The sclerobionts of the Bajocian Oolithe ferrugineuse de Bayeux Formation from Calvados (Paris Basin,Normandy, France)

Silicified fossils collected in ploughed fields at Gavrus (Calvados, France), mainly mollusc shells, are Bajocian in age, and come from the Oolithe ferrugineuse de Bayeux Formation. The entire formation is highly condensed and most fossils are reworked. Their silicification allowed treatment with dilute hydrochloric acid. This treatment brought to light numerous encrusting sclerobionts: Porifera (3 taxa), Bryozoa (n), Polychaeta (9), Brachiopoda (n), Bivalvia (5). The borings and bioerosional traces are described using the “categories of architectural design” as defined by Buatois et al. (2017). Among the 28 ichnotaxa described (corresponding to an ichnodisparity of 14), Planavolites wisshaki isp. nov., Kleithrichnus belemnophilus igen. nov., isp. nov. and Foggara foggara igen. nov., isp. nov. are new. Some Gastrochaenolites dijugus Kelly and Bromley, 1984 borings accommodate the shell of the presumed borer Lithophaga fabella J.-A. Eudes-Deslongchamps, 1838. A group of Nododendrina europaea (Fischer, 1875) on a belemnite rostrum provides a good instance of an ichnogenetic series. Encrusting sclerobionts and macroscopic boring and bioerosional trace-makers are dominantly suspension feeders. The ichnofacies is a peculiar Entobia-ichnofacies, found in deeper, lower-energy environments than the “classical” coastal Entobia-ichnofacies, on deep shell-grounds far from the coast, in the deep euphotic zone.     

Dung-Beetle Fossil Brood Balls: The Ichnogenera Coprinisphaera Sauer and Quirogaichnus (Coprinisphaeridae)

Coprinisphaera is one of the most common trace fossils in South American paleosols from the Eocene to the Recent. Nevertheless, an ichnotaxonomical review of this ichnogenus has been lacking until now. The revision of hundreds of specimens of Coprinisphaera from numerous formations in Argentina, Uruguay and Ecuador has resulted in the recognition of two ichnogenera. One of them, Coprinisphaera, includes isolated, spherical, subspherical, pear-shaped and bispherical structures grouped in five ichnospecies: C. ecuadoriensis Sauer, C. murguiai (Roselli), C. kraglievichi (Roselli), C. tonni isp. n., and C. kheprii isp. n. The new ichnogenus and ichnospecies, Quirogaichnus coniunctus, consists of Coprinisphaera-like structures clustered in a common excavated chamber. The ichnotaxobases for the taxonomical arrangement of Coprinisphaera are mostly related to the presence and position of a small chamber (interpreted as the original egg chamber) with respect to a large chamber (provision chamber) and emergence hole. The egg chamber may be included in (1) the provision chamber, leaving no remains in the trace fossils and resulting in cylindrical emergence holes; (2) the wall of the provision chamber, without additional structures; and (3) a mamillate to spherical additional structure, external to the provision chamber, resulting in pear-shaped to bispherical trace fossils.     

Upper Triassic (Keuper) non-marine trace fossils from the Haßberge area (Franconia, south-eastern Germany)

