Evolution of postcanine complexity in Gomphodontia (Therapsida: Cynodontia).

Gomphodonts form a Triassic radiation of small to medium-bodied (<0.5-2.5 m in length) quadrupedal cynodonts characterized by labiolingually expanded gomphodont postcanines. They were the dominant cynodont group in Middle and Late Triassic ecosystems from the Southern Hemisphere and the first predominantly herbivorous cynodonts to evolve. Gomphodonts were also the first therapsids to develop hypsodonty and a dentition with complex occlusal patterns, and their highly diagnostic upper and lower postcanines show many different morphologies. Here, we explored dental complexity in gomphodont cynodonts through time using geographic information system analysis and orientation patch count applied on 3D crown surfaces of upper and lower gomphodont postcanines belonging to 32 gomphodont taxa. This study reveals that the peak in postcanine complexity was reached early in the evolution of gomphodonts with the emergence in the Early Triassic of omnivorous or insectivorous forms with postcanines made of well-separated cusps and cingular cuspules. Traversodontids evolved simpler postcanines via coalescence of cusps into crests and the development of large occlusal basins, and the Middle Triassic radiation of traversodontids led to a sharp decrease in mean postcanine complexity. Simplification of the postcanines in traversodontids is interpreted as being related to a gradual increase in the consumption of plant material. Interestingly, the trend of insectivory/omnivory high postcanine complexity and herbivory low dental complexity in gomphodonts is opposite to the trend of dental complexity reported in some extant mammals, with omnivorous having low dental complexity and herbivorous higher. Postcanine complexity remained relatively stable throughout the evolution of traversodontids and only slightly diminished in the Late Triassic due to the presence of minute forms with particularly simple postcanines in the Rhaetian. The major phylogenetic diversity and taxonomic richness of Gomphodontia are represented in two periods of time: at the end of the Anisian, an age in which the postcanine complexity is simplifying, and at the early Carnian when the postcanine complexity in traversodontids, the only Gomphodontia represented, is stable.

Patterns of ontogenetic evolution across extant marsupials reflect different allometric pathways to ecomorphological diversity.

The relatively high level of morphological diversity in Australasian marsupials compared to that observed among American marsupials remains poorly understood. We undertake a comprehensive macroevolutionary analysis of ontogenetic allometry of American and Australasian marsupials to examine whether the contrasting levels of morphological diversity in these groups are reflected in their patterns of allometric evolution. We collate ontogenetic series for 62 species and 18 families of marsupials (n = 2091 specimens), spanning across extant marsupial diversity. Our results demonstrate significant lability of ontogenetic allometric trajectories among American and Australasian marsupials, yet a phylogenetically structured pattern of allometric evolution is preserved. Here we show that species diverging more than 65 million years ago converge in their patterns of ontogenetic allometry under animalivorous and herbivorous diets, and that Australasian marsupials do not show significantly greater variation in patterns of ontogenetic allometry than their American counterparts, despite displaying greater magnitudes of extant ecomorphological diversity.

Craniodental anatomy in Permian-Jurassic Cynodontia and Mammaliaformes (Synapsida, Therapsida) as a gateway to defining mammalian soft tissue and behavioural traits.

Mammals are diagnosed by more than 30 osteological characters (e.g. squamosal-dentary jaw joint, three inner ear ossicles, etc.) that are readily preserved in the fossil record. However, it is the suite of physiological, soft tissue and behavioural characters (e.g. endothermy, hair, lactation, isocortex and parental care), the evolutionary origins of which have eluded scholars for decades, that most prominently distinguishes living mammals from other amniotes. Here, we review recent works that illustrate how evolutionary changes concentrated in the cranial and dental morphology of mammalian ancestors, the Permian-Jurassic Cynodontia and Mammaliaformes, can potentially be used to document the origin of some of the most crucial defining features of mammals. We discuss how these soft tissue and behavioural traits are highly integrated, and how their evolution is intermingled with that of craniodental traits, thus enabling the tracing of their previously out-of-reach phylogenetic history. Most of these osteological and dental proxies, such as the maxillary canal, bony labyrinth and dental replacement only recently became more easily accessible-thanks, in large part, to the widespread use of X-ray microtomography scanning in palaeontology-because they are linked to internal cranial characters. This article is part of the theme issue 'The mammalian skull: development, structure and function'.

