David Martill

• BSc University of Leicester
• Lecturer at University of Portsmouth

The diets and feeding strategies of pterosaurs remain a poorly known although speculatively debated topic in vertebrate paleontology. Fossilized gut contents, which offer a crucial direct line of evidence to help decipher these elusive questions, are only known from a handful of pterosaur specimens in a few notable fossil Lagerstätten, such as the...

An incomplete, but large hollow bone is identified as pterosaurian on account of its pneumatic nature, thin bone wall, palaeohistology and the characteristic surface texture of the peridermal bone. Despite being incomplete it is tentatively identified as a ?left ulna, and is remarkable for its size. Coming from the lower Middle Jurassic Lincolnshir...

An enigmatic partial trackway from the Aalenian to Bathonian (Middle Jurassic) Ravenscar Group of Whitby, North Yorkshire bears a striking resemblance to the pterosaur trace Pteraichinus. It also displays some similarities with the xiphosuran trackway Kouphichnium. The trackway is assigned to a new ichnogenus and ichnospecies Pseudopteraichnus whit...

A partial caudal fin of a large-bodied asthenocormine pachycormiform (Pachycormiformes: Pachycormidae: Asthenocorminae) is described from the Upper Jurassic (lower Tithonian) Kimmeridge Clay Formation of Dorset, England. The specimen comprises associated fin rays, basal fulcra, ‘epurals’ and paired pre-caudal scutes, the combination and morphology...

Well-defined manus and pes print couple, likely from a trackway attributable to a sauropod dinosaur are revealed on a fallen block of Ravenscar Group sandstones in a cliff fall at Whitby, Yorkshire, England. The pes print displays four, possibly five toes and thus closely resembles that of the ichnogenera Brontopodus and Parabrontopodus. They are h...

Grey Seals, Halichoerus grypus, are abundant, native pinnipeds to the waters of the Isle of Wight and Solent, UK reappearing after an absence of ~three decades. The White-tailed Eagle, Haliaeetus albicilla, became extinct on the south coast and the Isle of Wight around 1780, but were re-introduced in 2019. Presently, only a few eagle pairs have rem...

An isolated jaw fragment from the Late Cretaceous (Cenomanian) Cambridge Greensand Member of the West Melbury Marly Chalk Formation previously identified as a cestraciontid shark fin spine is referred to the pterosaur clade Azhdarchoidea on account of its lateral and occlusal foramina and edentuly. The specimen differs from the azhdarchoid Ornithos...

The pterosaur assemblage of the mid-Cretaceous Kem Kem Group of Morocco is reviewed. This analysis examines their taxonomy, palaeoecology and palaeobiology with comments on taphonomy. New material permits the rediagnosis of the azhdarchoids Alanqa saharica and Afrotapejara zouhrii. Several specimens are reported that do not ft within the paradigms...

A B S T R A C T The Lower Cretaceous of England has produced a diverse assemblage of dinosaurs, including ornithischians, sauropods, and theropods. The origins of this assemblage are poorly understood. Here, we describe a new dromaeosaurid, Vectiraptor greeni gen. et sp. nov., from the Barremian Wessex Formation of the Isle of Wight. The animal is...

In our recent paper discussing the impacts of dragonfly behaviour on their taphonomy from the Lower Cretaceous Crato Formation of Brazil (Barling et al., 2021), we figured eight specimens (Fig. 3, p. 145) illustrating dragonflies in various states of preservation. In the caption to this figure we listed the temporary laboratory numbers assigned to...

The relative completeness of Odonata (dragonflies and damselflies) fossils from the Crato Formation is investigated and compared to other fossil insect groups from the same formation. Tagma completeness is measured as present, partial, or absent, with some additional subdivision of body components (head, thorax, limbs, individual wings, anterior an...

Azhdarchid pterosaurs, the largest flying vertebrates, remain poorly understood, with fundamental aspects of their palaeobiology unknown. X-ray computed tomography reveals a complex internal micro-architecture for three-dimensionally preserved, hyper-elongate cervical vertebrae of the Cretaceous azhdarchid pterosaur, Alanqa sp. Incorporation of the...