Various trace fossils from the Hassberge Formation and the Löwenstein Formation (Middle Keuper, Upper Triassic) of the Haßberge region are described. Twenty-three different forms have been identified, 17 of which are named, includingCruziana pascens n. isp.,Lockeia cunctator n. isp., andRusophycus versans n. isp.Lockeia siliquaria James, 1879,L. amygdaloides (Seilacher, 1953),L. triangulichnus Kim, 1994, andL. elongata (Yang, 1984) are revised and synonymized under the oldest available name,L. siliquaria James, 1879.Rusophycus eutendorfensis (Linck, 1942) andR. carbonarius Dawson, 1864 are revised. The diagnosis ofPolykladichnus Fürsich, 1981 is emended, and a diagnosis forHelminthoidichnites Fitch, 1850 is given for the first time. Among the described ichnotaxa,Skolithos ispp.,Rusophycus carbonarius, andTaenidium barretti are the most common forms. The trace fossil association is typical of theScoyenia ichnofacies, which indicates non-marine, periodically or completely inundated environments, such as floodplains and lake margins. Two palaeoichnocoenoses are identified. One ichnocoenosis, dominated byCruziana problematica, cf.Polykladichnus isp., andSkolithos isp. B characterizes margins of trough cross-bedded sandstones. Another ichnocoenosis, dominated byRusophycus versans n. isp.,Taenidium barretti,Scoyenia gracilis andSkolithos isp. A is related to ephemeral lake deposits. Taxonomic recommendations for the use of hitherto described and figured invertebrate Keuper trace fossils from Germany are given.     

Devonian Trace Fossils of the Horlick Formation,Ohio Range,Antarctica: Systematic Description and Palaeoenvironmental Interpretation

The clastic Horlick Formation contains an ichnofauna of 28 ichnotaxa dominated by burrowers. These are a mixture of simple vertical forms (Skolithos linearis, S. magnus, Bergaueria cf. langi, Rosselia socialis, Monocraterion isp.), U or arc-like forms (Diplocraterion parallelum, Arenicolites types A and B, Catenarichnus antarcticus, C. isp., aff. Lanicoidichna isp.), and complex, vertical spiral structures (Asterosoma isp., Spirophyton isp.). Horizontal burrows include Ancorichnus cf. capronus, Palaeophycus tubularis, and Psammichnites devonicus isp. nov. Surface traces comprise Haplotichnus isp., Cruziana problematica, C. rhenana, Rusophycus aff. carbonarius, R. isp., Protovirgularia rugosa, Lockeia ornata and cubichnia indet., while trackways include Diplichnites gouldi, D. isp., Maculichna? isp. and large imprints. The Horlick Formation (maximum 56 m) records an early Devonian transgression onto a deeply weathered land area that lay in the direction of Marie Byrd Land, spreading from the South Africa sector of Gondwana. Analysis of the trace fossils confirms their formation in near-shore to intertidal environments, with some ichnotaxa living close to the marine/fluvial boundary (e.g., Spirophyton, aff. Lanicoidichna, Cruziana problematica, Rusophycus aff. carbonarius, Maculichna?). The ichnofauna probably comprised a variety of marine suspension and deposit-feeding worms, shallow burrowing molluscs (bivalves and possibly bellerophontids) and several different types of arthropods, including trilobites.     

Lithostratigraphy of the Lower Cambrian metaclastics and their age based on trace fossils in the Sand?kl? region, southwestern Turkey

In the Sand?kl? region of the Taurus Range of Turkey, greater than 3000 m in thickness metamorphosed siliciclastics and volcanics (Kocayayla Group) underlies the trilobite-and conodont-bearing Middle-Late Cambrian Hudai Quartzite and Çaltepe Formation.The Kocayayla Group, previously regarded as Infracambrian or Precambrian, is dated for the first time as Early Cambrian on the basis of trace fossils. Cruziana ?fasciculata, C. ?salomonis, ?Cruziana isp., ?Diplichnites isp., Monomorphichnus isp., Petalichnus isp., Rusophycus ?avalonensis, R. ?latus, Arenicolites isp., cf. Altichnus foeyni, Planolites isp., Skolithos isp., and ?Treptichnus isp. have been recognised. These trace fossils are considered Tommotian or younger in age but older than the overlying, trilobite and conodont bearing Middle Cambrian limestones of the Çaltepe Formation. The trace fossils were likely produced by trilobites, suspension feeding annelids and deposit feeding “worms”, probably polychaetes. Sections bearing abundant Skolithos represent the Skolithos ichnofacies, which is typical of high energy environments with loose sandy, well sorted to slightly muddy substrates in intertidal to shallow subtidal zones. The other trace fossils represent the Cruziana ichnofacies, which is typical of subtidal, poorly sorted and soft substrates, from moderate energy to low energy environments between the fairweather and storm wave base.The Kocayayla Group was deposited at an early stage in a shallow marine stable shelf condition. The shelf subsided in a later stage and was affected by normal faults along which mafic and felsic volcanic rocks erupted. The volcanic activity had ceased and a shallow marine clastic sedimentation took place in the final stage of the shelf development. The Kocayayla Group was deformed and metamorphosed before the deposition of the trilobite-bearing Middle-Upper Cambrian succession.     

