Late Cretaceous Of Madagascar Research Articles

A new pelomedusoid turtle, Sahonachelys mailakavava, from the Late Cretaceous of Madagascar provides evidence for convergent evolution of specialized suction feeding among pleurodires.

The Maevarano Formation in northwestern Madagascar has yielded a series of exceptional fossils over the course of the last three decades that provide important insights into the evolution of insular ecosystems during the latest Cretaceous (Maastrichtian). We here describe a new genus and species of pelomedusoid turtle from this formation, Sahonachelys mailakavava, based on a nearly complete skeleton. A phylogenetic analysis suggests close affinities of Sahonachelys mailakavava with the coeval Madagascan Sokatra antitra. These two taxa are the only known representatives of the newly recognized clade Sahonachelyidae, which is sister to the speciose clade formed by Bothremydidae and Podocnemidoidae. A close relationship with coeval Indian turtles of the clade Kurmademydini is notably absent. A functional assessment suggests that Sahonachelys mailakavava was a specialized suction feeder that preyed upon small-bodied invertebrates and vertebrates. This is a unique feeding strategy among crown pelomedusoids that is convergent upon that documented in numerous other clades of turtles and that highlights the distinct evolutionary pathways taken by Madagascan vertebrates.

Dental morphology of Adalatherium hui (Mammalia, Gondwanatheria) from the LATE Cretaceous of Madagascar

ABSTRACT The dentition in the only known specimen of Adalatherium hui, a gondwanatherian mammal from the Late Cretaceous of Madagascar, is markedly different from that of any previously known mammaliaform, extinct or extant. The two preserved upper incisors and single lower incisor of Adalatherium are each very large, open-rooted, and bear a restricted band of enamel. A shallow-rooted, rudimentary upper canine is present. The upper postcanine dentition is composed of a small, simple, two-rooted tooth mesially and four large, multiple-rooted (five roots or more) teeth distally. The latter are quadrangular in outline, each with four major cusps and three perimetric ridges bordering a central valley that opens buccally. There are four lower postcanines, also with four major cusps each. The first postcanine has two roots, but the following ones each have (or likely had) at least four. The two distal lower postcanines bear a prominent mesiobuccal basin. The cusps and other topographic features of the postcanines cannot be unambiguously homologized with those of other known mammaliaforms. Diastemata are present in the upper dentition between the incisors and canines and between the canines and postcanines, and in the lower dentition between the incisors and postcanines. Adalatherium, like several gondwanatherians from the Late Cretaceous and Paleogene of Argentina, retains relatively plesiomorphic, ‘normal’ radial enamel (i.e., single-layered schmelzmuster, non-decussating small prisms, interprismatic matrix anastomosing around prisms). This stands in contrast to the modified radial enamel found in other gondwanatherians from the Late Cretaceous of India and Madagascar, which exhibit prominent interrow sheets of interprismatic matrix.

Lower jaw morphology of Adalatherium hui (Mammalia, Gondwanatheria) from the Late Cretaceous of Madagascar

ABSTRACT The lower jaw of the holotype of Adalatherium hui, from the Late Cretaceous of Madagascar, is the most complete yet known for a gondwanatherian mammal. It reveals for the first time the morphology of the character-rich ascending ramus of the dentary in a gondwanatherian. Each half of the lower jaw is composed of only one bone, the dentary, which is short and deep and houses only five teeth: an enlarged, procumbent incisor and four postcanine teeth. In comparable parts of its anatomy, the dentary of Adalatherium is strikingly similar to that of Sudamerica but differs slightly from that of Galulatherium (conformation anterior to first postcanine, mental foramen position), the only two other gondwanatherians represented by complete horizontal rami. Among other Mesozoic mammaliaform taxa, the dentary of Adalatherium is most similar to those of the largely Laurasian group Multituberculata, most notably in absence of postdentary trough and Meckelian sulcus; presence of short, deep dentary with sizable diastema and articulating with squamosal via mediolaterally narrow condyle that continues onto posterior surface (i.e., no distinct peduncle); possession of much reduced dentition; absence of angular process; possession of large pterygoid fossa and pterygoid shelf, ventral surface of which is flat; absence of coronoid bone; and possession of unfused mandibular symphysis. Most of these features are clearly derived and stand in stark contrast to the much more plesiomorphic morphology exhibited by the lower jaw of the haramiyaviid Haramiyavia. The lower jaws of euharamiyidans, although derived in their own right, are also relatively plesiomorphic.

