Despite more than a century of collecting, resulting in one of the best-studied vertebrate fossil records anywhere in the world, the Upper Jurassic Morrison Formation has produced surprisingly few examples of dinosaur eggs associated with embryonic remains. Even more puzzling, none of these seem to pertain to the theropod Allosaurus, one of the most common and best-understood dinosaur taxa in the formation. Here we report on a dinosaur nest site that has produced both abundant prismatoolithid eggshell and embryonic (or perinatal) bones of Allosaurus from Fox Mesa, Wyoming. This represents the first such discovery for any theropod in the Jurassic of North America. The nest is heavily weathered but contains a few ellipsoid eggshell clusters that suggest an egg size of about 8 x 6.5 cm. Study of the eggshell morphology and microstructure confirms that a single egg type is present throughout, which is indistinguishable from Prismatoolithus coloradensis. All of the identifiable embryonic materials pertain to theropods, and two premaxillae specimens show the five alveoli diagnostic for Allosaurus among Morrison theropods. This confirms the theropod origin of Prismatoolithus eggs and implicates Allosaurus as the specific Morrison parent taxon. As a result, it is now possible to assign several previous discoveries of dinosaur eggs and potential nests to Allosaurus, including the isolated egg from the Cleveland-Lloyd Quarry. This discovery also calls into question prior assignments of Prismatoolithus eggs to ornithopods, and suggests that more detailed study of such sites is warranted. Prismatoolithus eggshells are also associated with the Upper Jurassic theropod Lourinhanosaurus from Portugal, along with larger embryos that exhibit four premaxillary alveoli.

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The Upper Jurassic of Portugal has a rich vertebrate fauna well documented from both body and trace fossils. Although the occurrence of crocodyles and pterosaurs is well documented from body fossils, trace fossils from both groups were unknown until now. Here we describe an isolated crocodyle-like track from Praia da Peralta and pterosaur tracks from the Kimmeridgian of Pedreira do Avelino, Sesimbra (Azóia Fm.) and Porto das Barcas, Lourinhã (Lourinhã Fm.). An enigmatic track suggests the possible presence of a small, tail-dragging tetrapod. Possible track-makers are suggested based on the known Late Jurassic vertebrate fauna of Portugal.

THE CRETACEOUS SKELETON COAST OF ANGOLA JACOBS, Louis, SMU, Dallas, TX, USA; POLCYN, Michael, SMU, Dallas, TX, USA; MATEUS, Octávio, Museu da Lourinhã, Lourinhã, Portugal; SCHULP, Anne, Natuurhistorisch Museum Maastricht, Maastricht, Netherlands; NETO, André , Universidade Agostinho Neto, Luanda, Angola Cretaceous coastal sediments of Angola present a rich and diverse fauna of marine amniotes, including turtles, mosasaurs, and plesiosaurs. The abundance of mosasaurs in particular suggests a highly productive coastal area. Angola today lies at the northern limit of the Namibian Desert, the so-called Skeleton Coast, which results from prevailing southeasterly winds of the descending limb of the southern Hadley Cell sweeping across the African coast. The Benguela upwelling and a highly productive sea are found today off the Namibian Desert coast. However, the Benguela upwelling system, based on results of DSDP studies, is said to have originated in the late Neogene and therefore cannot explain the productivity found along the length of the West African coast. The explanation is found in the northward drift of Africa through the arid climate zone, and is demonstrated by the tracing of the paleogeographic position of fossil localities through time.

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In the early 20thcentury, Belgian naturalists reported Paleocene and Eocene sharks, the bothremydid pleurodiran turtleTaphosphrys(formerlyBantuchelys), and a neosuchian and the dyrosaurid crocodyliformCongosaurusfrom coastal outcrops near Landana in the northern province of Cabinda, Angola. In 1935, rare and fragmentary mammals were reported from strata at Malembo Point, south of Landana, and originally considered to be Miocene in age. Subsequent re-identification of the mammal fossils suggested that the Malembo deposits are Lower Oligocene based on the presence of an arsinoithere and hyracoids. An anthropoid canine, originally identified as a carnivore incisor, was also recognized. In four recent expeditions to Cabinda, Projecto PaleoAngola has traced and measured the stratigraphy between Landana and Malembo Point and sampled these deposits for pollen, stable isotopes, detrital zircons, paleomagnetic stratigraphy, and for U/Pb dating of bones, teeth, and coprolites. Although the faunas from all Cabinda localities are dominated by sharks and rays, new discoveries from Landana include a complete cheloniid cryptodire turtle skull, a small snake vertebra, and a bird bone. Discoveries from the Malembo level include a narrow-snouted crocodyliform similar toCongosaurusandEuthecodon, an arsinoithere anterior tooth, an upper molar similar to that of the ptolemaiidanKelba, an unidentified mammalian incisor, and a large primate-like premolar. Recent biostratigraphic advances in East Africa and the new fossil discoveries in Cabinda suggest similarity to late Oligocene faunas in Ethiopia and Kenya, although the large primate-like premolar is unique. To this day, the fossil localities of Malembo provide the only coastal, low latitude, low elevation record of West African Cenozoic terrestrial mammals. https://gsa.confex.com/gsa/2013AM/webprogram/Paper232115.html

