A new basilosaurid (Cetacea, Pelagiceti) from the Late Eocene to Early Oligocene Otuma Formation of Peru

Cynthiacetus peruvianus nov. sp. is a new basilosaurid species, from Late Eocene to Early Oligocene Otuma Formation of Peru. It is the first described archaeocete in South-America and is represented by a sub-complete skeleton. C. peruvianus differs from C. maxwelli (middle to Late Eocene of Egypt and United States) principally in having one cuspid less on both mesial and distal sides of p3 and p4. Cynthiacetus is among the largest basilosaurids. Its more characteristic features are located on its postcranial skeleton: large vertebrarterial foramina on cervical vertebrae and absence of ventral expansion of the transverse process on C3-C5. Besides, C. peruvianus presents the greatest number of thoracic vertebrae (20) and ribs observed in Cetacea and the first thoracics have an almost vertical neural spine. A preliminary parsimony analysis establishes the monophyly of the Basilosauridae on the basis of three unambiguous cranial synapomorphies. However, within the Basilosauridae, the most diagnostic characters are observed on the postcranial skeleton.     

The dentition of Amygdalodon patagonicus (Dinosauria: Sauropoda) and the dental evolution in basal sauropods

Amygdalodon patagonicus is the oldest record of Sauropoda from South America and is known from several teeth and fragmentary postcranial remains. Here we describe in detail its dental morphology, characterized by the presence of broad spatulated teeth (with low SI values) and the absence of denticles in their crowns. The enamel bears a particular wrinkling pattern composed only of apicobasally-aligned pits, which are frequently joined to each other by a continuous sulcus. Some worn teeth have large wear-facets that extend over only one of the crown's edges. This unique combination of characters provides a proper diagnosis for Amygdalodon. Despite the fragmentary available material, Amygdalodon is here interpreted as a non-eusauropod sauropod based on the results of a phylogenetic analysis. The presence of derived dental characters in Amygdalodon, such as the presence of tooth-tooth occlusion, shows several features previously thought to diagnose Eusauropoda or Gravisauria appeared earlier during the early evolution of sauropods.     

NEW GENUS OF DORUDONTINE ARCHAEOCETE (CETACEA) FROM THE MIDDLE-TO-LATE EOCENE OF SOUTH CAROLINA

A new archaeocete whale from the late middle or early late Eocene of South Carolina, Chrysocetus healyorum gen. et sp. nov., is described on the basis of a single subadult specimen. This individual includes: a partial skull; hyoid apparatus; lower jaws; teeth; all cervical, some thoracic and some lumbar vertebrae; ribs and sternum; left forelimb elements; and pelves. The specimen includes portions of much of the body, but while some of the bones are fairly complete, others are damaged, particularly the skull. The pelves resemble those of Basilosaurus , documenting a similar stage of hind limb reduction in dorudontines and suggesting that Chrysocetus was not able to support its body on land. The acetabulum for articulation of the femur is well formed and indicates that the hip joint was functional. Chrysocetus is distinguishable from other described dorudontines based on body size, characteristics of the teeth, and forelimb elements. Absence of deciduous teeth in a subadult individual of Chrysocetus may be indicative of an early stage of the evolution of monophyodonty.     

A new species of Azendohsaurus (Diapsida: Archosauromorpha) from the Triassic Isalo Group of southwestern Madagascar: cranium and mandible

Abstract: Here, we describe a new species of Azendohsaurus from the Middle–Late Triassic of Madagascar, extending the geographical range of a taxon known otherwise only by a single species from Morocco. Although Azendohsaurus has consistently been regarded as an early dinosaur (based on various advanced dental and gnathic features resembling those characterizing certain dinosaur subgroups), the relatively complete skeletal material, now available from Madagascar, argues strongly against its dinosaurian affinities. Rather, the retention of numerous primitive cranial and postcranial features indicates a surprisingly early divergence of Azendohsaurus within Archosauromorpha and an unusual mosaic of characters in this taxon. Features considered diagnostic of Sauropodomorpha thus are inferred to occur homoplastically in at least one clade of nondinosaurian archosauromorphs, indicating a complex evolution and distribution of features traditionally thought to be derived within archosaurs. Azendohsaurus has teeth resembling those of both early sauropodomorph and ornithischian dinosaurs, yet also possesses numerous inarguable basal archosauromorph cranial and postcranial attributes. This highlights the risk of uncritically referring isolated, Middle–Late Triassic (or even later), ‘leaf‐shaped’ teeth with denticles to the Dinosauria. Similarly, the occurrence of such teeth in an early diverging archosauromorph indicates that specializations for herbivory originated more frequently within this clade than conventionally assumed. For example, Azendohsaurus and numerous basal sauropodomorph dinosaur taxa share an array of convergently acquired features associated with herbivory, including tooth denticles, expanded tooth crowns, a downturned dentary and the articular located at the ventral margin of the mandible. Some of these features (denticles, expanded crowns and the ventrally deflected articular) are even more widespread among archosauromorphs, including aetosaurs, silesaurs and ornithischian dinosaurs. A downturned dentary also occurs in Trilophosaurus, a taxon further marked by unique specializations for herbivory, including transversely lophate, tricuspid teeth. An array of features associated with herbivory also occurs in rhynchosaurs and certain crocodilians (e.g. Simosuchus). This distribution suggests that craniodental features associated with herbivory were much more pervasive across the archosauromorph clade than previously recognized, possibly evolving at least six to eight times independently.     

