Extinct Clade Research Articles

Cretaceous and Paleocene fossils reveal an extinct higher clade within Cornales, the dogwood order.

Characterization and phylogenetic integration of fossil angiosperms with uncertain affinities is relatively limited, which may obscure the diversity of extinct higher taxa in the flowering plant tree of life. The order Cornales contains a diversity of extinct taxa with uncertain familial affinities that make it an ideal group for studying turnover in angiosperms. Here, we describe a new extinct genus of Cornales unassignable to an extant family and conduct a series of phylogenetic analyses to reconstruct relationships of fossils across the order. Two permineralized endocarps were collected from the Cedar District Formation (Campanian, 82-80 Ma) of Sucia Island, State of Washington, United States. Fossils were sectioned with the cellulose acetate peel technique and incorporated into a morphological dataset. To assess the utility of this dataset to accurately place taxa in their respective clades, we used a series of phylogenetic pseudofossilization analyses. We then conducted a total-evidence analysis and a scaffold-based approach to determine relationships of fossils. Based on their unique combination of characters, the fossils represent a new genus, Fenestracarpa washingtonensis gen. nov. et sp. nov. Pseudofossilization analyses indicate that our morphological dataset can be used to accurately recover taxa at the major clade to family level, generally with moderate to high support. The total-evidence and scaffold-based analyses recovered Fenestracarpa and other fossil genera in an entirely extinct clade within Cornales. Our findings increase the reported diversity of extinct Cornales and indicate that the order's initial radiation likely included the divergence of an extinct higher clade that endured the end-Cretaceous Mass extinction but perished during the Cenozoic.

Skeletal elements controlled soft-tissue preservation in echinoderms from the Early Ordovician Fezouata Biota

Exceptional fossils preserve non-biomineralized tissues in the geological record and provide crucial information on the evolution of life on Earth. Exceptionally preserved fossils are rarely discovered complete, challenging their morphological description and their palaeontological interpretation. Although decay experiments reconstructing degradation sequences in modern animals are necessary to better understand taphonomic processes involved in exceptional preservation, their applicability to some enigmatic and/or extinct fossil taxa remains limited. Here, based on a representative sample of 423 specimens collected from a single stratigraphic level from the Early Ordovician Fezouata Biota, we reconstruct the degradation sequence of both skeletal remains and soft tissues of stylophorans, an extinct clade of echinoderms. The rare preservation of the water vascular system can be explained by the rapid post-mortem opening of the cover plates resulting from the fast decay of associated muscles and the action of ligaments. In contrast, the proximal aulacophore and associated stylocone formed a particularly decay-resistant closed module, thus favouring the preferential preservation of included soft parts (fore-gut). The non-random location and frequency of pyritised intra-skeletal structures strongly suggest that skeletal elements dictated the preservation of underlying soft parts. As such, taphonomic investigations should not only focus on the environment surrounding a decaying animal, but also on the different environments created within a particular carcass.

What else is dentition telling us? A new specimen-level phylogeny of Mesotheriidae (Mammalia, Notoungulata).

Mesotheriidae (Panperissodactyla, Notoungulata) are an extinct clade (early Oligocene-Pleistocene) of small to medium-sized herbivorous mammals that were widely distributed in South America. Although two subfamilies traditionally have been recognized (Trachytheriinae and Mesotheriinae), recent cladistic analyses based on discrete characters have indicated that "Trachytheriinae" is a paraphyletic assemblage. Given the availability of a large number of specimens and the fact that dental characters are the most common characters used in mesotheriid phylogenies, we performed specimen-level cladistic analyses combining discrete, continuous and geometric morphometrics-based dental characters. The aim was to: (1) include new scored morphological characters to solve the phylogenetic relationships of Mesotheriidae; (2) compare the results of the upper and lower dentition analyses as different character partitions and in combination, to establish phylogenetic hypotheses; and (3) trace the evolution of dental traits. Phylogenetic analyses employing characters of associated upper and lower dentitions recovered one most parsimonious tree with Archaeohyracidae (outgroup) as the sister group of Pan-Mesotheriidae (= Mesotheriidae; converted clade name), this latter composed of trachytheriines (stem-mesotheriine) + Mesotheriinae (converted clade name). Within Mesotheriinae, we recovered two main lineages phylogenetically defined here as Bolivarini and Pampaini (new clade names). Analyses of isolated upper and lower dentition sub-datasets each resulted in one most parsimonious tree congruent with the associated dentition. Our study emphasizes the use of geometric morphometrics characters to resolve additional clades in phylogenetic analyses, provides information on the evolution of size and morphology of teeth, and exposes specimen assignment issues at a taxonomic level. The integration of osteological characters might be crucial to further understanding the evolution of Mesotheriidae.

