A new Oligocene Ctenodactylinae (Rodentia: Mammalia) from Ulantatal (nei Mongol): new insight on the phylogenetic origins of the modern Ctenodactylidae

The four living genera of Ctenodactylidae (Rodentia) are the only survivors of a flourishing Tertiary group. In this paper, we describe a new Oligocene species from Ulantatal (Chinese Mongolia) that highlights the origins of crown ctenodactylids. This species, Helanshania deserta gen. et sp. nov. , is lophodont and displays semi‐hypsodont teeth, a dental pattern that is somewhat transitional between that of primitive Oligocene ctenodactylids and the later hypsodont genera. We perform here a cladistic assessment of the dental evidence for species produced by the successive radiations of the group. In order to get new data, to decipher homologies for the dental pattern of modern ctenodactylids, and to specify their dental replacement, we describe additional dental material of Ctenodactylus, Massoutiera, and Felovia. The phylogenetic analysis (using PAUP) considered 45 characters (mainly dental) and 31 species. The performed heuristic searches yielded 596 equally most parsimonious trees. Protataromys and Karakoromys are stem ctenodactylids and appear as the earliest offshoots of the Ctenodactylidae clade, which represents a well‐supported family rank. Within this family, the Tataromyinae appear paraphyletic, whereas the Ctenodactylinae sensu lato are a clade including the new taxon Helanshania. As such, a revision of the Tataromyinae is envisaged and a new subfamily is erected (Yindirtemyinae). Amongst the Ctenodactylinae, a tribe Ctenodactylini encompassing the crown ctenodactylines is proposed. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 531–550.     

Morphological integration in the hominin dentition: evolutionary, developmental, and functional factors

As the most common and best preserved remains in the fossil record, teeth are central to our understanding of evolution. However, many evolutionary analyses based on dental traits overlook the constraints that limit dental evolution. These constraints are diverse, ranging from developmental interactions between the individual elements of a homologous series (the whole dentition) to functional constraints related to occlusion. This study evaluates morphological integration in the hominin dentition and its effect on dental evolution in an extensive sample of Plio- and Pleistocene hominin teeth using geometric morphometrics and phylogenetic comparative methods. Results reveal that premolars and molars display significant levels of covariation; that integration is stronger in the mandibular dentition than in the maxillary dentition; and that antagonist teeth, especially first molars, are strongly integrated. Results also show an association of morphological integration and evolution. Stasis is observed in elements with strong functional and/or developmental interactions, namely in first molars. Alternatively, directional evolution (and weaker integration) occurs in the elements with marginal roles in occlusion and mastication, probably in response to other direct or indirect selective pressures. This study points to the need to reevaluate hypotheses about hominin evolution based on dental characters, given the complex scenario in which teeth evolve.     

Geographic Variation in Nonmetric Dental Traits of the Deciduous Molars of Pan and Gorilla

Physical anthropologists often use nonmetric dental traits to trace the movement of human populations, but similar analysis of the teeth of nonhuman primates or the deciduous teeth is rare. Because nonmetric dental characteristics are manifestations of genetic differences among groups, they vary among geographically distant members of the same species and subspecies. We use 28 nonmetric dental traits in the deciduous molars to compare genetically and geographically distinct groups of extant African apes (Gorilla and Pan). Previous researchers have studied these traits in the adult or juvenile teeth of great apes and humans, and we score our observations according to established standards for hominins. We observe marked differences in trait frequencies between Gorilla and Pan, Pan troglodytes and P. paniscus, and two P. troglodytes subspecies but we find no significant differences between geographically isolated groups within the subspecies. Trait frequencies differ from those found in previous studies that contained fewer individuals. We find that the deciduous molars show similar variation to adult premolars and molars within Pan and Gorilla. This suggests that the deciduous dentition of these and other apes may contain diagnostic traits that are not currently in use.     

