the stratigraphic (temporal) dimension of tetrapod evolution.     

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.     

Ichnology and sedimentology of the early Permian deep-water deposits from the Lercara-Roccapalumba area (Western Sicily,Italy)

Summary Diverse and abundant trace fossils of the deep-waterNereites ichnofacies have been found in well-dated Early Permian deep-water turbidites (Lercara Formation) of western Sicily (Italy). Conodonts indicate a latest Artinskian to Cathedralian (late Early Permian) age. Microfossils (pelagic conodonts, albaillellid Radiolaria, paleopsychrospheric ostracods, foraminiferal associations dominated byBathysiphon), trace fossils (deep-bathyal to abyssalNereites ichnofacies) and sedimentologic data collectively indicate a deep-water environment for the Early Permian turbidites of the Lercara Formation. The dominance ofAgrichnium and of thePaleodictyon subichnogeneraSquamodictyon andMegadictyon suggests that this icnofauna is closely related in ichnotaxonomic composition to other late Paleozoic deep-water ichnofaunas. The occurrence ofAcanthorhaphe. Dendrotichnium andHelicoraphe, to date only reported from Cretaceous or Tertiary flysch deposits, suggests that the entire ichnofauna corresponds well to previously documented Silurian-Tertiary flysch ichnofaunas. Eight new ichnospecies and a new ichnosubgenus,Megadictyon, are described.     

Late Carboniferous-Early Permian Tetrapod Ichnofauna from the Khenifra Basin, Central Morocco

In terms of cumulative thickness and areal extent, the Khenifra Basin is one of the most important outcrops of Late Palaeozoic red-beds in central Morocco. Macro- and microfloral remains near the centre of the 1800 m-thick succession of interbedded conglomerates, sandstones, and mudstones are considered to be of middle to late Early Permian age. Here we give the first comprehensive analysis of the vertebrate ichnofossil record from the study area, based on 17 specimens of isolated footprints and incomplete step cycles collected at three localities that are lithostratigraphically equivalent to the plant-bearing horizons. The tetrapod ichnofauna comprises tracks of the plexus Batrachichnus Woodworth - Limnopus Marsh, Ichniotherium sphaerodactylum (Pabst), Dimetropus Romer and Price, and Dromopus Marsh which can be referred to temnospondyl, diadectomorph, synapsid (“pelycosaurian”) and early sauropsid trackmakers. This clearly Euramerican footprint assemblage, including the first occurrences of Ichniotherium and Dimetropus from outside Europe and North America, indicates a Late Carboniferous to Early Permian age of the fossiliferous strata. Judging from the relatively diverse ichnofauna and flora, the Khenifra Basin must have represented a well-established terrestrial ecosystem during that period. Its habitat could be specially important for the understanding of the phylogeny and dispersal of early tetrapods, inasmuch as we are able to report on an extremely rare type of diadectomorph footprint hitherto known only from the Early Permian of central Germany.     

Revision of Late Permian tetrapod tracks from the Dolomites (Trentino-Alto Adige,Italy)

The Val Gardena Formation of the Dolomites region in northern Italy preserves the most significant assemblage of Late Permian tetrapod footprints in the world. More than 120 years of collecting resulted in about 900 publicly accessible specimens from the study area. This huge amount of data is comprehensively revised in the light of recent advances in the study of Late Palaeozoic – Early Mesozoic tetrapod ichnofossils. According to our analyses, the Val Gardena Sandstone Formation includes tracks that can be assigned to cf. Batrachichnus isp. (temnospondyl amphibian), Capitosauroides isp. (amphibian), Dicynodontipus isp. (cynodont therapsid), Dolomitipes accordii n. igen. n. comb. (dicynodont therapsid), cf. Dromopus isp. (neodiapsid), Pachypes dolomiticus (pareiasaurian parareptile), Paradoxichnium problematicum (archosauromorph neodiapsid), Procolophonichnium tirolensis n. comb. (procolophonoid parareptile), cf. Protochirotherium isp. (archosauriform neodiapsid) and Rhynchosauroides pallinii (neodiapsid). The ichnoassociation is dominated by tracks of neodiapsid and parareptilian tetrapods, whereas synapsid and anamniote tracks are rather minor components. It includes 10 out of 12 tetrapod ichnogenera known from Lopingian deposits and thus it constitutes a reference for the Paradoxichnium biochron. It shows striking similarities with other low-latitude non-aeolian contemporaneous ichnoassociations of Europe and North Africa, differences may be linked to the palaeoenvironment. Moreover, it shows a clear Triassic affinity.

The new ichnogenus Dolomitipes was registered in Zoobank.org. urn:lsid:zoobank.org:act:5B4D871C-D16A-4E93-8211-CEE08019BA60     

A remarkable assemblage of terrestrial tetrapods from the Zechstein (Upper Permian: Tatarian) near Korbach (northwestern Hesse)

We present a preliminary report on the first diverse central European assemblage of Late Permian terrestrial tetrapods from a fissure-filling in marine sediments of the lower Zechstein near Korbach (northwestern Hesse, Germany). It includes therapsids (cynodonts and dicynodonts), archosauromorph diapsids, and pareiasaurs. Similar assemblages were previously known only from the Upper Permian of Russia, Scotland, and South and East Africa. The occurrence of the basal cynodontProcynosuchus is paleobiogeographically significant because this taxon was previously known only from theDicynodon lacerticeps-Whaitsia assemblage zone of South Africa and the Madumabisa Mudstone of Zambia. The geological context of the Korbach site permits a rather precise chronostratigraphic placement of the tetrapod assemblage in the interval between Zechstein 1 and 3. The Korbach tetrapod assemblage is late Tatarian in age.     

