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Vertebrate footprints and a mammal mud-bath trace fossil (Laspichnia) from the Mukdadiya Formation (Late Miocene-Pliocene), Chamchamal Area, Kurdistan Region, Northeast Iraq Vertebrate footprints and a mammal mud-bath trace fossil (Laspichnia) from the Mukdadiya Formation (Late Miocene-Pliocene), Chamchamal Area, Kurdistan Region, Northeast Iraq
Abstract
The Mukdadiya Formation (Late Miocene-Pliocene) consists of alternations of red to brown sandstone and claystone layers in the Zagros Mountains Belt, northeastern Iraq. Two track-sites preserving bird and mammal tracks were recorded from the base of the formation in the Chamchamal area of the Kurdistan region in northeast Iraq. Avian tracks are large footprints with an average length of 25 cm that belong to Avipeda filiportatis. Mammal footprints imprinted by terrestrial cetartiodactyls belong to Pecoripeda amalphaea and Bifidipes velox, with cervids the most likely track makers. An unfamiliar large trace fossil on the studied slab, has symmetrical to asymmetrical marks, kidney or number-8-shaped and strong wrinkles on the surface. We interpret trace as having been produced by cetartiodac-tyls wallowing on the soft sediment surface. To accommodate this class of behavioral trace fossils; thus, we introduce a new fossilized behavior class, named "Laspichnia", which includes a vertebrate mud-bathing imprint on a soft sediment surface.
... However, the guidelines for ichnotaxonomic discrimination are not universally agreed upon, and depend largely on the suggestions of various authors, and the interpretation of tracks and trackways with variable preservational quality (Belvedere & Farlow, 2016). As noted below, subsequent work in Iran has demonstrated that Iranipeda and Ardeipeda are very similar: see N. Abbassi et al. (2020, fig. 4) for useful side-byside comparisons of Iranipeda, Ardeipeda, and Avipeda filiportatis, here incorporated into Figure 4 in modified form, with additions. ...
... N. S. Abbassi et al. (2015), identified Oligocene tracks from Iran as Gruipeda intermedia and also identified the presence of Iranipeda. Subsequently, N. Abbassi et al. (2020) identified avian tracks with an average length of 25 cm that belonged to Avipeda filiportatis from the Miocene-Pliocene of Iraq. However, they overlooked the opinion of Lockley and Harris (2010, p. 25) that 'A. ...
... filiportatis is justifiably considered a synonym of Ardeipeda'. We follow N. Abbassi et al. (2020, fig. 4, and Figures 4 herein), in comparing the heron-like tracks assigned to Ardeipeda, Iranipeda and Avipeda filiportatis. ...
... Also, the ratio of footprint length to hallux length of Iranipeda abeli (FL / DL hallux = 5.7) is greater than that of Ardeipeda gigantea (FL / DL hallux = 2.8). Avipeda filiportatis is large, four-digit tracks up to 19 cm long (Vialov, 1965), and Abbassi et al. (2020) reported large bird footprints from the late Miocene-Pliocene in northern Iraq with 22 cm length and 17 cm width. Lockley and Harris (2010), however, transferred Avipeda filiportatis to Ardeipeda filiportatis. ...
... Lockley and Harris (2010), however, transferred Avipeda filiportatis to Ardeipeda filiportatis. Among these footprints, Iranipeda is large, tetradactyl avian footprints with prominent, broad, consistently diverging and acutely tapering digit impressions with circular heel imprints (Abbassi et al., 2020). Ardeipeda gigantea is characterized by connected and forward-directed digit imprints and backward-directed, long and isolated digit I imprints. ...
... Recent snake trails on creek sediments. Arrows show lateral pressures during the crawling of snakes (Abbassi, 2020). Length of hammer equals 30 cm. with braided to meandering rivers and ephemeral ponds under a warm climate during the middle to late Miocene. ...
An extensive vertebrate tracksite from the middle–late
Miocene Upper Red Formation in western Zanjan Province, northwestern Iran,
provides new records of paleobiogeographical significance. These are records
of common footprints of felids referred to as Felipeda lynxi Panin and Avram, canids
referred to as Canipeda longigriffa Panin and Avram and less common bird footprints referred to as
Iranipeda abeli Lambrecht. The Canipeda record establishes the late Miocene presence of canids on
the Iranian Plateau as part of the Eurasia-wide “Eucyon event”. The
Felipeda footprints are consistent with body fossil and footprint records elsewhere
in Eurasia that indicate a widespread distribution of felids by late Miocene
time. An unusual trace associated with the footprints is a large, shallow
grazing or locomotion trace similar to Megaplanolites in some features but distinctive in
various features and its occurrence in nonmarine facies, and it will be the
subject of further study.
... Agha Jari Formation deposited in the continental environment to shallow marine conditions (Motiei 1993b;Ghazban andMotiei 2009, 1993a;Bahrami 2009;Sahraeyan and Bahrami 2012), and its continental condition is confirmed by occurrence of terrestrial vertebrate footprints. Most Cenozoic vertebrate tracks from the Zagros Mountains and adjacent areas have been reported from the Agha Jari Formation and its equivalent rock units in the Persian Gulf region (Lambrecht 1938;Abbassi et al. 2020;Abbassi 2021;Karim et al. 2003;Bibi et al. 2012;Schulp et al. 2011;Kazemi and Abbassi in press) (Fig. 2). ...