How to weigh a fossil mammal? South American notoungulates as a case study for estimating body mass in extinct clades

How to weigh a fossil mammal? South American notoungulates as a case study for estimating body mass in extinct clades

Extended embryo retention and viviparity in the first amniotes

The amniotic egg with its complex fetal membranes was a key innovation in vertebrate evolution that enabled the great diversification of reptiles, birds and mammals. It is debated whether these fetal membranes evolved in eggs on land as an adaptation to the terrestrial environment or to control antagonistic fetal–maternal interaction in association with extended embryo retention (EER). Here we report an oviparous choristodere from the Lower Cretaceous period of northeast China. The ossification sequence of the embryo confirms that choristoderes are basal archosauromorphs. The discovery of oviparity in this assumed viviparous extinct clade, together with existing evidence, suggests that EER was the primitive reproductive mode in basal archosauromorphs. Phylogenetic comparative analyses on extant and extinct amniotes suggest that the first amniote displayed EER (including viviparity).

Permian trilobites and the applicability of the “living fossil” concept to extinct clades

Some taxa occupy our imaginations as “living fossils” because they were known from the fossil record before being discovered alive today. Other taxa are considered “living fossils” because modern relatives bear a strong morphological resemblance to fossil relatives, or because they occupy a contracted geographic range or have less diversity now than in the past, or because they represent phylogenetic diversity that requires conservation. A characterizing feature of living fossils–and thus an implicit assumption of all criteria–is that the “living fossil” of interest is extant. However, the general research questions that “living fossils” inspire–Why do rates of evolution vary across organisms, across traits, and across time? Why do some clades decline in diversity over extended periods?–may be applied to any clade, including completely extinct clades. We propose that there is nothing special about “now” when it comes to pursuing these questions and that it is unnecessarily limiting to restrict research programs to clades for which an extant member meets some conception of the “living fossil” moniker. To this end, we investigate the extent to which Permian trilobites might resemble “living fossils,” albeit from the perspective of 253 million years ago, when the last trilobites were still alive. We do so by comparing the taxonomic diversity, geographic range, and morphological disparity of trilobites living in the Permian to earlier time periods. We find that Permian trilobites meet most definitions of living fossils, although our assessment of morphological change and character retention depend on taxonomic scale.

Neuroanatomical Study and Three-Dimensional Cranial Reconstruction of the Brazilian Albian Pleurodiran Turtle Euraxemys essweini

Pleurodira represent one of the two clades that compose the crown Testudines, and their temporal range is Late Jurassic–present. However, knowledge about the neuroanatomy of extinct pleurodires is still very limited. In this context, scarce neuroanatomical information about the Cretaceous clade Euraxemydidae is currently available, limited to some characters of the Moroccan Cenomanian Dirqadim schaefferi. In the present work, we perform the detailed neuroanatomical study of its sister taxon, the Brazilian Albian Euraxemys essweini, based on the analysis of the skull of its holotype and only known individual of the species. The detailed virtual three-dimensional reconstruction of all its cranial bones is performed, also improving the information about its osseous anatomy. The different neuroanatomical cavities (i.e., cranial, nasal, and labyrinthic ones) and canals (i.e., nervous and circulatory ones) are compared with those identified thus far for other extinct and extant members of the Pleurodira in order to characterize the neuroanatomy of the extinct clade Euraxemydidae in detail.