New trace fossils of termites (Insecta: Isoptera) from the late Eocene‐early Miocene of Egypt,and the reconstruction of ancient isopteran social behavior

Three new ichnogenera and five new ichnospecies are described for new trace fossils of termitaria (including associated gallery systems) of subterranean termites from upper Eocene through lower Miocene rocks of northern Egypt. All but two ichnospecies (Krausichnus trompitus, ichnogen, and ichnosp. nov., and A’, altus, ichnosp. nov.) show varying degrees of affinity to nests of the extant subterranean termite species Sphaerotermes sphaerothorax (Termitidae, Macro‐termitinae).

Termitichnus qatranii (Bown, 1982) is divided into two ichnospecies, the previously named T. qatranii, and a more generalized form, T. simplicidens, ichnosp. nov. Vondrichnus obovatus, ichnogen. and ichnosp. nov., is named for simple, possibly macrotermitine nests with oblate form, and Fleaglellius pagodus, ichnogen. and ichnosp. nov., records a nest form similar to V. obovatus, but one in which successive vertical growth by chamber apposition has produced a subterranean, tower‐like structure, reminiscent of that built by extant, epigeous Cubitermes. Krausichnus trompitus, ichnogen. and ichnosp. nov., and A’, altus, ichnosp. nov., record two unique nest architectures probably produced by unknown but related species of humivorous termites. The nest architecture expressed by Krausichnus is of wholly unknown affinity and is only distantly related in form to the ichnofossils of other termite nests known from the Tertiary of Egypt.

Study of the architecture of the nests of these ancient termites reveals details important in reconstructing the phylogeny of termite nests and documents two novel blueprints for chamber expansion and society budding in what were probably primitive Macrotermi‐tinae. It also indicates that at least two constructed edifices, earlier known only as epigeous manifestations of extant termite species, were almost certainly first developed by species living underground.

The extant Macrotermitinae are a subfamily of termites originally believed to have evolved in the post‐Eocene of the Ethiopian biogeo‐graphic region. We offer fossil evidence of four distinct, possibly macrotermitine structures from upper Eocene rocks. This evidence suggests that this termite subfamily (or at least their peculiar mode of nest construction) might have evolved considerably earlier, as it is already well established and exhibits several variants by the late Eocene in Egypt.     

The ichnofamily Celliformidae for Celliforma and allied ichnogenera

This contribution undertakes a comprehensive revision, lacking until now, of all the ichnotaxa attributed to fossil bee cells, one of the most common traces in paleosols. These ichnotaxa are morphologically related to Celliforma and consequently grouped herein in the new ichnofamily Celliformidae. Two new ichnogenera are formulated, Cellical‐ichnns igen. nov., for several ichnospecies previously included in Celliforma, and a new ichnospecies, C. chubutensis isp. nov., belonging to this ichnofamily, and Brownichnus igen. nov., for the ichnospecies favosites, which is not morphologically related to Celliforma. The ichnogenera included in this ichnofamily involve traces in which Celliforma, the most simple trace of the group, is part of their structure. Paltniraichnus is akin to Celliforma with antechambers and discrete walls. Uruguay, Corim‐batichnus and Rosellichnus are clusters composed of adjacent rows of Celliforma, or Palmiraichnus‐like cells. Ellipsoideichnus and Cellicalichnus are different arrangements of Celliforma‐like cells attached to tunnels.