A slab with three tetradactyl avian footprints was excavated from the Agha Jari Formation (Mio-Pliocene) in the Zagros Mountains, southwest Iran, in the early last century and named as Urmiornis abeli. The name Urmiornis had been used for the didactyl bird fossil and ichnogenus name was later revised to Iranipeda. Validity of Iranipeda abeli has been discussed by authors in recent decades, and revision of the original material is necessary. Based on re-examination the holotype I. abeli reveals that this name is valid and it differs by long hallux, separate digit imprints and size of lateral digit II and IV traces with other large wading bird footprints. Iranipeda include three ichnospecies: I. abeli and I. millumi and I. intermedia by emended diagnosis of Iranipeda. Ciconiidae is the nearest track-maker candidate for Iranipeda. I. abeli type specimen has been collected from the lower part of the Agha Jari Formation and based on lithostratigraphic position and radiometric data, it is of Late Miocene age.
Agha Jari Formation (Miocene-Pliocene, 788 m) comprises alternations of sandstone, siltstone, and marl in red to brown color layers in the east Gotvand, north Khuzestan province, southwest Iran. Numerous vertebrate footprints have been found in the middle part of the Formation, which includes footprints of wetland small bird (Aviadactyla media, Gruipeda dominguensis), artiodactyls (Lamaichnum alfi, Lamaichnum isp., Pecoripeda satyri, Pecoripeda isp.), canids (Canipeda isp.) and large reptiles (Hatcherichnus sanjuanensis). Frequency of artiodactyl imprints are more than the other footprints. These footprints proof the living of camels and crocodiles in the Zagros basin for the first time. The ichnotaxa diversity of the footprints in the studied section is more than the other reports of Agha Jari Formation in Iran and its stratigraphic equivalent in the adjacent countries.
A diverse continental ichnofauna from the Lower Cretaceous (Aptian) Gething Formation occurs at the Ninesting Creek site in northeastern British Columbia. Vertebrate tracks including Wupus and Saurexallopus are inferred to represent medium-sized (footprint length ∼10 cm) tridactyl avian and larger (footprint length ∼15–17 cm), gracile, tetradactyl non-avian theropod trackmakers respectively. Shape, length-to-width and divarication ratios, suggest two morphotypes that are distinct from the poorly known, and infrequently-cited, Gething Formation ichnogenera Columbosauripus and Irenichnites (footprint lengths ∼12.5 and ∼15.0 cm respectively). This suggests a diverse theropod ichnofauna representing size classes which are quite distinct from the better-known large theropod ichnogenus Irenesauripus with footprint lengths in the range of ∼38.0–53.5 cm. Thus, current evidence suggests that the ichnodiversity of smaller avian and non-avian theropod tracks (footprint length ∼<15.0 cm) outnumbers large ‘megatheropod’ morphotypes by about 7:1. The implications of this distribution are consistent with recently proposed palaeoecological models. The assemblage also includes probable pterosaur tracks and distinctive arcuate compound Archaeonassa-Lockeia traces, attributed to freshwater unionid bivalves. The trace fossils comprise a composite Mermia-Scoyenia Ichnofacies assemblage. Palaeoenvironmentally the site represents the low energy margin of an open lake basin or floodplain pond, or a fluvial point bar.
The Ebro Basin (Spain) is one of the few worldwide areas where Palaeogene avian and mammal tracksites have been found. A new unpublished tracksite known as La Sagarreta is here described. The tracksite is located in a sandstone-dominated outcrop from the Early Oligocene Peralta Formation in the northern-central sector of the basin. Six different ichnotaxa have been identified. Four belong to mammals, including perissodactyl (cf. Plagiolophustipus isp), artiodactyl (Megapecoripeda isp. and cf. Pecoripeda isp.) and carnivoramorph tracks (cf. Canipeda isp.), and the other two to avian tracks (Aviadactyla vialovi and Gruipeda dominguensis). La Sagarreta presents a high ichnodiversity with the presence of a medium-sized member likely of the genus Plagiolophus (or relative), one medium- and one small-sized artiodactyl that probably belong to the family Entelodontidae, one medium-size unidentified carnivorous mammal and two types of birds. La Sagarreta tracksite is the most diverse Early Oligocene tracksite of the Ebro Basin and one of the few palaeontologic vertebrate records at the central area of the basin during this age.
The Mesozoic and Cenozoic track record of large birds (avian theropods) with footprint lengths (FL) > 10.0 cm is quite limited, whereas small tracks (FL < 10.0 cm) are abundant from the Early Cretaceous onwards. This large versus small threshold value is consistent with the track record of extant birds among which only ˜10% are large, and so is scaled appropriately to Class Aves. The proportion of large pachydactylous (thick-toed) or robust non-avian theropod ichnotaxa reported from the Mesozoic considerably exceeds the few large leptodactylous (thin-toed) or gracile forms such as Archaeornithipus and Magnoavipes, named, albeit controversially, to suggest avian affinity. The Cenozoic record of large avian tracks is quite different, with large pachydactylous tracks limited to a few Eocene-Oligocene (Palaeogene) ichnogenera (Ornithoformipes and Rivavipes), and rare unnamed Neogene and Holocene dinornithiform tracks. Thus, large flightless species are less well represented by tracks than body fossils. This suggests probable preservational bias in favour of waterbird or shorebird-like tracks in shoreline facies. However, the Cenozoic record of large leptodactylous tracks attributable to cranes, herons and related trackmakers is quite extensive, mostly confined to the Neogene and formally named to suggest representatives of extant avian clades capable of flight. These distribution patterns apparently reflect the dominance of large pachydactylous non-avian and avian theropods in the Mesozoic and early Palaeogene, and contrast with the rise of large, as well ongoing diversification of smaller leptodactylous Aequornithes in the Neogene. These distribution patterns in space and time likely reflect changing selection pressures as today's modern avifauna evolved.
