Three-dimensional Preservation Research Articles

Paleocene and Eocene deep-water benthic foraminifera at IODP Site U1511, Tasman Sea: Part 2

Deep-water benthic foraminifera are investigated from Paleocene to Eocene sediments recovered from IODP Hole U1511B in the northeastern Tasman Sea. The recovered foraminifera display exceptional three-dimensional preservation: they are relatively unaltered by sediment diagenesis and compaction. We examined 33 samples from Cores U1511B-33R to -42R, and recovered 79 species of deep-water agglutinated foraminifera (DWAF) and 13 species of calcareous benthic foraminifera. The assemblage consists entirely of “cosmopolitan” forms originally described from the Carpathians, Caucasus, Trinidad, and the western Tethys, and other DSDP/ODP sites, implying that there is little or no endemism among deep-water benthic faunas in the semi-isolated Tasman Sea. The Paleocene–Eocene interval is characterized by successive acmes of Spiroplectammina, Trochammina, ammodiscids, and Karrerulina, and therefore bears striking similarity to previously studied sections in the western Tethys and Boreal North Atlantic. The taxonomic turnover among the DWAF is minor across the P/E contact, with only three apparent extinctions and three originations from the uppermost cores of the Paleocene and the lowermost cores of the Eocene. Calcareous benthic foraminifera were not observed in the Eocene samples, possibly a result of the shoaling of the CCD.

Pachycormid fish fed on octobrachian cephalopods: new evidence from the ‘Schistes bitumineux’ (early Toarcian) of southern Luxembourg

A re-examination of the early Toarcian fish fossils preserved in public paleontological collections in Luxembourg revealed 70 specimens of large Toarcian pachycormid fish with an excellent three-dimensional preservation within calcareous nodules. Six of them are associated with octobrachian coleoid gladii in their oesophagus or stomach, an association not previously described from Luxembourg. The pachycormids are ascribed to Pachycormus macropterus (Blainville, 1818) and Saurostomus esocinus Agassiz, 1843 while the octobrachian gladii are ascribed to Teudopsis bollensis Voltz, 1836, Teudopsis sp. indet. and Loligosepiidae indet. The position and orientation of the gladii provide direct evidence of these fishes feeding on coleoids and thus a teuthophagous diet, rather than an accidental joint burial. Together with evidence from coeval deposits in Germany, these findings suggest that teuthophagy was a widespread feeding strategy at the base of the clade that contains the suspension-feeding pachycormid giants of the Jurassic–Cretaceous.

A new species of possible archipolypodan millipedefrom the Carboniferous of the Netherlandswith unusually long tergites

Millipedes have a long evolutionary history, with the oldest presumed fossils of Diplopoda de Blainville in Gervais, 1844 being from the Silurian and the first definite fossil record originating from Devonian deposits. The phylogeny of Diplopoda is not fully resolved yet, especially not concerning fossil representatives. At the same time, already in the Palaeozoic millipedes showed quite a morphological and presumably also ecological variety. We describe here a new species of a Carboniferous millipede from the Westphalian A of the Netherlands, Lauravolsella willemeni n. gen., n. sp., a possible representative of Archipolypoda (†Archipolypoda Scudder, 1882). The species is based on a single specimen, preserved with part and counterpart, which both show a three-dimensional preservation. The specimen has unusually long tergites, in normal life position covering most of the following segment. These long tergites might have been beneficial when performing defensive enrolling. In extant millipedes, enrolling is usually facilitated by softer areas between the sternites, allowing for a certain degree of ventral compression. In the new fossil, the sclerotic sternites occupy the entire length of the ventral side of the segment, not allowing for any type of compression. The new fossil therefore demonstrates another solution for the mechanical challenges during enrolment and increases the morphological diversity of Carboniferous millipedes.