The first juvenile specimen of Eolacerta (Squamata: Eolacertidae) from the early–middle Eocene of the Messel Pit (Germany)

We describe the first juvenile specimen of an eolacertid lizard. The material comes from one of the most important Eocene localities, the Messel Pit in Germany. The new specimen provides unique information on the early ontogeny of Eolacerta, the largest known lizard from Messel, with a maximum snout-vent length greater than 30 cm and a mass approaching 1 kg. The specimen described here, with a SVL of 11.3 cm and an estimated mass of 21 g, can be allocated more precisely to Eolacerta robusta based on the co-occurrence and the combination of following features: (1) the nasal process of premaxilla is long; (2) position of lacrimal (being more anteriorly located in Stefanikia siderea); (3) the postorbital process of jugal is broad; (4) the ratio of the anterior and posterior region of the frontal between the sulcus interfacialis; (5) a mid-parietal constriction of the parietal table is present; (6) the interparietal shield broadens anteriorly; (7) the transverse sulcus is straight anteriorly. The incipient character of the parietal constriction and the slightly lower number of maxillary teeth (28 vs. 30–32 in adults) are consistent with a juvenile animal. Very important is evidence for the presence of pterygoid dentition (pointed teeth arranged in a single line) and the absence of palatine dentition. Ceratobranchial I is observed for the first time for this species, and its shape and length are very similar to those of Lacertidae. There are 27 presacral vertebrae in the juvenile, as in adults. In the juvenile specimen, ventral keel on the centrum is present in all vertebrae. The ventromedial portion of the ischium is well preserved here and gives information on the exact shape of this portion, at least in juvenile form. The scalation, as far as it is preserved, is similar to that of Stefanikia, except that the rectangular subdigital scales are longer in comparison with their width, and therefore have a broader appearance.     

The dentition of a well-preserved specimen of <Emphasis Type="Italic">Camarasaurus</Emphasis> sp.: implications for function,tooth replacement,soft part reconstruction,and food intake

The basal macronarian genus Camarasaurus was the most common sauropod in the Upper Jurassic Morrison Formation of North America and is known from several complete and partial skeletons. The specimen used for this study is Camarasaurus sp. SMA 0002 from the Sauriermuseum Aathal, Switzerland. This specimen was found in the Howe-Stephens Quarry, Bighorn Basin, WY, USA. In this study, the dental morphology, characterized by the spatulate, broad-crowned teeth, the tooth replacement pattern, and the function of the dentition and its implications for food intake is described. Features such as the absence of denticles, the wrinkled pattern of the enamel, and the occurrence of large wear facets on older teeth are characteristic for Camarasaurus sp. A slab of sediment with soft tissue impressions ranging up to the middle part of the crown suggests the presence of a gingival soft tissue structure partially covering the teeth. The wrinkled enamel on the crown of the teeth of Camarasaurus sp. and other sauropods is interpreted as indication of this cover of gingival connective tissue. In addition, there possibly was a keratinous beak, which together with the gingiva held the teeth in the jaw and provided stability for teeth in which the root is almost completely resorbed.     

Living naked: first case of lack of skin-related structures in an elasmobranch,the blackmouth catshark (Galeus melastomus)

As far as is known, in this paper the first case of lacking of skin-related structures (epidermis, stratum laxum, dermal denticles and teeth) in a free-swimming elasmobranch, the blackmouth catshark, Galeus melastomus, is reported. The individual was caught by trawl in Sardinian waters (central-western Mediterranean) in July 2019 at a depth of 500 m. Although this kind of morphological abnormality is potentially fatal, the observations suggested that the specimen was in good health and well developed.     