A peirosaurid mandible from the Albian–Cenomanian (Lower Cretaceous) of Algeria and the taxonomic content of Hamadasuchus (Crocodylomorpha, Peirosauridae)

AbstractPeirosaurids form an extinct clade of terrestrial crocodylomorphs from the Cretaceous of Africa and South America. Here, we describe a new mandibular ramus attributable to Hamadasuchus cf. rebouli from the Albian–Cenomanian of La Gara Samani (Algeria). We propose an emended diagnosis for this taxon, originally described from a left dentary fragment from the Kem Kem Group of Morocco and discuss the assignment of the specimens currently referred to it. Using ontogenetic series of extant crocodylians, we show that several mandibular characters commonly used to differentiate between close taxa instead record intraspecific or ontogenetic variability. In contrast, reliable mandibular characters enable us to propose that the current taxonomic content of Hamadasuchus rebouli should be reduced to three specimens, pending future description of relatable cranial remains. Finally, we demonstrate the importance of mandibular characters in phylogenies, by recovering the new specimen from La Gara Samani as closely related to North African peirosaurids on the basis of a data matrix designed solely on mandibular characters.

A high latitude Gondwanan species of the Late Devonian tristichopterid Hyneria (Osteichthyes: Sarcopterygii).

We describe the largest bony fish in the Late Devonian (late Famennian) fossil assemblage from Waterloo Farm near Makhanda/Grahamstown, South Africa. It is a giant member of the extinct clade Tristichopteridae (Sarcopterygii: Tetrapodomorpha) and most closely resembles Hyneria lindae from the late Famennian Catskill Formation of Pennsylvania, USA. Notwithstanding the overall similarity, it can be distinguished from H. lindae on a number of morphological points and is accordingly described as a new species, H. udlezinye sp. nov. The preserved material comprises most of the dermal skull, lower jaw, gill cover and shoulder girdle. The cranial endoskeleton appears to have been unossified and is not preserved, apart from a fragment of the hyoid arch adhering to a subopercular, but the postcranial endoskeleton is represented by an ulnare, some semi-articulated neural spines, and the basal plate of a median fin. The discovery of H. udlezinye shows that Hyneria is a cosmopolitan genus extending into the high latitudes of Gondwana, not a Euramerican endemic. It supports the contention that the derived clade of giant tristichopterids, which alongside Hyneria includes such genera as Eusthenodon, Edenopteron and Mandageria, originated in Gondwana.

A new pipid from the Cretaceous of Africa (In Becetèn, Niger) and early evolution of the Pipidae

Pipimorpha and its crown-group Pipidae possess one of the most extensive fossil records among anurans, known since the Early Cretaceous in both Laurasia and Gondwana. Pipimorph diversification may have been driven by the breakup of West Gondwana during the Cretaceous. Numerous fossils from South America have been unearthed in the last decade, documenting this event. Unfortunately, Cretaceous pipimorphs from Africa have been limited to a few well-preserved taxa from sub-Saharan Africa and the Arabian Peninsula, which hinders our comprehension of pipimorph diversification during this key period. The site of In Becetèn, in south-east Niger, is one of the few mid-Late Cretaceous (Coniacian–Santonian) sites from which a pipid, Pachycentrata taqueti, is known. Here, we describe and name a second pipid from the same locality. This taxon is known by a relatively complete braincase. Phylogenetic analyses confirm its position as a pipid, with pipinomorph affinities. This makes In Becetèn the oldest site with at least two pipids. Phylogenetic results are congruent with recent pipimorph relationships, with the presence of an endemic extinct clade in South America, Shelaniinae. The phylogenetic results also allow us to review the proposed definition for Pipimorpha and its subclades and propose new systematic definitions for them. Temporal calibration of the phylogenetic tree based on the fossil record suggests that pipimorphs diversified in a western Gondwana block and confirms that South America separated from Africa around the mid-Cretaceous. Between these two events, pipids diverged in Africa, giving rise to major extant clades. This study highlights the importance of Africa for early pipid diversification during the Cretaceous and of the opening of the Southern Atlantic Ocean for anuran dispersion and diversification. http://zoobank.org/urn:lsid:zoobank.org:pub:D40F9A91-D57C-4ED8-B80A-9CA5246599E3