Inner ear morphology of Adalatherium hui (Mammalia, Gondwanatheria) from the Late Cretaceous of Madagascar

ABSTRACT The cochlear canal of Adalatherium hui, from the Late Cretaceous of Madagascar, exhibits a combination of features previously unknown among mammaliaforms. The cochlear canal of Adalatherium is short and ‘C’-shaped (curved through at least 210°). A presumptive lagenar nerve canal extends from the apex of the cochlea to the internal acoustic meatus. In conjunction with an apical expansion of the cochlear canal, this morphology suggests that Adalatherium retained a lagenar macula. Adalatherium is derived among mammaliaforms in possessing primary and secondary osseous laminae, a cribriform plate, and a distinct cochlear ganglion canal. Strikingly, Adalatherium shows a single-layered primary osseous lamina that lacks internal, radially oriented canaliculi for auditory nerve fibers. In this respect, the primary lamina of Adalatherium is structurally different from the double-layered primary lamina containing habenulae perforatae of extant therians. Furthermore, Adalatherium resembles the gondwanatherian Vintana in possessing a secondary canal running parallel to the cochlear ganglion canal. Although its exact role and homology are uncertain, the morphology of this secondary canal and its tributary canaliculi is most consistent with a vascular function. The inner ear of Adalatherium augments a growing list of characters (or character combinations) in which gondwanatherians are not only different from the inferred plesiomorphic condition for mammaliaforms but also distinct from the derived characters of therians. Collectively, the new morphological and phylogenetic data support homoplastic transformation of such structures as the cribriform plate and primary osseous lamina in mammals and independent loss of the lagenar macula within gondwanatherians.

Introduction to Adalatherium hui (Gondwanatheria, Mammalia) from the Late Cretaceous of Madagascar

ABSTRACT Adalatherium hui is a latest Cretaceous (Maastrichtian) gondwanatherian mammal based on only a single specimen, a virtually complete, articulated, and well-preserved skull and postcranial skeleton. The specimen is the most complete and best preserved of any mammaliaform from the Mesozoic of the southern supercontinent Gondwana. It was discovered in the Anembalemba Member of the Maevarano Formation in the Mahajanga Basin of northwestern Madagascar. The holotype specimen includes the only complete lower jaw and the only postcranial remains known for Gondwanatheria, which, other than the cranium of Vintana sertichi (also from the latest Cretaceous of Madagascar), are represented only by isolated teeth and fragmentary dentaries. Despite being represented by an immature individual, A. hui is third only to V. sertichi and Coloniatherium cilinskii as the largest Mesozoic mammaliaform (based on body fossils) from Gondwana. Here, we (1) review the paucity of mammaliamorph skull and postcranial skeletal material from the Mesozoic of Gondwana relative to the record from Laurasia; (2) review the systematic paleontology of A. hui; (3) provide an overview of the history of discovery of the holotype specimen; (4) detail the preservation of the holotype, its preparation history, and the imaging techniques used to study it; (5) provide an overview of the geological context of A. hui, which indicates that the species lived close to the end-Cretaceous extinction event in a highly seasonal, semiarid climate; and (6) estimate the body mass of A. hui in the context of other Mesozoic mammaliaforms.