The Early Cretaceous Wealden Group of Swanage Bay, Dorset, southern England, comprises two formations, a lower Wessex Formation and an overlying Vectis Formation. Presently only part of the former is exposed and here its stratigraphy, sedimentology and palaeontology are redescribed. Recent work on the Wealden Group of the Wessex Sub-basin has focus...

Several specimens from the Lias Group (Lower Jurassic) of Northern Ireland have been suspected as dinosaurian in origin. Bone histology and morphology demonstrates that two of these, both from the same locality in Co. Antrim, demonstrably are from dinosaurs. We interpret one as the proximal end of the left femur of a basal thyreophoran ornithischia...

The pterosaurs were the first vertebrates to evolve powered flight and they staged a major radiation in the Cretaceous. Cretaceous pterosaurs occupied many of the niches occupied today by birds, including aerial insect hawkers, piscivores, and filter feeders. The diversity of this radiation remains poorly known due to the uneven stratigraphic and g...

The sense of touch is important for hunting and feeding in vertebrates, especially when visual cues are unreliable. Foramina in the jaws and face, associated with nerves and sensory organs, may provide information about feeding. Pterosaurs, many of which had large, well-developed eyes, are often assumed to have been visual feeders. Here, we show th...

A new locality near Tarda on the northern margin of the Tafilalt, south eastern Morocco exposes extensive sequences of the Ifezouane and Aoufous formations of the fluvial Kem Kem Group (Cretaceous, ?Albian-Cenomanian) on the south western flank of Ikfh n’Oufza escarpment of the Hamada du Meski. The stratigraphic sequence here differs significantly...

The Crato Formation Lagerstätte of NE Brazil is well known for the exceptional preservation of its fossil assemblage. A wide variety of plants and animals are preserved with a high degree of fidelity, revealing structures of labile tissues at the micron scale, or even smaller. Chemical analyses and descriptions of replacement fabrics observed in in...

Remains of enigmatic spinosaurs from mid-Cretaceous North African strata have, for over a century, been the subject of taxonomic deliberations. The gigantic Spinosaurus aegyptiacus Stromer, 1915 has gained iconic status in the vertebrate palaeontological community and amongst the general public. Perhaps the largest predatory dinosaur to have lived,...

In recent decades, intensive research on non-avian dinosaurs has strongly suggested that these animals were restricted to terrestrial environments¹. Historical proposals that some groups, such as sauropods and hadrosaurs, lived in aquatic environments2,3 were abandoned decades ago4–6. It has recently been argued that at least some of the spinosauri...

The geological and paleoenvironmental setting and the vertebrate taxonomy of the fossiliferous, Cenomanian-age deltaic sediments in eastern Morocco, generally referred to as the “Kem Kem beds”, are reviewed. These strata are recognized here as the Kem Kem Group, which is composed of the lower Gara Sbaa and upper Douira formations. Both formations h...

A new pterosaur, Afrotapejara zouhri gen. et sp. is described on the basis of a partial rostral fragment from the Cretaceous Kem Kem beds of Takmout, near Erfoud in southern Morocco. The taxon is distinguished from all other Tapejaridae on the possession of a dorsal expansion of the rostral margin a short distance from the rostral tip. Tapejarid fe...

A new genus and species of edentulous pterodactyloid pterosaur with a distinctive partial rostrum from the mid-Cretaceous (?Albian/Cenomanian) Kem Kem beds of southeast Morocco is described. The taxon is assigned to Chaoyangopteridae based upon its edentulous jaws, elongate rostrum and slightly concave dorsal outline. The rostral cross-section is r...

Pterodactyloid pterosaurs underwent a diversification in the Late Jurassic and Early Cretaceous, followed by a major turnover event in the mid-Cretaceous, when ornithocheiroids and basal azhdarchoids were replaced by pteranodontids, nyctosaurids and azhdarchids in the latest Cretaceous. However, precise patterns of turnover are obscured by the inco...

A new ichthyosaur specimen (MJML K1885) from the Late Jurassic (Kimmeridgian) Kimmeridge Clay Formation of Dorset, United Kingdom is described. The specimen, expertly prepared on a slab of laminated coccolith limestone, comprises a near complete skull, in articulation with the anterior vertebral column and associated thoracic ribs, complete pectora...