Ecological divergence and calcaneal‐cuboid morphology in gorillas

AbstractObjectivesGorillas are diverse behaviorally and ecologically with higher frequencies of arboreality and frugivory observed in populations living at lower altitudes compared to those living at higher altitudes. Previous studies have shown that the more arboreal western lowland gorilla has hallucial tarsometatarsal and talocrural joint morphologies associated with enhanced hallucial abduction and foot inversion capabilities whereas the more terrestrial eastern gorilla does not. Here we examine how known ecological and behavioral differences among gorillas relate to variation in calcaneal and cuboid morphology. Specifically, we test whether gorillas that climb more frequently (i.e., western lowland and lowland grauer gorillas) have cuboid‐metatarsal and calcaneocuboid morphologies that facilitate increased foot mobility compared to those that climb less often (i.e., mountain and highland grauer gorillas).MethodsWe quantify the relative areas of articular and nonarticular surfaces, angles between articular surfaces, articular surface curvatures, the relative length and depth of the calcaneocuboid joint, and the relative size of the calcaneal tuber using three‐dimensional models of gorilla calcanei and cuboids (n = 90 and 81, respectively).ResultsOur results, based on univariate and multivariate analyses, show that western and eastern gorillas follow our predicted arboreal‐terrestrial morphocline in terms of their calcaneal and cuboid morphology; however, highland and lowland grauer gorilla populations do not.ConclusionsThis study has implications for understanding gorilla foot function and the evolutionary processes responsible for producing the morphological patterns among living gorilla populations. Future studies of gorilla morphological diversity may provide a useful comparative model for studies of locomotor evolution in extinct clades.

New data from the first discovered paleoparadoxiid (Desmostylia) specimen shed light into the morphological variation of the genus Neoparadoxia

Desmostylia is an extinct clade of marine mammals with two major sub-clades, Desmostylidae and Paleoparadoxiidae, known from Oligocene to Miocene strata of the North Pacific coastline. Within Paleoparadoxiidae, three genera have been identified: Archaeoparadoxia, Paleoparadoxia, and Neoparadoxia. The latter taxon is the geochronologically youngest palaeoparadoxiid and Neoparadoxia is characterized by a comparatively larger body size, although it is known only from a few specimens within a short temporal and geographic range. Here we report the discovery of an isolated tooth, which we identify as Neoparadoxia cf. N. cecilialina, constituting only the second individual specimen of Neoparadoxia with preserved dentition yet reported. This specimen was collected near Corona, California, USA, and we attribute it to the “Topanga” Formation, extending the geographic range of this taxon in Southern California. While the exact geographic locality was not recorded when it was collected in 1913, we establish two potential localities based on associated hand-written museum label and new stratigraphic information. Although initially identified as Desmostylus hesperus, this specimen of Neoparadoxia was collected 10 years before the first named paleoparadoxiid from Japan. We expect that description of more complete desmostylian material from elsewhere in Southern California will clarify the taxonomic richness and paleoecological role of this clade in Cenozoic marine mammal assemblages.

