Occlusal relief changes with molar wear in Pan troglodytes troglodytes and Gorilla gorilla gorilla

Most research on primate tooth form-function relationships has focused on unworn teeth. This study presents a morphological comparison of variably worn lower second molars (M(2)s) of lowland gorillas (Gorilla gorilla gorilla; n=47) and common chimpanzees (Pan troglodytes troglodytes; n=54) using dental topographic analysis. High-resolution replicas of occlusal surfaces were prepared and scanned in 3D by laser scanning. The resulting elevation data were used to create a geographic information system (GIS) for each tooth. Occlusal relief, defined as the ratio of 3D surface area to 2D planometric area of the occlusal table, was calculated and compared between wear stages, taxa, and sexes. The results failed to show a difference in occlusal relief between males and females of a given taxon, but did evince differences between wear stages and between taxa. A lack of significant interaction between wear stage and taxon factors suggests that differences in occlusal relief between chimpanzees and gorillas are maintained throughout the wear sequence. These results add to a growing body of information on how molar teeth change with wear, and how differences between primate species are maintained at comparable points throughout the wear sequence. Such studies provide new insights into form-function relationships, which will allow us to infer certain aspects of diet in fossils with worn teeth.     

Dental topography and molar wear in Alouatta palliata from Costa Rica

Paleoprimatologists depend on relationships between form and function of teeth to reconstruct the diets of fossil species. Most of this work has been limited to studies of unworn teeth. A new approach, dental topographic analysis, allows the characterization and comparison of worn primate teeth. Variably worn museum specimens have been used to construct species-specific wear sequences so that measurements can be compared by wear stage among taxa with known differences in diet. This assumes that individuals in a species tend to wear their molar teeth in similar ways, a supposition that has yet to be tested. Here we evaluate this assumption with a longitudinal study of changes in tooth form over time in primates. Fourteen individual mantled howling monkeys (Alouatta palliata) were captured and then recaptured after 2, 4, and 7 years when possible at Hacienda La Pacifica in Costa Rica between 1989-1999. Dental impressions were taken each time, and molar casts were produced and analyzed using dental topographic analysis. Results showed consistent decreases in crown slope and occlusal relief. In contrast, crown angularity, a measure of surface jaggedness, remained fairly constant except with extreme wear. There were no evident differences between specimens collected in different microhabitats. These results suggest that different individual mantled howling monkeys wear their teeth down in similar ways, evidently following a species-specific wear sequence. Dental topographic analysis may therefore be used to compare morphology among similarly worn individuals from different species.     

Technical note: Estimating original crown height in worn mandibular canines using aspects of dentin morphology

We present a novel method to estimate original crown height (OCH) for worn human mandibular canines using a cubic regression equation based on ratios of worn crown height and exposed dentin. This method may help alleviate issues frequently presented by worn teeth in dental analyses, including those in bioarchaeology. Mandibular canines (n = 28) from modern day New Zealand and English populations were selected. Crown height and dentin thickness were measured on dental thin sections (n = 19) and the resulting (log10) ratios were fitted to a cubic regression curve allowing OCH in worn crowns to be predicted. Variation in the dentin apex position was recorded and effects of angled wear slopes investigated allowing adjusted values to be generated. Our method is trialed for use on intact and sectioned teeth (n = 17). A cubic regression curve best describes the relationship between (log10) ratios and crown height deciles (R² = 0.996, df₁ = 3, df₂ = 336, p < 0.001). No significant differences were detected between OCH estimates using our method and digitally recreated cusp outlines of the same crowns (t = 1.024, df = 16, p > 0.05), with a mean absolute error of 0.171 mm and an adjusted coefficient of determination of 0.923. Our approach offers a quantitative method to estimate the percentage of OCH remaining on worn mandibular canines, and by extension, the OCH. Our estimates are comparable to digitally recreated cusps but less subjective and not limited to crowns with minimal wear.     

