A new specimen of Ichthyosaurus anningae from Asturias (Northern Spain) is described in detail, representing the first unambiguous record of this species outside the type locality Dorset (UK). The associated ammonite assemblage firmly constrains the stratigraphic position of the specimen within the Rodiles Formation, Jamesoni Zone (Pliensbachian, Lower Jurassic). It aligns the Asturian faunal succession with coeval northwestern European basins. The new material preserves diagnostic cranial and postcranial features that refine the morphological characterization of I. anningae, including previously undocumented details of the skull anatomy. Based on the length of the humerus, the specimen described here represents the largest known individual of I. anningae. Incorporation of the specimen into an updated phylogenetic dataset supports the monophyly of Ichthyosaurus and reinforces the distinctiveness of I. anningae relative to other Pliensbachian taxa. The co-occurrence of I. anningae and Leptonectes sp. in the Rodiles Formation provides new evidence for niche partitioning among Early Jurassic ichthyosaurs in the Asturian Basin, paralleling patterns documented in localities in southwestern England. Taken together, these discoveries extend the paleobiogeographic range of Early Jurassic ichthyosaurs, provide new insights into the diversity and complexity of the faunas of the Iberian margin, and offer further evidence of the close affinities of marine communities in Northern Spain with those of the Jurassic subboreal sea of the United Kingdom.
Dimetropus is one of the most widespread tetrapod ichnogenera of the Permian, known from several Cisuralian units of Euramerica and North Africa. Generally attributed to pelycosaur-grade synapsid trackmakers, two ichnospecies are currently considered valid: D. leisnerianus and D. osageorum. A revision of a well-preserved Dimetropus pes footprint from the lower Permian (upper Artinskian) Rabéjac Formation of the Lodève Basin highlights the first definitive presence of D. osageorum from the Variscan Belt. A synapomorphy-based track–trackmaker correlation suggests ophiacodontids, such as Ophiacodon, as potential tracemakers of D. osageorum. Therefore, this potentially extends the palaeobiogeographic distribution of ophiacodontids in the upper Cisuralian in both North American and European basins. Moreover, the presence of well-preserved epidermal scale imprints on the D. osageorum pes of Lodève precises the evolution of epidermal scalation in pelycosaur-grade synapsids and suggests the presence of both dermal and epidermal scales in ophiacodontids. This occurrence is time-equivalent with the Artinskian Warming Event, strengthening the scenario of epidermal scale evolution in light of palaeoclimatic changes in the Variscan Belt, from everwet-adapted biotas to seasonal drought-adapted biotas.
In the fields of functional morphology, paleoecology, and taxonomy, the Bryozoan Skeletal Index (BSI) is an increasingly useful metric for the degree of calcification in bryozoan colonies. The goal of this study is to quantify how it varies from proximal to distal through a colony’s growth transect. Colony fragments of the stenolaemate trepostome bryozoan Leptotrypella asterica from New York State, USA were reassembled into three large colonies. They are from the Kashong Shale Member of the Moscow Formation of the Hamilton Group deposited during the Givetian, Middle Devonian. Colonies were serially sectioned transversely from proximal to distal along the growth axis of the longest branches. To calculate BSI, three characters were measured: exozone width, zooecial wall thickness, and maximum autozooecial apertural diameter at the branch surface. Exozone width decreased significantly from proximal to distal through each colony’s growth transect. Zooecial wall thickness did not vary significantly. Maximum autozooecial apertural diameter decreased significantly in some of the colonies. BSI decreased significantly distally from their bases. We interpreted the distal colony branch tips with lower BSI values as being weaker and more susceptible to breakage. They could more easily break off and regrow without jeopardizing the success of the entire colony had the stronger proximal base with its higher BSI values broken.