Agha Jari Formation is one the extensive Cenozoic formations in the Zagros Mountains, that outcropped from the Bandar Abbas in southeast to Iraqi Kurdistan region in the northwest. This formation comprises of the siliciclastic and marl sediments. Agha Jari Formation is well exposed in the southern flanks of Zagros Mountains such as north Bushehr province. The formation has been deposited in the continental to shallow marine conditions, which includes lower member as red sandstone and marl alternations and upper red marl which named Lahbari Member. About 12 hyporelief mammal footprints have been found on the dislocated red sandstone slab (95 × 65 cm) from the west of Bibi Hakimeh anticline in the north of Bandar Deylam, Bushehr province. Based on the trackway ornamentations, it seems that footprints made by at last two mammals. Footprints have rounded outlines with three to four-digit imprints with claw drag marks in some of them. There are however, round footprints without digit imprints. All of trackways are same in direction. These footprints are tracks of Canidae (carnivores) based on morphology features, and this is first report of carnivore footprints from the Zagros Mountains
Numerous dinosaur footprints are reported from the Cretaceous of Iran for the first time. These footprints are small and they have been discovered in the one of uppermost layers of the Tirgan Formation (Neocomian-Aptian), Gelian section, south Shirvan, Northeast Iran. The Tirgan Formation comprises Orbitolina-bearing bioclastic and oolitic limestones (200 m), which were deposited in the tidal flat to shallow marine environments of a carbonate ramp. Footprints were impressed in the subaerial conditions of an intertidal environment. Based on microfossils and stratigraphic position, they are Aptian in age. Most footprints are weathered and remained as indistinct holes, but five distinctive trackways are distinguishable. One of the trackways belongs to small sauropod with a narrow gauge. The means of pes footprint length and footprint width are 27.8 cm and 37.7 cm, respectively. Means of footprint length and footprint width of the manus in this trackway are 14.1 cm and 19.3 cm. Well preserved plantigrade stegosaur footprints are circular to ellipsoidal in outline and include tridactyl pes and tetradactyl manus imprints. The means of footprint length and footprint width equal 15.5 cm and 12.8 cm in the pes and 6.4 cm and 10.3 in the manus. These stegosaur footprints are assigned to Deltapodus isp. Abundant, small, quadrupedal ornithopod footprints (footprint length of pes and manus equals 13.0 cm and 9.7 cm; footprint width of pes and manus equals 18.4 and 15.0 cm) are attributable to Caririchnium and Ornithopodichnus, though they have a lower footprint length/footprint width ratio than the well-known ichnospecies of these ichnotaxa. It seems that dwarfism of the Tirgan Formation's dinosaur track-makers was due to the position of the Iranian microplate in northern part of Tethys, which is explainable by the island rule.
Sedimentological study of Mukdadiya Formation (Late Miocene-Pliocene) indicates that its sediments were deposited in fluviatile meandering channels system. The sequence of the formation is composed of several fining-upward cyclJs in which three facier *"ri recognized: ' 4 The sandstones facies, represents channels deposits, the gravelly sandstones and conglomerates facies, represents a high energy environment reflecting a climatic or tectonic. .,, pulses in the source area, and the fine grained sediments facies, represents overbank deposits. The petrographic study of the sandstones of Mukdadiya Formation indicates that they are immature, mostly poorly sorted and classified as litharenite. They are composed mainiy of qtrartz and sedimenkry rock fragments; the predominant type of sedimentary fragments is , carbonate. The study of heavy minerals indicates that epidote group is forming the main heavy minerals, followed by amphibole, pyroxene and gamet. The so*ce ro"t q based on pelrographical and heavy minerals studies are interpreted to be composed essentially of sedimentary followed by igneous and metamorphic rocks
A new vertebrate skeleton of Late Miocene-Early Pliocene age has been discovered for the first time, near Kanisard village (Chamchamal area, Kurdistan/NE-Iraq). The carnivorous vertebrate remains comprised of the skull, teeth, backbone, and legs of a mammal of the Late Miocene Zagros orogenic belt. The remains are recognized and collected within pebbly sandstone horizons of Mukdadyiah Formation. The skeleton bearing sequence is comprised of rhythmic alternations of reddish brown claystone and greenish gray pebbly sandstone, which is also characterized by megaripple marks, channel deposits, flute casts and clay balls, associated with the dominance of Skolithos and Scoyiena trace fossils. The terrestrial skeleton with bioturbation and the associated sedimentary structures indicates continental fluvial environment. All conjugate lines of evidences, that is morphological criteria of skull, leg length and width, teeth types, size and numbers, Jaw curvatures outline and the overall vertebrate size and shape, specify that the carnivorous mammal is related to Canis family (dog-like carnivorous). Its age when the individual is died estimated to be about 5 years, while its death possibly occurs before 8-9 Million years ago. (That is Late Miocene).
Bird and mammal tracks from the White Narrows Formation (early Pliocene) in the Moapa area, near
Glendale, Nevada, have been known and collected for several decades, furnishing at least four museums with
representative specimens. This easily accessible site contains the best Nevadan examples of early Pliocene ungulate and carnivore tracks. One carnivore ichnotaxon (Platykopus) has its type specimen from the locality. Bird tracks are also very abundant and include tracks attributable to heron- and duck-like species (cf. Ardeipeda and Avipeda sensu lato). Because the track assemblage has been collected with impunity in the past, it is in need of further research, continual monitoring and stewardship. The site is currently within a federal ACEC and protected by regular BLM patrols.
