Tooth development and histology patterns in lamniform sharks (Elasmobranchii,Lamniformes) revisited

The dentition of lamniforme sharks exhibits several characters that have been used extensively to resolve the phylogenetic relationships of extant taxa, yet some uncertainties remain. Also, the development of different teeth of a tooth file within the jaws of most extant lamniforms has not been documented to date. High‐resolution micro‐computed tomography is used here to re‐evaluate the importance of two dental characters within the order Lamniformes, which were considered not to be phylogenetically informative, the histotype and the number of teeth per tooth file. Additionally, the development and mineralization patterns of the teeth of the two osteodont lamniforms Lamna nasus and Alopias superciliosus were compared. We discuss the importance of these dental characters for phylogenetic interpretations to assess the quality of these characters in resolving lamniform relationships. The dental characters suggest that (1) Lamniformes are the only modern‐level sharks exhibiting the osteodont histotype, (2) the osteodont histotype in lamniform sharks is a derived state in modern‐level sharks (Elasmobranchii), (3) the osteodont type, conversely is convergently achieved when the clade Chondrichthyes is considered and thus might comprise a functional rather than a phylogenetic signal, and (4) there is an increase in the number of teeth per file throughout lamniform phylogeny. Structural development of the teeth of L. nasus and A. superciliosus is congruent with a previous investigation of the lamniform shark Carcharodon carcharias. J. Morphol. 277:1584–1598, 2016. © 2016 Wiley Periodicals, Inc.     

Lamniform Shark Teeth from the Late Cretaceous of Southernmost South America (Santa Cruz Province,Argentina)

Here we report multiple lamniform shark teeth recovered from fluvial sediments in the (Campanian-Maastrichtian) Cerro Fortaleza Formation, Santa Cruz Province, Argentina. This small tooth assemblage is compared to various lamniform sharks possessing similar dental morphologies, including Archaeolamna, Cretalamna, Dwardius, Dallasiella, and Cretodus. Although the teeth share numerous morphological features with the genus Archaeolamna, including a developed neck that maintains a relatively consistent width along the base of the crown, the small sample size and incomplete nature of these specimens precludes definitive taxonomic assignment. Regardless, the discovery of selachian teeth unique from those previously described for the region broadens the known diversity of Late Cretaceous South American sharks. Additionally, the discovery of the teeth in fluvial sandstone may indicate a euryhaline paleobiology in the lamniform taxon or taxa represented by this tooth assemblage.     

NEW MULTITUBERCULATE MAMMALS FROM THE HAUTERIVIAN/BARREMIAN TRANSITION OF EUROPE (IBERIAN PENINSULA)

Abstract: New multituberculate mammals from the Hauterivian/Barremian transition of Europe are described. They were found in the late Hauterivian‐early Barremian fossiliferous locality of La Cantalera (Josa, Teruel, Spain), one of the Early Cretaceous sites in the Aragonese branch of the Iberian Ranges, in northeastern Iberia. The fossils have been assigned to at least three taxa on the basis of nine isolated teeth: a new pinheirodontid taxon, Cantalera abadi gen. et sp. nov.; a representative of the eobaatarid Eobaatar; a taxon described as Plagiaulacidae or Eobaataridae gen. et sp. indet.; and other as Plagiaulacida indet. These fossils have increased the resolution of European Early Cretaceous multituberculate mammalian biostratigraphy and palaeobiogeography: the oldest representative of Eobaatar is described here; a taxon is assigned to ?Plagiaulacidae, in which case it would be the first of this family in the Iberian Peninsula; and the discovery of a new late Hauterivian pinheirodontid taxon demonstrates greater biodiversity and a wider distribution for these multituberculates than was previously known. The mutituberculate fauna of La Cantalera consists of endemic taxa (Pinheirodontidae), which were restricted to what is now Western Europe, and others (Eobaataridae) which have also been described in Asia. Consistent with the Iberian record of late Barremian gobiconodontid mammals, the presence of Eobaatar in Iberia with representatives from the late Hauterivian to late Barremian, as well as in the Aptian or Albian of Mongolia, indicates that faunal exchanges between Europe and Asia could have existed for most of the Early Cretaceous, either sporadically or constantly.     