We provide a detailed description of the dental morphology of Gobiconodon zofiae based on the holotype and clarify its diagnostic features in comparison with other species in the genus. We also report a new species, Gobiconodon gongzhulingensis sp. nov., from the lower Upper Cretaceous Quantou Formation in Gongzhuling City, Jilin Province, based on a maxilla with five molariforms. The dental morphologies of both species allow us to reassess tooth assignments in Gobiconodon and support a dental formula of 2.1.3.4/2.1.3.5 for the genus. The molariforms of Gobiconodon exhibit a distinct type of tooth widening that is achieved primarily through cusp inflation, without pronounced cusp rotation or the addition of new cusps. In addition to the differences in occlusal pattern, embrasure shearing dominated by an orthal power stroke, accompanied by a labiolingually directed component of relative displacement of the lower teeth during mastication also distinguishes Gobiconodon from other eutriconodontans, particularly triconodontids, suggesting an insectivorous and/or omnivorous diet for Gobiconodon. The tooth shapes and wear patterns described here indicate diverse ecomorphological specializations and species diversification within eutriconodontans.
The genus Macrocera (Diptera: Keroplatidae) is highly diverse, comprising over 200 species distributed worldwide, yet it remains poorly documented in the extensive sciaroid fossil record. This study presents the results of our investigation of Macrocera in Baltic amber, which led to the discovery of five new species (Macrocera barteki Pełczyńska sp. nov., Macrocera hoffeinsorum Pełczyńska, Krzemiński, & Soszyńska sp. nov., Macrocera poseidonis Pełczyńska, Krzemiński, & Soszyńska sp. nov., Macrocera stuckinamberi Pełczyńska, Krzemiński, & Soszyńska sp. nov., and Macrocera tymoni Pełczyńska sp. nov.) and to the re-examination of the only previously described valid species, Macrocera electricornis. A diagnostic key for species identification is provided. Our results indicate that Macrocera had already attained its modern morphological body plan by the Eocene, highlighting Baltic amber as a largely unexplored source of information on the evolutionary history of certain insect groups. We further discuss the delimitation of the genus and emphasise the need for its comprehensive revision.
The leatherback sea turtle (Dermochelys coriacea) is the largest living sea turtle, but their evolutionary history remains poorly known due to the lack of any confirmed fossil record of the extant species. Here we describe a fossil humerus from the Pleistocene of Taiwan. Its preserved morphology shows a close match to the extant leatherback sea turtle, including the humeral head tilted to the intertubercular side, a slightly-curved secondary deltopectoral crest, and the absence of a deep intertubercular fossa. This Pleistocene humerus from Taiwan then presents the first fossil record of the extant leatherback sea turtle worldwide. The record suggests that the modern global migration pattern of the leatherback sea turtle has already included the western North Pacific. Our discovery of the first fossil leatherback sea turtle should encourage more fossil finds to reveal the detailed evolutionary history of the leatherback sea turtle and the ecosystem they utilize.
Documenting past trophic relationships is crucial to understand deep time changes in the ecology and geographic distribution of large marine predators. Though bite marks on cetacean bones are a useful source of information to assess shark-whale predator-prey relationships, in many cases the identity of the prey and/or predator cannot be precisely determined. In this work, we investigate two cetacean specimens from the Kattendijk Formation (Lower Pliocene, north of Belgium): (i) the partial cranium of a small balaenid (right whale) that we describe and refer to Balaenella brachyrhynus and (ii) the previously described partial skeleton of a monodontid (family of beluga and narwhal), attributed here to the genus Casatia. Shark bite marks observed on the bones of the two specimens are described and the tip of a shark tooth found embedded in each cranium is visualized through micro-CT imaging. A comparison with shark species recorded in the Kattendijk Formation allows for the identification of the author of at least one bite for each cetacean: the bluntnose sixgill shark Hexanchus griseus for B. brachyrhynus and the large lamnid shark Carcharodon plicatilis for Casatia sp. Based on the position and topology of the marks, the H. griseus bite on B. brachyrhynus may have occurred on an upside down, floating carcass, a hypothesis that could indicate a scavenging event. Bite marks on the monodontid cranium suggest an attempt by C. plicatilis to sever the head from the rest of the body and point to the forehead region as a main targeted area. These new Early Pliocene records of shark-cetacean trophic relationships constitute a first step towards the exploration of the possible link between changes through time in the availability of prey in the southern North Sea and the local loss of large predatory sharks, which occurred in the Late Pliocene to Pleistocene time interval.