Mastication and enamel microstructure in <Emphasis Type="Italic">Cambaytherium</Emphasis>, a perissodactyl-like ungulate from the early Eocene of India

The dentition of Cambaytherium was investigated in terms of dental wear, tooth replacement and enamel microstructure. The postcanine tooth row shows a significant wear gradient, with flattened premolars and anterior molars at a time when the last molars are only little worn. This wear gradient, which is more intensive in Cambaytherium thewissi than in Cambaytherium gracilis, and the resulting flattened occlusal surfaces, may indicate a preference for a durophagous diet. The tooth replacement (known only in C. thewissi) shows an early eruption of the permanent premolars. They are in function before the third molars are fully erupted. During the dominant phase I of the chewing cycle the jaw movement is very steep, almost orthal, with a slight mesiolingual direction and changes into a horizontal movement during phase II. The enamel microstructure shows Hunter-Schreger-bands (HSB) in the inner zone of the enamel. In some teeth the transverse orientation of the HSB is modified into a zig-zag pattern, possibly an additional indicator of a durophagous diet.     

Evolutionary and developmental dynamics of the dentition in Muroidea and Dipodoidea (Rodentia, Mammalia)

SUMMARY When it comes to mouse evo‐devo, the fourth premolar–first molar (P4–M1) dental complex becomes a source of longstanding controversies among paleontologists and biologists. Muroidea possess only molar teeth but with additional mesial cusps on their M1. Developmental studies tend to demonstrate that the formation of such mesial cusps could result from the integration of a P4 germ into M1 during odontogenesis. Conversely, most Dipodoidea conserve their fourth upper premolars and those that lost these teeth can also bear additional mesial cusps on their first upper molars. The aim of this study is to assess this developmental model in both Muroidea and Dipodoidea by documenting the morphological evolution of the P4–M1 complex across 50 Ma. Fourteen extinct and extant species, including abnormal and mutant specimens were investigated. We found that, even if their dental evolutionary pathways strongly differ, Dipodoidea and Muroidea retain common developmental characteristics because some of them can present similar dental morphological trends. It also appears that the acquisition of a mesial cusp on M1 is independent from the loss of P4 in both superfamilies. Actually, the progressive decrease of the inhibitory effect of P4, consequent to its regression, could allow the M1 to lengthen and mesial cusps to grow in Muroidea. Apart from these developmental explanations, patternings of the mesial part of first molars are also deeply constrained by morpho‐functional requirements. As there is no obvious evidence of such mechanisms in Dipodoidea given their more variable dental morphologies, further developmental investigations are needed.     

The anatomy and systematic position of the early Miocene proconsulid from Meswa Bridge, Kenya

A small collection of fossil catarrhines was recovered from the early Miocene locality of Meswa Bridge in western Kenya between 1978 and 1980. The associated fauna from Meswa Bridge indicates an age older than 20 Ma. Much of the material has been briefly described previously, and its taxonomic status considered. The material can be assigned to a minimum of four individuals, all of which are infants or juveniles. Although the specimens were shown to belong to a distinct species of Proconsul, the taxon was not named, primarily because many of the specimens belonged to immature individuals. Nevertheless, the combined morphological features of the deciduous and permanent teeth allow the diagnosis of a new species of Proconsul, which is formally named here as P. meswae. It is a large-sized species, similar in dental size to P. nyanzae. The main features distinguishing it from all other previously named species of Proconsul are: incisors and deciduous incisors relatively low crowned; upper deciduous canines relatively higher crowned and more robust; molars and deciduous premolars relatively broader and higher crowned, with a more pronounced degree of buccolingual flare and better developed cingula; size differential between molars not as marked; dP₄ with a longer mesial fovea and smaller hypoconulid and distal fovea; P₄ relatively broader, with a better developed buccal cingulum; lower molars less rectangular with a longer mesial fovea, smaller distal fovea, more restricted talonid basin, and a tendency for a smaller hypoconulid; dP⁴ and upper molars with strongly buccolingually splayed roots; mandibular corpus in infants relatively deeper and more slender; maxilla with a well developed canine jugum and fossa. The broader and more flared molars with better developed cingula indicate that the Meswa Bridge species is more primitive than other species of Proconsul. The inference that it is a stem member of the Proconsul clade is consistent with the estimated age of the material.     

Report of inaccuracies in the study of the dentition of mammals

After discussing the existing view points and on the base of author's own studies, the designation of premolars and molars is proposed in the deciduous and permanent dentition of placental mammals, corresponding to the function, form, and structure of the teeth. With the admittance of the molars in the deciduous dentition of mammals, the discrepancy between milk dentition of mammals and humans is eliminated. In this way, the molars of deciduous dentition appeared to be the precursors of premolars and molars of permanent dentition. The separate groups of teeth are differentiated depending on the way of life and kind of consumed food, according to position and function of the teeth in the dentition. Lacerating teeth, premolars are formed in carnivora mammals for tearing off the food, and in case of intensive masticatory function of herbivora and omnivora, very likely, from the same germs of premolars, the molars are formed.     