Neoselachians (Chondrichthyes,Elasmobranchii) from the Lower and lower Upper Cretaceous of north‐eastern Spain

Extensive sampling of several Barremian and Albian–Cenomanian levels across the Aguilón, Oliete and Aliaga subbasins of the Iberian Basin, north‐east Spain, yielded abundant material of new or so far poorly known neoselachians. The faunas consist of 16 different species, five of which represent new species and two new genera: Cantioscyllium brachyplicatum sp. nov. , Platypterix venustulus gen. et sp. nov. , Ptychotrygon pustulata sp. nov. , Ptychotrygon striata sp. nov. and Iberotrygon plagiolophus gen. et sp. nov. In addition, teeth of Heterodontus cf. H. carerens, Lamniformes indet., Pteroscyllium sp., Scyliorhinidae indet., Rhinobatos sp., Spathobatis sp., Belemnobatis sp., Ptychotrygon geyeri, Ptychotrygon sp. and Celtipristis herreroi are described. The new family Ptychotrygonidae is defined. The localities comprise palaeoenvironments ranging from lacustrine and shallow lake to open marine settings. Neoselachians are almost completely absent from continental settings in the Barremian, as a result of prevailing freshwater conditions, but became more abundant in marine strata. The Albian–Cenomanian selachian assemblage is the most profuse and diverse of the three assemblages studied. It is dominated by small, benthic and near‐coastal taxa, for instance Cantioscyllium and Ptychotrygon, and contains several new species, including an endemic batoid, Iberotrygon plagiolophus gen. et sp. nov. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 316–347.     

A Gigantic Shark from the Lower Cretaceous Duck Creek Formation of Texas

Three large lamniform shark vertebrae are described from the Lower Cretaceous of Texas. We interpret these fossils as belonging to a single individual with a calculated total body length of 6.3 m. This large individual compares favorably to another shark specimen from the roughly contemporaneous Kiowa Shale of Kansas. Neither specimen was recovered with associated teeth, making confident identification of the species impossible. However, both formations share a similar shark fauna, with Leptostyrax macrorhiza being the largest of the common lamniform sharks. Regardless of its actual identification, this new specimen provides further evidence that large-bodied lamniform sharks had evolved prior to the Late Cretaceous.     

A SPALACOLESTINE SPALACOTHERIID (MAMMALIA,TRECHNOTHERIA) FROM THE EARLY CRETACEOUS (BARREMIAN) OF SOUTHERN ENGLAND AND ITS BEARING ON SPALACOTHERIID EVOLUTION

Abstract: Bulk screening of Early Cretaceous (Barremian) Wealden Group strata of the Wessex Formation exposed on the south‐west and south‐east coasts of the Isle of Wight, southern England, has resulted in the recovery of fragmentary remains pertaining to a new spalacolestine spalacotheriid mammal, Yaverlestes gassoni gen. et sp. nov. These represent the first European record of the Spalacolestinae. The remains comprise a dentulous incomplete dentary and isolated upper and lower molariforms, the former representing the most substantial mammal remains yet recovered from the Wealden Group. Hitherto, six species of spalacotheriid mammal were known from the Lower Cretaceous of Europe. All are referred to the genus Spalacotherium but in the case of taxa diagnosed on the basis of isolated lower teeth and other specimens where the post‐canine dentition is incompletely known, it is now evident that these referrals should be treated with caution. Furthermore, the new Wessex Formation spalacotheriid and recently described spalacotheriids from the ?Barremian of Japan, and the Barremian and Aptian of China exhibit combinations of characters that suggest that spalacotheriids were more diverse and that their evolution was more complex than previously recognized. The systematic position of an isolated tooth from the basal Cretaceous Lourinhã Formation of Portugal is discussed and the tooth reassigned to the Spalacotheriidae. Together with the new Wessex Formation taxon, eight species are now known from the Lower Cretaceous of Europe. The discovery of a spalacolestine in the Barremian Wessex Formation supports the concept of faunal interchange between Europe, Asia and North America during the Early Cretaceous. It also supports derivation of North American spalacotheriids from a European or Eurasian ancestor.     

