Evolution of Teeth and Quadrate in Non-avian Theropoda (Dinosauria: Saurischia), with the Description of Torvosaurus Remains from Portugal

Theropods form a highly successful and morphologically diversified group of dinosaurs that gave rise to birds. They include most, if not all, carnivorous dinosaurs, yet many theropod clades were secondarily adapted to piscivory, omnivory and herbivory, and theropods show a large array of skull and dentition morphologies. This work aims to investigate aspects of the evolution of theropod dinosaurs by analyzing in detail both the anatomy and ontogeny of teeth and quadrates in non-avian theropods, and by studying embryonic and adult material of a new species of theropod. A standardized list of terms and notations for each anatomical entity of the tooth, quadrate, and maxilla is here proposed with the goal of facilitating descriptions of these important cranial and dental elements. The distribution of thirty dental characters among 113 theropod taxa is investigated, and a list of diagnostic dental characters is proposed. As an example, four isolated theropod teeth from the Lourinhã Formation (Kimmeridgian‒Tithonian) of Portugal are described and identified based on a cladistic analysis performed on a data matrix of 141 dentition-based characters coded in 60 taxa. Two shed teeth are referred to an abelisaurid, providing the first record of Abelisauridae in the Jurassic of Laurasia and the one of the oldest records of this clade in the world, suggesting a possible radiation of Abelisauridae in Europe well before the Upper Cretaceous. The consensus tree resulting from this phylogenetic analysis, the most extensive on theropod teeth, indicates that theropod teeth provide reliable data for identification at approximately family level, and this method will help identifying theropod teeth with more confidence. A detailed description of the dentition of Megalosauridae is also provided, and a discriminant analysis performed on a dataset of numerical data collected on the teeth of 62 theropod taxa reveals that megalosaurid teeth are hardly distinguishable from other theropod clades with ziphodont dentition. This study highlights the importance of detailing anatomical descriptions and providing additional morphometric data on teeth with the purpose of helping to identify isolated theropod teeth. In order to evaluate the phylogenetic potential and investigate the evolutionary transformations of the quadrate, a phylogenetic morphometric analysis as well as a cladistic analysis using 98 discrete quadrate related characters were conducted. The quadrate morphology by its own provides a wealth of data with strong phylogenetic signal, and the phylogenetic morphometric analysis reveals two main morphotypes of the mandibular articulation of the quadrate linked to function. As an example, six isolated quadrates from the Kem Kem beds (Cenomanian) of Morocco are determined to be from juvenile and adult individuals of Spinosaurinae based on phylogenetic, morphometric, and phylogenetic morphometric analyses. Morphofunctional analysis of the spinosaurid mandibular articulation has shown that the posterior parts of the two mandibular rami displaced laterally when the jaw was depressed due to a mediolaterally oriented intercondylar sulcus of the quadrate. Such lateral movement of the mandibular ramus was possible due to a movable mandibular symphysis in spinosaurids, allowing the pharynx to be widened. A new species of theropod from the Lourinhã Formation of Portugal, Torvosaurus gurneyi, is erected based on a right maxilla and an incomplete caudal centrum. This taxon supports the mechanism of vicariance that occurred in the Iberian Meseta during the Late Jurassic when the proto-Atlantic was already well formed. A theropod clutch containing several crushed eggs and embryonic material is also assigned to this new species of Torvosaurus. Investigation on the maxilla ontogeny in basal tetanurans reveals that crown denticles, elongation of the anterior ramus, and fusion of interdental plates appear at a posthatchling stage. On the other hand, maxillary pneumaticity is already present at an embryonic stage in non-avian theropods.