Ontogenetic growth in the crania of Exaeretodon argentinus (Synapsida: Cynodontia) captures a dietary shift.

Background: An ontogenetic niche shift in vertebrates is a common occurrence where ecology shifts with morphological changes throughout growth. How ecology shifts over a vertebrate's lifetime is often reconstructed in extant species-by combining observational and skeletal data from growth series of the same species-because interactions between organisms and their environment can be observed directly. However, reconstructing shifts using extinct vertebrates is difficult and requires well-sampled growth series, specimens with relatively complete preservation, and easily observable skeletal traits associated with ecologies suspected to change throughout growth, such as diet. Methods: To reconstruct ecological changes throughout the growth of a stem-mammal, we describe changes associated with dietary ecology in a growth series of crania of the large-bodied (∼2 m in length) and herbivorous form, Exaeretodon argentinus (Cynodontia: Traversodontidae) from the Late Triassic Ischigualasto Formation, San Juan, Argentina. Nearly all specimens were deformed by taphonomic processes, so we reconstructed allometric slope using a generalized linear mixed effects model with distortion as a random effect. Results: Under a mixed effects model, we find that throughout growth, E. argentinus reduced the relative length of the palate, postcanine series, orbits, and basicranium, and expanded the relative length of the temporal region and the height of the zygomatic arch. The allometric relationship between the zygomatic arch and temporal region with the total length of the skull approximate the rate of growth for feeding musculature. Based on a higher allometric slope, the zygoma height is growing relatively faster than the length of the temporal region. The higher rate of change in the zygoma may suggest that smaller individuals had a crushing-dominated feeding style that transitioned into a chewing-dominated feeding style in larger individuals, suggesting a dietary shift from possible faunivory to a more plant-dominated diet. Dietary differentiation throughout development is further supported by an increase in sutural complexity and a shift in the orientation of microwear anisotropy between small and large individuals of E. argentinus. A developmental transition in the feeding ecology of E. argentinus is reflective of the reconstructed dietary transition across Gomphodontia, wherein the earliest-diverging species are inferred as omnivorous and the well-nested traversodontids are inferred as herbivorous, potentially suggesting that faunivory in immature individuals of the herbivorous Traversodontidae may be plesiomorphic for the clade.

The emblematic South African therocephalian Euchambersia in China: a new link in the dispersal of late Permian vertebrates across Pangea.

Therapsids were widely distributed in Pangea in the late Permian. South Africa in Gondwana and Russia in Laurasia are the principal areas recording tetrapods (including therapsids) of this age. More recent field explorations have increased the importance of Chinese late Permian fossil assemblages. This is clearly reflected in the discovery of several new therocephalians from the Naobaogou Formation in Nei Mongol. Here, we report a therocephalian from that unit identified as a new species of the emblematic South African taxon Euchambersia. The new species, Euchambersia liuyudongi, is represented by a well-preserved skull and mandible showing a well-developed maxillary fossa and the absence of postcanine teeth. This is the third akidnognathid therocephalian recovered from the Naobaougou Formation, but oddly, the two basal Chinese akidnognathids previously known were recovered from a younger unit of the formation than the derived E. liuyudongi. This is the first time that the same therocephalian genus has been recorded in northern and southern continents, making the record of the Naobaougou Formation key to understanding the evolution of late Permian continental fauna in general, and of akidnognathid therocephalians in particular.

Tooth replacement patterns in the Early Triassic epicynodont Galesaurus planiceps (Therapsida, Cynodontia).

Sixteen specimens of the Early Triassic cynodont Galesaurus planiceps (including eight that were scanned using micro-computed tomography) representing different ontogenetic stages were assembled to study the dental replacement in the species. The growth series shows that the incisors and postcanines continue to develop and replace, even in the largest (presumably oldest) specimen. In contrast, replacement of the canines ceased with the attainment of skeletal maturity, at a basal skull length of ~90 mm, suggesting that Galesaurus had a finite number of canine replacement cycles. Additionally, the functional canine root morphology of these larger specimens showed a tendency to be open-rooted, a condition not previously reported in Mesozoic theriodonts. An alternating pattern of tooth replacement was documented in the maxillary and mandibular postcanine series. Both postcanine series increased in tooth number as the skull lengthened, with the mandibular postcanine series containing more teeth than the maxillary series. In the maxilla, the first postcanine is consistently the smallest tooth, showing a proportional reduction in size as skull length increased. The longer retention of a tooth in this first locus is a key difference between Galesaurus and Thrinaxodon, in which the mesial-most postcanines are lost after replacement. This difference has contributed to the lengthening of the postcanine series in Galesaurus, as teeth continued to be added to the distal end of the tooth row through ontogeny. Overall, there are considerable differences between Galesaurus and Thrinaxodon relating to the replacement and development of their teeth.