NEOSELACHIANS FROM THE UPPER CAMPANIAN AND LOWER MAASTRICHTIAN (UPPER CRETACEOUS) OF THE SOUTHERN PYRENEES, NORTHERN SPAIN

Abstract: Bulk sampling of upper Campanian to lower–middle Maastrichtian coastal and lagoonal deposits in five sections of the Tremp Formation in the south‐central Pyrenees yielded numerous neoselachian teeth. The fauna comprises nine taxa of which three species and one genus are new: Hemiscyllium sp., Lamniformes indet., Paratrygonorrhina amblysoda gen. et sp. nov., Coupatezia trempina sp. nov., Coupatezia sp., Coupatezia? sp., Rhombodus ibericus sp. nov. and Igdabatis indicus. The neoselachian fauna is dominated by small nectobenthic rays. This composition resembles assemblages known from the marine Upper Cretaceous, but differs from nearby localities of the Basque‐Cantabrian region and continental selachian associations of the French Pyrenees. The results indicate that Rhombodus might not be a reliable biostratigraphic marker for the Maastrichtian. The faunal composition suggests a shallow trans‐Tethyan connection between Eurasia and India at the end of the Cretaceous Period.     

Dental ontogeny of a white shark embryo

Unlike most viviparous vertebrates, lamniform sharks develop functional teeth during early gestation. This feature is considered to be related to their unique reproductive mode where the embryo grows to a large size via feeding on nutritive eggs in utero. However, the developmental process of embryonic teeth is largely uninvestigated. We conducted X‐ray microcomputed tomography to observe the dentitions of early‐, mid‐, and full‐term embryos of the white shark Carcharodon carcharias (Lamniformes, Lamnidae). These data reveal the ontogenetic change of embryonic dentition of the species for the first time. Dentition of the early‐term embryos (∼45 cm precaudal length, PCL) is distinguished from adult dentition by 1) the presence of microscopic teeth in the distalmost region of the paratoquadrate, 2) a fang‐like crown morphology, and 3) a lack of basal concavity of the tooth root. The “intermediate tooth” of early‐term embryos is almost the same size as the adjacent teeth, suggesting that lamnoid‐type heterodonty (lamnoid tooth pattern) has not yet been established. We also discovered that mid‐term embryos (∼80 cm PCL) lack functional dentition. Previous studies have shown that the maternal supply of nutritive eggs in lamnoid sharks ceases during mid‐ to late‐gestation. Thus, discontinuation of functional tooth development is likely associated with the completion of the oophagous (egg‐eating) phase. Replacement teeth in mid‐term embryos include both embryonic and adult‐type teeth, suggesting that the embryo to adult transition in dental morphology occurs during this period. J. Morphol. 278:215–227, 2017. © 2016 Wiley Periodicals,Inc.     

Teeth of Embryos in Lamniform Sharks (Chondrichthyes: Elasmobranchii)

The dentitions of lamniform sharks possess a unique heterodonty, the lamnoid tooth pattern. However, in embryos, there are 'embryonic' and 'adult' dentitions. The teeth in the embryonic dentition are peg-like and appear to be attached to the jaw in an acrodont fashion. The adult dentition is characterized by the presence of replacement tooth series with the lamnoid tooth pattern. The embryonic–adult transition in dentitions appears at around 30–60cm TL. Tooth replacement generally begins before birth in embryos with adult dentitions. The adult dentition becomes functional just before or after parturition. An embryo of one species (Lamna nasus) shows a tooth directly on the symphysis of the upper jaws, marking the first record of a medial tooth for the order Lamniformes.     

