Revisiting proboscidean phylogeny and evolution through total evidence and palaeogenetic analyses including Notiomastodon ancient DNA.

The extinct Gomphotheriidae is the only proboscidean family that colonized South America. The phylogenetic position of the endemic taxa has been through several revisions using morphological comparisons. Morphological studies are enhanced by paleogenetic analyses, a powerful tool to resolve phylogenetic relationships; however, ancient DNA (aDNA) preservation decreases in warmer regions. Despite the poor preservation conditions for aDNA in humid, sub-tropical climates, we recovered ∼3,000 bp of mtDNA of Notiomastodon platensis from the Arroyo del Vizcaíno site, Uruguay. Our calibrated phylogeny places Notiomastodon as a sister taxon to Elephantidae, with a divergence time of ∼13.5 Ma. Additionally, a total evidence analysis combining morphological and paleogenetic data shows that the three most diverse clades within Proboscidea diverged during the early Miocene, coinciding with the formation of a land passage between Africa and Eurasia. Our results are a further step toward aDNA analyses on Pleistocene samples from subtropical regions and provide a framework for proboscidean evolution.

The micro from mega: Dental calculus description and the first record of fossilized oral bacteria from an extinct proboscidean.

OBJECTIVE: This study explores the chemical composition and surface aspects of fossilized dental calculus from the South American Quaternary proboscidean Notiomastodon platensis and the first record of fossilized oral bacteria from extinct megafauna. MATERIALS: Blocks of dental calculus removed from the third molar of five specimens of Notiomastodon platensis collected from Brazil, Argentina, and Ecuador. METHODS: We analyzed five samples of dental calculus by SEM and SEM-EDS, following a rigid protocol to avoid bacterial contamination. RESULTS: The dental calculus surface is homogeneous, porous, with various crystals, and composed mainly by oxygen and calcium. One sample revealed a well-preserved mineralized biofilm, with several rods and cocci bacteria. CONCLUSIONS: This is the first fossilized record of oral bacterial communities associated with extinct proboscideans. SIGNIFICANCE: This record confirms the parasitism between oral bacteria and Notiomastodon platensis and will enable the study of paleogenomic aspects of oral microbiota of proboscideans. LIMITATIONS: Fossilization conditions of proboscidean teeth with dental calculus are variable among specimens. Although rare, the preservation of oral bacteria is expected because of the oral biofilm composition.

A new Mammutidae (Proboscidea, Mammalia) from the Late Miocene of Gansu Province, China.

The "Yanghecun specimen", a proboscidean specimen represented by a mandible from Miocene of China and previously described as Gomphotheriidae, is here reviewed and described as a new genus and species of Mammutidae: Sinomammut tobieni. This taxon is a longirostrine mastodon, lacking lower tusks, and bearing a wide last molar with oblique and non-inflated lophids, broad transverse interlophids, and yoke-like wear figures. Phylogenetic analysis of Mammutidae based on dental and mandibular features recovered S. tobieni as sister group of the mastodon Mammut. The longirostrine condition and the well-developed lower incisors seem to be primitive for Mammutidae, while the brevirostry is the derived condition, probably emerged during the middle Miocene (12-11 Mya). However, two derived conditions are recognized to the lower tusks: the absence of lower tusks (S. tobieni) and the occasional presence of vestigial lower tusks (Mammut).

The Dance of Tusks: Rediscovery of Lower Incisors in the Pan-American Proboscidean Cuvieronius hyodon Revises Incisor Evolution in Elephantimorpha.

The incisors of proboscideans (tusks and tushes) are one of the most important feature in conservation, ecology and evolutionary history of these mammals. Although the absence of upper incisors is rare in proboscideans (occurring only in deinotheres), the independent losses of lower incisors are recognized for most of its lineages (dibelodont condition). The presence of lower incisors in the Pan-American gomphothere Cuvieronius hyodon was reported a few times in literature, but it was neglected in systematic studies. We analyzed several specimens of Cuvieronius hyodon from the Americas and recognized that immature individuals had lower incisors during very early post-natal developmental stages. Subsequently, these are lost and lower incisors alveoli close during later developmental stages, before maturity. Moreover, for the first time in a formal cladistic analysis of non-amebelodontine trilophodont gomphotheres, Rhynchotherium and Cuvieronius were recovered as sister-taxa. Among several non-ambiguous synapomorphies, the presence of lower incisors diagnoses this clade. We recognize that the presence of lower incisors in Cuvieronius and Rhynchotherium is an unique case of taxic atavism among the Elephantimorpha, since these structures are lost at the origin of the ingroup. The rediscovery of the lower incisors in Cuvieronius hyodon, their ontogenetic interpretation and the inclusion of this feature in a revised phylogenetic analysis of trilophodont gomphotheres brought a better understanding for the evolutionary history of these proboscideans.