The Neogene Red Beds of the Tarom valley (north-western Iran) include conglomerate, sandstone, marl and gypsum. Avian and mammal footprints were discovered in one of the sandstone layers at the base of a third Miocene stratigraphical unit in the Gilankesheh area located in the east Tarom valley. The avian ichnia include Aviadactyla vialovi, Avipeda filiportatis, Charadriipeda disjuncta, Charadriipeda isp. A and B and cf. Ornithotarnocia lambrechti. Bird feeding traces are preserved as bilobate, loop-shaped, sinusoidal and ring-like traces. We have also identified a reticulate texture of sole scale imprints in some of the avian ichnia. Two mammal footprints of camelid-like artiodactyls are also present with the avian ichno-assemblage.
A diverse suite of vertebrate footprints documented along the coast of Lithuania offers insight into the distribution and preservation potential of tracks in sandy substrates. Whereas ichnotaxonomy is not conventionally applied to modern tracks, a case is made for using footprint classification based on Vialov’s (1966) nomenclature to both observed and predicted track distribution based on extramorphological characteristics of the tracemaker. Along a barrier spit (Curonian Spit) and mainland shoreline (Būtingė) sites, two general types of avian footprints (Avipeda sp), as well as three carnivore (Bestiopeda), four ungulate (Pecoripeda, Suipeda, Hippipeda), and two rodent (Rodentipeda) track morphotypes are described. The habitat preferences result in a predictable distribution of track assemblages with a potential of differentiating vertebrate ichnofacies (lagoon shoreline, open sea beach, dunes). The moisture content, possibility of freezing and rapid aeolian burial, and lithological anomalies (e.g., heavy-mineral concentrations) enhance the preservation potential of tracks. Once preserved, the attitude of trackways in respect to bedding surfaces determines their integrity and here a model of potential regional track distribution is proposed. Track formation and preservation in modern settings provides a useful guide to locating and studying vertebrate footprints and other traces in sand-dominated coastal and aeolian sequences.
The stratigraphy of the High Folded Zone (HFZ), in Iraq is reviewed. The oldest exposed rocks are Early Triassic in age, whereas the youngest are of Pliocene – Pleistocene age, which belong to the Bai Hassan Formation. The exposed stratigraphical column is represented by 36 formations, with 10 main types of Quaternary sediments, which have wide geographic extent and well preserved in the HFZ.
The Triassic, Jurassic, Cretaceous and Paleogene rocks are mainly of marine carbonates with some clastics. The Oligocene rocks form a complex of reef – backreef – forereef and are restricted almost in the eastern and western parts of the area, with restricted exposures in different parts. The Early and Middle Miocene rocks are mainly of marine origin, lagoonal carbonates and fairly developed evaporates, respectively, with restricted exposures. The Late Miocene – Pleistocene rocks, which represent the beginning of the continental environment, consist of molasse sediments, deposited in sinking foredeep, which had few separated and isolated basins. The Quaternary sediments are well developed, especially Pleistocene river terraces of different stages, alluvial fan sediments, calcrete and other different types. Glacier moraine sediments, may be present in Rawandooz – Galala vicinity.
For each exposed formation, the type locality, exposure areas, subsurface extension, main lithology (as described in form of members and/ or informal units), thickness, fossils, age, depositional environment, and the lower contact are described. The described lithologies of the formations by different authors from different localities are reviewed, with occasional remarks by the present authors. The main tectonic events and the paleogeography of each era or period are reviewed briefly. Each formation is discussed, for the majority of them the present authors' opinion are given, with many recommendations for future studies. Some new ideas dealing with many aspects for many formations including proposals for establishing new formations are given, too.
The Zagros Mountains of Iran formed by continental collision from closure of the Neo-Tethyan Ocean. The Zagros Orogen underlying the mountain range reflects a much longer history with the Pan-African basement and Phanerozoic successions. New mapping, radiometric ages, and stratigraphic analyses have enabled advances in our understanding of the Jurassic to Cenozoic tectonic history. The northwestern Zagros Orogen consists of three belts: (1) the Zagros Fold and Thrust Belt, divided into the outer Zagros Simply Folded Belt and the inner High Zagros Belt; (2) the Zagros Suture Zone including radiolarite, ophiolite, and Bisotun limestone thrust sheets; and (3) the Sanandaj–Sirjan Zone, which contains abundant metamorphic rocks. Late Cretaceous ophiolites of the Kermanshah region are part of the outer ophiolite belt of the Zagros Orogen and have formed in passive margin and supra-subduction zone settings. Major events include early Mesozoic rifting, Jurassic subduction followed by a more cryptic interval of subduction in the Cretaceous, multiple ophiolite emplacement on the Arabian margin in the Late Cretaceous to Eocene, and collision of central Iran and the Arabian margin in the Oligocene with final closure of the shallow Tethyan seaway in the mid-Miocene. A Middle to Late Jurassic plutonic belt, the Qorveh–Aligodarz Plutonic Belt, formed a magmatic arc with subdued topography related to a moderately NE-dipping subduction zone under the Sanandaj–Sirjan Zone. An Early Cretaceous unconformity reflects limited uplift followed by widespread marine deposition with intercalated volcanic rocks in the Sanandaj region. Subduction continued with a low-lying arc that underwent trenchward advance. In the Late Cretaceous to Oligocene interval, the Neo-Tethyan Ocean closed with ophiolite obduction over the Arabian Peninsula margin and major shortening affected the Sanandaj–Sirjan Zone with uplift and plutonism. Much of the forearc of the Jurassic to Cretaceous arc system has been lost by tectonic erosion along a low-angle Eocene subduction zone prior to collision. Flattening of the subducting slab in the Late Cretaceous and Palaeogene explains the inland retreat of the arc to central Iran. Continental collision initiated in the Oligocene but the Tethyan seaway remained open until the mid-Miocene.