The first fossil representatives of the harvestman subfamily Ortholasmatinae (Opiliones, Dyspnoi, Nemastomatidae) are described as Balticolasma wunderlichi gen. et sp. nov. One male is preserved in Eocene Baltic amber and a presumably conspecific female in Eocene Rovno amber (northwest Ukraine). Ortholasmatines are typically highly ornate arachnids, and for the first time with an amber harvestman we applied computed tomography using synchrotron radiation to investigate its three-dimensional morphology and surface structure in considerable detail. Some of its morphological characters, appear to be closer to extant Asian genera. In a wider biogeographic context, our amber record is a significant find for the Paleogene of Europe given that (i) it is another species apparently found in both Baltic and Rovno amber and (ii) all modern ortholasmatines are restricted to East Asia and North and Central America.
This paper describes three new species belonging to the genus Styringomyia, based on specimens preserved in Early Miocene (Burdigalian) Dominican amber: Styringomyia caridadi sp. nov., S. caribeana sp. nov., and S. grimaldii sp. nov. Previously, only five extinct species of Styringomyia were known, including two from Dominican amber. These new discoveries increase the total number of species known from fossils to eight. The genus Styringomyia is morphologically intriguing, characterized by the highly complex structure of the male and female terminalia, particularly the morphology of the gonostyles. The hypopygial features of the five Dominican amber species resemble those of certain extant Styringomyia species found in Australia. These findings contribute to our broader understanding of cranefly diversity and significantly enhance our knowledge of Miocene Styringomyia and this stage of evolution of the genus. Notably, despite the abundance of Styringomyia fossils in Dominican amber, the genus is not known to inhabit the island of Hispaniola today.
This study presents a study of biting midge (Diptera: Ceratopogonidae) fauna from early Eocene amber of Oise, Paris Basin, France. The fauna comprise eight species, all of them new to science: Eosphaeromias eocenicus Szadziewski, Santer, Nel, & Krzemi & nacute;ska gen. et sp. nov. Leptoconops oisensis Szadziewski, Santer, Nel, & Krzemi & nacute;ski sp. nov., Brachypogon (Isohelea) europaeus Szadziewski, Santer, Nel, & Krzemi & nacute;ski sp. nov.; Brachypogon (Isohelea) oisen sis Szadziewski, Santer, Nel, & Krzemi & nacute;ski sp. nov.; Brachypogon (Brachypogon) parisiensis Szadziewski, Santer, Nel, & Krzemi & nacute;ski sp. nov., Stilobezzia (Acanthohelea) eocenica Szadziewski, Santer, Nel, & Krzemi & nacute;ski sp. nov., Forcipomyia oisensis Szadziewski, Santer, Nel, & Krzemi & nacute;ski sp. nov., and Dasyhelea eocenica Szadziewski, Santer, Nel, & Krzemi & nacute;ski sp. nov. The fauna of biting midges from early Eocene Oise forests (53 Ma) shows a low diversity and has no common species with the well-studied middle Eocene fauna of Baltic amber forests (c. 40 Ma).