Evolutionary Implications of Dental Eruption in Dasypus (Xenarthra)

Late eruption of the permanent dentition was recently proposed as a shared anatomical feature of endemic African mammals (Afrotheria), with anecdotal reports indicating that it is also present in dasypodids (armadillos). In order to clarify this question, and address the possiblity that late eruption is shared by afrotherians and dasypodids, we quantified the eruption of permanent teeth in Dasypus, focusing on growth series of D. hybridus and D. novemcinctus. This genus is the only known xenarthran that retains two functional generations of teeth. Its adult dentition typically consists of eight upper and eight lower ever-growing (or euhypsodont) molariforms, with no premaxillary teeth. All but the posterior-most tooth are replaced, consistent with the identification of a single molar locus in each series. Comparison of dental replacement and skull metrics reveals that most specimens reach adult size with none or few erupted permanent teeth. This pattern of growth occurring prior to the full eruption of the dentition is similar to that observed in most afrotherians. The condition observed in Dasypus and many afrotherians differs from that of most other mammals, in which the permanent dentition erupts during (not after) growth, and is complete at or near the attainment of sexual maturity and adult body size. The suture closure sequence of basicranial and postcranial epiphyses does not correlate well with dental eruption. The basal phylogenetic position of the taxon within dasypodids suggests that diphyodonty and late dental replacement represent the condition of early xenarthrans. Additionally, the inferred reduction in the number of molars to a single locus and the multiplication of premolars represent rare features for any living mammal, but may represent apomorphic characters for Dasypus.     

Dentition development and budding morphogenesis

The development of functional teeth in the mouse has been widely used as a model to study general mechanisms of organogenesis. Compared with other mammals, in which three incisors, one canine, four premolars, and three molars may occur even in each dental quadrant, the mouse functional dentition is strongly reduced. It comprises only one incisor separated from three molars by a toothless gap diastema at the location of the missing teeth. However, mouse embryos also develop transient vestigial dental primordia between the incisor and molar germs in both the upper and lower jaws. These rudimental structures regress, and epithelial apoptosis is involved in this process. The existence of the vestigial dental structures allowed a better assessment of the periodicity in the mouse dentition, which extends opportunities for the interpretation of molecular data on tooth development. We compared the dentition development with tentative models of budding morphogenesis in other epithelial appendages lungs and feathers. We suggested how developmental control by signaling molecules, including bone morphogenetic protein (Bmp), sonic hedgehog (Shh), and fibroblast growth factor (Fgf), can be similarly involved during budding morphogenesis of dentition and other epithelial appendages. We propose that epithelial apoptosis plays an important role in achieving specific features of dentition, whose development involves both budding and its more complex variant branching. The failure of segregation of the originating buds supports the participation of the concrescence of several tooth primordia in the evolutionary differentiation of mammalian teeth.     

Metric and morphological variability in the dentition of Colobine monkeys

A morphological study was performed using 97 Colobus polykomos and 41 Nasalis larvatus. The premolars and molars of these two taxa of leaf-eating monkeys were similar in that they possessed deeply grooved and highly pointed cusps. Of 7 dental traits investigated, 4 were significantly different between the 2 forms. Indeed, with 2 of these, the position of the lingual cusp on PM₄ and the number of cusps on PM³ displayed highly contrasting structures.Metric analysis was conducted using 47 C. polykomos and 37 N. larvatus specimens. Mesiodistal and buccolingual measurements were taken on the upper canine and all postcanine teeth. Sexual dimorphism was detected in most maxillary dental measurements and all mandibular estimates of both species. In all cases, males exceeded females in tooth diameter. Testing between species by sex revealed that C. polykomos was significantly larger in several canine and premolar dimensions while N. larvatus had significantly larger molar dimensions. An additional 73 measurements were selected to represent occluding dental structures and a morphological integration analysis was conducted. Results of this analysis indicated that C. polykomos possessed a C¹-PM₃ complex which was more highly intercorrelated whereas the broad lophed N. larvatus molars constituted a more highly correlated functional unit.It was suggested that the above differences between these two groups of leaf-eating monkeys probably resulted from inbreeding, isolation and drift.     