The Zagros Basin is broadly defined as the palaeodepositional wedge of sediments along the present belt of the Zagros Mountains. A series of ten regional isopach maps trace the development of a portion of this basin from Permian to Recent. From the Permian to the middle Cretaceous the area occupied a position along the stable northeastern Atlantic-type shelf margin of the Afro-Arabian continent, bounded by a rift zone that evolved into a southern Tethys ocean. Late Cretaceous to Recent subsidence patterns are influenced by plate margin tectonics, obduction, and eventual continental collision along the Zagros Suture as this ocean closed. The late Alpine Zagros folding and faulting took place from the Miocene onwards.
The Late Miocene Sorbas Member of the Sorbas Basin, Almería Province, southeast Spain contains an extensive avian ichnofauna preserved in lagoonal marls. Three distinctive avian ichnotaxa can be identified: Antarctichnusfuenzalidae Covacevich and Lamperein, 1970; Iranipedamillumi n. ichnosp.; and Roepichnusgrahami n. ichnogen, n. ichnosp. In common with many other Cenozoic avian ichnofaunas, these traces are associated with shorebirds, including plovers, storks, ducks and/or gulls, respectively. Associated mammalian tracks include possible cat and artiodactyl footprints. The avian tracks are abundant and show a range of behavioural aspects in common with other recorded examples of Cretaceous–Recent shorebird tracks. These include both solitary and group activities consistent with their postulated avian tracemakers.
The most recent global "icehouse-hothouse" climate transition in earth history began during the Permian. Warmer polar conditions, relative to today, then persisted through the Mesozoic and into the Cenozoic. We focus here on two Permian stages, the Sakmarian (285-280 Ma) and the Wordian (267-264 Ma; also known as the Kazanian), integrating floral with lithological data to determine their climates globally. These stages postdate the Permo- Carboniferous glaciation but retain a moderately steep equator-to-pole gradient, judging by the level of floral and faunal differentiation. Floral data provide a particularly useful means of interpreting terrestrial paleoclimates, often revealing information about climate gradations between "dry" and "wet" end-member lithological indicators such as evaporites and coals. We applied multivariate statistical analyses to the Permian floral data to calibrate the nature of floral and geographical transitions as an aid to climate interpretation. We then classified Sakmarian and Wordian terrestrial environments in a series of regional biomes ("climate zones") by integrating information on leaf mor- phologies and phytogeography with patterns of eolian sand, evaporite, and coal distributions. The data-derived biomes are compared here with modeled biomes resulting from new Sakmarian and Wordian climate model simulations for a range of CO2 levels (one, four, and eight times the present levels), presented in our companion article. We provide a detailed grid cell comparison of the biome data and model results by geographic region, introducing a more rigorous approach to global paleoclimate studies. The simulations with four times the present CO2 levels (4#CO2) match the observations better than the simulations with 1#CO2, and, at least in some areas, the simulations with 8#CO2 match slightly better than those for 4#CO2. Overall, the 4#CO2 and 8#CO2 biome simulations match the data reasonably well in the equatorial and midlatitudes as well as the northern high latitudes. However, even these highest CO2 levels fail to produce the temperate climates in high southern latitudes indicated by the data. The lack of sufficient ocean heat transport into polar latitudes may be one of the factors responsible for this cold bias of the climate model. Another factor could be the treatment of land surface processes and the lack of an interactive vegetation module. We discuss strengths and limitations of the data and model approaches and indicate future research directions.
Two new sites with mammalian footprints in the early Oligocene of southeastern France are described here. They represent one of the best preserved and more numerous record of tracks and trackways in the world with more than 320 ichnites. Many of those are arranged in trackways and sometimes show pes-manus impressions, a quite rare feature in mammalian ichnology. The ichnotaxonomic study indicates the presence of perissodactyls tracks referred to as Rhinoceripeda voconcense (Demathieu et al., 198419.
Demathieu , G. , Ginsburg , L. , Guérin , C. and Truc , G. 1984 . Etude paléontologique, ichnologique et paléoécologique du gisement oligocène de Saignon (bassin d’Apt, Vaucluse) . Bulletin du Museum National d’Histoire Naturelle, Paris , 6 : 153 – 183 . View all references), artiodactyls footprints referred to as Megapecoripeda velox (Demathieu et al., 198419.
Demathieu , G. , Ginsburg , L. , Guérin , C. and Truc , G. 1984 . Etude paléontologique, ichnologique et paléoécologique du gisement oligocène de Saignon (bassin d’Apt, Vaucluse) . Bulletin du Museum National d’Histoire Naturelle, Paris , 6 : 153 – 183 . View all references) and a carnivore footprint referred to as Bestiopeda sp. Vialov (1966)55.
Vialov , O. S. 1966 . Sledy zhiznedeyatelnosti organizmow i ikh paleontologicheskoe znachenie . Naukova Dumka , : 219 (in Russian) View all references. They can be attributed to early Rhinocerotids, Lophiomerycids and/or Entelodonts and Mustelid-like carnivore, respectively. This study also aims at homogenizing the ichnotaxonomy used for mammal tracks where several genera were erected without a full review of the literature. All this taken together reveals a rich mammalian ichnofauna at a time period when no other fossils of mammals are known in the area and represents a good opportunity to provide state-of-the-art concerning the worldwide known sites that yielded mammalian footprints.