Isolated cervical vertebrae from the mid Cretaceous Kem Kem beds of south east Morocco are referred to the theropod dinosaur clade Abelisauroidea, and represent the first axial remains from this deposit referred to this group. An isolated axis is referred to Abelisauroidea on account of the invaginated spinopostzygapophyseal lamina; the extremely l...

The Cretaceous succession exposed in the cliffs and on the foreshore at Yaverland on the south-east coast of the Isle of Wight is one of the Wessex Sub-basin’s classic vertebrate fossil sites. Its history stretches back to the time of Dean William Buckland who in 1829 just five years after scientifically describing the World’s first dinosaur, Magal...

The Aldabra rail, Dryolimnas cuvieri subsp. aldabranus, endemic to the Aldabra Atoll, Seychelles, is the last surviving flightless bird in the Indian Ocean. Aldabra has undergone at least one major, total inundation event during an Upper Pleistocene (Tarantian age) sea-level high-stand, resulting in the loss of all terrestrial fauna. A flightless D...

A partial distal left metacarpal IV of a pterodactyloid pterosaur is notable for its size. The specimen, from the Early Cretaceous (Barremian)Wessex Formation exposed on the southwest coast of the Isle of Wight, southern England is identified as an ornithocheirid, perhaps close to the Ornithocheirus, Coloborhynchus, Caulkicephalus, Anhanguera compl...

Documented here is a variation of the widely used resin transfer technique, allowing for the preparation of fragile articulated fossil arthropod material preserved in laminated limestone. The extensive use of the resin transfer technique in palaeontology has traditionally been restricted to palaeobotany and vertebrate palaeontology. The parameters...

The Aldabra rail, Dryolimnas cuvieri subsp. aldabranus, endemic to the Aldabra Atoll, Seychelles, is the last surviving flightless bird in the Indian Ocean. Aldabra has undergone at least one major, total inundation event during an Upper Pleistocene (Tarantian age) sea-level high-stand, resulting in the loss of all terrestrial fauna. A flightless D...

The current understanding of UK Middle Jurassic pterosaur taxonomy is under-developed, leading to it being previously considered a time of low diversity. This is despite the presence of a productive but under-studied pterosaur-bearing horizon extending over parts of Oxfordshire and Gloucestershire. This unit, informally called the Stonesfield Slate...

A large, slender, recurved tooth with crenulated ornamentation and arcuate enamel border from the Oum ed Diab Member of the Aïn el Guettar Formation (Early Cretaceous, Albian) of southern Tunisia is assigned to the Ornithocheiroidea (Pterosauria, Pterodactyloidea). It is most similar to the teeth of Coloborhynchus Owen (1874) and Ludodactylus Frey...

[This corrects the article DOI: 10.1371/journal.pbio.2001663.].

The Pleistocene vertebrate assemblage of Aldabra Atoll has been comparatively well studied. Three Upper Pleistocene fossil localities have been described yielding birds, reptiles and terrestrial molluscs. Those of Bassin Cabri and Bassin Lebine on Ile Picard are undated but must be in excess of 136,000 YBP, whereas Point Hodoul on Malabar Island is...

Data deposition. All fossil material in the text is deposited and accessioned in an appropriate public repository, the United States National Museum (USNM). All fossil material was loaned by the USNM for study and will be returned for permanent depository and public availability. (DOCX)

Pterosaurs were the first vertebrates to evolve powered flight and the largest animals to ever take wing. The pterosaurs persisted for over 150 million years before disappearing at the end of the Cretaceous, but the patterns of and processes driving their extinction remain unclear. Only a single family, Azhdarchidae, is definitively known from the...

