Article

A new specimen of Camelops hesternus (Artiodactyla, Camelidae) from Valsequillo, Puebla, Mexico, with comments about their dietary preferences and the population density of the species

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  • Secretaría de Medio Ambiente e Historia Natural
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Abstract

Camelidae is one of the most diverse and successful artiodactyl families, with a long geological history and wide geographical range in North America. Camelops hesternus was the one of last camelids in North America during the Pleistocene (Rancholabrean) and its remains have been found from Alaska south into Mexico. Here, we describe a new specimen of this species based on a complete mandible from the Late Pleistocene sediments of the Cerro Grande de la Mesa Calderón monogenetic volcano in the Valsequillo Basin, Puebla, central Mexico. A mesowear analysis of the dentition indicates wear in the new specimen to be similar to that of the Klipspringer, Oreotragus oreotragus, indicating a browsing diet. The population density of C. hesternus in the Valsequillo Basin was estimated using a differential non-linear equations mathematical model under three scenarios (stress, optimal, and abundance conditions) a indicating the range in its population density of 0.51-1.8 ind/km2, and this species was not abundant in the region. Despite the low population density C. hesternus has been reported in Mexico from 27 Pleistocene localities, with a geographic range from north (29°40' latitude, Sonora) to south (16°14' latitude, Chiapas) at an altitude range of 0–200 m asl in Baja California Sur to 2,500 m asl in the Trans-Mexican Volcanic Belt.

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The La Brea Tar Pits are a highly fossiliferous late Pleistocene locality, with over 230 vertebrate species, and the potential to help clarify the ecology of Pleistocene megafauna prior to their extinction. Here, we analyze dental microwear textures, dental mesowear, and stable carbon isotopes to assess the dietary behavior of the three most abundant ungulates at La Brea. We examined Bison antiquus, Camelops hesternus, and Equus occidentalis teeth from pits spanning the latest Pleistocene, focusing on pit 77 (~ 35 Ka, glacial period), and pits 61/67 (~ 11.5 Ka, interglacial period). Dental microwear attributes demonstrate that B. antiquus and E. occidentalis consumed foods with similar textural properties to those preferred by mixed feeding and browsing bovids, as well as significantly different food textures than the diet of modern bison and horses. Camelops hesternus has attributes most similar to modern browsing bovids and giraffes, revealing that all taxa studied likely consumed at least some woody browse. Complexity values (Asfc) in B. antiquus and E. occidentalis are significantly greater at pit 77 than at pits 61/67 and modern populations, suggesting a dietary decrease in woody vegetation and shrubs from the glacial to interglacial period. Dental mesowear values further indicate that B. antiquus and C. hesternus consumed woody vegetation, whereas E. occidentalis grazed. The different dietary interpretation for E. occidentalis may reflect a discrepancy between its average diet inferred from mesowear and its most recent diet inferred from microwear. Lastly, stable carbon isotopes reveal diets composed mostly of C3 vegetation, with B. antiquus incorporating significantly more C4 vegetation than either of the other dominant herbivores during the accumulation of both assemblages. Collectively, these data suggest that the dietary ecology of the dominant La Brea herbivores was more nuanced than previously thought, shifting coincident with changing climates, and not necessarily predictable from morphology or any one proxy method.
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Wild members of Camelidae live in some of the most arid environments, including North Africa, Arabia, the Gobi Desert of China and Mongolia and high elevation environments in the Andes Mountains. A better understanding of the paleoecology of the three most abundant Pleistocene camelids (Camelops, Hemiauchenia, and Palaeolama) may clarify modern adaptations to arid environments. Mammalian tooth enamel δ¹³C values were used to compare diets of co-occurring species in California, Texas, and Florida and δ¹⁸O values were used to investigate climate. Carbon isotope analysis suggests Camelops was likely an opportunistic browser that consumed both C3 and C4 browse/CAM plants, potentially consuming C4 browse (e.g., saltbush). Hemiauchenia had an opportunistic and highly generalized diet, while Palaeolama was a specialized forest browser. Stable oxygen isotopes and aridity index values suggest that Ingleside was warmer than McKittrick Brea, but there are no significant differences in aridity between the two sites. Co-occurrence data from the Paleobiology Database suggest that Palaeolama was restricted to forested environments as it occurred with two browsers, Tapirus and Odocoileus at 90.5% of all sites. Camelops and Hemiauchenia both co-occurred with a broader range of taxa, further suggesting these camelids lived in diverse habitats. The generalized diet of Hemiauchenia, the likely ancestor of modern South American camelids, allowed for the adaptations of extant Lama and Vicugna to survive in the arid environments of the Andes Mountains. Collectively, these data clarify the dietary ecology of extinct camelids and provide insight into the potential importance of generalist diets for increased resilience to changing environments and/or climates.