A bibliography of 584 references describing Cenozoic or Holocene vertebrate tracks and trackways with a world-wide distribution is presented. We have analyzed the reports by decade; there has been a steady increase in the publication of such studies since the first verifiable report of a Cenozoic track in 1859. We show all described occurrences on two maps, one for the Paleocene to Miocene, and another for the Pliocene and Pleis-tocene, with map symbols for each Epoch. We provide an extensive cross-index by categories of age, geographical location and taxa. Finally, we list all described Cenozoic vertebrate ichnotaxa known to us.
The Zagros orogen provides a unique opportunity within the Alpine system to evaluate the interplay between a young Tertiary collision and earlier subduction/obduction processes. Within the Crush zone and the Sanandaj-Sirjan (internal) zone separating the Zagros Fold belt from Central Iran, we document several major tectonic events taking place at the end of the Cretaceous, of the Eocene and from the Mio-Pliocene onwards (ca. <20-15 ma). contrary="" to="" recent="" interpretations,="" our="" data="" (cross-sections="" and="" description="" of="" the="" overall="" deformation="" style)="" strongly="" suggest="" that="" the="" main="" zagros="" thrust="" (mzt)="" is="" deeply="" rooted,="" possibly="" to="" moho="" depths,="" and="" that="" the="" suture="" zone="" effectively="" runs="" along="" the="" mzt.="" field="" observations="" show="" that="" the="" final="" resorption="" of="" the="" oceanic="" domain="" took="" place="" slightly="" after="" 35 ma="" and="" that="" collision="" must="" have="" started="" before="" ca.="" 23-25 ma="" in="" northern="" zagros.="" the="" shortening="" rate="" across="" the="" crush="" zone="" since="" the="" mid-miocene="" (20-15 ma)="" is="" estimated="" at="" a="" minimum="" 3-4 mm/year.="" shear="" movements="" in="" the="" crush="" zone="" during="" the="" eocene-oligocene="" period="" and="" extensional/strike-slip="" movements="" in="" the="" internal="" zones="" during="" the="" late="" cretaceous="" point="" to="" an="" oblique="" setting="" early="" in="" the="" convergence="" history.="" a="" geotectonic="" scenario="" for="" convergence="" from="" the="" time="" of="" obduction="" to="" the="" present="" is="" finally=""
Petrographic, geochemical, and scanning electron microscope analyses of the sandstone and mudstone units of the Upper Miocene Injana Formation are presented. Furthermore, microprobe analysis for amphiboles, pyroxenes, garnet, and chromian spinels as common heavy mineral species present is done to support other results for better understanding of the provenance history of the Injana Formation. The sandstones of the Injana Formation consist of terrigenous carbonate lithic fragments as common type of sedimentary rock fragments in addition to chert, argillaceous, and rare sandstone fragments. They also include metamorphic and igneous lithic fragments, quartz, feldspars, and mica and generally, the sandstones are lithic arenites and immature. Scanning electron microscopic analysis for the heavy minerals shows that they have been affected by dissolution due to chemical etching and mechanical abrasion through several surface texture generated either in arid and semihumid environment or in diagenetic environment. Clay mineralogy of the mudstone units indicates the presence of illite, chlorite, kaolinite, palygorskite, and illite-smectite mixed layers. Bulk-rock and mineral phase geochemistry in addition to petrographic data suggest the derivation of the Injana Formation from a nearby sources with contribution from igneous, metamorphic, and sedimentary provenance mainly from the high lands in the northeastern parts of Iraq which comprise mainly the Zagros mountains and the older sedimentary formations.
Although dinosaur eggs were first discovered and identified in the late 1800s, limited attention was given to the scientific value of oological fossils in contrast to observations based on skeletal features. Here, we offer a review of Mesozoic saurischian egg materials, in comparison with extant crocodilians and avians, and their paleobiological interpretation based either on the presence of embryos in ovo or brooding adults on egg clutches. Our study focuses on the eggs of the oviraptorid Citipati osmolka (Mongolia), the troodontid Troodon formosus (North America), the theropod oospecies Macroelongatoolithus xixiaensis (China), the ornithothoracine bird (Argentina), an indeterminate theropod (Thailand), and titanosaurs (Argentina). Results show that (1) many oological characters and reproductive behaviors associated with modern birds are rooted among non-avian theropods, (2) there is a reproductive evolutionary cline from crocodilians to modern birds with (3) a noticeable pattern of coeval development between the accretion of eggshell layers, origination and size increased of larger air cells (inferred from egg polar asymmetry), and brooding/incubating behaviors. Most of these pre-adaptations are grouped in two main clades of the saurischian cladogram: one at the level of Oviraptorosauridae and the other at Troodontidae. Although undeniably these two theropod taxa seem to represent two important phases for the evolution of avian reproduction, the phylogenetic distance between these clades and Titanosauria cannot be ignored. As such, the reproductive features that appeared in concert in oviraptorids might have gradually evolved across more basal theropod clades. Although Troodon formosus by its egg shape and nesting behavior seems to be in this study the precursors of modern avian reproduction, the importance of small-bodied theropods such as those who laid the Phu Phok eggs cannot be dismissed and the eggs of such dinosaurs could suggest a closer phylogenetic ties to Aves than troodontids. At a higher level of inferences, there is a strong possibility that the evolution of these reproductive features is concurrent with profound physiological and metabolic changes that occurred in saurischian dinosaurs throughout their evolution.