The tetrapod fauna of the upper Permian Naobaogou Formation of China: 5. Caodeyao liuyufengi gen. et sp. nov., a new peculiar therocephalian.

The upper Permian Naobaogou Formation has been the goal of recent contributions that notably increased the knowledge of its terrestrial vertebrate fauna and unravelled a hidden late Permian therocephalian diversity in China. Two very different species of therocephalians have been documented in the Naobaogou Formation and they were recovered as basal akidnognathids in cladistic analyses. In this contribution we describe Caodeyao liuyufengi gen. et sp. nov., represented by a partial skull and mandible, and a humerus. The new taxon features a short, high snout and a wide temporal opening with the coronoid process of the mandible separated by a wide space medially to the zygomatic arch. The latter feature is only recognized in the Russian therocephalian Purlovia maxima and it is also characteristic of non-mammaliaform cynodonts. Phylogenetic analysis indicates a close relationship of the new Chinese taxon with Purlovia maxima, producing a monophyletic Laurasian group in therocephalian phylogeny. With the representation of three different species, the Naobaogou Formation is now the most prolific unit documenting therocephalian late Permian diversity in China.

A proposed terminology for the dentition of gomphodont cynodonts and dental morphology in Diademodontidae and Trirachodontidae.

Gomphodont cynodonts were close relatives of mammals and one of the Mesozoic lineages of cynodont therapsids that became extinct at the end of the Triassic. Gomphodonts were omnivorous to herbivorous animals characterized by labiolingually expanded postcanines, which allowed tooth-to-tooth occlusion. The morphology of the upper and lower postcanines presents important means of distinguishing among major lineages within Gomphodontia, that is, Diademodontidae, Trirachodontidae, and Traversodontidae, but the dentition of most Diademodontidae and Trirachodontidae remain poorly documented. Here, we present a comprehensive description of the dentition of each diademodontid and trirachodontid species, as well as detailed illustrations of each dental unit, after firsthand examination of material and 3D reconstructions of postcanine teeth. Based on dental morphology, Trirachodon berryi and "Trirachodon kannemeyeri," considered as separate taxa by some authors are here interpreted as representing different ontogenetic stages of the same species. Likewise, Sinognathus and Beishanodon, thought to belong to non-cynognathian cynodonts and traversodontids by some authors, are referred to Trirachodontidae and Gomphodontia based on dental characters, respectively. Finally, we propose a standardized list of terms and abbreviations for incisors, canines, and postcanines anatomical entities, with the goal of facilitating future descriptions and communication between researchers studying the gomphodont dentition.

The tetrapod fauna of the upper Permian Naobaogou Formation of China: 3. Jiufengia jiai gen. et sp. nov., a large akidnognathid therocephalian.

Recent field trips to Member III of the Naobaogou Formation, Nei Mongol, China yielded new fossil discoveries, increasing our knowledge of the late Permian continental fauna from China. We present here a new large therocephalian, Jiufengia jiai gen. et sp. nov., represented by a partial skull with mandibles and part of the postcranial skeleton. This is the second therocephalian recovered from the Naobaogou faunal association and, in turn, the second akidnognathid from this unit and from China. The new taxon shows clear differences from Shiguaignathus wangi, the akidnogathid previously reported from the Naobaogou Formation: the presence of four upper postcanines, of a large suborbital vacuity, and the flat ventral surface of the vomer, lacking a ventromedian crest. Updating a previous phylogeny of therocephalians, we recover the new species as a basal member of Akidnognathidae, above a basal polytomy including the other two Laurasian akidnognathids, Shiguaignathus and Annatherapsidus, adding support to the hypothesis that this group originated in Laurasia.

Dental ontogeny in extinct synapsids reveals a complex evolutionary history of the mammalian tooth attachment system.