Taphonomy of non-biomineralized trilobite tissues preserved as calcite casts from the Ordovician Walcott-Rust Quarry, USA

Trilobites with appendages from the Rust Formation of New York State were discovered in the 1870s and represent one of the earliest known cases of exceptional preservation of non-biomineralized tissues. The Rust Formation trilobites feature three-dimensionally preserved walking legs and delicate respiratory lamellae, but the mechanism behind their fossilization remains unknown. Here we show that after burial, carcass decay produced framboidal pyrite, while fibrous calcite precipitated on the visceral side of the body, followed by the widespread formation of sparry calcite crystal replicating non-biomineralized morphological features. Trilobites and co-occurring calcite veins show no chemical or petrographic differences, rejecting the hypothesis that exceptional preservation was caused by a local microenvironment within enrolled trilobites. These results suggest that fine-grained sediment provided support for the appendages and facilitated their fossilization through calcite replacement. Our findings carry broader implications for understanding the exceptional three-dimensional preservation of animal Paleozoic body fossils through calcite casts.

Investigating Mazon Creek fossil plants using computed tomography and microphotography

More than 20,000 siderite concretions from the Mazon Creek area of northern Illinois, United States are housed in the paleobotanical collections of the Field Museum. A large proportion contain fossil plants of Middle Pennsylvanian age that often have excellent three-dimensional morphology and sometimes anatomical detail. Approximately eighty plant taxa have been recognized from the Mazon Creek Lagerstätte, but few have been studied in detail, and in some cases the systematic affinities of these fossils need reevaluation. The three-dimensional (3D) preservation of Mazon Creek fossil plants makes them ideal candidates for study using x-ray micro-computed tomography (μCT), and here we apply these techniques to more accurately reconstruct the morphology of specimens of Tetraphyllostrobus Gao et Zodrow and Crossotheca Zeiller. The mineralogical composition of the fossil plant preservation was studied using elemental maps and Raman spectroscopy. In-situ spores were studied with differential interference contrast, Airyscan confocal super-resolution microscopy, and scanning electron microscopy, which reveal different features of the spores with different degrees of clarity. Our analyses show that μCT can provide excellent detail on the three-dimensional structure of Mazon Creek plant fossils, with the nature of associated mineralization sometimes enhancing and sometimes obscuring critical information. Results provide guidance for selecting and prioritizing fossil plant specimens preserved in siderite concretions for future research.

Preservation of Soft-Bodied Organisms in Early Cambrian Carbonates

Abstract —Preservation of soft-bodied organisms as casts is common in the Ediacaran fossil record and extremely rare in Cambrian rocks. Among the factors, limiting fossilization of soft tissues, there is bioturbation–sediment disturbance by burrowing organisms. It is the emergence of burrowing metazoans and associated sediment bioturbation that is thought to be one of the major causes for the disappearance of Ediacaran soft-bodied organisms from the fossil record around ~540 Ma. Here, we study an assemblage of fossils preserved as casts in the Fortunian carbonates of the Olenek Uplift (northeastern Siberian Platform) in association with a typical Fortunian ichnoassemblage. The overall morphology and preservation of the fossils reveal that they comprise microbially induced sedimentary structures and soft-bodied holdfasts. The latter are vaguely reminiscent of some Ediacaran holdfasts, though it is unlikely that they include their phylogenetic descendants. Three-dimensional preservation of the studied fossils was caused by authigenic crystallization of calcite and its further early-diagenetic dolomitization. Our study confirms the critical importance of specific environmental conditions ensuring preservation of soft-bodied organisms as three-dimensional molds and casts. This unique interplay of environmental factors became rare in the Cambrian Period, which was caused by intensification and expansion of bioturbation in marine basins.