Chondrichthyan teeth from the Early Triassic Paris Biota (Bear Lake County,Idaho, USA)

A new, diverse and complex Early Triassic assemblage was recently discovered west of the town of Paris, Idaho (Bear Lake County), USA. This assemblage has been coined the Paris Biota. Dated earliest Spathian (i.e., early late Olenekian), the Paris Biota provides further evidence that the biotic recovery from the end-Permian mass extinction was well underway ca. 1.3 million years after the event. This assemblage includes mainly invertebrates, but also vertebrate remains such as ichthyoliths (isolated skeletal remains of fishes). Here we describe first fossils of Chondrichthyes (cartilaginous fishes) from the Paris Biota. The material is composed of isolated teeth (mostly grinding teeth) preserved on two slabs and representing two distinct taxa. Due to incomplete preservation and morphological differences to known taxa, the chondrichthyans from the Paris Biota are provisionally kept in open nomenclature, as Hybodontiformes gen. et sp. indet. A and Hybodontiformes gen. et sp. indet. B, respectively. The present study adds a new occurrence to the chondrichthyan fossil record of the marine Early Triassic western USA Basin, from where other isolated teeth (Omanoselache, other Hybodontiformes) as well as fin spines of Nemacanthus (Neoselachii) and Pyknotylacanthus (Ctenachanthoidea) and denticles have been described previously.     

Paranthodon africanus (broom) a stegosaurian dinosaur from the Lower Cretaceous of South Africa

The holotype of Paranthodon africanus (Broom) from the Kirkwood Formation (Lower Cretaceous) of the Algoa Basin, Cape Province, South Africa consists of a partial skull, the bones of which are very similar to those of Stegosaurus. Both sides of the maxillary tooth crowns have a bulbous cingulum and a very prominent central vertical ridge above the large apical denticle that is bordered anteriorly and posteriorly by four to seven smaller denticles. Diagnostically stegosaurian material is also known from the Lower Cretaceous of England (Craterosaurus) and China (Wuerhosaurus) and the Upper Cretaceous of India (Dravidosaurus).     

A fossil record of the Eastern clade of Blanus (Amphisbaenia: Blanidae) from the late Miocene of Ukraine

The fossil history of worm lizards in Eastern Europe is very patchy because of the small number of localities and the fragmentary vertebral material available. A new record of the genus Blanus from the late Miocene of the Egorovka 1 and 2 localities in Ukraine is reported here. Based on its dental morphology, Blanus from Egorovka differs from all extinct species of the genus and from the extant B. cinereus complex. It is most closely related to the Eastern clade of Blanus (B. strauchi complex) in having enlarged second and small third dentary teeth. Blanus from Egorovka is the second fossil record of the B. strauchi complex and the first record of the Eastern clade outside of its current geographical range. It indicates a wider past range of the Eastern European Blanus, including the northern Black Sea coastal area.     

Tooth morphology of Notosuchus terrestris (Notosuchia: Mesoeucrocodylia): New evidence and implications

Notosuchia is a large and diverse group of Crocodyliforms, characterized, among other features, by a heterodont dentition. New information on the tooth anatomy of Notosuchus terrestris is presented, based on well-preserved specimens from the Late Cretaceous of Patagonia (southern Argentina). This allows a complete characterization of its dental anatomy (composed by incisiviform, caniniform, and molariform teeth) that includes autapomorphic features and derived features shared with Sphagesaurus and Mariliasuchus. This includes the extensive wear facets in molariforms, indicative of tooth–tooth occlusion and a sharp keel that bears rounded denticles. Notosuchus also shares with Mariliasuchus the presence of a tooth with a transitional morphology located at the premaxilla–maxilla contact and the absence of interalveolar septa in the entire premaxillary and maxillary dentition.     