Dental topography and diets of Australopithecus afarensis and early Homo

Diet is key to understanding the paleoecology of early hominins. We know little about the diets of these fossil taxa, however, in part because of a limited fossil record, and in part because of limitations in methods available to infer their feeding adaptations. This paper applies a new method, dental topographic analysis, to the inference of diet from fossil hominin teeth. This approach uses laser scanning to generate digital 3D models of teeth and geographic information systems software to measure surface attributes, such as slope and occlusal relief. Because it does not rely on specific landmarks that change with wear, dental topographic analysis allows measurement and comparison of variably worn teeth, greatly increasing sample sizes compared with techniques that require unworn teeth. This study involved comparison of occlusal slope and relief of the lower second molars of Australopithecus afarensis (n=15) and early Homo (n=8) with those of Gorilla gorilla gorilla (n=47) and Pan troglodytes troglodytes (n=54). Results indicate that while all groups show reduced slope and relief in progressively more worn specimens, there are consistent differences at given wear stages among the taxa. Early Homo shows steeper slopes and more relief than chimpanzees, whereas A. afarensis shows less slope and relief than any of the other groups. The differences between the two hominin taxa are on the same order as those between the extant apes, suggesting similar degrees of difference in diet. Because these chimpanzees and gorillas differ mostly in fallback foods where they are sympatric, results suggest that the early hominins may likewise have differed mostly in fallback foods, with A. afarensis emphasizing harder, more brittle foods, and early Homo relying on tougher, more elastic foods.     

Variation in dental wear and tooth loss among known‐aged,older ring‐tailed lemurs (Lemur catta): a comparison between wild and captive individuals

Tooth wear is generally an age‐related phenomenon, often assumed to occur at similar rates within populations of primates and other mammals, and has been suggested as a correlate of reduced offspring survival among wild lemurs. Few long‐term wild studies have combined detailed study of primate behavior and ecology with dental analyses. Here, we present data on dental wear and tooth loss in older (>10 years old) wild and captive ring‐tailed lemurs (Lemur catta). Among older ring‐tailed lemurs at the Beza Mahafaly Special Reserve (BMSR), Madagascar (n=6), the percentage of severe dental wear and tooth loss ranges from 6 to 50%. Among these six individuals, the oldest (19 years old) exhibits the second lowest frequency of tooth loss (14%). The majority of captive lemurs at the Indianapolis Zoo (n=7) are older than the oldest BMSR lemur, yet display significantly less overall tooth wear for 19 of 36 tooth positions, with only two individuals exhibiting antemortem tooth loss. Among the captive lemurs, only one lemur (a nearly 29 year old male) has lost more than one tooth. This individual is only missing anterior teeth, in contrast to lemurs at BMSR, where the majority of lost teeth are postcanine teeth associated with processing specific fallback foods. Postcanine teeth also show significantly more overall wear at BMSR than in the captive sample. At BMSR, degree of severe wear and tooth loss varies in same aged, older individuals, likely reflecting differences in microhabitat, and thus the availability and use of different foods. This pattern becomes apparent before “old age,” as seen in individuals as young as 7 years. Among the four “older” female lemurs at BMSR, severe wear and/or tooth loss do not predict offspring survival. Am. J. Primatol. 72:1026–1037, 2010. © 2010 Wiley‐Liss, Inc.     

Dental topography and diets of platyrrhine primates

Abstract

More than half a century ago, Percy Butler touted the importance of analyzing teeth to understand their function in an evolutionary context. There have been many advances in the study of dental functional morphology since that time. Here we review the various approaches to characterizing and comparing occlusal form that have been developed, especially dental topographic analysis. We also report on a new study of dental topography of platyrrhine primates (n = 341 individuals representing 16 species) with known differences in both dietary preferences and other food items eaten. Results indicate frugivores, gummivores, folivores, and seed eaters each have a unique combination of slope, relief, angularity, sharpness, and occlusal orientation patch size and count values. Likewise, among frugivores, those that supplement their diets with hard objects, insects, leaves, and seeds, also each have a distinctive suite of topographic features. We conclude that both primary and secondary diet choices select for occlusal form, and that functional morphology more reflects the types of foods and mechanical challenges they pose rather than the frequencies in which they are eaten.     

Comparison of dental measurement systems for taxonomic assignment of Neanderthal and modern human lower second deciduous molars