Les Plus Anciens Mastodontes Tetralophodontes (Langhien inférieur Vb), évolution et remarques sur la Tetralophodontie

This study deals with mastodont teeth found near Lisbon in Lower Langhian (lower Middle Miocene) fluviatile, feldspathic sands (Vb division). Conclusions are as follows: 1. Tetralophodont molars (even if at a still primitive stade of the tetralophodont condition) do exist at least since lower Langhian times, and not only since late Middle Miocene as was previously known. 2. Tri- and tetralophodont structures may (and indeed do) coexist in the same individual: such examples do not correspond to transitional forms, but instead to a mosaic of juxtaposed characters (however this does not mean there are no transitional forms in other instances). 3. So these structures coexisted in a population not yet genetically separated beyond fertile cross-breeding, i.e. beyond species' level. 4. Origin of the tetralophodont molar was due to some mutation (s). but without crossing species, limits and even more genus'ones. 5. At this times probably soon after the first appearance of tetralophodont mutants, animals with such characters were a small but significant minority among the population (17% if account is taken on D4's: only 2% after M2's). 6. There was not then any direct and clear correlation between number of lophs (transversal crests) and tooth size, even if the increase of such number goes along with length's increase. 7. Dimensions (length in special) in tetralophodont teeth tend to exceed those in «normal» trilophodont teeth, this being particularly clear in D4, even if there is no clear distinction: the situation is quite the same, maybe less marked, with the M2. 8. According to the preceding conclusions there are no reasons to segregate different taxa among such mastodont population on the grounds of the presence in D4, M1 and M2 of 3 or 4 crests (this character being regarded as diagnostic of the genus Tetralophodon). 9. On the contrary, if any natural (in biological sense) classification is disregarded and a morphological parataxonomy is adopted there should be considered both Gomphotherium angustidens and Tetralophodon sp.: however this is absolutely not our opinion.

Contributions à la Paléontologie du Miocène moyen continental du Bassin du Tage. II - Observations sur les dents pharyngiennes de poissons cyprinidés - Póvoa de Santarém.

The study of Cyprinid fish pharyngeal teeth, collected by M. Telles Antunes in continental "Helvetian" sediments from Póvoa de Santarém, makes possible to demonstrate the occurrence of two distinct species. One remains undetermined. The other belongs to the recent genus Leuciscus CUV. Several dental types of this genus are described and figured as Leuciscus antunesi nov. sp. Palaeoclimatical and palaeoecological interpretations are proposed.

Notes sur la géologie et la paleontology du Miocène de Lisbonne; (XX) les plus anciens mastodontes tetralophodontes (Langhien inférieur Vb), évolution et remarques sur la tetralophodontie

This study deals with mastodont teeth found near Lisbon in Lower Langhian (lower Middle Miocene) fluviatile, feldspathic sands (Vb division). Conclusions are as follows: 1. Tetralophodont molars (even if at a still primitive stade of the tetralophodont condition) do exist at least since lower Langhian times, and not only since late Middle Miocene as was previously known. 2. Tri- and tetralophodont structures may (and indeed do) coexist in the same individual: such examples do not correspond to transitional forms, but instead to a mosaic of juxtaposed characters (however this does not mean there are no transitional forms in other instances). 3. So these structures coexisted in a population not yet geneticaliy separated beyond fertile cross-breeding, i.e. beyond species'level. 4. Origin of the tetralophodont molar was due to some mutation (s). but without crossing species, limits and even more genus' ones. 5. At this times probably soon after the first appearance of tetralophodont mutants, animals with such characters were a small but signifiant minority among the population (17% if account is taken on D4's: only 2% after M2's). 6. There was not then any direct and clear correlation between number of lophs (transversal crests) and tooth size, even if the increase of such number goes along with length's increase. 7. Dimensions (length in special) in tetralophodont teeth tend to exceed those in «normal» trilophodont teeth, this being particularly clear in D4, even if there is no clear distinction: the situation is quite the same, maybe less marked, with the M2. 8. According to the preceding conclusions there are no reasons to segregate different taxa among such mastodont population on the grounds of the presence in D4, M1 and M2 of 3 or 4 crests (this character being regarded as diagnostic of the genus Tetralophodon). 9. On the contrary, if any natural (in biological sense) classification is disregarded and a morphological parataxonomy is adopted there should be considered both Gomphotherium angustidens and Tetralophodon sp.: however this is absolutely not our opinion.