Celliformidae are based on the morphology of the traces, all of them comprising cells, groups of cells and cells attached to tunnels. As such, its component ichnotaxa are based exclusively on morphological ichnotaxobases, which, in turn, were evaluated and selected with respect to the nest architecture of the trace makers, the bees. This paper analyzes some procedures and clues (i.e. behavioral homologies) that may be used to select the proper taxobases to erect ichnotaxa when the identity of the trace makers is known, as in this study case.     

CLEPTOPARASITISM AND DETRITIVORY IN DUNG BEETLE FOSSIL BROOD BALLS FROM PATAGONIA, ARGENTINA

Abstract:  Traces within traces is a new ichnological field that is meant to shed light on significative palaeoecological aspects. Dung beetle fossil brood balls ( Coprinisphaera ispp.), from the Middle Eocene – Lower Miocene Sarmiento Formation of Patagonia, Argentina, show two different trace fossils excavated in its infillings and/or wall that reveal the presence and relationships among different components of past dung communities. Tombownichnus pepei n. isp. is represented by elongated pits, circular to elliptical in cross-section, occurring in the centre or beside ovoid mounds in the internal surface of the Coprinisphaera wall. These traces record the activity of cleptoparasites, such as other dung beetles or flies, whose larvae were probably carried passively with the dung for provisions. Tombownichnus pepei would represent the pupation chambers excavated by full grown larvae in the Coprinisphaera wall after completing their development inside provisioned dung. The other trace fossil, Lazaichnus fistulosus is represented by circular to subcircular borings occurring in Coprinisphaera walls, in connection with an internal gallery in their infillings. Its connection also with meniscate burrows and chambers in the surrounding palaeosol attributable to aestivation chambers of earthworms revealed that these organisms would have been active cleptoparasites or detritivores in dung beetle fossil brood balls.     

Isotopic dietary reconstructions of Pliocene herbivores at Laetoli: Implications for early hominin paleoecology

Major morphological and behavioral innovations in early human evolution have traditionally been viewed as responses to conditions associated with increasing aridity and the development of extensive grassland-savanna biomes in Africa during the Plio-Pleistocene. Interpretations of paleoenvironments at the Pliocene locality of Laetoli in northern Tanzania have figured prominently in these discussions, primarily because early hominins recovered from Laetoli are generally inferred to be associated with grassland, savanna or open woodland habitats. As these reconstructions effectively extend the range of habitat preferences inferred for Pliocene hominins, and contrast with interpretations of predominantly woodland and forested ecosystems at other early hominin sites, it is worth reevaluating the paleoecology at Laetoli utilizing a new approach. Isotopic analyses were conducted on the teeth of twenty-one extinct mammalian herbivore species from the Laetolil Beds (∼ 4.3–3.5 Ma) and Upper Ndolanya Beds (∼ 2.7–2.6 Ma) to determine their diet, as well as to investigate aspects of plant physiognomy and climate. Enamel samples were obtained from multiple localities at different stratigraphic levels in order to develop a high-resolution spatio-temporal framework for identifying and characterizing dietary and ecological change and variability within the succession. In general, dietary signals at Laetoli suggest heterogeneous ecosystems with both C₃ and C₄ dietary plants available that could support grassland, woodland, and forested communities. All large-bodied herbivores analyzed yielded dietary signatures indicating mixed grazing/browsing strategies or exclusive reliance on C₃ browse, more consistent with wooded than grassland-savanna biomes. There are no clear isotopic patterns documenting shifting ecology within the Laetolil Beds or between the Laetolil and overlying Upper Ndolanya Beds, although limited data from the U. Ndolanya Beds constrains interpretations. Comparison of the results from Laetoli with isotopic enamel profiles of other African fossil and modern communities reveals significant differences in dietary patterns. Relative to extant taxa in related lineages, carbon isotopic ranges of a number of Laetoli fossil herbivores are anomalous, indicating significantly more generalized intermediate C₃/C₄ feeding behaviors, perhaps indicative of dietary niches and habitat types with no close modern analogs. Enamel oxygen isotope ranges of fossil taxa from Laetoli are consistently more ¹⁸O depleted than modern E. African herbivores, possibly indicating more humid conditions during that interval in the past. These data have important implications for reconstructing dietary trajectories of mammalian herbivore lineages, as well as the evolution of ecosystems in East Africa. Isotopic analyses of similar or related taxa at other hominin fossil sites yield signatures generally consistent with Laetoli, suggesting that mammalian communities in East Africa were sampling ecosystems with similar proportions of browse and grass. Collectively, the isotopic dietary signatures indicate heterogeneous habitats with significant wooded or forested components in the Laetoli area during deposition of the Laetolil and Upper Ndolanya Beds. Early hominin foraging activity in this interval may have included access to forest or woodland biomes within this ecosystem, complicating traditional interpretations linking early human evolutionary innovations with a shift to savanna habitats.     