Spine-like structures in Paleogene muricate planktonic foraminifera

Abstract. Muricate planktonic foraminifera comprise an extinct clade that was diverse and abundant in the Paleogene oceans and are widely used in palaeoclimate research as geochemical proxy carriers for the upper oceans. Their characteristic wall texture has surface projections called “muricae” formed by upward deflection and mounding of successive layers of the test wall. The group is generally considered to have lacked “true spines”: that is, acicular calcite crystals embedded in and projecting from the test surface such as occur in many modern and some Paleogene groups. Here we present evidence from polished sections, surface wall scanning electron microscope images and test dissections, showing that radially orientated crystalline spine-like structures occur in the centre of muricae in various species of Acarinina and Morozovella and projected from the test wall in life. Their morphology and placement in the wall suggest that they evolved independently of true spines. Nevertheless, they may have served a similar range of functions as spines in modern species, including aiding buoyancy and predation and especially harbouring algal photosymbionts, the function for which we suggest they probably first evolved. Our observations strengthen the analogy between Paleogene mixed-layer-dwelling planktonic foraminifera and their modern spinose counterparts.

Three-dimensional polygonal muscle modelling and line of action estimation in living and extinct taxa

Biomechanical models and simulations of musculoskeletal function rely on accurate muscle parameters, such as muscle masses and lines of action, to estimate force production potential and moment arms. These parameters are often obtained through destructive techniques (i.e., dissection) in living taxa, frequently hindering the measurement of other relevant parameters from a single individual, thus making it necessary to combine multiple specimens and/or sources. Estimating these parameters in extinct taxa is even more challenging as soft tissues are rarely preserved in fossil taxa and the skeletal remains contain relatively little information about the size or exact path of a muscle. Here we describe a new protocol that facilitates the estimation of missing muscle parameters (i.e., muscle volume and path) for extant and extinct taxa. We created three-dimensional volumetric reconstructions for the hindlimb muscles of the extant Nile crocodile and extinct stem-archosaur Euparkeria, and the shoulder muscles of an extant gorilla to demonstrate the broad applicability of this methodology across living and extinct animal clades. Additionally, our method can be combined with surface geometry data digitally captured during dissection, thus facilitating downstream analyses. We evaluated the estimated muscle masses against physical measurements to test their accuracy in estimating missing parameters. Our estimated muscle masses generally compare favourably with segmented iodine-stained muscles and almost all fall within or close to the range of observed muscle masses, thus indicating that our estimates are reliable and the resulting lines of action calculated sufficiently accurately. This method has potential for diverse applications in evolutionary morphology and biomechanics.

Combined data analysis of fossil and living mammals: a Paleogene sister taxon of Placentalia and the antiquity of Marsupialia.

The Cretaceous-Paleogene (KPg) boundary, one of Earth's five major extinction events, occurred just before the appearance of Placentalia in the fossil record. The Gobi Desert, Mongolia and the Western Interior of North America have important fossil mammals occurring just before and after the KPg boundary (e.g. Prodiacodon, Deltatheridium) that have yet to be phylogenetically tested in a character-rich context with molecular data. We present here phylogenetic analyses of >6000 newly scored anatomical observations drawn from six untested fossils and added to the largest existing morphological matrix for mammals. These data are combined with sequence data from 27 nuclear genes. Results show the existence of a new eutherian sister clade to Placentalia, which we name and characterize. The extinct clade Leptictidae is part of this placental sister clade, indicating that the sister clade survived the KPg event to co-exist in ancient ecosystems during the Paleogene radiation of placentals. Analysing the Cretaceous metatherian Deltatheridium in this character-rich context reveals it is a member of Marsupialia, a finding that extends the minimum age of Marsupialia before the KPg boundary. Numerous shared-derived features from multiple anatomical systems support the assignment of Deltatheridium to Marsupialia. Computed tomography scans of exquisite new specimens better document the marsupial-like dental replacement pattern of Deltatheridium. The new placental sister clade has both Asian and North American species, and is ancestrally characterized by shared derived features such as a hind limb modified for saltatorial locomotion.

Novel evolution of a hyper-elongated tongue in a Cretaceous enantiornithine from China and the evolution of the hyolingual apparatus and feeding in birds.