SURVEY PLANNING FOR DOCUMENTATION OF A MONUMENT FOR THE UNDERSTANDING, PRESERVATION AND RESTORATION

Abstract. The three-dimensional (3D) preservation and repair of historic sites are increasingly in practice using modelling and digital documentation. This study focuses on replacing conventional techniques of historical documentation by creating a digital documentation procedure employing laser scanning for 3D mapping of a monument located in Prayagraj, India. To quickly record the entire monument structure, four scanning stations were planned, where three for the facades and one for the interior. A 3D structure of the monument and its elements dimension that included structural, architectural, historical, and non-engineering information was the end product. Researchers, architects, and conservationists can use this laser scanning-based technique to analyze data in great detail to identify weaknesses and conservation requirements. In order to preserve the monument's cultural relevance, it can also be used for virtual tours. Digital documentation can also provide an accurate monument record for restoration needs, protecting the monument from human- or natural-caused damage. Overall, 3D Modelling and digital documentation are valuable tools in heritage conservation, providing comprehensive records of heritage sites and aiding in practical conservation and restoration plans while making cultural heritage accessible to a broader societies.

Three-dimensional anatomy of the early Eocene Whitephippus (Teleostei, Lampriformes) documents parallel conquests of the pelagic environment by multiple teleost lineages

ABSTRACT The early Eocene fossil assemblage of the London Clay (Southeastern England) is a key window to the early Paleogene diversification of teleost fishes in the open ocean. Despite their three-dimensional preservation that offers unique insight into skeletal anatomy, the London Clay fossils are still poorly described for the most part. †Whitephippus tamensis is a fossil teleost from this assemblage, known by several well-preserved specimens. Based on a complete description of the known material, including previously hidden structures (braincase, hyoid, and branchial arches) revealed through 3D microtomography, we reinterpret †Whitephippus as an early member of the teleost group Lampriformes. More specifically, the anatomy of †Whitephippus indicates that it is likely a member of the so-called ‘pelagic clade’ including modern opahs and oarfishes. This redescription of †Whitephippus provides the earliest definitive evidence of lampriforms conquering the pelagic environment, alongside numerous other teleost lineages.

Cretaceous pollen cone with three-dimensional preservation sheds light on the morphological evolution of cycads in deep time.

The Cycadales are an ancient and charismatic group of seed plants. However, their morphological evolution in deep time is poorly understood. While molecular divergence time analyses estimate a Cretaceous origin for most major living cycad clades, much of the extant diversity is inferred to be a result of Neogene diversifications. This leads to long branches throughout the cycadalean phylogeny that, with few exceptions, have yet to be rectified by unequivocal fossil cycads. We report a permineralized pollen cone from the Campanian Holz Shale located in Silverado Canyon, CA, USA (c. 80 million yr ago). This fossil was studied via serial sectioning, SEM, 3D reconstruction and phylogenetic analyses. Microsporophyll and pollen morphology indicate this cone is assignable to Skyttegaardia, a recently described genus based on disarticulated lignitized microsporophylls from the Early Cretaceous of Denmark. Data from this new species, including a simple cone architecture, anatomical details and vasculature organization, indicate cycadalean affinities for Skyttegaardia. Phylogenetic analyses support this assignment and recover Skyttegaardia as crown-group Cycadales, nested within Zamiaceae. Our findings support a Cretaceous diversification for crown-group Zamiaceae, which included the evolution of morphological divergent extinct taxa with unique traits that have yet to be widely identified in the fossil record.

Internal anatomy of a fossilized embryonic stage ofthe Cambrian-Ordovician scalidophoran Markuelia.

The Wangcun fossil Lagerstätte in Hunan, South China, has yielded hundreds of fossilized embryos of Markuelia hunanensis representing different developmental stages. Internal tissues have only rarely been observed, impeding further understanding of the soft tissue anatomy, phylogenetic affinity and evolutionary significance of Markuelia. In this study, we used synchrotron radiation X-ray tomographic microscopy (SRXTM) to study a new collection of fossil embryos from the Wangcun fossil Lagerstätte. We describe specimens exhibiting a spectrum of preservation states, the best of which preserves palisade structures underneath the cuticle of the head and tail, distinct from patterns of centripetal mineralization of the cuticle and centrifugal mineralization of hypha-like structures, seen elsewhere in this specimen and other fossils within the same assemblage. Our computed tomographic reconstruction of these mineralization phases preserves the gross morphology of (i) longitudinal structures associated with the tail spines, which we interpret as the proximal ends of longitudinal muscles, and (ii) a ring-shaped structure internal to the introvert, which we interpret as a ring-shaped brain, as anticipated of the cycloneuralian affinity of Markuelia. This is the first record of a fossilized nervous system in a scalidophoran, and the first instance in Orsten-style preservation, opening the potential for further such records within this widespread mode of high-fidelity three-dimensional preservation.