ABSTRACT: Type specimens (holotypes, neotypes, syntypes, etc.) are of crucial importance because they are the only tangible evidence of the nomenclatural act that supports the understanding of paleobiodiversity. The list of the vertebrate species whose type specimen is based on fossils from Portugal is presented here. We counted 206 species, of which there are 45 bony fishes, 39 Late Jurassic Mammaliamorpha, 33 Cenozoic mammals, 28 non-avian dinosaurs, 25 non-dinosaur reptiles, 11 Cenozoic Aves, 22 Chondrichthyes and three other vertebrates. Except for the Quaternary fossil record, the type specimens can be used as a shortcut for measuring the fossil record and paleobiodiversity through geological time and rock units because they correlate in 95% with the fossil record by epoch in the case-study of fossil vertebrates of Portugal.

Although rich in Cretaceous vertebrate fossils, prior to 2005 the amniote fossil record of Angola was poorly known. Two horizons and localities have yielded the majority of the vertebrate fossils collected thus far; the Turonian Itombe Formation of Iembe in Bengo Province and the Maastrichtian Mocuio Formation of Bentiaba in Namibe Province. Amniotes of the Mesozoic of Angola are currently restricted to the Cretaceous and include eucryptodire turtles, plesiosaurs, mosasaurs, pterosaurs, and dinosaurs. Recent collecting efforts have greatly expanded our knowledge of the amniote fauna of Angola and most of the taxa reported here were unknown prior to 2005.

NEW SPECIMENS OF ANGOLASAURUS BOCAGEI AND COMMENTS ON THE EARLY RADIATIONS OF PLIOPLATECARPINE MOSASAURS POLCYN, Michael, SMU, Dallas, TX, USA; JACOBS, Louis, SMU, Dallas, TX, USA; MATEUS, Octávio, Museu da Lourinhã, Lourinhã, Portugal; SCHULP, Anne, Natuurhistorisch Museum Maastricht, Maastricht, Netherlands New, well preserved material of the Turonian mosasaur Angolasaurus bocagei from the Tadi Beds of the Itombe Formation in northern Angola, allows detailed redescription of its morphology and reassessment of its phylogenetic relationships. Angolasaurus had been previously referred to the genus Platecarpus; however, phylogenetic analysis confirms the valid taxonomic status of A. bocagei, and reconstructs that taxon within a clade that also includes the genera Selmasaurus and Ectenosaurus. These forms are united by an elaborated infrastapedial process of the quadrate and a unique ridge-like descending process of the parietal forming the supraoccipital articulation, but also retain a relatively plesiomorphic configuration of the braincase. That clade is united with all other plioplatecarpines by a number of derived characters including the presence of a novel basicranial circulation pattern. In Africa, North and South America, early plioplatecarpines are known by the Middle Turonian and Angolasaurus and closely related forms appear by the Upper Turonian. Selmasaurus and Ectenosaurus are a rare faunal component of the Santonian and Campanian of North America. Platecarpus planifrons appears in the Coniacian of North America and represents the plesiomorphic condition of the clade containing the remaining species of Platecarpus and Plioplatecarpus, that appears in the Santonian and persist until the end of the Cretaceous, reaching global distribution. The temporal and geographic distribution of these radiations suggest influence of paleogeography and eustatic sea levels.