The Mesozoic Biogeographic History of Gondwanan Terrestrial Vertebrates: Insights from Madagascar's Fossil Record

The Mesozoic plate tectonic and paleogeographic history of Gondwana had a profound effect on the distribution of terrestrial vertebrates. As the supercontinent fragmented into a series of large landmasses (South America, Africa-Arabia, Antarctica, Australia, New Zealand, the Indian subcontinent, and Madagascar), particularly during the Late Jurassic and Cretaceous, its terrestrial vertebrates became progressively isolated, evolving into unique faunal assemblages. We focus on four clades that, during the Mesozoic, had relatively low ability for dispersal across oceanic barriers—crocodyliforms, sauropod dinosaurs, nonavian theropod dinosaurs, and mammals. Their distributions reveal patterns that are critically important in evaluating various biogeographic hypotheses, several of which have been informed by recent discoveries from the Late Cretaceous of Madagascar. We also examine the effects of lingering, intermittent connections, or reconnections, of Gondwanan landmasses with Laurasia (through the Caribbean, Mediterranean, and Himalayan regions) on the distributions of different clades. ▪ This article reviews the biogeographic history of terrestrial vertebrates from the Mesozoic of the southern supercontinent Gondwana. ▪ Relatively large, terrestrial animals—including crocodyliforms, sauropod and nonavian theropod dinosaurs, and mammals—are the focus of this review. ▪ Most patterns related to vicariance occurred during the Late Jurassic and Cretaceous, the intervals of most active Gondwanan fragmentation. ▪ Recent discoveries of vertebrates from the Late Cretaceous of Madagascar have played a key role in formulating and testing various biogeographic hypotheses.

A new mammal from the Upper Cretaceous (Turonian–Campanian) Galula Formation, southwestern Tanzania

We here establish a new mammaliaform genus and species, Galulatherium jenkinsi (Mammalia), from the Upper Cretaceous Galula Formation in the Rukwa Rift Basin of southwestern Tanzania. This represents the first named taxon of a mammaliaform from the entire Late Cretaceous of continental Afro-Arabia, an interval of 34 million years. Preliminary study of the holotypic and only known specimen (a partial dentary) resulted in tentative assignation to the Gondwanatheria, a poorly known, enigmatic clade of Late Cretaceous-Paleogene Gondwanan mammals (Krause et al. 2003). The application of advanced imaging (mu CT) and visualization techniques permits a more detailed understanding of key anatomical features of the new taxon. It reveals that the lower dentition consisted of a large, procumbent lower incisor and four cheek teeth, all of which were evergrowing (hypselodont). Importantly, all of the teeth appear devoid of enamel. Comparisons conducted with a range of Mesozoic and selected Cenozoic mammaliaform groups document a number of features (e.g., columnar, enamel-less and evergrowing teeth, with relatively simple occlusal morphology) expressed in Galulatherium that are reminiscent of several distantly related groups, making taxonomic assignment difficult at this time. Herein we retain the provisional referral of Galulatherium (RRBP 02067) to Gondwanatheria; it is most similar to sudamericids such as Lavanify and Bharratherium from the Late Cretaceous of Madagascar and India, respectively, in exhibiting relatively simple, high-crowned, columnar cheek teeth. Other features (e.g., enamel-less dentition) are shared with disparate forms such as the Late Jurassic Fruitafossor and toothed xenarthrans (e.g., sloths), here attributed to convergence. Revised analyses of the depositional context for the holotype place it as having lived sometime between the late Turonian and latest Campanian (roughly 91-72 million years ago). This enhanced geochronological context helps to refine the palaeobiogeographical significance of Galulatherium among Cretaceous mammals in general and those from Gondwanan landmasses specifically.