Vertébrés (crocodilians, poisons) du Miocène marin de l’Algarve occidentale

Karst fillings rich in fossils from carbonate series, Miocene in age, exposed on the coast of Algarve, yielded some thousands of teeth and other vertebrate remains. A crocodilian and more than fifty species of fishes (Selacians, Teleosteans) have been identified (see tableau 1). This fauna is consistent as a whole with a Burdigalian (maybe late) and/or Langhian age. It also indicates shallow waters, warmer than at present in the same region although not strictly tropical. This fauna shows more affinities with others of the same age from mediterranean region (i.e. Baleares) than with those from Lisbon.

Quelques mastodontes miocènes du Portugal

From a paleontological point of view previous determinations for some specimens as Tetralophodon longirostris can be confirmed from Azambujeira (upper level), Valverde and Vale de Matança. A so far undescribed and rare D3 (from Azambujeira, middle level) has been studied. For the first time in Portugal a Gomphotherium angustidens transitional to «Tetralophodon» grandincisivus is reported (from Portas do Sol). Teeth from Lisbon formerly reported (in part) to T. longirostris are now ascribed to G. angustidens. The presence of T. longirostris at Vale de Matança excludes a Pliocene age for Marateca Formation. This and some other evidence clearly points out towards an early Vallesian age. G. angustidens transitional to T. grandincisivus found at Portas do Sol is enough to ascribe this locality to the latest Middle Miocene or earliest Upper Miocene. Therefore it is possible to correlate overlying Santarém limestones to the Vallesian Cartaxo and Almoster ones which are better dated.

Goldra: premier gisement karstique en Algarve, faune et industries

A bone breccia from Goldra, near Loulé, is studied. It corresponds to the infilling of a karst depression, consisting of: rather worn and probably transported dolomite pebbles at the bottom; accumulations of frequently burnt bone scraps, much broken and with acute edges (no transport), certainly debris of human food, suggesting habitat level (s); in association with the former, stone (flint, quartz, quartzite, graywacke) rather uncharacteristic artifacts that seem compatible with middle and upper Paleolithic, or with Epipaleolithic; and small mammal teeth and bones. Fauna includes an extinct species, Microtus brecciensis recognized for the first time in Portugal. It is not older than Riss-Wiirm interglacial, and may be of this age or later, maybe that of one of wurm's first interstades. Fauna points out to a varied landscape with open country and woods; and to a rather warm and dry temperate, or dry subtropical mediterranean climate. Climate differences should not be significant in comparison with the extant situation. The presence of the mammal species found so far is consistent with modern distribution.

Iberosuchus et pristichampsus, crocodilians de l’éocène données complémentaires, discussion, distribution stratigraphique

Two eocene ziphodont crocodillans (Ilberosuchus and Pristichampsus) are dealt with. Their distinction seems possible, even with isolated teeth. The association of both in some localities may account for some previous identification difficulties. Geographical and stratigraphical distribution indicated: for Pristichampsus from Germany to Spain, Cuisan to to Upper Lutetian; for Iberosuchus from France to Portugal, lower Lutetian to Bartonian and maybe Ludian.

Petits mammifères du burdigalien infèrieur (Universidade Católica, Avenida do Uruguay)