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A new Late Cretaceous turtle, Angolachelys mbaxi gen. et sp. nov., from the Turonian (90 Mya) of Angola, represents the oldest eucryptodire from Africa. Phylogenetic analysis recovers Angolachelys mbaxi as the sister taxon of Sandownia harrisi from the Aptian of Isle of Wight, England. An unnamed turtle from the Albian Glen Rose Formation of Texas (USA) and the Kimmeridgian turtle Solnhofia parsonsi (Germany), are successively more distant sister taxa. Bootstrap analysis suggests those four taxa together form a previously unrecognized monophyletic clade of marine turtles, herein named Angolachelonia clade nov., supported by the following synapomorphies: mandibular articulation of quadrate aligned with or posterior to the occiput, and basisphenoid not visible or visibility greatly reduced in ventral view. Basal eucryptodires and angolachelonians originated in the northern hemisphere, thus Angolachelys represents one of the first marine amniote lineages to have invaded the South Atlantic after separation of Africa and South America.

The diversity of fossil crocodylomorphs in Portugal is high, with occurrence as old as Mystriosaurus (=Steneosaurus) bollensis from the Lower Jurassic. The Late Jurassic forms are the better documented, and include the following taxa: Machimosaurus hugii, Lisboasaurus estesi Seiffert, 1973, Lusitanisuchus mitrocostatus Seiffert, 1975; Schwarz & Fechner 2004, Theriosuchus guimarotae Schwarz and Salisbury 2005, Cf. Alligatorium, Goniopholis baryglyphaeus, and a crocodylomorph-like eggs in dinosaur nests (Mateus et al., 1998; Ricqlès et al., 2001). From the Lower Cretaceous were reported a few dinosaurs but its record is strangely scarce in crocodylomorphs (Mateus et al., 2011). The Upper Cretaceous crocodiles show a large diversity, but it is mostly based in fragmentary material that require revision, such as “Crocodylus” blavieri? Grey from the Upper Campanian - Maastrichtian of Viso, near Aveiro (initially reported by Sauvage 1897-98), Goniopholis cf. crassidens Owen 1841 and Oweniasuchus pulchelus Jonet 1981. Moreover there is a fascinating, but poorly understood, crocodylomorph diversity in the Cenomanian of Portugal, documented by fragmentary specimens that have been doubtfully assigned to Thoracosaurus Leidy 1852 of the Middle Cenomanian of Cacém, to the nomen dubium Oweniasuchus lusitanicus Sauvage 1897-98 (interpreted as a mesosuchian goniopholid) based in a fragmentary mandible from the Campanian-Maastrichtian, and also from the Middle Cenomanian of Portugal, Buffetaut and Lauverjat (1978) report an fragmentary unidentified possible dyrosaurid from Nazaré. All this specimens are too incomplete to be compared with the specimen here described. In contrast, Cenozoic crocodiles of Portugal are often known after complete skulls and several individuals. The taxa list include Iberosuchus macrodon (Lower to Middle Eocene), Tomistoma calaritanus (Early Miocene) and T. lusitanica (Burdigalian-Helvetian), and Diplocynodon sp. (Antunes, 1961, 1987, 1994).
At least, two different morphotypes of crocodylomorph eggs from the Late Jurassic of Lourinhã Formation are also known.
A new specimen here reported of crocodile based in a partial skull and mandible (ML1818) from the Uppermost Middle Cenomanian platform carbonates of Baixo Mondego, west central Portugal (Tentúgal Fm., Callapez, 2004). The taxon is phylogenetically positioned as a basal Eusuchia, due to the choanae enclosed by the pterygoid, and closely related with stem Crocodylia and Borealosuchus. This specimen represents the only well documented and valid eusuchian species in the Cenomanian of Europe and is the oldest representative of an eusuchian crocodylomorph, with the exception for the Barremian Hylaeochampsa vectiana.

Several well preserved clutches of dinosaurs have been discovered in the upper Kimmeridgian/ Tithonian of Lourinhã (Estramadur Province, Portugal). Some eggs of one clutch contained embryo elements of a theropod dinosaur. The egg-shell resembles that of eggs which have been discovered in the Upper Jurassic of Colorado