Bite force in the horned frog (Ceratophrys cranwelli) with implications for extinct giant frogs

Of the nearly 6,800 extant frog species, most have weak jaws that play only a minor role in prey capture. South American horned frogs (Ceratophrys) are a notable exception. Aggressive and able to consume vertebrates their own size, these “hopping heads” use a vice-like grip of their jaws to restrain and immobilize prey. Using a longitudinal experimental design, we quantified the ontogenetic profile of bite-force performance in post-metamorphic Ceratophrys cranwelli. Regression slopes indicate positive allometric scaling of bite force with reference to head and body size, results that concur with scaling patterns across a diversity of taxa, including fish and amniotes (lizards, tuatara, turtles, crocodylians, rodents). Our recovered scaling relationship suggests that exceptionally large individuals of a congener (C. aurita) and extinct giant frogs (Beelzebufo ampinga, Late Cretaceous of Madagascar) probably could bite with forces of 500 to 2200 N, comparable to medium to large-sized mammalian carnivores.

Endocranial and Inner Ear Morphology of Vintana Sertichi (Mammalia, Gondwanatheria) from the Late Cretaceous of Madagascar

ABSTRACTWe present the first digital reconstruction of the endocranial cavity and endosseous labyrinth of the Late Cretaceous gondwanatherian mammal Vintana sertichi from the Maevarano Formation of Madagascar. The Malagasy specimen is exceptionally well preserved and represents the only described cranium known for Gondwanatheria, an enigmatic clade from the Late Cretaceous and Paleogene of Gondwana. The endocranial cast of Vintana is relatively small for an animal of its estimated body mass. Its encephalization quotient is 0.28–0.56 for a range of body mass estimates, which is similar to that of basal mammaliaforms. The olfactory bulbs are very large, occupying over 14% of the endocranial volume. The cerebral hemispheres are only slightly expanded, more similar to the condition in Morganucodon than to that in multituberculates and monotremes. Unlike the condition in other Mesozoic mammaliaforms, the endocast is greatly flexed at the circular fissure. The osseous labyrinth displays a mixture of derived and primitive features. The cochlear canal is only slightly curved and short compared with that of extant therians. The ratio between total cochlear canal length and maximum cranial length is smaller than in basal mammaliaforms and approximates that of non-mammaliaform cynodonts. By contrast, the presence of both primary and secondary osseous laminae, the tractus foraminosus, and Rosenthal's canal represent derived characteristics of the mammalian inner ear typical of cladotherians. A modern innervation of the cochlea has either evolved independently in Vintana and cladotherians or was already present in the last common ancestor of both clades.SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP

Introduction, Systematic Paleontology, and Geological Context of Vintana Sertichi (Mammalia, Gondwanatheria) from the Late Cretaceous of Madagascar

ABSTRACT—Vintana sertichi is a sudamericid gondwanatherian mammal known only from the Upper Cretaceous (Maastrichtian) Maevarano Formation in the Mahajanga Basin of northwestern Madagascar. It is based on a single specimen, a well-preserved and virtually complete cranium discovered in 2010 near Lac Kinkony. The cranium is superficially bizarre and constitutes the only cranial remains of the poorly known, phylogenetically enigmatic Gondwanatheria, which are otherwise known only from isolated teeth and fragmentary dentaries. Gondwanatheria are represented by seven other monotypic genera assigned to two families (Ferugliotheriidae and Sudamericidae). Historically, this clade was assigned to Xenarthra, Paratheria (as a sister group to Xenarthra), Multituberculata, Allotheria (as a sister group to Multituberculata), Mammalia incertae sedis, and, most recently, back to Multituberculata or a close relative of Multituberculata. The craniodental evidence provided by Vintana supports inclusion in Sudamericidae, the monophyly of Gondwanatheria, and the position of Gondwanatheria as nested within or sister to Multituberculata. In addition to briefly reviewing the taxonomic composition and phylogenetic history of Gondwanatheria, this introductory chapter sets the stage for the other chapters in the volume by (1) briefly summarizing the inferred life habits of gondwanatherians; (2) reviewing the systematic paleontology of V. sertichi; and (3) providing overviews of the discovery of the holotypic specimen, its preservation, its preparation, and the imaging and measurement techniques used to study it. The chapter closes with an overview of the geological context of V. sertichi, which indicates that the species lived in a coastal floodplain environment and in a highly seasonal, semiarid climate.

