The revision of the Rhombodontidae from the Maastrichtian of Morocco led us to the description of a new species: Rhombodus andriesi.

The biometrical study of populations of R. binkhorsti and R. microdon, species sometimes considered as synonymous, supports the conclusion that they represent two distinct species and not young and adult specimens of a single species. The stratigraphical range of these two species confirms this
result.

The reexarnination of type-series studied by C. Ararnbourg led us to revise the generic status of Rhombodus bondoni which is now ranged, because his dental features, in the new genus Dasyrhombodus showing a less derived dentition than Rhombodus. The stratigraphical range of each species is clarified within the phosphatic series. 

Pangolins have never shown a high taxic diversity and their fossil record is scarce. We report here the first discovery of a partial humerus from late Oligocene deposits in Pech du Fraysse (MP28, France). The new specimen from Pech du Fraysse is described and compared to various extant and extinct species of pangolins. It shows a suite of morphological features very similar to the humeri discovered in Saint-André (MP 26), Solnhofen (Burdigalien), and Saulcet (Aquitanian), attributed here to Necromanis franconica. The description of the specimen from Pech du Fraysse allowed us to discuss the systematics of Paleogene and Neogene pholidotans. The differences between PFY 4051 and N. franconica on the one side, and N. quercyi on the other side, might be sufficiently important to justify a generic distinction. A comparison with extant species showed that N. franconica was likely terrestrial and fossorial based on its humeral morphology. 

Tetraceratops insignis is known from a single, crushed skull from the Lower Permian of Texas. Its unique proportions and osteological details gained central meaning in the question of the origins of Therapsida since this early synapsid has been determined as the oldest and less derived therapsid. Apart from Tetraceratops, the ‘mammal-like’ Therapsida and their sister, the pelycosaur-grade Sphenacodontidae, are separated by one of the longest ghost lineages in tetrapod fossil record. However, the minor, though well justified critique faced insistent publication regarding the therapsid hypothesis. A carefull re-evaluation of the holotypic skull reveals that therapsid traits cannot be supported, including a rejection of the formerly supposed adductor shelf in the temporal fenestra. Increased understanding of ‘pelycosaur’ character variation underlines a haptodontine-grade or, less likely, sphenacodontid position for Tetraceratops. 

The East African Miocene Macroscelidea, lnsectivora and Chiroptera are revised on the basis of new material. New taxa proposed are: Miorhynchocyon, .n. gen. (Macroscelididae): Míorhynchocyon meswae, n. sp.: Pronasílío ternanensis. n. gen.. n. sp. (Macroscelididae); Hiwegicyon juvenalis, n. gen. n. sp. (Macroscelididae); Parageogale, n. gen. (Tenrecidae): Prochrysochlorinae, n. subfam. (Chrysochloridae): Propottininae, n. subfam, (Pteropodidae); Chamtwaria pickfordi, n. gen., n. sp. (Vespertilionidae). Gymnurechnínus songhorensis is synonymised with G. camptolophus. The new material provides additional information on the dentition, especially of Myohyrax oswaldi. Galerix africanus. Amphechínus rusingensis, Protenrec tricuspis and Parageogale aletris. Partial skulls are described of Amphechinus rusingensis, Protenrec tricuspis, Prochrysochloris míocaenicus and Taphozous incognita. The oldest member of the Macroscelidinae (Pronasilio) is described from Fort Ternan. Galerix africanus is closely related to G. exilis from Europe. Amphechinus rusingenesis is compared with Asiatic Oligocene Erinaceinae. The Miocene age of Crocidura is rejected. On the evidence of humeri, the following families of Chiroptera are newly reported: Pteropodidae, Nycterididae, Vespertilionidae, Molossidae. Propotto is regarded as an offshoot from the Pteropodidae, not ancestral to modern forms. Chamtwaria is a primitive vespertilionoid, provisionally placed in the Kerivoulinae. Erinaceidae probably entered Africa at the beginning of the Miocene, before 20 Ma. Faunistic differences between deposits are largely to be ascribed to differences in local environment. 