With high organic content and abundant microbiota, feces are typically only preserved for more than a couple of days, weeks or months after deposition and most never enter the fossil record. The range of depositional conditions favorable to the feces preservation is narrow, but depends on a large number of biotic and abiotic factors. Moreover, because the chemistry and microbiology of feces often promotes mineralization, discriminating between coprolites and some inorganic sedimentary structures is often difficult. We propose a protocol to identify coprolites in the fossil record that encompasses all the criteria previously defined. We then apply this protocol to identify putative coprolites from the lower Carnian (Upper Triassic) Chañares Formation of northwestern Argentina. Using a variety of analytical methods and several different criteria, we were able to identify them as carnivore coprolites. Based on extant analogs, the most probable producer would be a small carnivorous cynodont or archosauriforms, and the prey is likely a tiny therapsid or archosauriform. This is the first detail report of carnivore coprolites from this unit, and provides direct evidence of trophic links in the Chañares ecosystem.
Vertebrate coprolites are common in the Upper Triassic Chinle Group of the western United States. Two new ichnotaxa (Dicynodontocopros maximus, Heteropolacopros texaniensis, both ichnogen. and ichnosp. nov.) have stratigraphic ranges restricted in the late Carnian. Heteropolacopros occurs throughout the Otischalkian and Adamanian whereas Dicynodontocopros is restricted to the Adamanian. Coprolite acme zones occur in the upper Carnian, lower Norian and Rhaetian portions of the Chinle Group. The distribution of some Chinle Group coprolites is facies controlled.
Mammalian and avian footprints documented from the Oligocene of the Khaf area, northeastern Iran, represent the first reported Oligocene vertebrates in the country. Avian andmammalian footprints are located on the upper bedding plane of two successive, medium-bedded, brownish litharenite sandstone layers. Layer one has more than 97 concave epirelief avian footprints that are arranged in 29 straight or curved trackways. Layer two contains dispersed mammalian footprints. The avian footprints include four slender digits with sharp tips, three oriented forward (II–IV), and one of them oriented backward (I). Metapodium imprints are visible in some footprints as isolated circular concavities. The avian footprints are attributed to Gruipeda intermedia. In order to establish this identification, we re-affirm the diagnostic features of Gruipeda, Iranipeda and Charadriipeda, and also re-evaluate the ichnospecies assigned to these ichnogenera. The mammalian footprints from the Khaf locality include Khafipus khadari nov. igen and nov. isp.,which are large plantigrade footprints that have five dispersed digit imprints in the pes or manus and heel or digit imprints that are enclosed by circular ridges, which are disconnected from each other. Palaeotheriipus isp. is a tridactyl mammalian footprint that is large in the heel with forward oriented toe imprints. Platykopus isp. includes one footprint as a broad plantigrade impression with imprints of five connected digits. Avian footprints are attributed to anisodactyl shore birds and mammalian footprints are those of perissodactyls and large mammals such as ursid carnivores. The footprints from the Oligocene of Iran documented here add to a growing, but much understudied global record of Cenozoic tetrapod footprints and suggest a broad paleogeographic distribution of various, particularly avian, ichnogenera.
The discovery of new fossils in the Late Miocene of Injana, Jebel Hemrin (Iraq) enables us to complete the Injana vertebrate faunal list, which now includes 25 vertebrate taxa, 19 of them belonging to mammals. The Injana paleoenvironment corresponds to a vast fluviatile system where, under hot and wet conditions, there existed a mosaic of landscapes ranging from grassland to woodland and forest, with swampy zones. It is similar to that of the Middle Siwaliks of the Indian Subcontinent and differs from that of the Irano-Turkish Plateau.The Injana fauna fo Iraq established the existence, during the Late Miocene, of a biogeographical area which had preferential links with the Indian Subcontinent in the east and Africa in the west. This area was more or less isolated from the adjacent northern regions (Irano-Turkish Plateau) by the southern branch of the Asian Alpine Range (Taurids and Zagrids); this acted as a geographical barrier which many vertebrates did not cross.RésuméLa découverte de nouveaux fossiles dans le Miocène supérieur à Injana, Jebel Hemrin en Irak, permet de compléter la liste faunique d'Injana qui comporte à présent 25 taxons de Vertébrés dont 19 Mammifères. Le paléoenvironnement d'Injana correspond à un vaste système fluviatile où, sous un climat chaud et humide, existait une mosaïque de paysages allant de la prairie à la forêt claire et dense, avec des zones marécageuses; il ressamble à celui du Siwalik moyen du Sous-continent indien et diffère de celui de la Plateforme irano-turque.Au cours du Miocène supérieur, la faune d'Injana en Irak témoigne de l'existence d'une aire biogéographique présentant des relations privilégiées avec le Sous-continent indien à l'Est et l'Afrique à l'Ouest. Cette aire est plus ou moins isolée des régions limitrophes septentrionales (Plateforme irano-turque) par la branche Sud de la Chaîne alpine asiatique (Taurides et Zagrides) qui joue un rôle de barrière géographique que de nombreux Vertébrés n'ont pas franchi.
Vertebrate coprolites are common in the Upper Triassic Chinle Group of the western United States. Two new ichnotaxa (Dicynodontocopros maximus, Het‐eropolacopros texaniensis, both ichnogen. and ichnosp. nov.) have stratigraphic ranges restricted in the late Carnian. Heteropolacopros occurs throughout the Oti‐schalkian and Adamanian whereas Dicynodontocopros is restricted to the Adamanian. Coprolite acme zones occur in the upper Carnian, lower Norian and Rhaetian portions of the Chinle Group. The distribution of some Chinle Group coprolites is facies controlled.