Sensory Anatomy and Sensory Ecology of Vintana Sertichi (Mammalia, Gondwanatheria) from the Late Cretaceous of Madagascar

ABSTRACT—Living mammals are distinguished from other extant tetrapods by adaptations for improved senses of hearing, touch, and smell. These adaptations, and concomitant reductions in visual anatomy, evolved during the Mesozoic in the mammalian and therian stem lineages. Here, we present a comparative study of the sensory anatomy of the Late Cretaceous gondwanatherian mammaliaform Vintana sertichi in order to draw inferences regarding its sensory abilities and sensory ecology. Our analyses demonstrate that Vintana has relatively large orbits that may have accommodated large eyes. Vintana also possessed cochlear primary and secondary osseous laminae and a cochlear canal that was relatively longer than in non-mammaliaform cynodonts but shorter than in extant therians. These features suggest that Vintana had some capacity for high-frequency hearing (i.e., >20 kHz), but that its cochlea may have encoded a more limited range of frequencies than the cochleas of most extant therians. The semicircular canals of Vintana have large radii of curvature and are nearly orthogonal, suggesting high sensitivity to angular head accelerations. These vestibular features may have evolved in order to stabilize large eyes during rapid and/or agile locomotion. Combined with evidence for large olfactory bulbs and a large trigeminal endocast, these data reveal that Vintana possessed a unique suite of sensory adaptations that distinguish it from other Mesozoic mammaliaforms. If these inferences are correct, then Vintana was probably a large-eyed and agile species, with a keen sense of smell and better high-frequency hearing than most other Mesozoic mammaliaforms.SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP

Dental Function and Diet of Vintana Sertichi (Mammalia, Gondwanatheria) from the Late Cretaceous of Madagascar

ABSTRACT—The nearly complete and well-preserved cranium of Vintana sertichi provides an opportunity to investigate its dietary adaptations. We used a combination of comparative morphological and biomechanical analyses to reveal the direction of its power stroke during mastication, reconstruct the positions and relative sizes of its muscles of mastication, and predict its capacity to produce bite forces during incision and molariform occlusion. The orientation of dental wear striations on the upper molariform teeth of Vintana, in combination with the orientation of the enamel islets and synclines and the position of leading and trailing edges, demonstrates that the power stroke was primarily palinal (distally directed), with a significant buccally directed component that is absent in multituberculates, haramiyidans, and other gondwanatherians. The large palinal component of jaw movement was corroborated by biomechanical analyses of the moments generated by the primary jaw adductors around the dentary-squamosal joint axis. Similar analyses also confirmed the previously documented proal (mesially directed) power stroke of the extant rodent Myocastor. Finite element analyses predict that Vintana was capable of producing bite forces that were more than twice as high as the similarly sized Myocastor. Vintana was almost certainly an herbivore, as claimed previously for other sudamericid gondwanatherians. Its size and capacity to generate high bite forces at both the incisors and the molariform teeth suggest that it was a mixed feeder whose diet may have included relatively large, hard food items such as roots, seeds, twigs, or nut-like fruits.