Population structure of the gomphothere Stegomastodon waringi (Mammalia: Proboscidea: Gomphotheriidae) from the Pleistocene of Brazil.

The Quaternary fossil record of Águas de Araxá (Q AA) is represented mainly by an accumulation of skeletal elements of several sizes, which are assigned to a population of Stegomastodon waringi. We analyzed 97 molars according to the ear stages of Simpson and Paula-Couto (1957), and developed a morphometric ear index. The population structure (proportion of immature, subadult, adult, mature adult and senile adult individuals) was identified, and these five age classes were compared to those of extant elephant populations and defined with social implications. The analysis made possible to establish that the population is largely composed of adults: 14.89% are immature individuals, 23.04% subadults, 27.65% adults, 17.21% mature adults and another 17.21% senile adults. Based on population structure, we do not discard the possibility that the fossil population was stable or in recovery, and/or was experiencing a high-predation period on younger individuals. The number of individuals composing the past population studied here could suggest that the occupied environment was open due to comparisons to populations of extant elephants. We consider this population as an aggregation of family units, which suggests a time of low environmental humidity. Based on literature and our findings, their extinction appears to be regional and probably related to a catastrophic event.

Population structure of the gomphothere Stegomastodon waringi (Mammalia: Proboscidea: Gomphotheriidae) from the Pleistocene of Brazil

The Quaternary fossil record of Águas de Araxá (Q AA) is represented mainly by an accumulation of skeletal elements of several sizes, which are assigned to a population of Stegomastodon waringi. We analyzed 97 molars according to the ear stages of Sipson and Paula-Couto (1957), and developed a orphoetric ear index. The population structure (proportion of immature, subadult, adult, mature adult and senile adult individuals) was identified, and these five age classes were compared to those of extant elephant populations and defined with social implications. The analysis made possible to establish that the population is largely composed of adults: 14.89 percent are immature individuals, 23.04 percent subadults, 27.65 percent adults, 17.21 percent mature adults and another 17.21 percent senile adults. Based on population structure, we do not discard the possibility that the fossil population was stable or in recovery, and/or was experiencing a high-predation period on younger individuals. The number of individuals composing the past population studied here could suggest that the occupied environment was open due to comparisons to populations of extant elephants. We consider this population as an aggregation of family units, which suggests a time of low environmental humidity. Based on literature and our findings, their extinction appears to be regional and probably related to a catastrophic event.

O registro fossilífero do Quaternário de Águas de Araxá (QAA) é representado principalmente pelo acúmulo de restos dentários e de esqueleto de diversos tamanhos, atribuídos a uma população pretérita do mastodonte Stegomastodon waringi. Foram analisados 97 molares de acordo com os estágios de desgaste propostos por Simpson e Paula-Couto 1957, e desenvolveu-se um índice morfométrico de desgaste. A estrutura populacional (proporção entre indivíduos imaturos, subadultos, adultos, adultos maduros e adultos senis) foi identificada e comparada com populações de elefantes atuais. Foi possível estabelecer que esta população era amplamente composta por adultos: 14,89 por cento eram indivíduos imaturos; 23,04 por cento eram subadultos; 27,65 por cento eram adultos; 17,21 por cento eram adultos maduros e outros 17,21 por cento eram adultos senis. Baseados na estrutura populacional observada, não se descartou a possibilidade desta população estar estável ou em recuperação e/ou de ter passado por um período de predação dos indivíduos mais jovens. O número de indivíduos encontrados nesta população sugere que esta ocupava um ambiente aberto, de acordo com comparações com populações de elefantes atuais. Considerou-se a assembléia de mastodontes do Quaternário de Águas de Araxá como uma agregação de unidades familiares, o que sugere um momento de baixa umidade ambiental. Baseados em literatura e nos resultados encontrados, a sua extinção parece ser regional e possivelmente relacionada a um evento catastrófico.