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.     

Controls on trace fossil diversity in an Early Cambrian epeiric sea: new perspectives from northwest Scotland

The Lower Cambrian Eriboll Formation of northwest Scotland is renowned for the high density, low diversity trace fossils (Skolithos ichnofacies) found in its upper Pipe Rock Member. Ichnofabric analysis of the member indicates that relatively small examples of Skolithos terminating at the same foreset boundary were formed during a brief colonization window after a single depositional event, that particularly long Skolithos specimens are equilibrichnia, and that palimpsests of Skolithos represent the marginal, slightly deeper water fringes of the Pipe Rock Member depositional environment. Nearest neighbour analysis, however, suggests that such palimpsests were uncommon. A much more diverse trace fossil assemblage is present in the overlying Fucoid Member (An t‐Sròn Formation), comprising Cruziana barbata, Dactylophycus, Didymaulichnus, Halopoa imbricata, ?Margaritichnus, Monocraterion, Monomorphichnus, Palaeophycus striatus, P. tubularis, ?Phycodes, Planolites montanus, ?Polarichnus, Rusophycus ramellensis, ?Psammichnites, Skolithos and various unidentified traces, and represents the Cruziana ichnofacies. Above the Fucoid Member, the Salterella Grit Member ichnofauna is more impoverished, yielding only Cruziana, Monocraterion, Rusophycus, Skolithos and ?Spirophyton. The ichnological variations between the Pipe Rock, Fucoid and Salterella Grit members are interpreted as being driven by changes in sea level. The low trace fossil diversity in the Pipe Rock Member indicates opportunistic colonization of laterally extensive, shoreface sediments deposited by regular influxes of terrigenous material, which were overlain by more distal, ichnologically diverse sediments (Fucoid Member) as sea level rose. A minor regression then caused an increase in terrigenous sediment input, producing an impoverished, proximal Cruziana ichnofacies (Salterella Grit Member).     

Taphonomic signatures,ichnofacies analysis and depositional dynamics of fossil macro-invertebrate assemblages of the San Juan Raya Formation,Zapotitlán Basin,Puebla, Mexico

In this paper, we describe the taphofacies and ichnofacies from Aptian strata of the San Juan Raya Formation in the Santa Ana Teloxtoc area, Puebla, Mexico. A composite stratigraphic section was analysed on a bed-by-bed scale up to a total thickness of 765.5 m. Our results show the presence of 10 taphofacies and 3 ichnofacies. The taphofacies and ichnofacies interpretation, and its correlation with the sedimentary lithofacies, enabled the determination of a palaeoenvironmental model for the study area that corresponds to a shallow marine, open-coast, clastic system with episodic sedimentation as a product of storm events. This system had several variations in sub-environments, from foreshore to offshore. Foreshore and shoreface environments are characterised by taphofacies Tf1, Tf2, Tf3 and Tf10 and PsI and SkI ichnofacies, representing lower faunal diversity moments (with the exception of Tf3 taphofacies). Meanwhile, taphofacies Tf4, Tf5, Tf6, Tf7, Tf8 and Tf9 were representative of shelf environments and are described as moments of medium-to-high faunal diversity (with the exception of Tf5 taphofacies).     

Marine influence and incursion in the Gondwana basins of Orissa, India: A review

The Permian–Triassic succession of the Indian Gondwana Sequence was previously considered to have been deposited in a fluviatile-lacustrine environment. Similarly, earlier Lower Gondwanas of Orissa State (a major part of the Mahanadi Master basin) were considered entirely fresh water deposits. Faunal evidence is still scanty in this master basin. Ichnology and palynology along with a few sedimentary records are reviewed and analysed for inferring marine signature. The marine nature of the Talchir, Karharbari, Barakar, Barren Measures and Kamthi sediments of three major basins (Talcher, Ib River and Athgarh) in Orissa State was predicted on the basis of typical marine ichnofossils. Most of these sediments also contain acritarchs reflecting marine marginal environment throughout the Permian. Moreover, evidence of wave activity, salinity raise and discovery of phosphorite in Permian sediments also strengthen this view.Hence, the previous model of continental facies for the Lower Gondwanas is found to be incorrect. The ichnofossils (Skolithos and Cruziana ichno-facies), acritarchs (Foveofusa, Leiosphaeridia, Greinervillites, etc.) and other palynofossils of marine origin can be utilized as a tool for palaeoenvironmental reconstruction. In the Gondwana basins of Orissa (Mahanadi Master basin), consistent occurrence of marine acritarchs and trace fossils with some typical sedimentary structures such as wave ripples has been studied and reviewed from the Talchir (Early Permian) to Upper Kamthi (Triassic) formations at various time intervals. Here marine incursion could have occurred due to the well known global transgressions during Permian and Triassic.