Beyond Coprinisphaera: fossil nests of dung beetles

Forty‐two remains of fossil nests of dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae), which were recorded in four formations of the Cenozoic of South America, are described herein for the first time. Most of them are represented by nesting chambers containing a fossil brood ball (Coprinisphaera). However, in the most remarkable cases, parts of the burrows constructed by the parents and/or the vertical emergence burrows constructed by the offspring are preserved too. The preservation of the dung beetle nests is very unusual but more recurrent in younger formations probably because of the short‐term action of the diagenetic processes. Considering that the construction of brood balls always involves the construction of nests, the ichnotaxonomical proposal is that the remains of fossil nests should be considered as ‘structures associated with Coprinisphaera’ and added to the diagnosis to avoid the proliferation of names. The study of the fossil nests provides new palaeoetological inferences for dung beetles, such as the Nesting Pattern of the trace makers of Coprinisphaera tonnii and Coprinisphaera akatanka, and how phylogenetically related were they with the extant necrophagous species of the genera Coprophanaeus and Canthon, respectively. Additionally, the fossil evidence suggests that simple nests of dung beetles predate compound nests.     

New Miocene scarabeid and hymenopterous nests and early miocene (santacrucian) paleoenvironments,patagonian Argentina

Three new trace fossils are described from Miocene paleosols of southern Argentina. Celliforma pinturensis, n. ichnosp. and Celliforma rosellii, n. ichnosp. are interpreted as cells of digging bees, possibly Anthophoridae, and Coprinisphaerafrenguellii, n. ichnosp. are brood balls of dung‐beetles. Both burrowing bees and dung‐beetles are common nesters in relatively open areas, confirming previous reconstructions of the paleoenvironment of the Pinturas Formation. A brief review of scarabeid and bee fossil nests from South America is presented, and we propose that constructed nests have a higher preservation potential than excavated nests. This fact explains their more common occurrences as trace fossils in paleosols. A new ethological category, calichnia, is proposed for hymenopterous and coleopterous traces, in which adult individuals make nests exclusively for larvae.     

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.     

Evidence of Bioerosive Structures in Quaternary Glaciomarine Sediments from Southwestern Iceland

The Late Weichselian marine and glaciomarine sediments occurring in many places along the coast of Iceland are rich in invertebrate fossils, especially mollusks and barnacles. A diverse ichnofauna, constituted especially by bioerosional traces documenting the activity of predators and animals that use the host shell for attachment or the construction of dwellings, was identified at the localities of Ósmelur, Saurbær, and Brekkubakkar in southwestern Iceland. The ichnotaxa identified include Anellusichnus circularis (Santos, Mayoral, and Muñiz), Caulostrepsis isp., Centrichnus concentricus (Bromley and Martinell), Clionolithes isp., Finichnus peristroma (Taylor, Wilson, and Bromley), Finichnus isp., Sedilichnus asperus (Nielsen and Nielsen), Sedilichnus cf. excavatus (Donovan and Jagt), Sedilichnus gradatus (Nielsen and Nielsen), Sedilichnus smiley isp. nov., Sedilichnus ovalis (Bromley), Sedilichnus paraboloides (Bromley), Sedilichnus simplex (Bromley), and Sedilichnus spongiophilus (Müller).     