The globally distributed extinct clade Enantiornithes comprises the most diverse early radiation of birds in the Mesozoic with species exhibiting a wide range of body sizes, morphologies, and ecologies. The fossil of a new enantiornithine bird, Brevirostruavis macrohyoideus gen. et sp. nov., from the Lower Cretaceous Jiufotang Formation in Liaoning Province, northeastern China, preserves a few important skeletal features previously unknown among early stem and extant birds, including an extremely elongate bony hyoid element (only slightly shorter than the skull), combined with a short cranial rostrum. The long hyoid provides direct evidence for the evolution of specialized feeding in this extinct species, and appears similar to the highly mobile tongue that is mobilized by the paired epibranchials present in living hummingbirds, honeyeaters, and woodpeckers. The likely linkage between food acquisition and tongue protrusion might have been a key factor in the independent evolution of particularly elongate hyobranchials in early birds.

Apuliadercetis gonzalezae sp. nov., a North American Campanian dercetid fish (Teleostei, Aulopiformes) from Tzimol, Chiapas, Mexico

A new member of the family Dercetidae is named in this work as Apuliadercetis gonzalezae sp. nov., based on specimens from the marine Campanian deposits of the Angostura Formation, exposed in the Tzimol quarry, next to Ochuxhob, Tzimol Municipality, southeastern Chiapas. The osteology of this serpentiform and longirostrine new fish shows a diagnostic character of the order Aulopiformes, the occurrence of epipleurals bones associated with abdominal centra extends up to the centrum 2. This species is a member of the family Dercetidae because it shows the most outstanding characters of this extinct clade, two pairs of triangular and expanded transverse processes in the abdominal centra, the low and elongated neural arches on the abdominal centra bearing short neural spines, and a row of triradiate flank scutes on each side of the body. Additionally, this fish belongs to Apuliadercetis because it shares the synapomorphy of this genus, the neural spines of the centra located below the dorsal fin are extremely reduced. This fish represents the third species in the genus and the first from America; such a discovery also shows that this genus had a wide geographical distribution that at least comprised Europe and the far south of North America.

A second peirosaurid crocodyliform from the Mid-Cretaceous Kem Kem Group of Morocco and the diversity of Gondwanan notosuchians outside South America.

Notosuchians are an extinct clade of terrestrial crocodyliforms with a particularly rich record in the late Early to Late Cretaceous (approx. 130–66 Ma) of Gondwana. Although much of this diversity comes from South America, Africa and Indo-Madagascar have also yielded numerous notosuchian remains. Three notosuchian species are currently recognized from the early Late Cretaceous (approx. 100 Ma) Kem Kem Group of Morocco, including the peirosaurid Hamadasuchus rebouli. Here, we describe two new specimens that demonstrate the presence of at least a fourth notosuchian species in this fauna. Antaeusuchus taouzensis n. gen. n. sp. is incorporated into one of the largest notosuchian-focused character-taxon matrices yet to be compiled, comprising 443 characters scored for 63 notosuchian species, with an increased sampling of African and peirosaurid species. Parsimony analyses run under equal and extended implied weighting consistently recover Antaeusuchus as a peirosaurid notosuchian, supported by the presence of two distinct waves on the dorsal dentary surface, a surangular which laterally overlaps the dentary above the mandibular fenestra, and a relatively broad mandibular symphysis. Within Peirosauridae, Antaeusuchus is recovered as the sister taxon of Hamadasuchus. However, it differs from Hamadasuchus with respect to several features, including the ornamentation of the lateral surface of the mandible, the angle of divergence of the mandibular rami, the texture of tooth enamel and the shape of the teeth, supporting their generic distinction. We present a critical reappraisal of the non-South American Gondwanan notosuchian record, which spans the Middle Jurassic–late Eocene. This review, as well as our phylogenetic analyses, indicate the existence of at least three approximately contemporaneous peirosaurid lineages within the Kem Kem Group, alongside other notosuchians, and support the peirosaurid affinities of the ‘trematochampsid’ Miadanasuchus oblita from the Maastrichtian of Madagascar. Furthermore, the Cretaceous record demonstrates the presence of multiple lineages of approximately contemporaneous notosuchians in several African and Madagascan faunas, and supports previous suggestions regarding an undocumented pre-Aptian radiation of Notosuchia. By contrast, the post-Cretaceous record is depauperate, comprising rare occurrences of sebecosuchians in north Africa prior to their extirpation.