Traditional morphometric approaches for taxonomic assignment of Neanderthal and modern human dental remains are mainly characterized by caliper measurements of tooth crowns. Several studies have recently described differences in dental tissue proportions and enamel thickness between Neanderthal and modern human teeth. At least for the lower second deciduous molar (dm₂), a three-dimensional lateral relative enamel thickness index has been proposed for separating the two taxa. This index has the advantage over other measurements of being applicable to worn teeth because it ignores the occlusal aspect of the crown. Nevertheless, a comparative evaluation of traditional crown dimensions and lateral dental tissue proportion measurements for taxonomic assignment of Neanderthal and modern human dm₂s has not yet been performed.In this study, we compare various parameters gathered from the lateral aspects of the crown. These parameters include crown diameters, height of the lateral wall of the crown (lateral crown height = LCH), lateral enamel thickness, and dentine volume of the lateral wall, including the volume of the coronal pulp chamber (lateral dentine plus pulp volume = LDPV), in a 3D digital sample of Neanderthal and modern human dm₂s to evaluate their utility in separating the two taxa.The LDPV and the LCH allow us to discriminate between Neanderthals and modern humans with 88.5% and 92.3% accuracy, respectively. Though our results confirm that Neanderthal dm₂s have lower relative enamel thickness (RET) index compared with modern humans (p = 0.005), only 70% of the specimens were correctly classified on the basis of the RET index. We also emphasize that results of the lateral enamel thickness method depend on the magnitude of the interproximal wear. Accordingly, we suggest using the LCH or the LDPV to discriminate between Neanderthal and modern human dm₂s. These parameters are more independent of interproximal wear and loss of lateral enamel.     

Dental wear among cercopithecid monkeys of the Taï forest,Côte d'Ivoire

Studies of dental macrowear can be useful for understanding masticatory and ingestive behavior, life history, and for inferring dietary information from the skeletal material of extinct and extant primates. Such studies to date have tended to focus on one or two teeth, potentially missing information that can be garnered through examination of wear patterns across the tooth row. Our study measured macrowear in the postcanine teeth of three sympatric cercopithecid species from the Taï Forest, Côte d'Ivoire (Cercocebus atys, Procolobus badius, and Colobus polykomos), whose diets have been well‐described. Inter‐specific analyses suggest that different diets and ingestive behaviors are characterized by different patterns of wear across the molar row, with Cercocebus atys emphasizing tooth use near P₄‐M₁, P. badius emphasizing a large amount of tooth use near M₂‐M₃, and Colobus polykomos exhibiting wear more evenly across the postcanine teeth. Information regarding differential tooth use across the molar row may be more informative than macrowear analysis of isolated teeth for making inferences about primate feeding behavior. Am J Phys Anthropol 150:655–665, 2013. © 2013 Wiley Periodicals, Inc.     

Unravelling the Functional Biomechanics of Dental Features and Tooth Wear

Most of the morphological features recognized in hominin teeth, particularly the topography of the occlusal surface, are generally interpreted as an evolutionary functional adaptation for mechanical food processing. In this respect, we can also expect that the general architecture of a tooth reflects a response to withstand the high stresses produced during masticatory loadings. Here we use an engineering approach, finite element analysis (FEA), with an advanced loading concept derived from individual occlusal wear information to evaluate whether some dental traits usually found in hominin and extant great ape molars, such as the trigonid crest, the entoconid-hypoconulid crest and the protostylid have important biomechanical implications. For this purpose, FEA was applied to 3D digital models of three Gorilla gorilla lower second molars (M₂) differing in wear stages. Our results show that in unworn and slightly worn M₂s tensile stresses concentrate in the grooves of the occlusal surface. In such condition, the trigonid and the entoconid-hypoconulid crests act to reinforce the crown locally against stresses produced along the mesiodistal groove. Similarly, the protostylid is shaped like a buttress to suffer the high tensile stresses concentrated in the deep buccal groove. These dental traits are less functional in the worn M₂, because tensile stresses decrease physiologically in the crown with progressing wear due to the enlargement of antagonistic contact areas and changes in loading direction from oblique to nearly parallel direction to the dental axis. This suggests that the wear process might have a crucial influence in the evolution and structural adaptation of molars enabling to endure bite stresses and reduce tooth failure throughout the lifetime of an individual.     

Tooth Wear Difference Between the Yuanmou Hominoid and <Emphasis Type="Italic">Lufengpithecus</Emphasis>