A new genus and species of the extinct family Saurodontidae (Ichthyodectiformes) is described from the Upper Cretaceous deposits of Dakhla Formation, Western Desert, Egypt. The specimen is identified as Wadiichthys anbaawyi gen. et sp. nov. The holotype comprises complete skull with maxillae, premaxillae, dentaries and predentary. The specimen was embedded in a compacted mudstone layer of the lower part of Dakhla Formation, Western Desert, Egypt. The new taxon Wadiichthys anbaawyi differs from the others saurodontid genera by (i) the dorsal margin of the predentary is longer than the ventral margin. (ii) the longitudinal foramina below each alveolus of the dentary, premaxilla and maxilla; (iii) a small diastema between the teeth; (iv) a convex contact between the rostrodermethmoid and the frontal; (v) the posterior margin of the maxilla is triangular shaped; (vi) the teeth are more deeply embedded in the premaxilla, maxilla and dentary. Wadiichthys anbaawyi represents the first formal description of a saurodontid from Africa that contributes to the increase of the knowledge of the diversity and the paleogeographic distribution of this family. The new taxon sheds light on the taxonomy, morphology and evolutionary history of the Cretaceous marine vertebrates of the Tethys Sea. To preserve and digitally document the fossil, a terrestrial laser scanner Trimble TX6 was employed. Each of the five fossil parts was scanned separately and processed with Trimble RealWorks software. This created a high-resolution 3D point cloud in true color, grayscale, and intensity mode. This method protects the fossil from possible damage or loss. It also allows for remote collaboration and morphometric analysis without physical handling.
This study examines the bird ichnoassemblage of the Miocene Vinchina Formation (La Rioja Province, Argentina), recorded in fluvio-lacustrine environments. Both unpublished material and previously collected specimens were analysed across three transverse sections cropping out from north to south in the Quebrada Pedregal Negro, the Quebrada de la Troya, and the Quebrada del Yeso. The Pedregal Negro creek has yielded only Aviadactyla vialovi, recently synonymised with Ornithotarnocia lambrechti. The low ichnodiversity observed is likely due more to limited exploration efforts and poor outcrop exposure in the creek than to palaeoenvironmental constraints. Exposures at Quebrada de la Troya yield six ichnotaxa preserved in fluvial and lacustrine settings: Aramayoichnus rheae (a large rheid and the oldest rheid track in southern South America), Phoenicopterichnus rector (Phoenicopteridae), Ardeipeda isp. (Ciconiiformes), cf. Jindonornipes, cf. Aquatilavipes (small aquatic birds, possibly Recurvirostridae or Charadriidae), and Rionegrina isp. (a functional didactyl phorusrhacid). At Quebrada del Yeso creek, five ichnospecies of birds were recognised and preserved in a playa-lake environment: Phoenicopterichnus rector, Gragliavipes gavenskii (Threskiornithidae), Avipeda isp. (likely Scolopacidae or Charadriidae), together with a ralliform morphotype and a Cariamidae morphotype. Overall, the trackmakers represented in the three ichnoassemblages correspond well with the globally known Miocene avian body fossil record. While the Pedregal Negro section warrants further exploration, differences in ichnodiversity between Quebrada del Yeso and Quebrada de la Troya likely reflect ecological factors, as the track-bearing levels in the latter represent environments with greater freshwater availability, which may have supported more diverse bird communities.
A new genus and species of emperor butterfly (Nymphalidae, Apaturinae) is described based on an exceptionally well-preserved specimen from the “Calcaire de Campagne Calavon” limestone of southern France (early Oligocene age, approximately 34–28 Mya). Most parts of the right wings and upper parts of the left wings are visible, with complete venation and even wing patterns. The head and thorax are clearly visible from both sides, and most of the abdomen is preserved. Apaturoides monikae gen. et sp. nov., is compared to all genera of the subfamily and based on the form of the wings and venation is considered most similar to species of Apatura, today occurring widely in Palearctic, excluding North Africa and Middle East. This discovery provides an important calibration point based on direct fossil evidence for estimating relationships and divergence estimates among emperor butterflies.