Article
We review the history of investigations at the Valsequillo archaeological area south of Puebla, Mexico, from the early 1960s to 2010. Evidence from diatoms, (U-Th)/ He measurements, early uranium-series dates, later zircon fission-track dates, mineral weathering, tephra hydration dates, and vertebrate fossils imply that the principal archaeological site, Hueyatlaco, could be older than 250,000 years. Hueyatlaco rests unconformably on Xalnene Tuff (basaltic ash) dated at 1.3 Ma by whole-rock argon-argon analysis. This finding differs greatly from a recent interpretation that the site is 40,000 years old.
Article
Whilst in Paris in the month of September last, I was favoured by the Marchioness of Hastings with information of the discovery of the fossils that form the chief subject of the present communication. Her ladyship wrote,—“My search in a particular part of the Eocene beds of the Isle of Wight, where formerly I found that Lophiodon or Palæotherium bone figured in your ‘British Fossil Mammalia,’ has been eminently successful. I have got two portions of jaw and many other bones. I have sketched them for you. Are they Coryphodon or Anoplotherium?” The pen-and-ink sketches, executed with the skill and accuracy of an accomplished artist, showed the fossils to belong to the Anthracotherioid family of Ungulata, with an evident resemblance to that species in the upper molars of which Cuvier had detected a closer resemblance to the Anoplotherium than the same teeth of the typical genus Anthracotherium present
Article
Camelops is a giant llama from the Pliocene and Pleistocene of western North America. Of the 17 species (most, if not all, from the Irvingtonian and Rancholabrean North American Land Mammal Ages) that have been referred to Camelops, only 6 are currently recognised as valid. This review examines Camelops from the Irvingtonian and Rancholabrean (Middle and Late Pleistocene) and describes for the first time material from the latest Wisconsinan of the Nueces River valley of South Texas. During this interval, there are two valid previously named species: the smaller mainly Irvingtonian Camelops minidokae and the larger, mainly Rancholabrean Camelops hesternus. Camelops hesternus, Camelops sulcatus, Camelops huerfanensis and Camelops traviswhitei are junior synonyms of Camelops hesternus. In addition, there are possibly two additional species: one with short, broad metapodials and one with short, slender metapodials.
Article
A new approach of reconstructing ungulate diets, the mesowear method, was introduced by Fortelius, Solounias (2000). Mesowear is based on facet development on the occlusal surfaces of the teeth. Restricting mesowear investigation on the M2 as has previously been suggested would limit application of the mesowear methodology to large ungulate assemblages. Most of the fossil, subfossil and recent ungulate assemblages that have been assigned to a single taxon have a smaller number of individuals. This results in the demand to extend the mesowear method to further tooth positions in order to obtain stable dietary classifications of fossil taxa. The focus of this paper is to test, if a consistent mesowear classification is obtainable for the remaining positions of the upper cheek tooth dentition (P2, P3, P4, M1 and M3) and for combinations of these tooth positions. For statistical testing, large assemblages of isolated cheek teeth of the Vallesian hipparionine horse Hippotherium primigenium Meyer, 1829 and of two populations of the recent zebra Equus burchelli Gray, 1824 are employed as models. Subsequently, all single cheek tooth positions and all possible combinations of these tooth positions are tested for their consistency in classification of the mesowear variables compared to the M2, the model tooth of Fortelius, Solounias (2000). As the most consistent model for the proposed "extended" mesowear method, the combination of four tooth positions P4, M1, M2, and M3 is identified, which allows to include the largest number of isolated tooth specimens from a given assemblage, and fulfills the demand of being consistent in the dietary mesowear classification with the "original" mesowear method. We propose the "extended" mesowear method to be particularly well suited for the reconstruction of paleodiets in hypsodont equids. © Publications Scientifiques du Muséum national d'Histoire naturelle, Paris.