The Late Miocene hominoids recovered from Lufeng (Lufengpithecus) and Yuanmou of Yunnan Province, China, are among the most numerous hominoid fossils in Eurasia. They have yielded critical evidence for the evolutionary history, biogeography and paleobiology of Miocene hominoids. We examined and compared the wear pattern and differences of 804 molars of the Yuanmou hominoid and Lufengpithecus. Our results indicate that both the upper and lower molars of the Yuanmou hominoids were more heavily worn than those of Lufengpithecus. The wear patterns of the individual molars between the Yuanmou hominoid and Lufengpithecus also are different. The heaviest wear of lower molars of the Yuanmou hominoid occur in M₂, followed by M₁ and M₃. In Lufengpithecus, M₁ and M₃ were more heavily worn than M₂. There are differences in wear between the upper and lower molars for the two hominoids. Among the various factors related to tooth wear, we suggest that the main reason for the tooth wear differences between the Yuanmou hominoid and Lufengpithecus may be that they had different diets. More soft dietary items like leaves and berries were probably consumed by Lufengpithecus, and the Yuanmou hominoid may mainly have feed on harder or frugivorous diets. This result complements findings from previous studies of tooth size proportion, and the development of lower molar shearing crests in the 2 samples. Enamel thickness, living environment, behavior patterns, and population structure also might account for dental wear differences between the Yuanmou hominoid and Lufengpithecus.     

Dental anomalies in the gray foxUrocyon cinereoargenteus and the red foxVulpes vulpes

We examined dental anomalies, including oligodonty, polydonty, connation, rotation, and misalignment in 510 gray foxes and 150 red foxes from southern Illinois (USA). Dental anomalies were significantly more common (x ² = 11.5, df = 1,p < 0.001) in gray foxes (n = 177; 34.7% of sample) than red foxes (n = 25; 16.6% of sample), and more common in male than female gray foxes (x ² = 3.88, df = 1,p < 0.05). Polydonty was very uncommon, as expected for species in which the normal dental complement is close to the primitive eutherian number. In both species, the most prevalent anomaly was loss of the last lower molar. Loss of the upper or lower first premolar was also common. Thus, oligodonty almost always involved the smaller anterior (P¹ and P₁) or posterior (M₃) teeth of the dental arcade. Conversely, the large carnassial teeth, with complex occlusal patterns and shearing surfaces, appeared to be highly conserved with only three anomalous individuals (0.4%) among all specimens.     

Enamel thickness and the helicoidal occlusal plane

In the present study 38 unworn maxillary molars (M¹ = 16, M²= 12, M³ = 10) of modern humans from a Slavic necropolis were sectioned through the mesial cusps in a plane perpendicular to the cervical margin of the crown. Five slightly worn M¹s and one slightly worn M³ were also used thus increasing the total sample to 44, but measurements made on the worn areas were coded as missing values. Seven measurements of enamel thickness as well as the heights of the protocone and the paracone dentine horns were recorded in order to analyze whether changes in these dimensions in anteroposterior direction can be related to the helicoidal occlusal plane. Uni- and multivariate analyses revealed that the distribution of enamel thickness within and between maxillary molars corresponds to a helicoidal occlusal wear pattern. Enamel thickness along the occlusal basin increases from anterior to posterior, which may lead to rapid development of a reverse curve of Monson in first molars when compared to posterior teeth. However, although these overall differences together with the serial, especially delayed eruption pattern of human molars, contribute to the marked expression of the helicoidal occlusal plane in Homo, differences in enamel patterning between molars indicate that a helicoidal plane is a structural feature of the orofacial skeleton. In contrast to first upper molars, second and third molars show absolutely and relatively thicker enamel under the Phase I wear facet of the paracone, i. e., the lingual slope of the paracone, than under the Phase II facet of the protocone, i. e., the buccal slope of that cusp. These proportional differences are most pronounced in M³, as evidenced by uni- and multivariate statistics. It thus appears that the pattern of enamel thickness distribution from M¹ to M³ follows a trend towards providing additional tooth material in areas that are under greater functional demands, that is, corresponding to a lingual slope of wear anteriorly and to a flat or even buccal one posteriorly. In addition, the heights of the dentine horns in anteroposterior direction change in a way that lends support to the hypothesis that the axial inclination of teeth could be one of the most important factors for the development of the helicoidal occlusal plane. Finally, the changes in morphology and enamel thickness distribution from first to third upper molars found in this study suggest that molars could be “specialized” in their function, i. e., from performing proportionally more shearing anteriorly to increased crushing and grinding activities posteriorly. © 1994 Wiley-Liss, Inc.     