Enamel Microstructure of Vintana Sertichi (Mammalia, Gondwanatheria) From the Late Cretaceous of Madagascar

ABSTRACTThe enamel microstructure of the molariform cheek teeth of Vintana sertichi (Gondwanatheria, Mammalia) from the Late Cretaceous of Madagascar retains many of the plesiomorphic characteristics of mammalian prismatic enamel (e.g., single-layered schmelzmuster; non-decussating, small prisms). However, V. sertichi exhibits a relatively derived condition and resembles Lavanify miolaka (Late Cretaceous, Madagascar), Bharattherium bonapartei (Late Cretaceous, India), and an indeterminate sudamericid (Late Cretaceous, Madagascar) in possessing modified radial enamel, consisting of prisms separated by prominent interrow sheets of interprismatic matrix. A second group of gondwanatherians (Ferugliotherium windhauseni, Gondwanatherium patagonicum, Sudamerica ameghinoi, Greniodon sylvaticus), all from the Late Cretaceous and Paleogene of Argentina, retains relatively primitive, ‘normal’ radial enamel, in which the interprismatic matrix anastomoses around the prisms and does not form interrow sheets. The prisms of gondwanatherian molariforms generally become wider from the enamel-dentine junction to the outer enamel surface, dramatically so in Gondwanatherium. Seams are very rare; tubules are more common. Prism decussation has not been found in any gondwanatherian enamel. Gondwanatherian molariforms lack or have only a thin layer of external prismless enamel, although it is thick in the incisors of Sudamerica. Despite their apparent utility in differentiating among gondwanatherian taxa, or gondwanatherians as a whole from other clades, many of the derived characteristics of gondwanatherian enamel microstructure are found in other mammalian groups and may reflect a high degree of homoplasy.

New snakes from the Upper Cretaceous (Maastrichtian) Maevarano Formation, Mahajanga Basin, Madagascar

ABSTRACTWe describe three new fossil snakes on the basis of recently discovered vertebrae collected from the Upper Cretaceous (Maastrichtian) Maevarano Formation of Madagascar. One represents a new genus and species of madtsoiid, Adinophis fisaka, the third member of this family recognized from the Maevarano Formation. It exhibits dorsoventrally compressed centra and dorsally placed synapophyses. The second taxon is a new species of the nigerophiid genus Indophis, I. fanambinana, representing the second nigerophiid taxon known from the Maevarano Formation. It shares numerous features with the Indian nigerophiid I. sahnii, including small synapophyses positioned ventrolaterally on distinct synapophyseal processes and a unique cotylar shape in posterior trunk vertebrae. The discovery of I. fanambinana demonstrates a strong biogeographic link with penecontemporaneous snake faunas from India. A third new taxon is represented by a partial centrum that cannot be assessed thoroughly due to its incompleteness; this specimen is conservatively assigned to Serpentes incertae sedis, gen. et sp. indeterminate. It is distinct from other snakes known from the Late Cretaceous of Madagascar in possessing a strongly depressed condyle and relatively large lateral foramina. These new discoveries, together with previous descriptions of snakes from the Maevarano Formation, make the Maevarano snake fauna one of the most taxonomically diverse snake assemblages known from the Mesozoic and the most diverse in terms of body size range.

Vahiny depereti, gen. et sp. nov., a new titanosaur (Dinosauria, Sauropoda) from the Upper Cretaceous Maevarano Formation, Madagascar

ABSTRACTHere we describe Vahiny depereti, gen. et sp. nov., a new titanosaur dinosaur from the Upper Cretaceous Maevarano Formation of northwestern Madagascar. Vahiny is distinguished from other titanosaurs by characteristics of the basal tubera, basipterygoid processes, parasphenoid, and cranial nerve foramina. Diagnostic cranial material of Vahiny formalizes the long-standing hypothesis that two titanosaur genera were present in the Late Cretaceous of Madagascar. The other titanosaur, Rapetosaurus krausei, is the most common dinosaur in the fauna and is known from hundreds of bones, including multiple partial skeletons and skulls. In contrast, non-Rapetosaurus elements are extremely rare. Of these elements, which have been called ‘Malagasy Taxon B,’ we identify a partial braincase as the holotype of Vahiny depereti. Although Vahiny and Rapetosaurus coexisted on the island of Madagascar, differences in their braincases indicate that they are not closely related to one another. Instead, Vahiny is most similar to Jainosaurus from the Late Cretaceous of India, and shares similarities with the South American taxa Muyelensaurus and Pitekunsaurus.