Nutrient capsules maintain tear film homeostasis for human corneal lenticule transplantation

Corneal lenticule transplantation can be applied in the correction of refractive errors and various corneal diseases. However, current techniques to preserve cornea are limited due to their short preservation time and the structures and optical retention of cornea tissues. Inspired by the composition of the natural human tear film, a nutrient hydrogel capsule that could maintain the homeostasis of tear film was constructed for preserving human corneal lenticules in this study. The nutritional preservation solution containing sodium alginate was coordinated with calcium ions by rapid diffusion to form a stereoscopic capsule, in which chondroitin sulfate interacted with the corneal lenticule surface electrostatically to form a lubricating layer for three-dimensional preservation and nutritional supply. The light transmittance, cell activity, and collagen fiber density of human corneal lenticules after mid-term preservation in nutrient capsules were higher than those traditional preservation methods. Furthermore, after human allogeneic transplantation, the implanted lenticules remained transparent without displacement or graft rejection, and the corrected distance visual acuity increased by two or more lines in 70% of the patients’ operated eyes after surgery. Nutrient capsules could provide not only a simple, safe, and controlled strategy for three-dimensional preservation, but also new approach for other living tissues preservation.

A Permian fish reveals widespread distribution of neopterygian-like jaw suspension

The actinopterygian crown group (comprising all living ray-finned fishes) originated by the end of the Carboniferous. However, most late Paleozoic taxa are stem actinopterygians, and broadly resemble stratigraphically older taxa. The early Permian †Brachydegma caelatum is notable for its three-dimensional preservation and past phylogenetic interpretations as a nested member of the neopterygian crown. Here, we use computed microtomography to redescribe †Brachydegma, uncovering an unanticipated combination of primitive (e.g., aortic canal; immobile maxilla) and derived (e.g., differentiated occipital ossifications; posterior stem of parasphenoid; two accessory hyoidean ossifications; double jaw joint) dermal and endoskeletal traits relative to most other Paleozoic actinopterygians. Some of these features were previously thought to be restricted to the neopterygian crown. The precise phylogenetic position of †Brachydegma is unclear, with placements either on the polypterid stem or as an early-diverging stem neopterygian. However, our analyses decisively reject previous placements of †Brachydegma in the neopterygian crown. Critically, we demonstrate that key endoskeletal components of the hyoid portion of the suspensorium of crown neopterygians appeared deeper in the tree than previously thought.

Upside-down in volcanic ash: crown reconstruction of the early Permian seed fern Medullosa stellata with attached foliated fronds.