Late Cretaceous crocodile remains from Naskal (India): comparisons and biogeographic affinities

Crocodile teeth from the Maastrichtian inter-trappean beds of Naskal (peninsular India) are described here. Because of isolated denticles visible on sufficiently preserved carinae, the presence of a strong heterodonty (in size and shape), and by comparison to crocodile teeth from various taxa, they are considered as representing a ziphodont form with a heterodont dentition. The difference between ziphodont, “false ziphodont” and non-ziphodont dentitions is evaluated. With the help of scanning electron microscope photographs, it is shown that only precise characteristics of the denticles and not the tooth shape, allow to distinguish the three categories. These three categories do not correspond to monophyletic groups. It is also shown that the “alligatorid” heterodonty, meso- or eusuchian in grade, exists in each category. Although the ziphodont dentition is not sufficient to allow a taxonomical definition, the peculiarities that it often presents, depending on the taxa as well as the teeth shape, enable systematic approaches. An examination of previous works on the possible ziphodont crocodiles from the Tertiary deposits of the Indian subcontinent and on Naskal teeth demonstrate that the latter are closer to those of some Gondwanan crocodiles of mesosuchian grade, known from the early Cretaceous of Africa and possibly a form from the late Cretaceous of Madagascar. They are excluded from eusuchian Laurasiatic as well as Paleogene forms of the Indian subcontinent, either ziphodont or not. Contrary to the earlier works on the inter-trappean crocodiles, the present study removes this group as one of the evidences in support of an early (late Cretaceous-early Tertiary) India/Asia collision model. In fact, it provides an additional support for the existence of possible Cretaceous biogeographic links between India, Madagascar, Africa, and South America.     

Development of the pharyngeal teeth in the big head,Aristichthys nobilis (Cyprinidae)

The development of pharyngeal dentition was observed in the big head,Aristichthys nobilis, which is one of the hypophthalmichthyines of the cyprinids. This fish has the C-type larval dentition, in which no teeth ever occur at the position An3, and in which the first tooth at the position An2 is on the third replacement wave. So the positions Pol, Ce0, Ani and An2 in the larval dentition correspond to the positions A4, A3, A2 and A1 in the adult dentition, respectively. The initial tooth at each position is a conical one. The conical teeth are then changed to ones bearing a narrow grinding surface with a hook at the tip and some denticles on the margins. These teeth are of theLeuciscus stage. Tn the following teeth, the grinding surface is expanded, and the denticles are increased in number and distributed on not only the margins but also the whole grinding surface. These teeth bearing a very broad grinding surface characterize the hypophthalmichthyines. At the positions A2 to A4, the teeth become the hypophthalmichthyine type in the larval period. But the tooth at the position A1 becomes the hypophthalmichthyine type in the juvenile period. The morphological change of teeth in this species is simple although their teeth are highly specialized. We think that this phenomenon gives a hint on their phylogeny.     

Dysomma alticorpus,a new species of cutthroat eel from the Gulf of Aqaba,Red Sea (Teleostei: Synaphobranchidae)

The cutthroat eel Dysomma alticorpus n. sp. is described based on a single specimen collected in a trammel net at a depth of 350 m off Eilat, Israel, Gulf of Aqaba, Red Sea. The new species belongs to the Dysomma anguillare species complex, which comprises species possessing a well-developed pectoral fin, intermaxillary teeth, a uniserial row of 7–15 large compound teeth in the lower jaw (which may be followed by a few smaller teeth), and an anteriorly situated anus with the trunk shorter than the head length. It is characterised by a combination of the following characters: origin of the dorsal fin well anterior to the base of the pectoral fin, predorsal length 13.8% TL; preanal length 22.8% TL; three compound teeth on the vomer; head pores: IO 4, SO 3; M 6; POP 0; AD 1, F 0, ST 0; lateral-line pores: predorsal 4, prepectoral 8, preanal 14, total 57–58, the last at the posterior two-thirds of the total length; MVF 7–16–115; total vertebrae 115. Dysomma alticorpus n. sp. is compared with other species of the genus. A revised key to the species of the genera Dysomma and Dysommina is provided.     

On the spines of the Stethacanthidae (Chondrichthyes), with a description of a new genus from the Mississippian Bear Gulch limestone

Several specimens of an elasmobranch from the upper Mississippian Bear Gulch limestone of Montana are described. The first dorsal spine and fin closely resemble Stethacanthus erectus and to a lesser extent Stethacanthus altonensis. The new genus and species Orestiacanthus fergusi is proposed for the specimens, and it is suggested that recent attempts to synonymize the species of Stethacanthus on the basis of spines are unjustified. The specimens also share the enlarged cranial denticles, general form of the palatoquadrate, cladodont teeth and pectoral fin with S. altonensis. They differ from S. altonensis in proportions of jaws, numbers of tooth families, numbers of prearticular basals and axial radials, morphology of the second dorsal fin, and in the presence of generalized squamation in some specimens (but not the female). The pelvic fins differ greatly in having a prominent metapterygium.     