Functional cues in the development of osseous tooth support in the pig, Sus scrofa

Alveolar bone supports teeth during chewing through a ligamentous interface with tooth roots. Although tooth loads are presumed to direct the development and adaptation of these tissues, strain distribution in the alveolar bone at different stages of tooth eruption and periodontal development is unknown. This study investigates the biomechanical effects of tooth loading on developing alveolar bone as a tooth erupts into occlusion. Mandibular segments from miniature pigs, Sus scrofa, containing M₁ either erupting or in functional occlusion, were loaded in compression. Simultaneous recordings were made from rosette strain gages affixed to the lingual alveolar bone and the M₂ crypt. Overall, specimens with erupting M₁s were more deformable than specimens with occluding M₁s (mean stiffness of 246 vs. 944 MPa, respectively, p=0.004). The major difference in alveolar strain between the two stages was in orientation. The vertically applied compressive loads were more directly reflected in the alveolar bone strains of erupting M₁s, than those of occluding M₁s, presumably because of the mediation of a more mature periodontal ligament (PDL) in the latter. The PDL interface between occluding teeth and alveolar bone is likely to stiffen the system, allowing transmission of occlusal loads. Alveolar strains may provide a stimulus for bone growth in the alveolar process and crest.     

Metrical dental analysis on golden monkey (Rhinopithecus roxellana)

Dental variation in the Chinese golden monkey (Rhinopithecus roxellana) is here evaluated by univariate, bivariate, and multivariate analyses. Allometric analyses indicate that canines and P3s are positively, but other dimensions negatively scaled to mandible and maxilla, and to body size. With the exception of the mesiodistal dimensions of I₁ and M₃, and the buccolingual dimension of P₄, mandibular dental variables show similar scaling relative to body size. Analysis of residuals shows that males have significantly larger canine, P³ and buccolingual dimensions of the postcanine teeth (M² and M³) than females. A significant difference in shape between the sexes is found in the buccolingual dimension of the upper teeth, but not in the mandible. Unlike the situation in some other species, female golden monkeys do not exhibit relatively larger postcanine teeth than males. In fact, the reverse is true, expecially for M²s and M³s. The fact that most of the dental variables show low negative allometry to body size might be related a cold environment that has led to the development of larger body size with reduced energy loss. When the raw data are examined by Discriminant Function Analysis the sexes are clearly distinguishable.     

Occlusal and morphogenetic field evolution in the dentition of European Nyctitheriidae (Euarchonta,Mammalia)

Abstract

The nyctithere genera Saturninia, Cryptotopos and members of the subfamily Amphidozotheriinae are well-represented by dental remains in the European later Eocene. Their molar occlusal relations are described in detail, demonstrating a diversity of adaptations to insectivory, including dilambdodonty, minor trends in zalambdodonty, development of a large talon shelf in upper first and second molars and a step in the trigon-trigonid shearing surface that provides an extra facet in buccal phase. Minor lingual phase wear is recognized for the first time in the family, but only in relatively worn teeth. Nyctithere molars differ from those of most insectivorous mammals today in having well-developed paraconules and metaconules on upper molars, which in most cases lack a marked bucco-lingual tilt, associated with a more transverse jaw action. Amphidozotherium, however, shows tilting and a basally shifted M¹ hypocone associated with M₁ talonid exodaenodonty and a minor trend in zalambdodonty. Nyctitheres primitively have three deeply notched lobate lower incisors and a large but procumbent premolariform lower canine. Amphidozotheriines have modified their I₃ into a premolariform tooth, by shifting the premolarization field mesially. Amphidozotherium has also shifted this field distally, reducing P_2–3 in size and their roots from two to one.     

Measurements of tooth wear among Australian Aborigines: I. Serial loss of the enamel crown

The dental casts made from Aboriginal children during the course of a longitudinal growth study in Central Australia provided material for analyzing tooth wear under known environmental conditions. The wear facets produced on the occlusal surfaces were clearly preserved on the dental stone casts and recorded the progress of enamel attrition from ages 6 to 18. These casts were photographed and traced by electronic planimetric methods that automatically recorded the location and size of wear facets on the first and second permanent molars. These areas of worn tooth surface were compared to the total tooth surface. The worn surface was regressed on age to calculate wear rates of each tooth. Discriminant analyses were also performed to determine the significance of dental attrition differences between the sexes at each age group. The total wear on each tooth was highly correlated with age as expected but females wore their teeth at a significantly higher rate than males. The mandibular molars wore more rapidly than maxillary teeth in both sexes. The discriminant analysis successfully grouped 91% of the cases according to age and sex. Pattern of wear, the location, and size of wear facets also differed between age groups and sex. The questions of why there is a difference between male and female wear or why there is greater wear on one arch or arch region have no ready answers. The differing rates and pattern of dental wear do suggest that arch shape and growth rates may be the answer though it has yet to be tested. However, the occlusal surface wear is useful for age estimation in a population and provides a record of shifting masticatory forces during growth.     