The mammalian dentition is uniquely characterized by a combination of precise occlusion, permanent adult teeth and a unique tooth attachment system. Unlike the ankylosed teeth in most reptiles, mammal teeth are supported by a ligamentous tissue that suspends each tooth in its socket, providing flexible and compliant tooth attachment that prolongs the life of each tooth and maintains occlusal relationships. Here we investigate dental ontogeny through histological examination of a wide range of extinct synapsid lineages to assess whether the ligamentous tooth attachment system is unique to mammals and to determine how it evolved. This study shows for the first time that the ligamentous tooth attachment system is not unique to crown mammals within Synapsida, having arisen in several non-mammalian therapsid clades as a result of neoteny and progenesis in dental ontogeny. Mammalian tooth attachment is here re-interpreted as a paedomorphic condition relative to the ancestral synapsid form of tooth attachment.

The tetrapod fauna of the upper Permian Naobaogou Formation of China: 1. Shiguaignathus wangi gen. et sp. nov., the first akidnognathid therocephalian from China.

The Permian from China has a well-known terrestrial record where approximately 30 tetrapod taxa, including several therapsids, have been described. However, the record of therocephalians in China has remained elusive. Shiguaignathus wangi gen. et sp. nov., discovered in the Member III of the Naobaogou Formation, Nei Mongol, China, is here described. This is the first therocephalian recovered from this fauna and only the second from the Permian of China. It is represented by a well-preserved robust snout of a medium-sized animal. This is the first akidnognathid reported from the Chinese Permian and only the second genus from Laurasia as one genus is known from Russia whereas the remaining members of the group are from the South African Karoo Basin. A phylogenetic analysis of therocephalians supports a basal position of S. wangi within Akidnognathidae, followed by the Russian Annatherapsidus. Akidnognathidae is the latest major group of therocephalian appearing in the fossil record, and one of the few that does not have species from South Africa representing its most basal members.

The mystery of a missing bone: revealing the orbitosphenoid in basal Epicynodontia (Cynodontia, Therapsida) through computed tomography.

The basal non-mammaliaform cynodonts from the late Permian (Lopingian) and Early Triassic are a major source of information for the understanding of the evolutionary origin of mammals. Detailed knowledge of their anatomy is critical for understanding the phylogenetic transition toward mammalness and the paleobiological reconstruction of mammalian precursors. Using micro-computed tomography (µCT), we describe the internal morphology of the interorbital region that includes the rarely fossilized orbitosphenoid elements in four basal cynodonts. These paired bones, which are positioned relatively dorsally in the skull, contribute to the wall of the anterior part of the braincase and form the floor for the olfactory lobes. Unlike procynosuchids and the more basal therapsids in which the orbitosphenoids are well developed, dense, and bear a ventral keel, the basal epicynodonts Cynosaurus, Galesaurus, and Thrinaxodon display cancellous, reduced, and loosely articulated orbitosphenoids, a condition shared with many eucynodonts. The hemi-cylindrical orbitosphenoid from which the mammalian condition is derived re-evolved convergently in traversodontid and some probainognathian cynodonts.

Oxygen isotopes suggest elevated thermometabolism within multiple Permo-Triassic therapsid clades.

The only true living endothermic vertebrates are birds and mammals, which produce and regulate their internal temperature quite independently from their surroundings. For mammal ancestors, anatomical clues suggest that endothermy originated during the Permian or Triassic. Here we investigate the origin of mammalian thermoregulation by analysing apatite stable oxygen isotope compositions (δ18Op) of some of their Permo-Triassic therapsid relatives. Comparing of the δ18Op values of therapsid bone and tooth apatites to those of co-existing non-therapsid tetrapods, demonstrates different body temperatures and thermoregulatory strategies. It is proposed that cynodonts and dicynodonts independently acquired constant elevated thermometabolism, respectively within the Eucynodontia and Lystrosauridae + Kannemeyeriiformes clades. We conclude that mammalian endothermy originated in the Epicynodontia during the middle-late Permian. Major global climatic and environmental fluctuations were the most likely selective pressures on the success of such elevated thermometabolism.

Aggregations and parental care in the Early Triassic basal cynodonts Galesaurus planiceps and Thrinaxodon liorhinus.