Variation in enamel thickness within the genus Homo

Recent humans and their fossil relatives are classified as having thick molar enamel, one of very few dental traits that distinguish hominins from living African apes. However, little is known about enamel thickness in the earliest members of the genus Homo, and recent studies of later Homo report considerable intra- and inter-specific variation. In order to assess taxonomic, geographic, and temporal trends in enamel thickness, we applied micro-computed tomographic imaging to 150 fossil Homo teeth spanning two million years. Early Homo postcanine teeth from Africa and Asia show highly variable average and relative enamel thickness (AET and RET) values. Three molars from South Africa exceed Homo AET and RET ranges, resembling the hyper thick Paranthropus condition. Most later Homo groups (archaic European and north African Homo, and fossil and recent Homo sapiens) possess absolutely and relatively thick enamel across the entire dentition. In contrast, Neanderthals show relatively thin enamel in their incisors, canines, premolars, and molars, although incisor AET values are similar to H. sapiens. Comparisons of recent and fossil H. sapiens reveal that dental size reduction has led to a disproportionate decrease in coronal dentine compared with enamel (although both are reduced), leading to relatively thicker enamel in recent humans. General characterizations of hominins as having ‘thick enamel’ thus oversimplify a surprisingly variable craniodental trait with limited taxonomic utility within a genus. Moreover, estimates of dental attrition rates employed in paleodemographic reconstruction may be biased when this variation is not considered. Additional research is necessary to reconstruct hominin dietary ecology since thick enamel is not a prerequisite for hard-object feeding, and it is present in most later Homo species despite advances in technology and food processing.     

Unique pattern of dietary adaptation in the dentition of Carnivora: its advantage and developmental origin

Carnivora is a successful taxon in terms of dietary diversity. We investigated the dietary adaptations of carnivoran dentition and the developmental background of their dental diversity, which may have contributed to the success of the lineage. A developmental model was tested and extended to explain the unique variability and exceptional phenotypes observed in carnivoran dentition. Carnivorous mammalian orders exhibited two distinct patterns of dietary adaptation in molars and only Carnivora evolved novel variability, exhibiting a high correlation between relative molar size and the shape of the first molar. Studies of Bmp7-hetero-deficient mice, which may exhibit lower Bmp7 expression, suggested that Bmp7 has pleiotropic effects on these two dental traits. Its effects are consistent with the pattern of dietary adaptation observed in Carnivora, but not that observed in other carnivorous mammals. A molecular evolutionary analysis revealed that Bmp7 sequence evolved by natural selection during ursid evolution, suggesting that it plays an evolutionary role in the variation of carnivoran dentition. Using mouse experiments and a molecular evolutionary analysis, we extrapolated the causal mechanism of the hitherto enigmatic ursid dentition (larger M₂ than M₁ and M₃). Our results demonstrate how carnivorans acquired novel dental variability that benefits their dietary divergence.     

Perinatal dental development in the chimpanzee (Pan troglodytes)

The synthesis of both ontogenetic and phylogenetic data should provide the ideal explanation of morphologic variation, but for the primate dentition, this has not yet occurred. Information concerning growth and development of primate teeth is lacking, in part because of the paucity of specimens. We have therefore examined the deciduous second molars (dm2) and tooth buds of the permanent first molar (M1) of 12 chimpanzees (Pan troglodytes), aged 6.5 months of gestation to 4 postnatal months. The ordering of cusp calcification was identical to that of other primates. By regression analysis, correlations of mesial and distal widths with buccal, lingual, and mesiodistal lengths were low, most probably because of decreased rates of change (slopes) and the relatively small sample size. Correlations were, however, greater for mandibular than for maxillary dentition and higher for age than for body weight; for both the dm2 and M1, distal moieties increased faster and were more highly correlated with other dental variables and age than were mesial ones. Comparison with data from humans revealed both differences and similarities in the absolute size and growth rate of dental moieties during the perinatal period. As the reasons for ontogenetic variation become understood for individuals, species, and higher taxa, the phylogenetic implications of differential growth should become clearer as well.     