Pathologic tooth deformities in modern and fossil chondrichthians: a consequence of feeding-related injury

Deformed teeth are found as rare components of the dentitions of both modern and fossil chondrichthians. Tooth deformities occur as bent or twisted tooth crowns, missing or misshaped cusps, atypical protuberances, perforations, and abnormal root structures. Deformed tooth files consisting of unusually overlapped or small teeth, or teeth misaligned in the jaw also occur in modern forms, but deformed tooth files generally are not recognizable in fossils due to post-mortem dissociation of teeth and jaws. A survey of 200 modern lamniform and carcharhiniform sharks as well as literature sources indicate that such deformities are produced by feeding-related injury to the tooth-forming tissue of the jaws, particularly by impaction of chondrichthian and teleost fin and tail spines. Tooth counts for several late Cretaceous genera, based on material recovered from coastal plain sites from New Jersey to Alabama, suggest that the frequency of occurrence of deformed teeth in a species varies from about 0.015% in Squalicorax kaupi to about 0.36% in Paranomotodon sp. Tooth counts for modern lamniform and carcharhiniform sharks yield a comparable range in frequency of tooth deformities. Variation in frequency of tooth deformity may reflect interspecific differences in feeding behavior and dietary preferences. There is no suggestion in our data of any strong patterns of temporal variation in tooth deformity frequency, or of patterns ­reflecting chondrichthian phylogenetic history and evolution. Skeletal components of the probable prey of the Cretaceous species are preserved in the same horizons as the deformed teeth, and also are found within co-occurring chondrichthian coprolites.     

Morphology and evolution of the jaw suspension in lamniform sharks

The morphology of the jaw suspension and jaw protrusion mechanism in lamniform sharks is described and mapped onto a cladogram to investigate how changes in jaw suspension and protrusion have evolved. This has revealed that several evolutionary modifications in the musculoskeletal apparatus of the jaws have taken place among lamniform sharks. Galeomorph sharks (Carcharhiniformes, Lamniformes, Orectolobiformes, and Heterodontiformes) have paired ethmopalatine ligaments connecting the ethmoid process of the upper jaw to the ethmoid region of the cranium. Basal lamniform sharks also acquired a novel single palatonasal ligament connecting the symphysis of the upper jaw to the cranium mid-ventral to the nasal capsule. Sharks in the family Lamnidae subsequently lost the original paired ethmopalatine ligament while retaining the novel palatonasal ligament. Thus, basal lamniform taxa (Mitsukurina owstoni, Carcharius taurus, Alopias vulpinnis) have increased ligamentous support of the lateral region of the upper jaw while derived species (Lamnidae) have lost this lateral support but gained anterior support. In previous studies the morphology of the jaw suspension has been shown to play a major role in the mechanism of upper jaw protrusion in elasmobranchs. The preorbitalis is the primary muscle effecting upper jaw protrusion in squalean (sister group to galeomorphs) and carcharhiniform (sister group to lamniforms) sharks. The preorbitalis originates from the quadratomandibularis muscle and inserts onto the nasal capsule in squalean and carcharhiniform sharks. Carcharhiniform sharks have evolved a subdivided preorbitalis muscle with the new division inserting near the ethmoid process of the palatoquadrate (upper jaw). Alopid sharks have also independently evolved a partially subdivided preorbitalis with the new division inserting at the base of the ethmoid process and surrounding connective tissue. Lamnid sharks have retained the two preorbitalis divisions but have modified both of the insertion points. The original ventral preorbitalis division now inserts onto the connective tissue surrounding the mid-region of the upper jaw, while the new dorsal preorbitalis division inserts onto the surrounding connective tissue and skin at a more posterior position on the upper jaw. The retractor muscle of the jaws, the levator hyomandibularis, has also been modified during the evolution of lamniform sharks. In most sharks, including basal lamniforms, the levator hyomandibularis inserts onto the hyomandibula and functions to retract the jaws after protrusion. In alopid and lamnid sharks the levator hyomandibularis inserts primarily onto the upper and lower jaws around the jaw joint and is a more direct route for retracting the jaws. Thus, there has been at least one instance of character loss (ethmopalatine ligament), acquisition (palatonasal ligament), subdivision (preorbitalis), and modification (ventral preorbitalis, dorsal preorbitalis, and levator hyomandibularis) in the ligaments and muscles associated with the jaw suspension and jaw protrusion mechanism in lamniform sharks. While derived lamniform sharks (Lamna nasus, Carcharodon carcharius, and Isurus oxyrinchus) lost the ancestral passive lateral support of the ethmoid articulation of the upper jaw, they simultaneously acquired muscular support by way of the levator hyomandibularis, which provides a dynamic mechanism for lateral support. The evolution of multiple divisions of preorbitalis insertions onto the palatoquadrate and modification of the levator hyomandibularis insertion directly onto the jaws provides an active mechanism for multiple protractions and retractions of the upper jaw, which is advantageous in those sharks that gouge or saw pieces from large oversized prey items.     