Extinct large colobines from the Plio-Pleistocene of Africa

Three genera and six species of extinct African Plio-Pleistocene Colobinae are discussed. One new genus, Rhinocolobus and three new species, R. turkanaensis, Cercopithecoides kimeui and Paracolobus mutiwa are named. These colobines show a diversity in postcranial and dental morphology not seen among extant species. Rhinocolobus was most similar to extant Colobus in postcranial morphology and had similar high-cusped shearing teeth. Cercopithecoides shows a number of postcranial skeletal features typical of terrestrial cercopithecid species and has lower cusped teeth. Paracolobus, while generally more similar to Rhinocolobus than to Cercopithecoides, is intermediate in some features of its postcranial morphology. The distribution of the various taxa among East and South African sites with different palaeoenvironments is generally consistent with the morphological interpretation. With the exception of Cercopithecoides kimeui, which persisted a little longer, these large colobines disappear from the fossil record about 1.8 million years ago. Their disappearance coincides with an interval of increasing aridity documented at Olduvai Gorge, the Omo Valley, and East Turkana.     

The Evolutionary Paradox of Tooth Wear: Simply Destruction or Inevitable Adaptation?

Over the last century, humans from industrialized societies have witnessed a radical increase in some dental diseases. A severe problem concerns the loss of dental materials (enamel and dentine) at the buccal cervical region of the tooth. This “modern-day” pathology, called non-carious cervical lesions (NCCLs), is ubiquitous and worldwide spread, but is very sporadic in modern humans from pre-industrialized societies. Scholars believe that several factors are involved, but the real dynamics behind this pathology are far from being understood. Here we use an engineering approach, finite element analysis (FEA), to suggest that the lack of dental wear, characteristic of industrialized societies, might be a major factor leading to NCCLs. Occlusal loads were applied to high resolution finite element models of lower second premolars (P₂) to demonstrate that slightly worn P₂s envisage high tensile stresses in the buccal cervical region, but when worn down artificially in the laboratory the pattern of stress distribution changes and the tensile stresses decrease, matching the results obtained in naturally worn P₂s. In the modern industrialized world, individuals at advanced ages show very moderate dental wear when compared to past societies, and teeth are exposed to high tensile stresses at the buccal cervical region for decades longer. This is the most likely mechanism explaining enamel loss in the cervical region, and may favor the activity of other disruptive processes such as biocorrosion. Because of the lack of dental abrasion, our masticatory apparatus faces new challenges that can only be understood in an evolutionary perspective.     

Dental topographic and three-dimensional geometric morphometric analysis of carnassialization in different clades of carnivorous mammals (Dasyuromorphia,Carnivora, Hyaenodonta)

The evolution of carnassial teeth in mammals, especially in the Carnivora, has been subject of many morphometric and some dental topographic studies. Here, we use a combination of dental topographic analysis (Dirichlet normal energy) and 3D geometric morphometrics of less and high carnassialized lower teeth of carnivoran, dasyuromorph and hyaenodont taxa. Carnassial crown curvature, as indicated by Dirichlet normal energy, is high in lesser carnassialized teeth and low in higher carnassialized teeth, where it is influenced by the reduction of crown features such as cusps and crests. PC1 of the geometric morphometric analysis is linked to enlargement of the carnassial blade, reduction of the talonid crushing basin and an increasingly asymmetric cervix line with an enlarged mesial flexure in more carnassialized teeth. Distribution of PC1 values further indicates that along the tooth row of dasyuromorphs (m2–m4) and hyaenodonts (m1–m3) the most distal carnassial is the most carnassialized (principal carnassial), and in most taxa with overall higher carnassialized teeth, carnassialization successively increases from the anterior to the posterior tooth position along the tooth row. PC2 indicates that a longitudinal elongated carnassial is present in caniforms and in unspecialized feliforms, which separates these taxa in morphospace from all dasyuromorphs, hyaenodonts and specialized feliforms. An ancestral state reconstruction shows that this longitudinal elongation may be a plesiomorphic ancestral state for the Carnivora, which is different from the Dasyuromorphia and the Hyaenodonta. This elongation, enabling the presence of a longitudinally aligned carnassial blade as well as a complete talonid basin, might have provided the Carnivora with an advantage in terms of adaptive versatility.