Ceboid models for the evolution of hominoid dentition

Platyrrhine and catarrhine Primates have independently evolved such features of the teeth and jaws as U-shaped dental arches, reduced cingula on molars and premolars, and modifications for small tough-object feeding. The presence of specialization for tough-object feeding in arboreal cebids such as Cebus apella and Callicebus indicates that hominids may have acquired such a specialization in an arboreal habitat before becoming terrestrial.     

A New Early Miocene Chinchilloid Hystricognath Rodent; an Approach to the Understanding of the Early Chinchillid Dental Evolution

Chinchilloidea is an emblematical group of caviomorph rodents characterized by euhypsodont, laminated cheek teeth. Recent molecular analyses proposed that the extant Dinomys (and implicitly its fossil allies) is also part of this group. Their relationships with fossil caviomorphs with less derived dental features are still obscured by the deficiency of the fossil record documenting its early dental evolution. The new genus and species Garridomys curunuquem, from the early Miocene deposits of the Cerro Bandera Formation, northern Patagonia, is here described. It is represented by numerous mandible and maxillary remains with dentition. This species has protohypsodont cheek teeth with three transverse crests in all ontogenetic stages arranged in a transitory S-shaped pattern, resembling putative early dinomyids. Garridomys curunuquem is here interpreted as the sister group of the clade including the living and fossil chinchillids; both chinchillas and viscaccias would have diverged from a Garridomys-like ancestor and acquired hypsodonty independently. Garridomys and other chinchilloids would have diverged from the lineage leading to chinchillids in pre-Oligocene times, suggesting a very early, still poorly documented chinchilloid radiation.     

Dental evolution in Xenodontomys and first notice on secondary acquisition of radial enamel in rodents (Rodentia, Caviomorpha, Octodontidae)

Rodents of the subfamily Ctenomyinae differentiated during the Late Miocene in relation to the development of open biomes in southern South America. This subfamily displays a peculiar and derived dental morphology characterized by euhypsodont molars with simplified occlusal figure. We analyze both adaptive and evolutionary significance of the gross molar morphology and the enamel microstructure of ctenomyines. In accordance with the basal position of the Xenodontomys lineage, dental changes experimented by this lineage illustrate a probable evolutionary pattern of acquisition of molar design for the subfamily. We propose that morphological trends in the lower molars of the Xenodontomys lineage would include changes arisen as a by-product from hypsodonty, and other adaptive ones. These latter comprise the acquisition of a crescent-shaped occlusal morphology, and the secondary acquisition of an external layer of radial enamel in the leading edge, which would favor the development of cutting edges. Such a secondary acquisition of radial enamel had not been found so far in other rodents. The evolutionary pattern of the dental changes in Xenodontomys reinforces the idea that anagenesis is frequent in the adaptive evolution of rodents.     

Deciduous dentition and eruption pattern in late Miocene Pachyrukhinae (Hegetotheriidae,Notoungulata) from La Pampa Province,Argentina

The study of juvenile remains of Paedotherium Burmeister from Cerro Azul Formation (La Pampa Province, Argentina; late Miocene) is presented. Upper and lower deciduous dentition (or permanent molars supposed to be associated with non-preserved deciduous teeth) are recognised. Several ontogenetic stages are distinguished among juveniles, according to the degree of wear and the replaced deciduous teeth. Besides, some morphological and metrical differences are observed along the crown height. Deciduous cheek teeth are high-crowned and placed covering the apex of the corresponding permanent tooth. The height of the crown and the degree of wear allow establishing the pattern of dental replacement of deciduous and permanent premolars in a posterior–anterior direction (DP/dp4–2 and P/p4–2), as well as the eruption of M/m3 before DP/dp4 is replaced. Some of the studied remains are recognised as young individuals of Tremacyllus Ameghino, but with complete permanent dentition, which leads to propose a different timing in the dental replacement with respect to Paedotherium; they also allow the establishment of an opposite premolar eruption pattern, from P/p2 to P/p4. This knowledge of the deciduous dentition of Paedotherium suggests the need of revising the morphological and metrical characters previously used for defining species within this taxon.     