Non-mammaliaform cynodonts gave rise to mammals but the reproductive biology of this extinct group is still poorly known. Two exceptional fossils of Galesaurus planiceps and Thrinaxodon liorhinus, consisting of juveniles closely associated with an adult, were briefly described more than 50 years ago as examples of parental care in non-mammaliaform cynodonts. However, these two Early Triassic fossils have largely been excluded from recent discussions of parental care in the fossil record. Here we re-analyse these fossils in the context of an extensive survey of other aggregations found in these two basal cynodont taxa. Our analysis revealed six other unequivocal cases of aggregations in Thrinaxodon, with examples of same-age aggregations among immature or adult individuals as well as mixed-age aggregations between subadult and adult individuals. In contrast, only one additional aggregation of Galesauruswas identified. Taking this comprehensive survey into account, the two previously described cases of parental care in Galesaurus and Thrinaxodon are substantiated. The juveniles are the smallest specimens known for each taxon, and the size difference between the adult and the two associated juveniles is the largest found for any of the aggregations. The juveniles of Thrinaxodon are approximately only 37% of the associated adult size; whereas in Galesaurus, the young are at least 60% of the associated adult size. In each case, the two juvenile individuals are similar in size, suggesting they were from the same clutch. Even though parental care was present in both Galesaurus and Thrinaxodon, intraspecific aggregations were much more common in Thrinaxodon, suggesting it regularly lived in aggregations consisting of both similar and different aged individuals.

Cranial Ontogeny of the Early Triassic Basal Cynodont Galesaurus planiceps.

Ontogenetic changes in the skull and mandible of thirty-one specimens of Galesaurus planiceps, a basal non-mammaliaform cynodont from the Early Triassic of South Africa, are documented. The qualitative survey indicated eight changes in the craniomandibular apparatus occurred during growth, dividing the sample into three ontogenetic stages: juvenile, subadult, and adult. Changes in the temporal region, zygomatic arch, occiput, and mandible occurred during the transition from the subadult to adult stage at a basal skull length of 90 mm. At least four morphological and allometric differences divided the adult specimens into two morphs, indicating the presence of sexual dimorphism in Galesaurus. Differences include extensive lateral flaring of the zygomatic arches in the "male" morph resulting in a more anterior orientation of the orbits, and a narrower snout in the "female". This is the first record of sexual dimorphism in a basal cynodont, and the first time it is quantitatively documented in a non-mammaliaform cynodont. An ontogenetic comparison between Galesaurus and the more derived basal cynodont Thrinaxodon revealed differences in the timing and extent of sagittal crest development. In Galesaurus, the posterior sagittal crest, located behind the parietal foramen, developed relatively later in ontogeny, and the anterior sagittal crest rarely formed suggesting the anterior fibres of the temporalis were less developed than in Thrinaxodon. In contrast, craniomandibular features related to the masseters became more developed during the ontogeny of Galesaurus. The development of the adductor musculature appears to be one of the main factors influencing skull growth in these basal non-mammaliaform cynodonts. Anat Rec, 300:353-381, 2017. © 2016 Wiley Periodicals, Inc.

Mineralized periodontia in extinct relatives of mammals shed light on the evolutionary history of mineral homeostasis in periodontal tissue maintenance.

AIM: Dental ankylosis is a rare pathological condition in mammals, however, it is prevalent in their extinct relatives, the stem mammals. This study seeks to compare the mineralized state of the periodontal attachment apparatus between stem and crown mammals and discuss its implications for the evolution of non-mineralized periodontal attachment in crown mammals, including humans. MATERIALS AND METHODS: Thin sections of a fossil mammal and three stem mammals were compared to reconstruct periodontal tissue development across distantly related lineages. RESULTS: Comparisons revealed that the extinct relatives of mammals possessed the same periodontal tissues as those in mammals, albeit in different arrangements. The ankylotic condition in stem mammals was achieved through extensive alveolar bone deposition, which eventually contacted the root cementum, thus forming a calcified periodontal ligament. CONCLUSIONS: Dental ankylosis was part of the normal development of the stem mammal periodontium for millions of years prior to the evolution of a permanent gomphosis in mammals. Mammals may have evolved a permanent gomphosis by delaying the processes that produced dental ankylosis in stem mammals. Pathological ankylosis may represent a reversion to the ancestral condition, which now only forms via advanced ageing and pathology.

Tiarajudens eccentricus and Anomocephalus africanus, two bizarre anomodonts (Synapsida, Therapsida) with dental occlusion from the Permian of Gondwana.