Our understanding of fossil floras through geological time is mainly based on various differently preserved plant parts, often found isolated under restricted taphonomic circumstances. Preservation of whole plants is exceptionally rare in the geological record but provides the most reliable proof of ancient plants, especially those lacking a nearest living relative like the late Paleozoic pteridosperms. Among them, the medullosan seed ferns represent the largest and most essential systematic group. Medullosaleans are well known from the large Euramerican tropical swamp forests of Carboniferous age, but also from seasonally dry, clastic-soil habitats of Late Pennsylvanian–early Permian intramontane basins in central Europe. An exceptional taphonomic window that offered three-dimensional preservation of early Permian plants is located in Chemnitz, eastern Germany. Here, a forest ecosystem had been buried by pyroclastic deposits in a geological instant, 291 ± 2 Ma ago. Medullosaleans are both abundant and diverse in this striking autochthonous assemblage. The upper part of a Medullosa stellata var. typica individual broke at its top resulting from the overload of volcanic ash and was buried upside-down in the basal pyroclastics. The tree crown consists of the anatomically preserved apical stem, ten attached Alethopteris schneideri foliated fronds with Myeloxylon-type petioles and rachises. Though already discovered during the scientific excavation in 2010, the remarkable find required several years of preparation work, documentation, and reconstruction. The fronds were up to 3.5 m long, bifurcating, and bore numerous bipinnately compound pinnae preserved pulvinated in life position. The apical stem vascular system consists of a cylindrical peripheral vascular segment and up to 87 central accessory strands, each surrounded by manoxylic wood and secondary phloem. The reconstructed tree is supposed to have been self-supporting and of slender stature. Its architectural model is comparable to modern tree ferns or cycads and adapted to have grown in light-deficient lower-storey (sub-) tropical forests, usually not exceeding 15 m in height. Apical meristematic growth dominated, whereas only minor secondary growth occurred during ontogenesis. The densely attached frond bases followed a 3/8 phyllotaxis and were most likely abscised shortly after becoming photosynthetically inactive. A high water-conducting potential is assumed due to the tree’s cauline, petiolar and leaf vascular anatomies. Concerning the extensive leaf surface of the densely foliated fronds, considerable transpiration is hypothesised. Alethopteris schneideri foliage is stratigraphically significant for lower Permian (Asselian–Sakmarian) continental strata of central Europe, preferring habitats of wet clastic soils in sub-humid, seasonal palaeoclimate. The new insights provide a substantial step towards the first whole-plant concept of intramontane medullosaleans.

Geological Characteristics of Deep Shale Gas in the Silurian Longmaxi Formation in the Southern Sichuan Basin, China

To investigate the geological characteristics and exploration potential of shale gas in the southern Sichuan Basin, we analyze the coupling relationship between the hydrocarbon generation and storage conditions of the Longmaxi Formation and discuss the preservation conditions from the lateral and vertical migration mechanisms of shale gas. According to the results, the organic-rich shale at the bottom was formed in a strongly euxinic and anoxic reducing environment, the oxygen content increased in the upper water body in which the Longmaxi Formation was deposited, and the water body became oxidized. The organic matter type in the shale is dominantly type I kerogen and a small amount of type II1 kerogen. The organic matter content is more than 3.0% and is in the high-to postmature stage. The enrichment of siliceous organisms increases the organic matter and enhances the brittleness of shale, resulting in “superior hydrocarbons and a favorable reservoir”. Pyrolysis of organic matter promotes the formation of organic matter pores and dissolution pores, resulting in a coupled “source-reservoir” accumulation control system. The high vertical formation pressure guarantees the sealing of shale and restrains the lateral escape of shale gas. The high-angle intersection of the highly filled fractures and the current crustal stress can effectively enhance the fracture sealing and inhibit the vertical escape of shale gas, forming a three-dimensional effective closure. Hence, the area featuring a short tectonic uplift time, small amplitudes, large-scale underdeveloped fractures, and a high formation pressure coefficient is a favorable area for shale gas exploration, according to the analysis of three-dimensional preservation conditions.

The Paleocene of IODP Site U1511, Tasman Sea: A lagerstatte deposit for deep-water agglutinated foraminifera

Deep-water agglutinated Foraminifera (DWAF) are investigated from Paleocene sediments recovered from IODP Hole U1511B in the northeastern Tasman Sea. The recovered foraminifera display exceptional three-dimensional preservation: they are relatively unaltered by sediment diagenesis and compaction.We examined 27 samples fromCores U1511B-45R to -47R, and recovered over 70 species of DWAF. The assemblage consists entirely of “cosmopolitan” forms originally described from the Carpathians, Caucasus, Trinidad, and the western Tethys, implying that there is no provinciality among DWAF faunas in the world ocean.