A new,three‐dimensionally preserved enantiornithine bird (Aves: Ornithothoraces) from Gansu Province,north‐western China

In recent years, the Lower Cretaceous (Aptian) Xiagou Formation has yielded approximately 100 avian partial skeletons, many with soft‐tissue traces, from sites in the Changma Basin of Gansu Province, north‐western China. The most abundant taxon amongst these is the ornithuromorph Gansus yumenensis, but enantiornithines have also been identified in the sample. Here we describe two incomplete, semi‐articulated appendicular skeletons, the first consisting of a partial left pelvic girdle and complete pelvic limb, and the second comprised of a nearly complete right pelvic limb. Both specimens bear characteristics diagnostic of Enantiornithes, and are referred to a new taxon, Qiliania graffini gen. et sp. nov. The exceptional, three‐dimensional preservation of these specimens (compared to the crushed, nearly two‐dimensional condition of most other Early Cretaceous avian fossils) reveals new information regarding enantiornithine anatomy, evolution, and diversity. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 201–219.     

A basal sphenodontian (Lepidosauria) from the Jurassic of Patagonia: new insights on the phylogeny and biogeography of Gondwanan rhynchocephalians

Herein we describe a new rhynchocephalian taxon from the Middle Jurassic of Patagonia, Argentina, representing the first Jurassic record of the group in South America. The new taxon, consisting of a complete dentary, is ascribed to Sphenodontia based on the presence of a deep and wide Meckelian groove, long posterior process, well‐developed coronoid process, and acrodont teeth showing dental regionalization including successional, alternate hatchling, and additional series. This allocation is reinforced by a phylogenetic analysis that places the new taxon in a basal position within a clade of sphenodontians that excludes Diphydontosaurus and Planocephalosaurus. Additionally, the new taxon clusters within a Gondwanan clade with the Indian Godavarisaurus from the Jurassic Kota Formation, sharing the presence of recurved and relatively large posterior successional teeth that are ribbed and bear a peculiar anterolingual groove. This sister‐group relationship is intriguing from a palaeobiogeographical viewpoint, as it suggests some degree of endemism during the initial stages of the breakup of Pangaea. We also discuss the ontogenetic stage of the new taxon and provide insights on the evolution of successional dentition in rhynchocephalians. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 342–360.     

A new basal ornithuromorph bird (Aves: Ornithothoraces) from the Early Cretaceous of China with implication for morphology of early Ornithuromorpha

Ornithuromorpha is the most derived avian group in the Early Cretaceous, advanced members of which encompass all living birds (Neornithes). Here we report on a new basal ornithuromorph bird, Bellulia rectusunguis gen. et sp. nov., represented by a nearly complete skeleton from the Early Cretaceous Jehol Biota in northeastern China. A comprehensive phylogenetic analysis resolved the new taxon in a basal position that is only more derived than Archaeorhynchus and Jianchangornis among ornithuromorphs, increasing the morphological diversity of basal ornithuromorphs. The new specimen has a V‐shaped furcula with a short hypocleidium, a feature otherwise known only in Schizooura among Cretaceous ornithuromorphs. We discuss the implications of the new taxon on the evolution of morphology of primitive ornithuromorphs, particularly of pectoral girdle, sternum and limb proportion pertaining to powered flight. The preserved gastroliths and pedal morphology indicate herbivory and lakeshore adaption for this new species. © 2015 The Linnean Society of London     