Early development of rostrum saw-teeth in a fossil ray tests classical theories of the evolution of vertebrate dentitions

In classical theory, teeth of vertebrate dentitions evolved from co-option of external skin denticles into the oral cavity. This hypothesis predicts that ordered tooth arrangement and regulated replacement in the oral dentition were also derived from skin denticles. The fossil batoid ray Schizorhiza stromeri (Chondrichthyes; Cretaceous) provides a test of this theory. Schizorhiza preserves an extended cartilaginous rostrum with closely spaced, alternating saw-teeth, different from sawfish and sawsharks today. Multiple replacement teeth reveal unique new data from micro-CT scanning, showing how the ‘cone-in-cone’ series of ordered saw-teeth sets arrange themselves developmentally, to become enclosed by the roots of pre-existing saw-teeth. At the rostrum tip, newly developing saw-teeth are present, as mineralized crown tips within a vascular, cartilaginous furrow; these reorient via two 90° rotations then relocate laterally between previously formed roots. Saw-tooth replacement slows mid-rostrum where fewer saw-teeth are regenerated. These exceptional developmental data reveal regulated order for serial self-renewal, maintaining the saw edge with ever-increasing saw-tooth size. This mimics tooth replacement in chondrichthyans, but differs in the crown reorientation and their enclosure directly between roots of predecessor saw-teeth. Schizorhiza saw-tooth development is decoupled from the jaw teeth and their replacement, dependent on a dental lamina. This highly specialized rostral saw, derived from diversification of skin denticles, is distinct from the dentition and demonstrates the potential developmental plasticity of skin denticles.     

Bone-Breaking Bite Force of Basilosaurus isis (Mammalia,Cetacea) from the Late Eocene of Egypt Estimated by Finite Element Analysis

Bite marks suggest that the late Eocence archaeocete whale Basilosaurus isis (Birket Qarun Formation, Egypt) fed upon juveniles of the contemporary basilosaurid Dorudon atrox. Finite element analysis (FEA) of a nearly complete adult cranium of B. isis enables estimates of its bite force and tests the animal’s capabilities for crushing bone. Two loadcases reflect different biting scenarios: 1) an intitial closing phase, with all adductors active and a full condylar reaction force; and 2) a shearing phase, with the posterior temporalis active and minimized condylar force. The latter is considered probable when the jaws were nearly closed because the preserved jaws do not articulate as the molariform teeth come into occulusion. Reaction forces with all muscles active indicate that B. isis maintained relatively greater bite force anteriorly than seen in large crocodilians, and exerted a maximum bite force of at least 16,400 N at its upper P³. Under the shearing scenario with minimized condylar forces, tooth reaction forces could exceed 20,000 N despite lower magnitudes of muscle force. These bite forces at the teeth are consistent with bone indentations on Dorudon crania, reatract-and-shear hypotheses of Basilosaurus bite function, and seizure of prey by anterior teeth as proposed for other archaeocetes. The whale’s bite forces match those estimated for pliosaurus when skull lengths are equalized, suggesting similar tradeoffs of bite function and hydrodynamics. Reaction forces in B. isis were lower than maxima estimated for large crocodylians and carnivorous dinosaurs. However, comparison of force estimates from FEA and regression data indicate that B. isis exerted the largest bite forces yet estimated for any mammal, and greater force than expected from its skull width. Cephalic feeding biomechanics of Basilosaurus isis are thus consistent with habitual predation.     

First discovery of the Miocene northern Atlantic sperm whale Orycterocetus in the Mediterranean

An incomplete mandible of a fossil sperm whale (Odontoceti; Physeteridae) is described. It is a portion of elongated and narrow symphysis with cylindrical and slightly curved teeth without enamel. The specimen, collected from the Miocene “Pietra leccese” sediments of South Italy, appears to represent a species of Orycterocetus, and is the first such record from the Mediterranean. Considering the wide diffusion of Orycterocetus in the North Atlantic, we believe its rare presence in the Mediterranean as occasional and due to a certain degree of interchange between the Mediterranean and the North Atlantic cetacean fauna during the Miocene.     

The fin spine of a new holocephalan from the lower jurassic of Lyme Regis,Dorset, England

A fin spine previously described as ‘Myriacanthus paradoxus’ Agassiz from the Lower Lias (Lower Jurassic) of Lyme Regis, is allocated to the new genus and species Recurvacanthus uniserialis. The spine is unusual in possessing a single median row of four large, hook-like, downturned denticles on the distal part of the posterior wall. It is concluded that the unique specimen should be classified with the myriacanthoid chimaeriform holocephalans since it possesses a tubercular ornament on the lateral walls.