Article
Hundreds of fossils of middle Pleistocene age mammals have been obtained from the tufa, sands, and gravels of the Tacubaya Formation just east of the City of Aguascalientes, in central Mexico. The fossils are probably of Illinoian age. This is the only large, systematically-collected, local fauna known from the Pleistocene of Mexico. Few remains of small mammals were found, but the Cedazo local fauna includes at least 39 species, three here described as new. The Cedazo local fauna is a unit fauna, representing the kinds of mammals that lived together in one small area during one period of time. The Cedazo local fauna lived on plains or grasslands with brush and trees along watercourses but with no tropical forest element in the environment.
Article
Camelops traviswhitei, from the Cedazo local fauna, Tacubaya Formation, medial Pleistocene age, differs from described species of Camelops in sharply "V"-shaped lakes in both upper and lower cheek teeth, with moderately thick enamel on the labial sides of the lakes but very thin enamel on the lingual sides, and lacking cementum in the lakes. Other species of Camelops have bent-oval lakes, bordered on both lingual and labial sides by thick enamel, and filled with cementum.
Article
Extension of the mesowear method to include the lower cheek teeth of ruminants will dramatically increase sample sizes and thus the statistical power of paleodietary inferences. However, the mesowear method of Fortelius and Solounias, which was designed for application to the upper molars, does not effectively separate ruminant species by diet when applied to the lower teeth. Upper and lower mesowear scores have sometimes been compared among non-analogous cusps (i.e. the buccal cusps of the maxillary teeth, which experience incursion and the buccal cusps of the mandibular teeth, which experience excursion during the chewing stroke). We therefore compare mesowear scores between the buccal cusps of maxillary cheek teeth and the lingual cusps of mandibular cheek for a large sample of ruminants because both cusps experience incursion during the chewing stroke. Using the original mesowear scoring method, we find dietary signal in both the maxillary and mandibular cheek teeth and a high correlation between them using both non-phylogenetic and phylogenetic comparative methods. Noting unique patterns of mesowear among the mandibular teeth, we also propose a new scoring method with additional wear categories that improves dietary inference when applied to the lower teeth and is highly repeatable. We also find that mandibular mesowear scores are consistently lower than for their maxillary counterparts. Although differential wear among the upper and lower teeth is much less apparent when applying our new scoring method, wear differences might relate to anisodonty (i.e. mandibular cheek teeth are narrower). Overall, we recommend our new scoring method for application to the lingual cusps of the lower second molars of fossil ruminants.
Article
Presented here is a cladistic analysis of the South American and some North American Camelidae. This analysis shows that Camelini and Lamini are monophyletic groups, as are the genera Palaeolama and Vicugna, while Hemiauchenia and Lama are paraphyletic. Some aspects of the migration and distribution of South American camelids are also discussed, confirming in part the propositions of other authors. According to the cladistic analysis and previous propositions, it is possible to infer that two Camelidae migration events occurred in America. In the first one, Hemiauchenia arrived in South America and, this was related to the speciation processes that originated Lama and Vicugna. In the second event, Palaeolama migrated from North America to the northern portion of South America. It is evident that there is a need for larger studies about fossil Camelidae, mainly regarding older ages and from the South American austral region. This is important to better undertand the geographic and temporal distribution of Camelidae and, thus, the biogeographic aspects after the Great American Biotic Interchange.
Article
On a review of the various facts and considerations discussed in the preceding pages, it seems clear that the Mammalian fauna of the Fluvio-marine Crag is of a Pliocene age. The undoubted association of M. (Tetraloph.) Arvernensis and of E. (Loxodon) meridionalis in this deposit admits of no other inference. The mixed contents of the Red Crag, including mammalian remains of different strata from the Eocene period upwards, are inferred to have been deposited in the reconstructed strata also within the Pliocene period, since M. (Tetraloph) Arvernensis, which occurs so abundantly in the Red Crag, has not been met with anywhere on the Continent of Europe except in deposits of a Pliocene age. The Red Crag sea appears to have breached a previously established and populated Pliocene land, and to have buried the bones referable to various epochs in the same sea-bottom.