Distal humerus of Tethydraco and Pteranodon. Comparison of Pteranodon YPM 1175 in ventral (A) and dorsal (C) views to T. regalis FSAC-OB 1 in ventral (B) and dorsal (D) views, showing the different degree of development of the entepicondyle and ectepicondyle. (JPG)

Femora of Tethydraco and Pteranodon. Femora of (A) T. regalis FSAC 201 and Pteranodon (B) YPM 2597 and (C) YPM 1175. (JPG)

Vectorized version of Fig 20. (EPS)

Catalogue of specimens examined in the course of this study. (CSV)

Functional diversity character-taxon matrix and references. (XLSX)

Humeri of Tethydraco and Pteranodon. T. regalis FSAC-OB 1 (A) compared to Pteranodon (YPM 2709) (B). Arrows denote the position of the base of the deltopectoral crest and the ulnar crest. (JPG)

Ulnae of Tethydraco and Pteranodon. (A) T. regalis FSAC-OB 199 and Pteranodon (B) YPM 2499, (C) YPM 2497, and (D) YPM 2409. (JPG)

Variation in the humeri of Alcione. (JPG)

Character-taxon matrix for phylogenetic analysis. (TXT)

Humeri of Alcione, Simurghia, and Barbaridactylus. (A) A. elainus FSAC-OB 5 to (B) S. robusta FSAC-OB 7, and (C) B. grandis FSAC-OB 8. (TIF)

Notes on provenance and stratigraphy of the phosphate pterosaurs; systematics and taxonomy; discussion of affinities of P. barbarulna; and age of “N.” lamegoi. (DOCX)

Two tribosphenic teeth recently recovered from the Purbeck Group ‘Mammal Bed’ (Lower Cretaceous, Berriasian) exposed in Durlston Bay, east Dorset, UK, have been shown to be those of highly derived eutherian mammals. They have been named Durlstotherium newmani and Durlstonodon ensomi. The ‘Mammal Bed’ was, until the latter part of the twentieth cent...

2017. Highly derived eutherian mammals from the earliest Cretaceous of southern Britain. Acta Palaeontologica Polonica 62 (4): 657-665. Eutherian mammals (Placentalia and all mammals phylogenetically closer to placentals than to marsupials) comprise the vast majority of extant Mammalia. Among these there is a phenomenal range of forms and sizes, bu...

Pterosaurs were the first vertebrates to evolve powered flight and occupied the skies of the Mesozoic for 160 million years. They occurred on every continent, evolved their incredible proportions and anatomy into well over 100 species, and included the largest flying animals of all time among their ranks. Pterosaurs are undergoing a long-running sc...

Eutherian mammals (Placentalia and all mammals phylogenetically closer to placentals than to marsupials) comprise the vast majority of extant Mammalia. Among these there is a phenomenal range of forms and sizes, but the origins of crown group placentals are obscure. They lie within the generally tiny mammals of the Mesozoic, represented for the mos...

A new genus and species, Xericeps curvirostris gen. et sp. nov., is erected for a highly distinctive pterosaur mandible from the mid-Cretaceous (?Albian to lower Cenomanian) Kem Kem beds of south east Morocco. The new taxon is referred to Azhdarchoidea based on the absence of teeth, slenderness of its mandible with sulcate occlusal surface, presenc...

A pterosaur humerus (PRC 64) from the Upper Jurassic of Thailand was initially assigned to the Azhdarchoidea, an important clade of edentulous Cretaceous pterodactyloid pterosaurs. Here it is reassigned, on the basis of morphological comparisons and a phylogenetic analysis, to the Rhamphorhynchidae, a widely distributed clade of small to medium-siz...

The Solnhofen pterosaurs Pterodactylus antiquus, Aerodactylus scolopaciceps, Diopecephalus kochi, Germanodactylus cristatus and Germanodactylus rhamphastinus all have complicated taxonomic histories. Species originally placed in the genus Pterodactylus, such as Aerodactylus scolopaciceps, Ardeadactylus longicollum, Cycnorhamphus suevicus and German...

The Eyam Limestone Formation of Steeplehouse Quarry, Wirksworth, Derbyshire, UK yields a diverse assemblage of Lower Carboniferous vertebrate remains. The assemblage is dominated by dermal denticles of the enigmatic selachian Petrodus patelliformis M’Coy, 1848, but also contains teeth of petalodonts, hybodonts and neoselachians. Actinopterygian rem...