A new Late Jurassic species of the rare synechodontiform shark, <Emphasis Type="Italic">Welcommia</Emphasis> (Chondrichthyes,Neoselachii)

A new Jurassic species of the very rare and incompletely known synechodontiform shark, Welcommia, is described. The new species, Welcommia cappettai, is represented only by a single tooth, precluding reconstruction of its dentition in detail. Nevertheless, this specimen provides sufficient information and characteristics to establish its taxonomic status. Welcommia cappettai n. sp. occurs in the middle Oxfordian (Upper Jurassic) of south-western Germany. This is the first unambiguous record and named species of Welcommia from the Late Jurassic, substantially reducing the rather large gap in the fossil record of this synechodontiform taxon. So far, two Welcommia species from the Lower Jurassic of Belgium and the Lower Cretaceous of southern France have been described. An additional, still unnamed species seemingly occurs in the Oxfordian of southern France. The new species has plesiomorphic and apomorphic characteristics and, probably, an intermediate dental pattern that tentatively enables reconstruction of evolutionary trends in the dentition of this shark from small and compact teeth with broad, almost triangular cusps, to mesio-distally lengthened teeth with elongated mesial heels resulting in an extremely extended mesial cutting edge in addition to more delicate cusp and cusplets in advanced forms. These differences might be related to improved feeding mechanisms. It is hypothesized that Welcommia was predominantly a component of the Mediterranean faunal province. The disappearance of Welcommia in the Early Cretaceous remains ambiguous and might be related to competition by other sharks, for example hexanchiforms, or might represent a collecting bias and/or taxonomic misidentification of isolated teeth.     

Ontogeny and homology of the dentition in dasyurid marsupials: Development inSminthopsis virginiae

Histological analysis of an ontogenetic series of the dasyurid marsupial,Sminthopsis virginiae, from birt to 60 days old, was undertaken to assess the developmental homologies of the deciduous and successional teeth. This period covers the time from the initiation of all teeth as epithelial buds up until the time of early eruption of some teeth. In addition, two older specimens, aged 81 and 97 days, were examined to provide additional information on the state of differentiation of the unerupted third premolar. In the postcanine dentition, only a single tooth position, dP3, was characterized by the later development of a replacing successional tooth (P3), following developmental pathways identical to those in eutherian mammals. In contrast, the anterior dentition is characterized by the formation of rudimentary, nonerupting deciduous incisors and canines, and by the accelerated development of normal, erupting successional incisors and canines in both jaws. Comparison of relative developmental stages for each tooth position throughout its preeruptive ontogeny suggests thatheterochrony (both developmental acceleration and retardation) has played an important role in the evolutionary history of the dasyurid dentition. Differing aspects of this phenomenon are identified and discussed for the anterior dentition, the anterior two premolars, P3, and the lower molars. Further evidence is presented to corroborate the identification of the anterior two premolars in the adult as dP1 and dP2, based on the relative retardation of their initiation and their lack of successor tooth germs. This developmental heterochrony has probably occurred in all three-premolared marsupials.     

Milk molars or extra premolars in Mesotheriinae (Mesotheriidae,Notoungulata): New insights into an old controversy

A recently recovered specimen of Mesotheriinae (Mesotheriidae, Notoungulata) from the late Miocene-early Pliocene of La Rioja Province (Argentina), CRILAR Pv 433, corresponds to an individual with three upper premolars, which appears to be an “anomaly” among mesotheriines. The detailed study of this specimen, however, brings up an old controversy on the interpretation of different mesotheriine specimens with three upper or two lower premolars. After being described as different taxa, these were later considered to be juvenile representatives of other known species. The three upper or two lower teeth were interpreted as the milk molars DP2–4 and dp3–4, respectively, which would be replaced in adult life by two upper (P3–4) and one lower (p4) permanent premolars. The new material leads us to set up a different interpretation. In our opinion, all these specimens actually preserve the permanent dentition, corresponding to different ontogenetic stages of more or less young individuals. This consideration implies the necessity of a deep systematic revision of the whole subfamily, keeping in mind this new point of view and the ontogenetic variation within a species. Therefore, the presence of P2/p3 is not enough to define a different taxon at this moment. Pending this taxonomic revision, the dental morphology of CRILAR Pv 433 resembles both Typotheriopsis (e.g., upper premolars with one labial sulcus) and Pseudotypotherium (e.g., P4 with lingual groove, wide median lobe of M3) as these two late Miocene genera are currently characterized. Furthermore, P2/p3 could be expelled soon in the lifetime of individuals, and even the presence of P2/p3 could be a variable character within the same taxon; if so, this might reflect an evolutionary trend to the loss of a dental element within mesotheriines, but the revision of a large sample is necessary to support or reject these hypotheses.