Late Jurassic Mammals from Tendaguru, Tanzania, East Africa

Records of Mesozoic mammals are extremely rare in Africa. The only previous record from the Upper Jurassic of Africa is a fragmentary mandible without teeth of Brancatherulum tendagurense. Here I report the discovery of two new mammals from the Upper Jurassic of Tendaguru, Tanzania. The fossils were recovered from the Middle Saurian Bed of the Tendaguru Series. A lower molar of a triconodontid mammal is described as Tendagurodon janenschi gen. et sp. nov., and a fragmentary dentary of a eupantothere as Tendagurutherium dietrichi gen. et sp. nov. The eupantothere in particular contributes to documenting the evolution of mammals during the Mesozoic. The posterior portion of the mandible of Tendagurutherium dietrichi gen. et sp. nov. shows that the angular (tympanic) bone was not yet completely separated from the dentary, a previously undocumented stage of eupantotherian middle ear evolution.     

The Oldest Beetle and Bee Ichnofossils from Mexico and their Paleoenvironmental Implications

ABSTRACT

Specimens of Fictovichnus gobiensis, Celliforma curvata, Celliforma rosellii, Celliforma germanica and Celliforma isp. were collected from paleosol horizons in a fluviolacustrine succession near Santiago Yolomécatl town, in northwestern Oaxaca, southern Mexico. These ichnofossils represent the oldest evidence of beetles and bees in Mexican localities. K–Ar ages and the record of the equid Miohippus assiniboiensis, an index fossil, indicate that the age of the deposits is late Eocene. Based on the presence of the insect ichnofossils, pedogenic carbonate isotopes and mammal proxies, the inferred type of vegetation present in the study area was scrubland/woodland, within subhumid to subarid conditions.     

新疆吐鲁番盆地桃树园晚二叠世非海相腹足类

提要新疆吐鲁番盆地桃树园地区晚二叠世地层称作下仓房沟群,自下而上分为泉子街组、梧桐沟组和锅底坑组,为河–湖相碎屑沉积,剖面连续,层序清楚,动、植物化石丰富。文中研究的腹足类标本产于桃东沟剖面梧桐沟组下部和中部的介壳灰岩层和灰岩透镜体中,见有2层,计有2科4属6种:Xinjiangospira rotundata Yu et Zhu,Xinjiangospira habita sp.nov.、Hydrobia turpanensis Wei、Hydrobia orientalis sp.nov.、Pseudamnicola taodonggouensis sp.nov.和Valvata complanusa sp.nov.;与腹足类共生的有双壳类、叶肢介、介形类,以及植物和脊椎动物化石等,其中双壳类主要是Palaeanodonta,Palaeomutela和Anthraconauta等属。该腹足动物群由Hydrobiidae和Valvatidae的属种组成,标本数量多,壳体小,保存完好,是迄今已知属种最丰富的古生代淡水腹足类动物群。当前梧桐沟组腹足动物群面貌与准噶尔盆地大龙口剖面小龙口组...     

Upper Silurian microplankton of the Leintwardine Group, Ludlow Series, in the type Ludlow area and adjacent regions