Improved bromoform concentration as developped at CEPUNL allowed better recovery of small mammals'teeth. At Universidade Católica and Avenida do Uruguay 19 taxa (and a further one with doubt) were recognized. Some are new for the level and for Tagus basin: Lagopsis cadeoti and Melissiodon dominans (1st reference for the genus); Glirudinus modestus (formerly under another name); Armantomys (1st reference for this level); Peridyromys murinus (referred before under another name); Microdyromys legidensis (1st ref. of gen. and sp. for this level); and Heteroxerus rubricati, formerly reported to other species of the same genus. Both localities share the same position viz marine levels under and above. This allows us to correlate them with NS or N6 Blow's zones. Both are distinctly younger-than glauconite in underlying beds about 21 MY old (K-Ar). Small mammals point out to MN3a Neogene subunit. Fauna is much alike Lower Burdigalian ones in Spain, France, Germany and Austria. Terrestrial, maybe steppe forms predominate. Land environment was open, with scant plant cover but not devoid of trees. Peridyromys murinus numerical importance and other data suggest a not so warm climate in correspondance to a minimum temperature event. This is corroborated by associated marine fish fauna entirely without warm water stenotherm species, and by paleobotanical/palynological data. Results are in close agreement with Central Northern Spain. The localities studied here are even more interesting as direct correlations between marine and continental stratigraphical scales are possible.

Suidés et tayassuidés du burdigalien V-a et du langhien V-b de lisbonne

Teeth and astragali were used for a biometrical study concerning suids from V-a (upper Burdigalian) and V-b (Langhian) divisions from Lisbon's Miocene series. The Hyotherium (V-b) are identical to those from french localities, hence they all belong in the same species H. soemmeringi. Bunolistriodon populations from V-a and V-b are homogenous; no significant difference between them has been found, inspite of different geological age. Both may be ascribed to B. lockharti. No evolutive trend was detected. The presence of another form close to the north african B. massai could not be confirmed either. French localities' Bunolistriodon populations also seem homogenous and conspecific with those from Lisbon. Notwithstanding its essentially homogenous character, there can be distinguished two sets in both V-a and V-b populations according to M3 size; this remains to be explained, since the last molars are the most likely to show a broad range of variation and are not unequivocally related to sexual dimorphism. Classification of the rare Tayassuidae has been confirmed. All known taxa are shown (see tableau I).

Castor fiber na gruta do Caldeirão. Existência, distribuição e extinção do castor em Portugal

Beaver only had been found in Portugal in a Chalcolithic locality, the Vila Nova de S. Pedro castrum. It has now been identified in the Upper Paleolithic (Solutrean) from Gruta do Caldeirão, near Tomar. The species has been found recently at «Gruta do Almonda»; 4 teeth were collected in bed C, older than a Solutrean sequence (see Anexo for details). The species seems to have been rare, as it was also the case with portuguese Miocene Castoridae Enroxenomys minutus and Chalicomys jaegeri. If account is taken of the presence in the Middle Ages until Castille of words meaning beaver (relared to the popular latin Fiber/Biber), it is obvious that these animais still existed then. Such nouns were largely predominant over rhe rather erudite latin (greek deríved) words as Castor,-óris and derived ones, as it could be expected. This allowed us to recognize that veiro should be the corresponding word with Fiber affinities in archaic portuguese. It was previously supposed to mean only expensive furs then imported into Portugal. Indeed it was also a zoonym. Anywày, beaver should be scarce by XIIIth century since it is not included in the quite detailed price list imposed by the «Lei da Almotaçaria» from December 26, 1253 (see Quadro II). Toponyms in veiro and derived words (fig. 2; Quadro III) (plural, feminines, diminutives, inhabited places) give a resrrictive view of rhe Middle Age distribution. Some of them are certainly older than Portugal itself (firsr half of XIlth cenrury); others existed by the XIVth century bur were probably older. Some rare toponyms seem to be derived from the erudite latin Castor,-óris. Nothing suggests that these words were still in use as zoonyms during the Middle Ages. All toponyms are located in regions near rivers and other freshwaters ecologically suitable for beavers, so we can approximately retrace its former, Middle Age disrribution in Portugal (fig. 2; Quadro III). Most of them are locared in the Center-West and Northwest of Portugal, with a suitable c1imate (rainfall in general over 800 mílimerers per year); the only sure geographical exception is Veiros, in Alto Alentejo province, in a region with comparable precipitations and less dry climate conditions than mosr of the territories South of rhe Tagus. There are less and less of these toponyms towards rhe South and the inner part of the country, and they are enrirely lacking in all drier regions from Trás-os-Montes, Beira, Alentejo beyond Tagus' basin, and in Algarve. Nothing suggests beavers lived there, No post-medieval toponym is known, nor any reference after middle XVth century. No such locality was at, or close by to, any frontier. Hence the hypothesis of veiro (et al.} as meaning but points where expensive furs (supposedly known as veiros in general but without c1early saying from what animal they were obrained from) is to be discarded. During the Middle Ages, beaver discriburion concerned all the main river basins from Minho to Tagus ones. Quite rarefied in rhe XIIIth, the beavers may have disappeared from Portugal during the XVth century. Ecological requiremenrs restricted their former distriburion. Vulnerability to natural causes (i.e., severe drought) and to human pressure may have accounted heavily for this species' extinction. Last (1446) reference for Portugal known to us suggests the species was by then almost extinct.