The Causses" is a near 3400 km2 large plateau located in the south of France. Here the first dinosaur footprints where found in 1935. After this, this area has yielded an ever-increasing number of ichnites now in excess of 500 specimens. These latter, 15 to 50 cm long, tridactyl or tetradactyl footprints of generally biped animals, were discovered at the surface of Hettangian to lower Sinemurian dolomite layers within 4 distinct stratigraphic units. The 35 sites bearing ichnites are located on the plateau margin. For the first time, morphologic characters studied through descriptive statistic methods with the usual parameters and classical Student and Snédecor tests, allowed us, to divide the whole set of biped traces into 6 ichnospecies. Their definitions are further constrained by multivariate statistical results using Principal Component Analysis (PCA), Factor Analysis of correspondances (FAC) and Discriminant Analysis (DA). All have confirmed the morphologic observations. So that now, the following taxa are identified : Grallator variabilis, G. lescurei, G. sauclierensis, G. minusculus, Eubrontes giganteus, Dilophosauripus williamsi, cf. Moraesichnium, Orníthopus fabrei nov ichnosp. The more immediately visible differences relate to the interdigital II-IV divarication and the digit length ratio. To this panel, we must add Batrachopus deweyi and shapes suggesting Trisauropodichnus and/or Anomoepus. Among all ichnite associations described in the lower Liasic, the New England assemblage presents the most affinities with ours. It shows the ichnotaxa Grallator, Dilophosauripus, Eubrontes, Batrachopus without forgetting Ornithopus fabrei nov. ichnosp. which is close to Ornithopus gallinaceus from the Massachusetts and Connecticut basins. On comparing the present early Jurassic ichnofauna of the Causses with the ones of the Middle and Upper Triassic formations of the eastem border of the Massif Central (France), it appears that tridactyl footprints become more and more numerous and large from Triassic to Early Jurassic. In the Causses, these latest are prevalent but in Quercy (France), Poland, Italy, USA, they are also associated with Omithopoda, Thyreophora and Sauropoda ichnites. Footprint areas considered here were generaly under an arid climate. Animals that passed by were heavy and bulky possible Megalosaur trackmakers, and lighter and slender Coelophysids or Ceratosaurs. For all, these areas were pathways as the orientations of the trackways seem point out. The directions followed by these reptiles were without any important variation during the Hettango-Sinemurian stages. These areas were also used from time to time by Crocodilomorpha and may be tetradactyl (I-IV) bipedal avian Theropods. However, the number of such trackways in sites, sometimes substantial, should not lead us to overestimate the trackmakers populations. These last were probably relatively moderately abondant in this inter-supratidal swamp environment. In the Causses, ichnites are connected with former algo-laminated deposits (Algal mats) which were rapidly hardened by means of calcitisation of cyanobacteria. The result has been a moderate depth of footprints; autopodia disturbing only a few cm of the carbonate substrate. Other fossils have been discovered : invertebrates with thin bivalve and gastropod shells, crustaceans tests and plants. These latter suggest the existence of paleomangroves like environments but also continental vegetation periodically overruning the swamp environment during regression/transgression cycles. At these times, wooded parts of it, could become protecting, feeding, resting and nesting places.

  

Near "la Lieude", in the Lodève basin, more than a thousand of footprints are distributed in a twenty of trackways which amounts to 220 m length. They have been found on calcareous siltstone level in the B site named also "Réserve Naturelle Volontaire". This last is located in the Saxonian summit dated Upper Permian. "La Lieude" tracks are described by using statistical methods then they are compared with others from the world Permian. What allows to distinguish 4 following ichnotaxa: Lunaepes ollierorum nov.ichnosp., Merifontichnus thalerius nov. ichnogen. and nov. ichnosp., Planipes brachydactylus nov.ichnosp. and Brontopus circagiganteus nov. ichnosp. All these traces are attributed with possibility or probability to Therapsida or to Therosauria, except Brontopus circagiganteus nov. ichnosp. that could be due to Caseamorpha. All these animals whose sizes have been estimated between l and 5 m lived probably in a playa environment.The biological and sedimentological data from "la Lieude" footprints levels compared with informations provided by the tracks orientations, suggest the following scenario. Animals coming from the North have crossed a sandy channel bank with plants zones by directing to the South for the majority. Maybe, they were going to the lacustrine part of the playa, close to "la Lieude" footprints that they have just trampled on. 