The locomotion of extinct secondarily aquatic tetrapods

The colonisation of freshwater and marine ecosystems by land vertebrates has repeatedly occurred in amphibians, reptiles, birds and mammals over the course of 300 million years. Functional interpretations of the fossil record are crucial to understanding the forces shaping these evolutionary transitions. Secondarily aquatic tetrapods have acquired a suite of anatomical, physiological and behavioural adaptations to locomotion in water. However, much of this information is lost for extinct clades, with fossil evidence often restricted to osteological data and a few extraordinary specimens with soft tissue preservation. Traditionally, functional morphology in fossil secondarily aquatic tetrapods was investigated through comparative anatomy and correlation with living functional analogues. However, in the last two decades, biomechanics in palaeobiology has experienced a remarkable methodological shift. Anatomy-based approaches are increasingly rigorous, informed by quantitative techniques for analysing shape. Moreover, the incorporation of physics-based methods has enabled objective tests of functional hypotheses, revealing the importance of hydrodynamic forces as drivers of evolutionary innovation and adaptation. Here, we present an overview of the latest research on the locomotion of extinct secondarily aquatic tetrapods, with a focus on amniotes, highlighting the state-of-the-art experimental approaches used in this field. We discuss the suitability of these techniques for exploring different aspects of locomotory adaptation, analysing their advantages and limitations and laying out recommendations for their application, with the aim to inform future experimental strategies. Furthermore, we outline some unexplored research avenues that have been successfully deployed in other areas of palaeobiomechanical research, such as the use of dynamic models in feeding mechanics and terrestrial locomotion, thus providing a new methodological synthesis for the field of locomotory biomechanics in extinct secondarily aquatic vertebrates. Advances in imaging technology and three-dimensional modelling software, new developments in robotics, and increased availability and awareness of numerical methods like computational fluid dynamics make this an exciting time for analysing form and function in ancient vertebrates.

Diversity of gobioid fishes in the late middle Miocene of northern Moldova, Eastern Paratethys \u2013 part I: an extinct clade of Lesueurigobius look-alikes

Studies of otoliths suggest that Gobioidei, which are among the most species-rich groups of modern-day vertebrates, were prominent elements of late middle Miocene (early Sarmatian sensu lato) fish faunas in Europe and Western Asia. However, few complete skeletons have come to light. Here, we report an assemblage of six marine gobiid species, based on skeletons preserved with otoliths in situ, from the lower Volhynian (lower Sarmatian s.l.) of Karpov Yar, Naslavcea, northern Moldova (Eastern Paratethys). Previously only one of these species had been reported from the Central Paratethys, based on its otoliths alone. Five new species representing four new genera are described: †Katyagobius prikryli gen. et sp. nov., Pseudolesueurigobius manfredi gen. et sp. nov., †Sarmatigobius compactus gen. et sp. nov., †Yarigobius decoratus gen. et sp. nov., and †Y. naslavcensis gen. et sp. nov. All six species share the following set of characters, suggesting that they represent a monophyletic clade: 27–29 vertebrae (of which 10 are abdominal); spines of first dorsal fin distally filamentous; second dorsal fin with spine and 14–16 soft rays; anal fin with spine and 13–15 soft rays; caudal fin longish-to-lanceolate; otoliths (sagittae) with rounded, trapezoid-to-squarish shape. Their skeletal features suggest that they are closely related to Lesueurigobius Whitley, 1950, but the otoliths preserved in situ do not support such a classification. The new fossils most likely represent a stem lineage of the European Aphia lineage, and indicate that the diversity of gobiid lineages 12 million years ago differed clearly from that observed today.