Anomodontia was a highly successful tetrapod clade during the Permian and the Triassic. New morphological information regarding two bizarre basal anomodonts is provided and their palaeoecological significance is explored. The osteology of the recently discovered Tiarajudens eccentricus Cisneros et al. 2011, from the Brazilian Permian, is described in detail. The taxon exhibits unusual postcranial features, including the presence of gastralia. Additional preparation and computed tomography scans of the holotype of Anomocephalus africanus Modesto et al. 1999 discovered in the Karoo Basin of South Africa allow a reappraisal of this genus. Anomocephalus is similar to Tiarajudens with regard to several traits, including a battery of large, transversally expanded, palatal teeth. Molariform teeth are present in the mandible of the African taxon, providing additional insight into the function of the earliest tooth-occlusion mechanism known in therapsids. At least two waves of tooth replacement can be recognized in the palate of Anomocephalus. The outsized, blade-like caniniforms of the herbivorous Tiarajudens allow several non-exclusive ecological interpretations, among which we favour intraspecific display or combat. This behaviour was an alternative to the head-butting practised by the contemporary dinocephalians. Combat specializations that are considered typical of Cenozoic herbivores likely evolved during the Middle Permian, at the time the first communities with diverse, abundant tetrapod herbivores were being assembled.

Physiological implications of the abnormal absence of the parietal foramen in a late Permian cynodont (Therapsida).

The third eye (pineal eye), an organ responsible for regulating exposure to sunlight in extant ectotherms, is located in an opening on the dorsal surface of the skull, the parietal foramen. The parietal foramen is absent in extant mammals but often observed in basal therapsids, the stem-group to true mammals. Here, we report the absence of the parietal foramen in a specimen of Cynosaurus suppostus, a Late Permian cynodont from South Africa (SA). Comparison with Procynosuchus delaharpeae, a contemporaneous non-mammalian cynodont from SA, demonstrates that the absence of this foramen is an abnormal condition for such a basal species. Because seasonality was marked during the Late Permian in SA, it is proposed that the third eye was functionally redundant in Cynosaurus, possibly due to the acquisition of better thermoregulation or the evolution of specialized cells in the lateral eyes to compensate for the role of the third eye.

Radiographic study on the anatomical characteristics of the proximal femur in Brazilian adults.

OBJECTIVE: To ascertain the geometry of the femur in the Brazilian population by means of a radiographic study and to correlate the values with regard to sex and right/left side. METHODS: Five hundred anteroposterior radiographs of the pelvis of skeletally mature patients (250 of each sex) who did not present any osteoarthrosis, fractures or tumoral or infectious lesions were analyzed. The length and width of the femoral neck, length of the femoral axis, neck-shaft angle and femoral offset were measured. RESULTS: The following means were observed: 36.54 mm for the length of the femoral neck; 37.48 mm for the width of the femoral neck; 108.42 mm for the length of the femoral axis; 130.47° for the neck-shaft angle; and 44.4 mm for the femoral offset. CONCLUSION: The mean values for the main measurements on the proximal femur in Brazilians differed from those of previous studies. It could also be shown that there was a statistically significant mean difference between men and women for all the variables, both on the left and on the right side, and that the men had greater means than the women.

OBJETIVO: Verificar a geometria do fêmur da população brasileira por meio de estudo radiográfico e correlacionar os valores quanto ao sexo e ao lado direito/esquerdo. MÉTODOS: Foram analisadas 500 radiografias anteroposteriores de bacia de pacientes esqueleticamente maduros, 250 de cada sexo, sem a presença de osteoartrose, fraturas e lesões tumorais ou infecciosas. Foram mensurados os valores do comprimento e da largura do colo do fêmur, do eixo femoral, do ângulo colodiafisário e do offset femoral. RESULTADOS: Observou-se uma média de 36,54 mm do comprimento do colo do fêmur, 37,48 mm da largura do colo do fêmur, 108,42 mm do comprimento do eixo femoral, 130,47° do ângulo colodiafisário e 44,4 mm do offset femoral. CONCLUSÃO: Os valores médios das principais medidas do fêmur proximal dos brasileiros diferem de trabalhos prévios. Foi possível evidenciar também que existe diferença média estatisticamente significante entre homens e mulheres para todas as variáveis, tanto do lado esquerdo quanto do direito, e que os homens tiveram média maior do que as mulheres.