Preservation of erniettomorph fossils in clay-rich siliciclastic deposits from the Ediacaran Wood Canyon Formation, Nevada: Ediacaran preservation in sandstone beds

Author(s): Hall, JG; Smith, EF; Tamura, N; Fakra, SC; Bosak, T | Abstract: Three-dimensionally preserved Ediacaran fossils occur globally within sandstone beds. Sandy siliciclastic deposits of the Ediacaran Wood Canyon Formation (WCF) in the Montgomery Mountains, Nevada, contain two fossil morphologies with similar shapes and sizes: one exhibits mm-scale ridges and a distinct lower boundary and the other is devoid of these diagnostic features. We interpret these as taphomorphs of erniettomorphs, soft-bodied organisms with uncertain taxonomic affinities. We explore the cast-and-mould preservation of both taphomorphs by petrography, Raman spectroscopy, X-ray fluorescence microprobe and X-ray diffraction. All fossils and the surrounding sedimentary matrix contain quartz grains, iron-rich chlorite and muscovite. The ridged fossils contain about 70% larger quartz grains compared to the ridgeless taphomorph, indicating a lower abundance of clay minerals in the ridged fossil. Chlorite and muscovite likely originated from smectite and kaolinite precursors that underwent lower greenschist facies metamorphism. Kaolinite and smectite are inferred to have been abundant in sediments around the ridged fossil, which enabled the preservation of a continuous, distinct, clay- and kerogen-rich bottom boundary. The prevalence of quartz in the ridged fossils of the WCF and in erniettomorphs from other localities also suggests a role for this mineral in three-dimensional preservation of erniettomorphs in sandstone and siltstone deposits.

Preservation of erniettomorph fossils in clay-rich siliciclastic deposits from the Ediacaran Wood Canyon Formation, Nevada.

Three-dimensionally preserved Ediacaran fossils occur globally within sandstone beds. Sandy siliciclastic deposits of the Ediacaran Wood Canyon Formation (WCF) in the Montgomery Mountains, Nevada, contain two fossil morphologies with similar shapes and sizes: one exhibits mm-scale ridges and a distinct lower boundary and the other is devoid of these diagnostic features. We interpret these as taphomorphs of erniettomorphs, soft-bodied organisms with uncertain taxonomic affinities. We explore the cast-and-mould preservation of both taphomorphs by petrography, Raman spectroscopy, X-ray fluorescence microprobe and X-ray diffraction. All fossils and the surrounding sedimentary matrix contain quartz grains, iron-rich chlorite and muscovite. The ridged fossils contain about 70% larger quartz grains compared to the ridgeless taphomorph, indicating a lower abundance of clay minerals in the ridged fossil. Chlorite and muscovite likely originated from smectite and kaolinite precursors that underwent lower greenschist facies metamorphism. Kaolinite and smectite are inferred to have been abundant in sediments around the ridged fossil, which enabled the preservation of a continuous, distinct, clay- and kerogen-rich bottom boundary. The prevalence of quartz in the ridged fossils of the WCF and in erniettomorphs from other localities also suggests a role for this mineral in three-dimensional preservation of erniettomorphs in sandstone and siltstone deposits.

Taphonomy of lacustrine fish fossils of the Parnaíba Basin, northeastern Brazil: Spatial and causative relations of Konservat Lagerstätten in West Gondwana during Jurassic-Cretaceous