A bivalve association reported herein was recently discovered from the Xiazhen Formation (upper Katian, Upper Ordovician) at Zhuzhai, southern Yushan County, northeastern Jiangxi Province, East China. Nine genera and ten species are systematically documented including four new species: Palaeoneilo rectus sp. nov. (most abundant), Palaeoneilo cf. constricta, Goniophorina volvens, Modiolopsis elegantulus, Cyrtodontula sp., Cuneamya? sp., Trigonoconcha brevis sp. nov., Similodonta minor sp. nov., Concavodonta varius sp. nov., and Paulinea cf. parva. The association is mainly composed of Protobranchia (Nuculiformii and Nuculaniformii) and subordinate Pteriomorphia, suggesting a relatively low palaeolatitudinal setting. The dominance of infaunal protobranch bivalves is notable, and against the trend of rapid diversification of pteriomorphs during the Late Ordovician. This discrepancy is likely attributable to a cold water tongue extending from then South Pole to South China palaeoplate, and the ecological pressure from those much more abundant brachiopods on the substrate surface, both hindering the development of epifaunal pteriomorphs in the region. Significantly, this bivalve association shares several common genera with its coeval bivalve fauna from the Oslo region, Norway.
The fossil genus Yuskaichthys, previously known only from Eocene deposits of the Maíz Gordo site in Argentina, is here identified based on three isolated parieto-supraoccipital bones collected in the Tiupampa locality, Cochabamba-Potosí Basin, Bolivia. This discovery notably extends both the paleogeographic and temporal distribution of the genus into the early Paleocene. Morphological traits including the bifid posterior margin of the supraoccipital process and the transverse pit lines closely resemble those described for Yuskaichthys. Futhermore, these fish share a longitudinal groove along the entire supraoccpital process, supporting a provisional assignment to Yuskaichthys. The occurrence of Yuskaichthys in both Tiupampa and Maíz Gordo is consistent with regional fluvial systems associated with the Andean foreland basin evolution. This finding supports an early diversification of modern catfish lineages, including extinct genera displaying long time range.
Meiolaniformes are a group of chelonians including the famous horned-turtles Niolamia argentina (Patagonia) and Meiolania platyceps (Australia). In South America, the Late Cretaceous meiolaniforms are represented by two named taxa: Patagoniaemys gasparinae coming from Campanian-Maastrichtian beds of the La Colonia Formation, Chubut province, and Trapalcochelys sulcata from the Campanian-Maastrichtian beds of the Allen Formation, Rio Negro Province. The aim of the present contribution is to describe a new meiolaniform chelonian coming from the Upper Cretaceous (Maastrichtian) Los Alamitos Formation, at Rio Negro Province, Argentina. The material is represented by a partial basicranium, incomplete carapace and fragmentary postcranial bones (MPMIK 1839/P/33) belonging to a new species Patagoniaemys aeschyli. To this new species we refer all the material previously referred as indeterminate meiolaniid, meiolaniform and cf. Niolamia sp., coming from the same site and locality, and described by different authors since the 1980s. This new species differs from the type species P. gasparinae, particularly by the presence of longitudinal ridges in the nuchal bone. We also include brief comments about meiolaniform palaeobiogeography and the impact of K/Pg extinction event in Patagonian chelonians.
The fossil elephant remains from the lower Middle Pleistocene site of Contrada Monticelli (Apulia, southern Italy) are described here in detail for the first time. The material, consisting of dental and postcranial bones from a single individual, was analyzed both morphologically and morphometrically. Estimates of shoulder height and body mass were calculated for the specimen. Based on dental characters, the Contrada Monticelli elephant is assigned to Palaeoloxodon antiquus. A comparative analysis with Mammuthus and Palaeoloxodon specimens from the Lower and Middle Pleistocene of Italy indicates the Contrada Monticelli individual represents a medium-sized, old male straight-tusked elephant. The comparative study also demonstrated that several morphological traits of the humerus, ulna, femur, and tibia, previously proposed as diagnostic in distinguishing between Mammuthus and Palaeoloxodon, are not consistent when intraspecific variability is taken into account. Furthermore, the morphometric analyses revealed no significant differences in size and proportions of the studied limb bones between Mammuthus meridionalis and Palaeoloxodon antiquus that would allow for a reliable discrimination between the two taxa. The Contrada Monticelli elephant represents one of the few known European straight-tusked elephant skeletons dated to the interval between 0.8 and 0.5 million years ago, corresponding to the dispersal of the genus into Eurasia, and contributes to a better understanding of the biochronological, paleoecological, and paleobiogeographical context of the early evolution of Palaeoloxodon in Europe.