A new specimen of the neosuchian crocodylomorph, Susisuchus sp. from the Lower Cretaceous (Aptian) Crato Formation of Ceará, North East Brazil is remarkable for extensive preservation of the epidermis and limb musculature. The specimen comprises incomplete post-cranial remains, including an articulated sequence of thirteen thoracic vertebrae, a dis...

Complete or near-complete skeletons of the herbivorous dinosaur Hypsilophodon foxii occur frequently in a metre-thick band of mudstone and sandstone in the Lower Cretaceous Wessex Formation of the Isle of Wight. The reasons for this accumulation have been the subject of some debate. This article examines new sedimentological clues that provide a pl...

A juvenile specimen of the ammonite Pseudaspidoceras from the Early Turonian of the Goulmima area in the Province of Er-Rachida in south-eastern Morocco shows clear evidence of predation by a tooth-bearing vertebrate. Most of the body chamber is missing, as a result of post-burial compactional crushing. The adapertural part of the shell on the left...

A specimen of a pterodactyloid pterosaur from the Upper Tunbridge Wells Sand Formation (Early Cretaceous, Valanginian) of Bexhill, East Sussex, southern England is described. It comprises a small fragment of jaw with teeth, a partial vertebral column and associated incomplete wing bones. The juxtaposition of the bones suggests that the specimen was...

Six specimens accessioned to the Bavarian State Collection for Palaeontology and Geology in Munich, Germany, in 1966 are identified as coming from a gigantic pterodactyloid pterosaur. The previously undescribed material was obtained in 1955 by Jean Otto Haas and compares favourably in size with the type specimen of the Late Cretaceous (Maastrichtia...

The Lower Jurassic (Toarcian) pterosaur Parapsicephalus purdoni Newton, 1888 from the Whitby Mudstone Formation of North Yorkshire is known from a three-dimensionally preserved skull with a three-dimensional endocast of the brain. Since Newton’s original description, its taxonomic status has been contentious. Several cladistic studies have placed i...

Fossil snakes are relatively well represented in the Upper Cretaceous of northern Africa, with material known from Morocco, Sudan, Egypt, Libya, Algeria, and Niger. The Moroccan Kem Kem beds yield a particularly diverse snake assemblage, with Simoliophiidae, Madtsoiidae, ?Nigerophiidae and several unnamed taxa co-occurring. These fossils are import...

Carbonate concretions hosted within organic carbon-rich shale sequences represent unique archives of often exceptionally preserved fossil biota. Besides providing high-fidelity preservation, their geochemical signatures can provide insight into the physical and chemical processes during early and later-stage concretion growth. Here, two fossilifero...

Invertebrate bioerosion including unusually large borings are reported for the first time in dinosaur bone from the Vectis Formation (Early Cretaceous, Barremian-Aptian) of Compton Bay, Isle of Wight. Sub-circular tunnels of up to 15mm diameter and 60mm maximum length penetrate trabecular bone from a large theropod (cf. Baryonyx sp.) pelvis. The sp...

The late Middle Jurassic (Callovian) Peterborough Member of the Oxford Clay Formation of eastern England yields a rich variety of marine vertebrate fossils, including a diverse assemblage of neoselachian elasmobranchs. Here we report the first record of the small Jurassic orectolobiform shark Akaimia Rees, 2010, otherwise known only from Poland and...

A well preserved middle caudal vertebra from middle Cretaceous (?Albian–lower Cenomanian) deposits informally known as the “Kem Kem beds” exposed in the Gara Sbaa region of Morocco is attributed to a large-bodied titanosaurian sauropod dinosaur. It represents one of the best-preserved and most complete skeletal elements reported for this sauropod g...

Approximately 40% of a skeleton including cranial and postcranial remains representing a new genus and species of basal neotheropod dinosaur is described. It was collected from fallen blocks from a sea cliff that exposes Late Triassic and Early Jurassic marine and quasi marine strata on the south Wales coast near the city of Cardiff. Matrix compari...

Snakes are a remarkably diverse and successful group today, but their evolutionary origins are obscure. The discovery of snakes with two legs has shed light on the transition from lizards to snakes, but no snake has been described with four limbs, and the ecology of early snakes is poorly known. We describe a four-limbed snake from the Early Cretac...