New material of Beelzebufo, a hyperossified frog (Amphibia: Anura) from the late cretaceous of Madagascar.

The extant anuran fauna of Madagascar is exceptionally rich and almost completely endemic. In recent years, many new species have been described and understanding of the history and relationships of this fauna has been greatly advanced by molecular studies, but very little is known of the fossil history of frogs on the island. Beelzebufo ampinga, the first named pre-Holocene frog from Madagascar, was described in 2008 on the basis of numerous disarticulated cranial and postcranial elements from the Upper Cretaceous (Maastrichtian) Maevarano Formation of Madagascar. These specimens documented the presence of a hyperossified taxon that differed strikingly from extant Malagasy frogs in its large size and heavy coarse cranial exostosis. Here we describe and analyse new, articulated, and more complete material of the skull, vertebral column, and hind limb, as well as additional isolated elements discovered since 2008. μCT scans allow a detailed understanding of both internal and external morphology and permit a more accurate reconstruction. The new material shows Beelzebufo to have been even more bizarre than originally interpreted, with large posterolateral skull flanges and sculptured vertebral spine tables. The apparent absence of a tympanic membrane, the strong cranial exostosis, and vertebral morphology suggest it may have burrowed during seasonally arid conditions, which have been interpreted for the Maevarano Formation from independent sedimentological and taphonomic evidence. New phylogenetic analyses, incorporating both morphological and molecular data, continue to place Beelzebufo with hyloid rather than ranoid frogs. Within Hyloidea, Beelzebufo still groups with the South American Ceratophryidae thus continuing to pose difficulties with both biogeographic interpretations and prior molecular divergence dates.

A new, richly fossiliferous member comprised of tidal deposits in the Upper Cretaceous Maevarano Formation, northwestern Madagascar

A new member of the Upper Cretaceous (Maastrichtian) Maevarano Formation is proposed to accommodate a distinctive succession of strata exposed along the shores of Lac Kinkony in northwestern Madagascar. The new Lac Kinkony Member overlies fully terrestrial sandstones of the Anembalemba Member of the Maevarano Formation, and is capped by marine dolostones of the Berivotra Formation. In the stratotype section, the base of the Lac Kinkony Member consists of siltstone interbeds that host networks of Ophiomorpha. Siltstone facies pass up-section to distinctive white sandstones packed with dolomitic mud matrix that exhibit rhythmic clay drapes, flaser and wavy bedding, and oppositely-oriented ripples developed on the toes of larger foresets. Thin flat interbeds of microgranular dolostone and claystone comprise the uppermost facies of the Lac Kinkony Member, and a laterally traceable ravinement bed mantled by cobbles of rounded dolostone marks the contact with the superjacent Berivotra Formation. Deposits of the Lac Kinkony Member are interpreted to represent siliciclastic and carbonate tidal flats dissected by tidally-influenced rivers. Vertebrate fossils are abundantly preserved in these coastal deposits, and are locally concentrated in microfossil bonebeds that have the potential to yield thousands of small identifiable specimens. In addition to many taxa already known from the Maevarano Formation, the Lac Kinkony Member has yielded a wealth of phyllodontid albuloid fish skull elements, the distal humerus of a new frog taxon, five vertebrae representing two new snakes, a tooth of a possible dromaeosaurid, and a complete skull of a new mammal. The discovery of several new vertebrate taxa from this new member reflects the fact that it samples a previously unsampled nearshore, peritidal paleoenvironment in the Late Cretaceous of Madagascar.