The acritarchs and prasinophyte algae from the Upper Bringewood, Lower Leintwardine, Upper Leintwardine and Lower Whitcliffe formations of the Ludlow type area and surrounding regions are described. The following new taxa are proposed: Cymatiosphaera pumila sp. nov., Melikeriopalla pustula sp. nov., Cheleutochroa beechenbankensis sp. nov., Cymbosphaeridium molyneuxii sp. nov., Flammulasphaera bella gen. et sp. nov., Percultisphaera incompta sp. nov., Salopidium aldridgei sp. nov. and Umbellasphaeridium? wicanderi sp. nov. A holotype is defined for Lophosphaeridium galeatum Hill and a further 34 new taxa are described under open nomenclature. Two biozones, identified by the first appearance of the zone taxon, are defined in the Sunnyhill section, Ludlow [Global Stratotype Section and Point (GSSP) for the base of the Ludfordian Stage, Ludlow Series]. The base of the Leoniella vilis Biozone is identified at 4.52m below the top of the Upper Bringewood Formation at Beechenbank, Aymestrey. The base of the Triangulina sanpetrensis Biozone is identified at 15.49m above the base of the Lower Leintwardine Formation and Ludfordian Stage at the Sunnyhill section. These biozones allow correlation with sections in north–west Spain, Podolia and Gotland. The possibility of using Visbysphaera whitcliffense and U.? wicanderi as biozonal indicators is suggested.     

Late Norian (Triassic) Radiolaria from the Kotel’nyi Island (New Siberian Islands,Arctic, Russia)

Two radiolarian assemblages were recovered from upper Norian strata of the Kotel’nyi Island (Russia); the first assemblage, from the Monotis zabaikalica Subzone (lower part of Upper Norian), is represented by Betraccium inornatum Blome, Dumitricaella (?) parva Sugiyama, Ferresium titulense Blome and 24 other species; the second assemblage, from the Monotis subcircularis Subzone (upper part of Upper Norian), is represented by Crucella sp. cf. C. angulosa Carter, Kahlerosphaera acris Bragin, K. sp. cf. K. parvispinosa Kozur & Mostler, Pseudohagiastrum crassum (Carter) and 11 other species. Both assemblages have common taxa with Upper Norian and Rhaetian radiolarian assemblages of British Columbia and they display clear Boreal features: low taxonomic diversity, abundance of taxa known from high-latitude regions, absence or rare presence of taxa known from low-latitude areas. The presence of early representatives of nassellarian genera Droltus and Parahsuum is very distinctive. Six new species are described: Pseudohagiastrum spinosum nov. sp., Cantalum boreale nov. sp., Plafkerium carteri nov. sp., Droltus guttaeformis nov. sp., Laxtorum blomei nov. sp., L. glacialis nov. sp.     

The earliest burst of necrophagous dung beetles in South America revealed by the Cenozoic record of Coprinisphaera

New ichnological, sedimentological and palaeobotanical information from a Miocene palaeosol succession from Patagonia bearing abundant fossil brood balls of dung beetles (Coprinisphaera) allow inferring novel aspects of the evolutionary history, biology and feeding habits of Scarabaeinae, along with the palaeoenvironmental and palaeoclimatic conditions in which they nested. Coprinisphaera tonnii and Coprinisphaera akatanka, both attributed to necrophagous dung beetles, represent 40.2% of the specimens. Considering their scarce or null record in older Cenozoic units from South America, these high values reveal that the first burst of necrophagous Scarabaeinae took place during the Miocene. Some Coprinisphaera preserve characters that indicate the developmental stage of the ball occupant and the adult emergence success. Both ichnospecies show the egg chamber isolated from the provision chamber, which was associated with a higher preference of cleptoparasites for necrophagous balls. The comparison among the abundance of traces of nest intruders in Coprinisphaera attributed both to necrophagous and to coprophagous beetles from different units of South America indicates that the presence of such intruders would be independent of the nature of the organic matter contained within the balls. Phytolith analyses performed in Coprinisphaera and extant necrophagous and coprophagous balls indicate that the comparison between the relative abundance of phytoliths in the wall of the brood ball, their infilling and the bearing palaeosol is a useful tool for inferring the feeding behaviour of the trace makers and support the attribution of Coprinisphaera tonnii and Coprinisphaera akatanka to necrophagous Scarabaeinae. Sedimentary and palaeosol analyses indicate that the beetles would have nested in well-drained soils showing mollic-like features, in grass-dominated habitats, under a seasonal sub-humid, temperate–warm climate.