Fossil insects from the Lower Cretaceous (Aptian) Crato Formation of north-east Brazil are preserved as goethite replacements in laminated limestones of lacustro-lagoonal origin. They display remarkable degrees of morphological detail down to the macromolecular level in some examples. We document the fidelity of preservation and reveal an astonishi...

The evolution of pterodactyloid pterosaurs occurred in a ‘modular’ fashion with ‘pterodactyloid’-type crania and cervical vertebrae evolving in pterodactyloid sister taxa – early monofenestratan pterosaurs – before later postcervical modifications marked the development of the true pterodactyloid condition. This means of evolution creates problems...

The second pterosaur genus to be established, Rhamphorhynchus von Meyer, 1847, has historically been used as a wastebasket material. Several species have been erected for fossils found in Europe and Africa, the majority of which are based on non-diagnostic material. Following Bennett's (1996) review of its taxonomy, Rhamphorhynchus is generally reg...

A new specimen comprising a fragmentary pterosaur rostrum can be referred to the ornithocheirid genus Coloborhynchus sp. on account of its anteriorly directed anterior-most teeth located on an upturned palatal surface. The new specimen from the Early Cretaceous Wessex Formation, Wealden Group of the Isle of Wight, southern England is the first reco...

Discovered over a century ago in Cretaceous rocks in Egypt, Spinosaurus aegyptiacus has remained an enigma, as all of the original associated bones were destroyed in a WW2 air raid and all that has come to light since are isolated specimens. In 2008, a partial skeleton was discovered by local collectors from the lower unit of the mid-Cretaceous Kem...

The taxonomy of the Late Jurassic pterodactyloid pterosaur Pterodactylus scolopaciceps Meyer, 1860 from the Solnhofen Limestone Formation of Bavaria, Germany is reviewed. Its nomenclatural history is long and complex, having been synonymised with both P. kochi (Wagner, 1837), and P. antiquus (Sömmerring, 1812). The majority of pterosaur species fro...

Bulk screening of Lower Cretaceous Wessex Formation (Wealden Group, Barremian) plant debris beds exposed on the south-east and south-west coasts of the Isle of Wight, southern England, has resulted in the recovery of large quantities of isolated chondrichthyan and osteichthyan fish remains. Among these are teeth representing five species of hybodon...

We describe adaptations for a semiaquatic lifestyle in the dinosaur Spinosaurus aegyptiacus. These adaptations include retraction of the fleshy nostrils to a position near the mid-region of the skull and an elongate neck and trunk that shift the center of body mass anterior to the knee joint. Unlike terrestrial theropods, the pelvic girdle is downs...

Pterosaur fossils are among the rarest and their eggs are even rarer. How then can we get an insight into the lifestyle of these flying reptiles? A wealth of new pterosaur fossils, including eggs, from China now provides exactly that.

We describe an extensive ichnofossil assemblage from the likely Cenomanian-age 'lower' and 'upper' units of the 'Kem Kem beds' in southeastern Morocco. In the lower unit, trace fossils include narrow vertical burrows in cross-bedded sandstones and borings in dinosaur bone, with the latter identified as the insect ichnotaxon Cubiculum ornatus. In th...

Pterosaur pelvic girdles are complex structures that offer a wealth of phylogenetic and biomechanical information, but have been largely overlooked by pterosaur anatomists. Here, we review pterosaur pelvic morphology and find significant differences that correlate well with pterosaur clades identified in some phylogenetic analyses. We find that the...

A framed water colour of a noctivagous pterosaur by the Reverend G. E. Howman displayed on a wall on the first floor of the Philpot Museum, Lyme Regis, Dorset is the earliest depiction of a restored pterosaur in its life environment. The image is a naïve effort based more on mythology than on fact, but its haunting mood was a harbinger of antediluv...

The Early Cretaceous ornithocheiroid pterosaur Istiodactylus latidens (Pterosauria, Archosauria, Reptilia) from the Wealden Supergroup of the Isle of Wight, southern England, has a distinctive dentition of labio-lingually compressed, sub-triangular tooth crowns with a triangular osseous projection at the dentary symphysis that occluded into a simil...