Gondwanatheria and ?Multituberculata (Mammalia) from the Late Cretaceous of Madagascar

In addition to four isolated mammalian teeth from the Late Cretaceous (Maastrichtian) of Madagascar described previously and allocated to Gondwanatheria, Marsupialia, and Mammalia incertae sedis, here I put on record five more specimens. Four of these, a virtually complete lower molariform cheek tooth, two fragmentary cheek teeth, and a fragmentary lower incisor, are referred to the Sudamericidae (Gondwanatheria). The internal structure of the hypsodont cheek teeth, as revealed by micro-computed tomography scans to simulate different stages of wear, is highly variable. Limited knowledge of intra-individual morphological and size variability in the dentition of sudamericids, which are known almost exclusively from isolated teeth, precludes a conclusive assessment of whether some or all of the new specimens belong to Lavanify miolaka, the only previously described sudamericid from Madagascar, or to a new taxon. Conservatively, therefore, pending the recovery of better material, all four specimens are referred to Sudamericidae gen. et sp. indet. The fifth specimen, a molar fragment, is tentatively allocated to the Multituberculata, thereby adding to the controversial specimens that comprise the very sparse and questionable record of this clade on the southern supercontinent Gondwana. The new specimens do not provide any profound insight into the origins of the highly endemic and imbalanced extant mammalian fauna on Madagascar; rather, they provide only more negative evidence. Like those previously described, they do not represent the basal stocks of any of the five mammalian clades that live on the island today and therefore further support the growing consensus that representatives of the extant clades arrived in the Cenozoic.

An articulated pectoral girdle and forelimb of the abelisaurid theropod Majungasaurus crenatissimus from the Late Cretaceous of Madagascar

ABSTRACT Abelisaurid theropods are common members of Cretaceous Gondwanan faunas and are characterized by a bizarre, highly reduced forelimb. Unfortunately, forelimb elements are rarely preserved and thus the basic structure of the abelisaurid forelimb remains poorly understood. Until recently, the Upper Cretaceous Maevarano Formation of northwestern Madagascar has produced numerous exceptional specimens of the abelisaurid theropod Majungasaurus crenatissimus but comparatively little forelimb material. A recently discovered articulated skeleton of Majungasaurus preserves a virtually complete pectoral girdle and forelimb, which, along with additional isolated forelimb elements, affords important new insights into the structure of these elements. New specimens of the scapulocoracoid and humerus allow more detailed description of their morphology, and antebrachial and manual elements are described for the first time. The radius and ulna are approximately one-quarter the length of the humerus and both have expanded proximal and distal articular surfaces relative to their narrow diaphyses. The manus consists of four digits, each composed of a short metacarpal and one (digits I and IV) or two (digits II and III) phalanges. No ossified carpals are present. The proportions of the brachium and antebrachium are stout, more similar to the condition in Carnotaurus than in Aucasaurus. We reinterpret manual digit identities in Aucasaurus and Carnotaurus based on new information provided by the manus of Majungasaurus. Overall, the morphology of the forelimb in Majungasaurus reveals that abelisaurids share an extremely reduced, unique morphology that is dissimilar to the more typical theropod condition seen in other ceratosaurs.

Sauropod dinosaur osteoderms from the Late Cretaceous of Madagascar

Osteoderms are bones embedded within the dermis, and are common to select members of most major tetrapod lineages. The largest known animals that bear osteoderms are members of Titanosauria, a diverse clade of sauropod dinosaurs. Here we report on two titanosaur osteoderms recovered from the Upper Cretaceous Maevarano Formation of Madagascar. Each osteoderm was discovered in association with a partial skeleton representing a distinct ontogenetic stage of the titanosaur Rapetosaurus krausei. Combined, these specimens provide novel insights into the arrangement and function of titanosaur osteoderms. Taphonomic data confirm that Rapetosaurus developed only limited numbers of osteoderms in its integument. The adult-sized osteoderm is the most massive integumentary skeletal element yet discovered, with an estimated volume of 9.63 litres. Uniquely, this specimen possesses an internal cavity equivalent to more than half its total volume. Large, hollow osteoderms may have functioned as mineral stores in fecund, rapidly growing titanosaurs inhabiting stressed environments.