This paper describes the first hexanchid tooth from the Tithonian (Late Jurassic) of New Zealand. For the moment, this tooth represents the earliest representative of the fossil genus Notidanodon in the world and one of the most ancient neoselachians in the Southern Hemisphere. Despite the perfect state of preservation of the unique tooth, the species is left in open nomenclature, pending the discovery of additional specimens. Few nominal species have been assigned to the genus Notidanodon. Four from Cretaceous deposits: N. antarcti Grande & Chatterjee, 1987, Notidanodon dentatus (Woodward, 1886), Notidanodon lanceolatus (Woodward, 1886), Notidanodon pectinatus (Agassiz, 1843) and only two from Paleocene: Notidanodon brotzeni Siverson, 1995, and Notidanodon loozi (Vincent, 1876). Considering the important morphological variations observed between some of these species, it seems obvious that the genus Notidanodon is not monophyletic and will need a revision in the future. 
  

Comparison of various measurements of taxonomic evolution using stratigraphic range data for orders, families and genera of land mammals indicates several means by which deficiencies of the South American fossil record (e.g., presence of hiatuses, unequal temporal and geographic representation of ages, unequal systematic treatment) may be normalized, thus permitting a less distorted appreciation of diversity pattern and trend. Initial radiation of native taxa resulted in a relative equilibrium by early Eocene time. Subsequent increases in absolute diversity were apparently induced by immigration at the family level and by environmental factors at the generic level. Miocene through Pleistocene phases of faunal stability, herein characterized as chronofaunas, are punctuated by rapid turnover events resulting from a complex of factors, including adaptive radiation of immigrant taxa into unoccupied eco-space; environmental and concomitant habitat change induced by orogenic events of the Andes; and biotic interactions between native and immigrant taxa, including competition and prey naivete. The first two factors account for major faunal transitions in the South American middle and late Tertiary; immigration-induced turnover may have been of greater importance in shaping the character of the fauna upon the Great American Interchange and the arrival of man in the Neotropics 

A new Propalaeotherium species, clearly distinct from the genus Eurohippus, is described. It is characterized by having a similar size as P. voigti from the German Geiseltal localities (MP 11 to MP 13 reference-level), but differs in several features suggesting a slighty more derived morphology. It presents indeed less brachyodont crowns with less prominent and less elevated cingula, slightly larger relative surface of premolars, and a more marked metaconid splitting on cheek teeth. This new species is unknown from other European localities except the nearby Saint-Martin de Londres locality which has been considered older than the MP 13 level. 

Abstract
 The validity of Propachynolophus Lemoine, 1891, supposedly an intermediate between Hyracotherium Owen, 1841 and Pachynolophus Pomel, 1847, has been questioned for a long time. A detailed analysis of features on which this genus is based further supported by a formal cladistic analysis demonstrates that Propachynolophus is not a valid taxon. The type species, “Propachynolophus gaudryi Lemoine, 1891” shall be assigned to Propalaeotherium Gervais, 1849, under the new combination Propalaeotherium gaudryi (Lemoine, 1891). “Pachynolophus maldani Lemoine, 1878”, later assigned to Propachynolophus, typifies the new genus Orolophus, under the binomen Orolophus maldani (Lemoine, 1878). The other referred species, “Propachynolophus levei Hooker, 1994” and “P. remyi Checa-Soler, 1997” are poorly documented, and both species shall be provisionally referred to as “Hyracotherium” levei (Hooker, 1994) and “Hyracotherium” remyi (Checa-Soler, 1997), pending new discoveries.
 
  

Mammal-bearing localities have been discovered in the marine and lacustrine series of the middle Ilerdian (Lowermost Eocene) from Southem France (Minervois and Corbières). In the localities of Fordones, Monze, Fournès, and La Gasque, thirty mammal species have been identified. Among others, they include ischyromyid rodents (Microparamys and Pseudoparamys), paromomyid and adapid primates (Arcius and Donrussellia), new insectivores, condylarths, and a dyspternine pantolestid. These faunas provide new informations on the early Eocene Mesogean faunas of Rians and Palette. The assemblages of primates and rodents from Fordones support good  correlations with Palette which was recently placed near the standard-level of Dormaal (MP 7). In fact, Palette and Fordones could be even older than Dormaal. Consequently, there seems to be a relatively important temporal gap between the late Paleocene of Cernay and the Sparnacian of Dormaal. This gap could be partly filled with the Mesogean faunas of Palette, Fordones, and Silveirinha. On the basis of these new mammal faunas the marine middle Ilerdian is proved to be older than the Cuisian stage of the Paris Basin. With regards to the position of the Fordones fauna at the top of the NP 10 calcareous nannoplankton biozone, the westem European paleomammalogists Paleocene/Eocene boundary could be situated between the NP 9 and NP 10 biozones. 