Skeletal anatomy of the Late Cretaceous shark,Squalicorax (Neoselachii: Anacoracidae)

The paleobiology of the Cretaceous neoselachian shark,Squalicorax, has largely been based on isolated teeth. We examined partial and nearly complete skeletons of three species ofSqualicorax, S. falcatus (Aoassiz),S. kaupi (Agassiz), andS. pristodontus (Agassiz), that were collected from the U.S.A. These specimens suggest that the total body length (TL) ofS. falcatus typically measured 1.8–2.0 m, and probably did not exceed 3 m. Moderatesized individuals ofS. kaupi andS. pristodontus perhaps measured about 3 m TL. AlthoughS. pristodontus was the largest form among the three species examined, this taxon possessed a set of large jaws (with large but fewer teeth) relative to its body size compared toS. falcatus orS. kaupi. This suggests that tooth size is not an accurate indicator of the TL if one compares oneSqualicorax species to another. Neurocranial features suggest that the vision ofSqualicorax was not as acute as that of a contemporaneous macrophagous lamniform shark,Cretoxyrhina mantelli (Agassiz) , but olfaction ofSqualicorax may have been better thanC. mantelli. The morphology of placoid scales suggests thatSqualicorax was capable of fast swimming. New skeletal data support the view that the feeding dynamics ofSqualicorax was similar to the modern tiger shark (Galeocerdo Müller & Henle). The present data do not allow for exact ordinal placement, but, contrary to some previous interpretations,Squalicorax can be excluded from the Hexanchiformes and Orectolobiformes. The taxon should more appropriately be placed within the Lamniformes or Carcharhiniformes.      

Dental homologies in lamniform sharks (Chondrichthyes: Elasmobranchii).

The dentitions of lamniform sharks are said to exhibit a unique heterodonty called the "lamnoid tooth pattern." The presence of an inflated hollow "dental bulla" on each jaw cartilage allows the recognition of homologous teeth across most modern macrophagous lamniforms based on topographic correspondence through the "similarity test." In most macrophagous lamniforms, three tooth rows are supported by the upper dental bulla: two rows of large anterior teeth followed by a row of small intermediate teeth. The lower tooth row occluding between the two rows of upper anterior teeth is the first lower anterior tooth row. Like the first and second lower anterior tooth rows, the third lower tooth row is supported by the dental bulla and may be called the first lower intermediate tooth row. The lower intermediate tooth row occludes between the first and second upper lateral tooth rows situated distal to the upper dental bulla, and the rest of the upper and lower tooth rows, all called lateral tooth rows, occlude alternately. Tooth symmetry cannot be used to identify their dental homology. The presence of dental bullae can be regarded as a synapomorphy of Lamniformes and this character is more definable than the "lamnoid tooth pattern." The formation of the tooth pattern appears to be related to the evolution of dental bullae. This study constitutes the first demonstration of supraspecific tooth-to-tooth dental homologies in nonmammalian vertebrates.     

Development and microstructure of tooth histotypes in the blue shark,Prionace glauca (Carcharhiniformes: Carcharhinidae) and the great white shark,Carcharodon carcharias (Lamniformes: Lamnidae)

Elasmobranchs exhibit two distinct arrangements of mineralized tissues in the teeth that are known as orthodont and osteodont histotypes. Traditionally, it has been said that orthodont teeth maintain a pulp cavity throughout tooth development whereas osteodont teeth are filled with osteodentine and lack a pulp cavity when fully developed. We used light microscopy, scanning electron microscopy, and high‐resolution micro‐computed tomography to compare the structure and development of elasmobranch teeth representing the two histotypes. As an example of the orthodont histotype, we studied teeth of the blue shark, Prionace glauca (Carcharhiniformes: Carcharhinidae). For the osteodont histotype, we studied teeth of the great white shark, Carcharodon carcharias (Lamniformes: Lamnidae). We document similarities and differences in tooth development and the microstructure of tissues in these two species and review the history of definitions and interpretations of elasmobranch tooth histotypes. We discuss a possible correlation between tooth histotype and tooth replacement and review the history of histotype differentiation in sharks. We find that contrary to a long held misconception, there is no orthodentine in the osteodont teeth of C. carcharias. J. Morphol. 276:797–817, 2015. © 2015 Wiley Periodicals, Inc.     