Spinosaurids are some of the most enigmatic Mesozoic theropod dinosaurs due to their unique adaptations to aquatic environments and their relative scarcity. Their taxonomy has proven to be especially problematic. Recent discoveries from Western Europe in general, specifically Iberia, provide some of the best specimens for the understanding of their phylogeny, leading to the description of the spinosaurid Vallibonavenatrix cani and the recognition of the Iberian dinosaur Camarillasaurus cirugedae as one of them. Portuguese associated spinosaurid remains (ML1190) from the Papo Seco Formation (early Barremian) were previously assigned to Baryonyx walkeri but new material recovered in 2020 along with new phylogenetic analyses suggests a different phylogenetic placement, making their revision necessary. Here we show that these remains are not attributable to Baryonyx walkeri, but to a new genus and species, Iberospinus natarioi, gen. et sp. nov. The new taxon is characterized by the presence of a single Meckelian foramen in the Meckelian sulcus, a straight profile of the ventral surface of the dentary and a distal thickening of the acromion process of the pubis between other characters. Iberospinus natarioi is recovered as a sister taxon of the clade formed by Baryonyx and Suchomimus, and outside Spinosaurinae when Vallibonaventrix cani is excluded from the analysis. The description of this taxon reinforces Iberia as a hotspot for spinosaur biodiversity, with several endemic taxa for the region. As expected for the clade, the dentary displays a highly vascularized neurovascular network. The morphometric analysis of parts of the skeleton (pedal phalanx and caudal vertebrae, among others) shows an intermediate condition between basal tetanurans and spinosaurines.

Portugal is rich on dinosaur remains (bones, eggs, and tracks) from Early Jurassic to Late
Cretaceous ages, but mainly from the Late Jurassic, in which dozen of tracksites have been reported.
Here are reported new or poorly known track localities:
1) Five tracksites share the preservation substrate (marine carbonated limestone), age (late Jurassic), geographic area (Leiria district of Portugal), kind of preservation (true tracks), and completeness (trackways of multiple individuals):
i) Praia dos Salgados includes eight trackways, mostly ornithopods and theropods, and one wide gauge sauropod, made in very soft sediment; some preserve the hallux impression.
ii) Serra de Mangues is mostly covered with vegetation but seems to include dozens of tracks comprising theropods, thyreophorans, ornithopods and sauropods.
iii) Sobral da Lagoa (Pedreira do Rio Real) include six trackways but poorly preserved;
and
iv) Serra de Bouro that preserves four sauropod trackways in one single layer.
v) Pedrógão, preserved, at least, one theropod trackway and several isolated tracks of
theropods and ornithopods were found in different layers in the Early Oxfordian.
2) The locality in Praia de Porto das Barcas yielded natural casts of stegosaur tracks
(including pes print with skin impression) and a very large sauropod pes print with about
1.2 m long pes.
3) A new pterosaur tracksite was found in the Late Jurassic of Peralta, Lourinhã (Sobral Member, Lourinhã Fm.; Late Kimmeridgian/Early Tithonian). More than 220 manus and pes tracks have been collected in about five square meters, all ascribed to pterosaurs. The tracks were produced in a thin mud layer that has been covered by sand which preserved them as sandstone mould infill (natural casts). The manus of the largest specimens is 13 cm wide and 5.5 cm long and the pes measures 14.5 cm in length and 9 cm in width. This shows the occurrence of very large pterosaurs in the Late Jurassic. Other pterosaur tracksites in the Late Jurassic of Portugal are: Porto das Barcas (Lourinhã Municipality), South of Consolação (Peniche Municipality), and Zambujal de Baixo (Sesimbra Municipality).

Stegosaurian dinosaurs have a quadrupedal stance, short forelimbs, short necks, and are generally considered to be low browsers. A new stegosaur, Miragaia longicollum gen. et sp. nov., from the Late Jurassic of Portugal, has a neck comprising at least 17 cervical vertebrae. This is eight additional cervical vertebrae when compared with the ancestral condition seen in basal ornithischians such as Scutellosaurus.
Miragaia has a higher cervical count than most of the iconically long-necked sauropod dinosaurs. Long neck length has been achieved by ‘cervicalization’ of anterior dorsal vertebrae and probable lengthening of centra. All these anatomical features are evolutionarily convergent with those exhibited in the necks of
sauropod dinosaurs. Miragaia longicollum is based upon a partial articulated skeleton, and includes the only known cranial remains from any European stegosaur. A well-resolved phylogeny supports a new clade that unites Miragaia and Dacentrurus as the sister group to Stegosaurus; this new topology challenges the common view of Dacentrurus as a basal stegosaur.

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Eleven new tracks from the Upper Jurassic of Portugal are described and attributed to the stegosaurian ichnogenus Deltapodus. One track exhibits exceptionally well−preserved impressions of skin on the plantar surface, showing the stegosaur foot to be covered by closely spaced skin tubercles of ca. 6 mm in size. The Deltapodus specimens from the Aalenian of England represent the oldest occurrence of stegosaurs and imply an earlier cladogenesis than is recognized in the body fossil record.