Based on fossil discoveries and phylogenetic studies, an Eocene Asian origin for hystricognathous rodents and anthropoid primates has gained strong support in recent years. The two groups then invaded both Africa and South America, which promoted their evolutionary success. However, the fossil record has so far failed to constrain the nature and precise timing of these pivotal dispersal events. In Africa, given the apparent absence of hystricognaths and anthropoids in early to early middle Eocene localities, it is suggested that these mammal groups dispersed from Asia to Africa sometime during the middle Eocene. In this paper, we report the discovery of several isolated teeth of a rodent from a new vertebrate locality situated in central Tunisia (Djebel el Kébar, KEB-1), dating from the late middle Eocene (Bartonian, ~39.5 Myr). These fossils document a diminutive new species of Protophiomys (P. tunisiensis nov. sp.), a basal genus of hystricognathous rodents which is well known from several North African mammalian-bearing localities of the end of the Eocene. The teeth of P. tunisiensis display a suite of anatomical details comparable with those observed in the other species of the genus, but with a lesser degree of development. Such an apparent primitive evolutionary stage is corroborated by the greater antiquity of this Tunisian species. P. tunisiensis nov. sp. is so far the most ancient representative of hystricognaths in Africa. However, it can be expected that hystricognaths were already present on that landmass given the new data on early caviomorphs recently reported from South America (at ~41 Myr). The arrival of hystricognaths in Africa from South Asia certainly predates the depositional period of the Kébar sediments, but perhaps not by much time. 

Confident attribution of bipedal tridactyl dinosaur tracks to theropods or ornithopods can be challenging. Here we describe trackways produced by tetanuran dinosaurs, previously attributed to hadrosaurs, from Coahuila State, northeastern Mexico. Multiple trackways headed in the same direction suggest gregarious behaviour in these late Campanian theropods.  

Rhinoceros Giants: the Paleobiology of Indricotheres. Donald R. Prothero. Life of the Past Collection, Indiana University Press; 160 pp. (66 b&w illustrations). Hardback (7x10”): USD 42.00 plus shipping. ISBN: 978-0-253-00819-0. E-book: USD 34.99. ISBN: 978-0-253-00826-8.
  

The Late Miocene site of Gritsev (MN 9, Ukraine) has yielded a very rich bat fauna, the remains of which are well preserved. Compared to other Neogene bat assemblages of Europe, the Gritsev bat community is unique in preserving plecotine bats, which are rare from Neogene sites. Some peculiar and new bat species, including a large plecotin Otonycteris, already were described from the Gritsev mammal site. Here we report new records of small plecotin bats from Gritsev, including a new taxon, Barbastella maxima nov. sp. This is the earliest reliable fossil record of this genus and it differs from more recent species of Barbastella in being considerably larger. The evolutionary patterns in the odontology within the tribe Plecotini, supported by biostratigraphical distribution of fossil records of Plecotus are discussed. The morphological peculiarities of the new fossils of plecotine bats from Gritsev are discussed in connection with its possible taxonomical affinity. 

A brief historical account of fossil vertebrate discoveries in the Eastern part of the Paris Basin between the beginning of the nineteenth century and 1950 is given. Other localities discovered since then are presented. A reconstruction of past landscapes is briefly elaborated. A biozonation based on mammals is proposed, from the Late Thanetian to the Middle Bartonian. Paleogeographical considerations are added. Suggestions regarding the search for new marnmal localities are made. 

The Las Águilas site near Porvenir de Jalpa, Coahuila, Mexico, is extremely rich in tetrapod remains comprising both bones and trackways of several dinosaur taxa of late Campanian age. Within a 50 m thick section we identified at least nine layers with dinosaur bone assemblages. In one of these the dinosaur bones are associated with remnants of eusuchian crocodilians, turtles, plesiosaurs, pterosaurs, tyrannosaurids, dromaeosaurids, parksosaurid, hadrosaurids, ceratopsids, and ankylosaurs. This layer is also rich in coprolites of turtles, crocodilians and likely theropods, thus providing evidence for the wealth of Late Cretaceous vertebrate life in the area. 