Osteology of the Late Jurassic Portuguese sauropod dinosaur Lusotitan atalaiensis (Macronaria) and the evolutionary history of basal titanosauriforms

Titanosauriforms represent a diverse and globally distributed clade of neosauropod dinosaurs, but their inter‐relationships remain poorly understood. Here we redescribe Lusotitan atalaiensis from the Late Jurassic Lourinhã Formation of Portugal, a taxon previously referred to Brachiosaurus. The lectotype includes cervical, dorsal, and caudal vertebrae, and elements from the forelimb, hindlimb, and pelvic girdle. Lusotitan is a valid taxon and can be diagnosed by six autapomorphies, including the presence of elongate postzygapophyses that project well beyond the posterior margin of the neural arch in anterior‐to‐middle caudal vertebrae. A new phylogenetic analysis, focused on elucidating the evolutionary relationships of basal titanosauriforms, is presented, comprising 63 taxa scored for 279 characters. Many of these characters are heavily revised or novel to our study, and a number of ingroup taxa have never previously been incorporated into a phylogenetic analysis. We treated quantitative characters as discrete and continuous data in two parallel analyses, and explored the effect of implied weighting. Although we recovered monophyletic brachiosaurid and somphospondylan sister clades within Titanosauriformes, their compositions were affected by alternative treatments of quantitative data and, especially, by the weighting of such data. This suggests that the treatment of quantitative data is important and the wrong decisions might lead to incorrect tree topologies. In particular, the diversity of Titanosauria was greatly increased by the use of implied weights. Our results support the generic separation of the contemporaneous taxa Brachiosaurus, Giraffatitan, and Lusotitan, with the latter recovered as either a brachiosaurid or the sister taxon to Titanosauriformes. Although Janenschia was recovered as a basal macronarian, outside Titanosauria, the sympatric Australodocus provides body fossil evidence for the pre‐Cretaceous origin of titanosaurs. We recovered evidence for a sauropod with close affinities to the Chinese taxon Mamenchisaurus in the Late Jurassic Tendaguru beds of Africa, and present new information demonstrating the wider distribution of caudal pneumaticity within Titanosauria. The earliest known titanosauriform body fossils are from the late Oxfordian (Late Jurassic), although trackway evidence indicates a Middle Jurassic origin. Diversity increased throughout the Late Jurassic, and titanosauriforms did not undergo a severe extinction across the Jurassic/Cretaceous boundary, in contrast to diplodocids and non‐neosauropods. Titanosauriform diversity increased in the Barremian and Aptian–Albian as a result of radiations of derived somphospondylans and lithostrotians, respectively, but there was a severe drop (up to 40%) in species numbers at, or near, the Albian/Cenomanian boundary, representing a faunal turnover whereby basal titanosauriforms were replaced by derived titanosaurs, although this transition occurred in a spatiotemporally staggered fashion. © 2013 The Linnean Society of London     

New snakeflies (Insecta: Raphidioptera) from the Lower Cretaceous of the UK,Spain and Brazil

Abstract: Several new taxa of snakeflies (Raphidioptera) are described from the Lower Cretaceous deposits of the Wealden, UK (Barremian), Montsec, Spain (Barremian) and the Crato Formation, Brazil (Aptian). Mesoraphidia ednae sp. nov. and M. hilli sp. nov. are described from the Wealden; Nanoraphidia lithographica sp. nov. and Iberoraphidia dividua gen. et sp. nov. are described from Montsec, and Baissoptera lisae sp. nov. is described from the Crato Formation. The geographical range of Nanoraphidia has potentially been extended.