Castor fiber na gruta do Caldeirão Existência, distribuição e extinção do castor em Portugal

Beaver only had been found in Portugal in a Chalcolithic locality, the Vila Nova de S. Pedro castrum. It has now been idenrified in the Upper Paleolithic (Solurrean) from Gruta do Caldeirão, near Tomar. The species has been found recently at «Gruta do Almonda»; 4 teeth were collected in bed C, older than a Solutrean sequence (see Anexo for details). The species seems to have been rare, as it was also the case with portuguese miocene Castoridae Enroxenomys minutus and Chalicomys jaegeri. If account is taken of the presence in the Middle Ages until Castille of words meaning beaver (related to the popular latin Fiber/Biber), it is obvious that these animals still existed then. Such nouns were largely predominant over the rather erudite larin (greek derived) words as Castor, -óris and derived ones, as it could be expected. This allowed us to recognize that veiro should be the corresponding word with Fiber affinities in archaic portuguese. It was previously supposed to mean only expensive furs then imported into Portugal. Indeed it was also a zoonym. Anyway, beaver should be scarce by XIIIth century since it is not included in the quite detailed price list imposed by the «Lei da Almotaçaria» from December 26, 1253 (see Quadro II). Toponyms in veiro and derived words (fig. 2; Quadro III) (plural, feminines, diminutives, inhabited places) give a restrictive view of the Middle Age distribution. Some of them are certainly older than Portugal itself (first half of XIIth century); others existed by the XIVth century but were probably older. Some rare toponyms seem to be derived from rhe erudite latin Castor, -óris. Nothing suggests that these words were still in use as zoonyms during the Middle Ages. All toponyms are located in regions near rivers and other freshwaters ecologically suitable for beavers, so wecan approximately retrace irs former, Middle Age distribution in Portugal (fig. 2; Quadro III). Most of them are located in the Center-West and Northwest of Portugal, with a suitable c1imate (rainfall in general over 800 milimeters per year); the only sure geographical exception is Veiros, in Alto Alentejo province, in a region with comparable precipitations and less dry climare conditions than most of the territories South of the Tagus. There are less and less of these toponyms towards the South and the inner part of the country, and they are enrirely lacking in ali drier regions from Trás-os-Montes, Beira, Alentejo beyond Tagus' basin, and in Algarve. Nothing suggests beavers lived there, No pose-medieval toponym is known, nor any reference after middle XVth century. No such locality was at, or close by to, any frontier. Hence the hypothesis of veiro (e: al.) as meaning but points where expensive furs(supposedly known as veiros in general but without clearly saying from what animal they were obtained from) is to be discarded. During the Middle Ages, beaver distribution concerned all the main river basins from Minho to Tagus ones. Quice racefied in the XIIIth, the beavers may have disappeared from Portugal during the XVth century. Ecological requirements restricted their former distribution. Vulnerability to natural causes (i.e., severe drought) and to human pressure may have accounted heavily for this species extinction. Last (1446) reference for Portugal known to us suggests the species was by then almost extinct.