Fossil bonanzas called Konservat Lagerstätten provide geological windows to the ecology of ancient life communities. These deposits often occur as cluster of fossil localities with similar geographic locations, geological ages and facies, as the fossils were generally preserved during special periods and large-scale events in Earth history within environments with exceptional taphonomic conditions. This work applied petrological and geochemical analyses to evaluate the taphonomic pathway of a Konservat Lagerstätten in the Muzinho Shale, Upper Jurassic-Lower Cretaceous of the Parnaíba Basin, northeastern Brazil. These lacustrine deposits include interbedded limestones and fossiliferous black shales deposited in anoxic, non-euxinic and saline environment. Black shales contain articulated skeletons of fish fossils, characterised by three-dimensional preservation of the skeletal tissue. The polytypical assemblage includes diverse ontogenetic stages and is concentrated in a specific stratigraphic level. Mass mortality was triggered by anoxia probably established by stratified water column, high C flux and thermocline demise. The specimens are encased in kerogen-bearing siliciclastic laminae and cementstone microfacies, both microbially-induced. The fossils were cemented by eodiagenetic poikilotopic calcite that filled the skeletal articulation, including bone trabeculae and voids originally filled by bone marrow. The short distance, similar fossil assemblages, ages and coincident palaeoenvironments suggest that Muzinho Shale (Parnaíba Basin) and Crato Formation (Araripe Basin) compose a cluster of exceptional preservation in West Gondwana. Lava flows and outgassing related to the Central Atlantic Magmatic Province (CAMP) resulted in the subaereal exposure of volcanic plains, acid rains and subsequent CO2 greenhouse crisis. CAMP-magmatism was especially voluminous in northern Brazilian basins, enhancing nutrient-rich waters and lacustrine eutrophication due to intensified chemical weathering in this area. Probably, other Konservat Lagerstätten remain unreported in West Gondwana. Despite similar causative events, the individual taphonomic pathways of Muzinho Shale and Crato Formation are very different, suggesting that in these cases local phenomena affected authigenesis and diagenesis.

A new species of large-sized pterosaur from the Maastrichtian of Transylvania (Romania)

The Upper Cretaceous “Hațeg Island” is not only renowned mostly for its peculiar assemblage of dwarf dinosaurs, but also for its Azhdarchidae pterosaurs including giant specimens (e.g. Hatzegopteryx thambema) and medium-sized ones (e.g. Eurazhdarcho langendorfensis), discovered in the Hațeg and Transylvanian basins of Romania. Here, we report a new species of azhdarchid pterosaur, Albadraco tharmisensis gen. et sp. nov. The material refers to two well-preserved “beak” fragments and a cervical vertebra. These fossils were discovered together in a fossil assemblage in the Maastrichtian Şard Formation, located in the southwestern area of the Transylvanian Basin (Alba County, Romania). The association of a premaxilla and a mandibular symphysis from the same specimen is the first ever reported in Europe. The fourth cervical exhibits the best three-dimensional preservation of any azhdarchid mid-cervical vertebra in Transylvania, as all specimens reported previously are poorly preserved (e.g. in E. langendorfensis) or incomplete like the specimen from Pui with a broken condyle. Albadraco tharmisensis represents a new species of a large–sized azhdarchid from the “Hațeg Island”. Its size fits between that of E. langendorfensis and H. thambema, hence confirming the co-existence of medium, large, and giant–sized azhdarchids during the Maastrichtian in Transylvania. The possibility of Ad. tharmisensis being a young Hatzegopteryx is also discussed.

Earliest record of exophytic insect oviposition on plant material from the latest Pennsylvanian (Gzhelian, Stephanian C) of the Saale Basin, Germany

The Late Palaeozoic fossil record of plant-insect interactions contains few records of oviposition. Here we report the earliest evidence of exophytic oviposition, where eggs are laid directly on the outer surface of plants. The fossil structures consist of small, circular impressions on plant fossils from the late Pennsylvanian (Gzhelian, Stephanian C) of the Saale Basin in Germany. Their interpretation as exophytic insect eggs is supported by the oviposition pattern, their three-dimensional preservation and the undamaged nature of the plant host tissue. The tight arrangement of the eggs in curved arcs suggests a producer with a short abdomen such as roachoids, the ancestral group of the Dictyoptera sensu stricto. A diagnostic feature of roachoids is their long ovipositor, which was well suited for deposition of single eggs in a distinct pattern.