Résumé :
L’étude des rongeurs Theridomorpha permet de suivre le déroulement d’une radiation adaptative pendant toute sa durée (Eocène moyen-Oligocène terminal), sur un territoire restreint à l’extrémité ouest de l’Europe Occidentale. Dans ce papier, la phylogénie de ce groupe est établie à partir d’une analyse cladistique reposant sur l’examen de 315 caractères (310 dentaires). Le groupe d’intérêt comprend 110 des 132 espèces (24 genres) de Theridomyoidea et deux genres encore inclus jusqu’ici dans les Reithroparamyinae qui rejoignent les Theridomorpha. Les groupes externes comprennent des Glires basaux, Cocomys, Tanquammys et 16 Ischyromyiformes. Un cadre phylogénétique robuste est produit, qui permet de clarifier la systématique des Theridomorpha. La position des Remyoidea (nov. sup.fam.) au sein des Ischyromyiformes, extérieure aux Theridomorpha, est confortée. Les Protadelomys et Tardenomys sont à la base des Theridomyoidea, avant la séparation en deux clades correspondant aux familles Pseudosciuridae et Theridomyidae. Les sous-familles sont consolidées : Pseudosciurinae et Sciuroidinae pour les Pseudosciuridae ; Issiodoromyinae, Oltinomyinae, Columbomyinae, Theridomyinae, auxquelles s’ajoute au moins une nouvelle sous-famille (Patriotheridomyinae), pour les Theridomyidae. La topologie des chrono-espèces (sensu Simpson), traitées antérieurement comme lignées évolutives, apparaît dans la plupart des cas sous forme de clades successifs dans lesquels les espèces sont le plus souvent arrangées de manière pectinée, émergeant dans l’ordre stratigraphique. L’analyse des caractères aux principaux nœuds permet de consolider les caractères diagnostiques des taxons et les tendances évolutives, ainsi que de discuter des divers parallélismes et convergences dans l’évolution des structures et patrons dentaires (e.g., émail des incisives unisérié chez les Issiodoromyinae et les Patriotheridomyinae, ou pseudo-multisérié chez les Blainvillimys les plus hypsodontes, les Protechimys et Archaeomys ; patrons dentaires téniodontes ; allongement des dents déciduales chez les Patriotheridomyinae, Issiodoromyinae et Theridomyidae ; sélénodontie ou lophodontie). Les dynamiques évolutives traduites par les changements morphologiques sont mises en relation avec les variations environnementales. Enfin, les implications biochronologiques de l’évolution des Theridomyoidea sont discutées.
Abstract:
The adaptive radiation of the rodents Theridomorpha occurred during a limited time window (middle Eocene to late Oligocene), on an area restricted to Western Europe. In this paper, the phylogeny of this group is established via a cladistic assessment of 315 morphological characters (310 dental). The group of interest encompasses 110 upon 132 species (24 genera) of Theridomyoidea, and two genera formerly included within the Reithroparamyinae, and which are included here within the Theridomorpha. The outgroups include basal Glires, Cocomys, Tanquammys and 16 Ischyromyiformes. A robust phylogenetic frame is produced, which allows clarifying the systematics of the Theridomorpha. Within the Ischyromyiformes, the Remyoidea (nov. supfam.) are set apart from the Theridomorpha. Protadelomys and Tardenomys represent the earliest offshoots of the Theridomyoidea, before the dichotomy between Pseudosciuridae and Theridomyidae. It supports the former subfamilies Pseudosciurinae and Sciuroidinae within the Pseudosciuridae; and for the Theridomyidae: the Issiodoromyinae, Oltinomyinae, Columbomyinae, Theridomyinae, with at least one new subfamily (Patriotheridomyinae). The topologies of the chronospecies (sensu Simpson), formerly considered as evolutionary lineages, appear in most cases as successive clades, in which the species are generally pectinately arranged and emerging in the stratigraphic order. The analysis of characters supporting the main nodes allow consolidating the diagnosic characters of the taxa and their evolutionary trends, as well as discussing the various cases of parallelism and convergence in the evolution of structures and dental patterns (e.g., uniserial incisor enamel for Issiodoromyinae and Patriotheridomyinae, or pseudo-multiserial for the most hypsodont Blainvillimys, Protechimys and Archaeomys; taeniodont dental patterns; lengthening of deciduous premolars for Patriotheridomyinae, Issiodoromyinae and Theridomyidae; selenodonty or lophodonty).
Evolutionary dynamics are analysed with respect to environmental changes. Finally, biochronological implications of the evolution of Theridomyoidea are discussed.
  