Relationships between ichnology and uniformitarianism are perhaps less complicated than those of their sister subdisciplines, paleontology and paleoecology. Trace fossils are manifestations of benthic behavior; and these traits, although evolving in significant ways, have remained stable over longer spans of time than individual species of invertebrates. The fossil record of behavior in fact originated earlier than the fossil record of invertebrate body parts. Although macroinvertebrates and their traces exhibit tremendous diversity of form and function, these fit into a relatively small number of behavioral patterns. The patterns, in turn, may correlate with prevailing environmental conditions, resulting in gradients among trace fossil assemblages, or ichnofacies. Behavioral patterns and characteristic ichnofacies therefore constitute the main basis for uniformity in ichnology. Thus, the most fundamental questions are: What is the specific function represented by the trace? How will it change as the tracemaker is influenced by other genetic, physiologic, or ecologic stimuli? In which facies will it likely occur? and what preservational biases are apt to modify the fossil record of this behavior and its environmental distribution? Approached from this standpoint, the present is indeed a key to the ichnologic past, and vice versa. In practice, however, the present has been studied considerably less than its importance would dictate.
The latest Neoproterozoic through Phanerozoic stratigraphy of the Zagros fold-thrust belt of Iran has been revised in the light of recent investigations. The revised stratigraphy consists of four groups of rocks, each composed of a number of unconformity-bounded megasequences representing various tectonosedimentary settings. In the lowest group, ranging in age from latest Precambrian to Devonian(?), the uppermost Neoproterozoic to middle Cambrian rocks constitute a megasequence of evaporites, siliciclastic deposits, and interlayered carbonates, which were deposited in pull-apart basins that developed by the Najd strike-slip fault system. This mega-sequence is overlain by a second one, Middle to Late Cambrian in age, which consists of shallow, marine siliciclastic and carbonate rocks representing deposition in an epicontinental platform. The overlying shales, siltstones, and partly volcanogenic sandstones of Ordovician, Silurian, and Devonian(?) age are local remnants of stratigraphic units that were extensively eroded during development of several major unconformities. The second group consists of two megasequences, one Permian and the other Triassic, composed of widespread, transgressive basal siliciclastic rocks and overlying evaporitic carbonates of an equatorial, epi-Pangean, very shallow platformal sea. The third group is composed of four megasequences formed of shallow-and deep-water carbonates with some siliciclastic and evaporite deposits, which accumu-lated on a Neo-Tethyan continental shelf during earliest Jurassic through late Turonian time. The fourth group comprises siliciclastic and carbonate deposits of a largely underfilled, NW-to SE-trending, forward and backward migrating, late Cretaceous to Recent proforeland basin, which has evolved as an integral part of the Zagros orogen. This last group consists of three megasequences (IX, X, and XI) with distinctive lateral and vertical facies variations, which reflect specific tectonic events. Megasequence IX comprises uppermost Turonian to middle Maastrichtian prograding and retrograding siliciclastic and carbonate deposits, whose accumulations reflect emplacement ("obduc-tion") of ophiolite slivers and subsequent collisional events in the Zagros orogen. Megasequence X consists of uppermost Maastrichtian to upper Eocene siliciclastic and carbonate rocks, which deposited first progradationally in front of the Zagros orogenic wedge with reduced contractional tectonic activity, and then retrogradationally due to intensified thrust stacking in the interior parts of the orogen. Megasequence XI consists of Oligocene and lower Miocene carbonate strata deposited retrogradation-ally shortly after a period of intensified late Eocene thrust faulting in the deforma-tional wedge, and an overlying succession of upward-coarsening, northeasterly-derived siliciclastic deposits of lower Miocene to Recent age which are composed of erosional byproducts of the southwest-vergent Zagros thrust sheets.
The Copper Canyon Formation consists of over 3000 m of basalt and alluvial fan and lacustrine strata deposited in a small Neogene basin. Lacustrine strata are of two typs: fine-grained dolomitic carbonates deposited in a playa-lake environment and calcitic bioclastic carbonates deposited in a freshwater lake environment. Bedding surfaces of the playa-lake carbonates deposited in shoreline environments commonly contain avian and mammalian footprints. Tracks of artiodactyls are most abundant; those of birds, horses, carnivores and proboscideans are less common. Using the binomial avian and mammalian footprint classification scheme of Vialov (1966), four different mammalian ichnogenera containing thirteeen ichnospecies, as well as six ichnospecies of avian tracks are recognized.
This paper reviews the coprolites (fossil excrements) of terrestrial vertebrates, giving particular attention to examples that may have originated from dinosaurs. Factors affecting the preservation of corprolites and briefly surveyed, and the internal and extent morphology of these fossils is examined in detail. Coprolites furnish small, but sometimes important, pieces of evidence about dinosaur biology: such evidence is rarely conclusive or unbiased, though in favourable circumstances it can be used to ascertain the identity, behaviour, habitats and diets of dinosaurs. Information derived from coprolites is best appraised in the context of broad-based studies which also consider the anatomical adaptations of dinosaurs, their gut contents and their feeding traces, along with any associated fossils that might betray the potential sources of food exploited by dinosaurs.
The ichnofauna of the Woodbine Formation (Cenomanian), Denton County, Texas includes bird and dinosaur tracks. A new bird trackway,Magnoavipes loweiichnogen. et ichnosp. nov., appears to represent the largest bird tracks known from the Mesozoic. A theropod trackway,Fuscinapedis woodbinensisichnogen. et ichnosp. nov., is characterized by long digits of uniform width and pointed small claws. Six hadrosaurid trackways,Caririchnium protohadrosaurichnosichnosp. nov., are the oldest hadrosaurid tracks associated with skeletal elements. They include one isolated small footprint, a medium sized quadrupedal, and five large bipedal hadrosaurid trackways.
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Dust Bathing Behaviours of Elephants, Zebras and Wildebeest and the Potential Risk of Inhalational Anthrax in Etosha National Park
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