The present paper describes a bryozoan fauna from the Devonian of Holy Cross Mountains (Poland) which includes nine species. Six species were described from the Dobruchna Member of the Skały Formation (Eifelian) of Skały section: Cyclotrypa sp., Fistuliphragma gracilis, Leioclema passitabulatum, Intrapora leunisseni, Laxifenestella sp., Hemitrypella nodulosa. The cryptostome species Bigeyella indigena was identified from the Laskowa Góra Beds (upper Givetian) of Józefka Hill at Górno. The fenestrate Rectifenestella sp. is described from the lower Frasnian Kowala Formation (Kadzielnia Member) of Bolechowice and Kielce. The new monotypic fenestrate genus Juanopora elegans gen. et sp. nov. is described from the Wietrznia Beds (upper Givetian) of Wietrznia-I quarry at Kielce. The described bryozoans show palaeobiogeographical connections to the Middle–Upper Devonian of Germany, Spain, and Armenia. The bryozoan faunas of the Holy Cross Mountains are in accordance with the worldwide acme of the sessile filter-feeding benthos during the Givetian age, followed by a two-stage decline in diversity due to the Taghanic and Frasnian crises.
The first fossil insect from the Cisuralian (Lower Permian) Arroyo Totoral Formation, south of Sierra de los Llanos, Paganzo Basin, southeastern La Rioja Province, Argentina is reported. A new species of Phyloblattidae (Dictyoptera), Anthracoblattina macucai sp. nov., is described from an almost complete forewing and based on its general venation scheme (e.g., distinct narrowing of the strip-like costal field near wing base, widened areas between the main veins, Sc pectinated, R and M slightly sigmoidal, twigs provide fan-like covering of the wing apex, the first forks of both R and M located between the first third and the half of the wing length, CuA straight, ends at the transition of the wing tip into the posterior wing border, CuP fluently curved, broad interspace between CuP and AA, cross-venation anastomosing- striate to reticulate). Additionally, we discuss the taxonomic position of A. macucai sp. nov. in relation to other phyloblattid species described from Carboniferous–Permian South American outcrops. Finally, we analyze this new fossil “cockroachoids” within its paleofloristic context and depositional environment. A. macucai sp. nov. represents the first known Permian insect found for both the La Rioja Province and the Arroyo Totoral Formation, pointing the unit as promising for the search of fossil insects in Argentina.
The extinct elasmobranch Protospinax is an intriguing shark known mostly from isolated teeth and rare complete skeletons. Most previous studies focused on differences in isolated teeth for taxonomic assignments, with little to no considerations of dental variations. In this study we provide a detailed morphological evaluation of the dentition of the squalomorph shark, Protospinax based on three skeletal remains of Protospinax annectans from the famous Upper Jurassic lithographic limestone of the Solnhofen area (Bavaria, Germany) with partially preserved dentitions and isolated teeth from the Kimmeridgian of Mahlstetten (Baden-Württemberg, Germany). The aim of this study is to clarify ambiguities in dental morphologies and to establish heterodonty patterns, allowing to taxonomically reassess species previously assigned to Protospinax. Accordingly, we consider Protospinax annectans (Callovian–Aptian?), Protospinax carvalhoi (Bathonian), Protospinax lochensteinensis (Oxfordian), and Protospinax planus (Kimmeridgian) as valid species. The species of Protospinax bilobatus is considered a junior synonym of Protospinax magnus. Furthermore, our results show that the dental morphologies of P. magnus and Protospinax? muftius are very different from those of other Protospinax species and rather resemble those of orectolobiforms. Consequently, we introduce two new orectolobiform genera, Jurascyllium gen. nov. and Archaeoscyllium gen. nov., to accommodate these species. The review of the species confirms a stratigraphic range of Protospinax extending from the Toarcian (Lower Jurassic) to the Valanginian (Lower Cretaceous).