The principles and practices employed in establishment and recognition of South American land mammal ages are reviewed along with previous and present concepts of distinguishing time, rock, and faunal units. Previous chronological arrangements of South American Tertiary land mammal faunas are appraised on the basis of recent geological and paleontological data. Twelve South American Tertiary land mammal ages are here recognized [from oldest to youngest, Riochican (middle to late Paleocene); Casamayoran (early Eocene); Mustersan (middle Eocene); Divisaderan (late Eocene); Deseadan (early [to middle?] Oligocene); Colhuehuapian (late Oligocene); Santacrucian (early Miocene); Friasan (middle Miocene); Chasicoan (late Miocene); Huayquerian (latest Miocene); Montehermosan (early to middle Pliocene); and Chapadmalalan (late Pliocene)]. As all except the Friasian were originally defined on the basis of Argentine faunas, these are discussed first and at length, and each is reviewed with discussion of type locality, stratigraphy, type fauna, and faunal correlations. Non-Argentine faunas are then discussed country by country in alphabetical order.

     A review is given of radioisotope dates obtained on volcanic rocks (i.e., basalts, tuffs) associated with mammalbearing beds in Argentina. Based on these age determinations and on correlation of the late Tertiary land mammals involved in the interchange between North and South America, a chronology of South American land mammal ages correlated with North American land mammal ages and European marine stages is proposed.

It is concluded that South America was an island continent through most of the Tertiary Period (ca 65 to about 3 Ma). As a result, the land mammal fauna of South America developed in isolation and was dominated by autochthonous endemic groups. Toward the end of the Tertiary (i.e., middle Miocene) a unique faunal balance had been achieved by the descendants of the ancient inhabitants (notoungulates, litopterns, condylarths, astrapotheres, edentates, marsupials) and of later (late Eocene) waif immigrants (caviomorph rodents, platyrrhine primates). A prominent feature of this mammal fauna was the combination of carnivorous and omnivorous marsupials with native placental herbivorous ungulates, subungulates, and edentates.

Sometime during the late Miocene, a limited but important interchange of mammalian taxa between North and South America took place. Procyonids (raccoons and their allies), a group of North American origin, first appear in South America in strata of Huayquerian Age, while members of the extinct South American ground sloth families Megalonychidae and Mylodontidae first appear in North America in early Hemphillian time. These groups dispersed along island arcs before the appearance of the Panamanian land bridge in the Pliocene (ca 3.0 Ma). Cricetine rodents, a group of North American origin, are first known in South America in strata of Montehermosan Age. The known taxa are too advanced and diversified to be considered the first of this group to invade South America. lt is believed by some workers that these rodents arrived before the Montehermosan, possibly in the late Miocene or earlier, by waif dispersal from North America.

The isolation of South America ended with the appearance of the Panamanian land bridge, which provided a direct, dry land connection between the two Americas. Across this portal an extensive interchange of terrestrial faunas occurred, and the fossil record documents an intermingling of these long-separated land mammals faunas.

      The beginning of this interchange by land route in South America is marked by the appearance of mammals which evolved from North American emigrants in the Chapadmalal Formation of Argentina. These include a mustelid (Conepatus), a tayassuid (Argyrohyus), and four genera (Akodon, Dankomys, Graomys, Reithrodon) of cricetine rodents. The appearance of this contingent of northern animals favors the existence of the Panamanian land bridge by this time. Likewise, a large number of terrestrial vertebrates of South American origin appear in North America in beds of late Blancan Age date around 2.7 Ma. Among the mammals are Neochoerus, Erethizon, Glyptotherium, Glossotherium, Kraglievichia, and Dasypus. 

A new carcharhinid shark species, Physogaleus hemmooriensis sp. nov., is described from the Lower Hemmoorian (Behrendorfian, late Burdigalian, early Miocene) of Werder, Lower Saxony, Germany. P. hemmooriensis also occurs in the Edegem and Antwerpen Sands Members of the Berchem Formation, Belgium, and in the Miste Bed, Aalten Member of the Breda Formation, The Netherlands, which have an early to middle Miocene age. In the Western Atlantic region, the taxon is present in the early Miocene Calvert Formation of Delaware, U.S.A, which is largely contemporaneous with the Hemmoorian.