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Journal of South American Earth Sciences 109 (2021) 103296

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Journal of South American Earth Sciences 109 (2021) 103296 Journal of South American Earth Sciences 109 (2021) 103296 Contents lists available at ScienceDirect Journal of South American Earth Sciences journal homepage: www.elsevier.com/locate/jsames Paleoenvironments and paleoecology of the Santa Cruz Formation (early-middle Miocene) along the Río Santa Cruz, Patagonia (Argentina) Richard F. Kay a,*, Sergio F. Vizcaíno b,c, M. Susana Bargo b,d, Jackson P. Spradley e, Jos´e I. Cuitino~ f a Department of Evolutionary Anthropology and Division of Earth and Ocean Sciences, Duke University, Durham, NC, 27708, USA b Division´ Paleontología Vertebrados, Facultad de Ciencias Naturales y Museo, Unidades de Investigacion,´ Anexo Museo, Av. 60 y 122, 1900, La Plata, Argentina c Consejo Nacional de Investigaciones Científicas y T´ecnicas (CONICET), Argentina d Comision´ de Investigaciones Científicas, Provincia de Buenos Aires (CICPBA), Argentina e North Carolina State University - College of Veterinary Medicine, Department of Molecular Biomedical Sciences. Raleigh, NC, USA f Instituto Patagonico´ de Geología y Paleontología Centro Nacional Patagonico,´ Puerto Madryn (U9120), Argentina ARTICLE INFO ABSTRACT Keywords: The continental early-middle Miocene Santa Cruz Formation (SCF) in Austral Patagonia contains the best record Ecometrics of South American mammalian faunas prior to the Great American Biotic Interchange (GABI) and is of particular Paleobiology interest because it is the best preserved high-latitude continental biotic record in the Southern Hemisphere Paleoclimate spanning the mid-Miocene Climatic Optimum. Through intensive fieldwork we recovered numerous fossil ver­ Mammals tebrates, mostly mammals, from the SCF along the Río Santa Cruz (RSC), the type area for the formation and its Neogene South America fauna. We examine whether the SCF fauna differed among three distinct temporal intervals of the SCF spanning, Frugivore problem from the oldest to youngest, the Atlantic coastal suite of localities Fossil Levels (FL) 1–7, at about 17.4 Ma, through localities in the RSC Barrancas Blancas (BB), between ~17.2 and ~16.3 Ma, and Segundas Barrancas Blancas (SBB), between ~16.5 and ~15.6 Ma. With the objective of reconstructing paleoenvironmental and community structure of these RSC faunas, we compared them with 55 extant lowland mammalian localities ◦ ◦ across South America from 8 N to 55 S latitude representing a wide range of seasonality and, annual rainfall and temperature, as well as canopy height and net primary productivity, sampling communities ranging from tropical rainforest to semi-arid steppe. Extant nonvolant mammalian genera at each locality were assigned a body size interval and niche parameters reflecting diet and substrate use, from behavioral data in the literature. Extinct genera were assigned similar niche metrics on the basis of their morphology. From the generic niche parameters, we compiled indices and ratios that express vectors of the community structure of each fauna, including the total number of genera, the pervasiveness of arboreality, frugivory, and browsing, and the relative richness of predators to their prey. The community structure variables were used to model community structure of the fossil localities based on uniformitarian principles. The fossil sample includes 44 genera of mammals from FL 1–7, 38 genera from BB, and 44 genera from SBB. The Simpson Coefficients of faunal similarity among the fossil localities are no greater than expected on the basis of the geographic distances among them, and do not suggest any apparent climatic differences. Based on the models we obtained no significant differences in MAP (Mean Annual Precipitation) for FL 1–7, BB and SBB, with mean estimates of 1635 mm, 1451 mm, and 1504 mm, with the confidence intervals for the estimates overlapping widely. MAT (Mean Annual Temperature) estimates ◦ ◦ ◦ are between ~21 C and ~22 C for FL 1–7 and SBB, possibly lower at 16 C for BB, but with a wide and ◦ ◦ overlapping range of estimates. Temperature seasonality is modest (3 C to 4 C) and similar for all localities. Canopy heights exceed 20 m for all sites. Despite these geographic and inferred climatic similarities, the presence of certain key taxa (e.g., the caviomorph rodent Prolagostomus and the typothere Pachyrukhos) together with an increased overall abundance and richness of rodents with ever-growing cheek teeth suggests a trend to aridifi­ cation in the upper part of the SCF at SBB compared with FL 1–7 and BB. Taken together, we propose that the SCF paleoenvironment consisted largely of semi-deciduous forests ranging into savannas with gallery-forest com­ ponents. This range of habitats occurs today where the mesic inland Atlantic forests of Southern Brazil, * Corresponding author. E-mail addresses: [email protected] (R.F. Kay), [email protected] (S.F. Vizcaíno), [email protected] (M.S. Bargo), [email protected] (J.P. Spradley), [email protected] (J.I. Cuitino).~ https://doi.org/10.1016/j.jsames.2021.103296 Received 9 December 2020; Received in revised form 23 March 2021; Accepted 23 March 2021 Available online 29 March 2021 0895-9811/© 2021 Elsevier Ltd. All rights reserved. R.F. Kay et al. Journal of South American Earth Sciences 109 (2021) 103296 northeastern Argentina and eastern Paraguay give way northwestward into the more xeric Paraguayan Gran Chaco. These interpretations are in general agreement with other sources of evidence from sedimentology, paleosols, isotopes, paleobotany and other faunal elements. We highlight the value of focusing paleoenvir­ onmental and paleocological studies of the SFC on stratigraphically and geographically confinedsamples instead of on the entire temporal and geographic distribution of the SCF based on historical collections with limited provenance. The Santacrucian can be considered a model to the study of South American faunas after the arrival of hystricomorph rodents and anthropoid primates but before GABI. 1. Introduction a renewed background and vision. From a paleoecological perspective, in the last decade there have The continental early-middle Miocene (Burdigalian-early Langhian) been two different approaches for the understanding of the biota and fossil record of the Santa Cruz Formation (SCF) from Austral Patagonia, environments of the SCF: either considering stratigraphically and represents the biota that has most impacted historically and conceptu­ geographically restricted samples obtained through exhaustive field ally the understanding of the Cenozoic biotic evolution of South America work efforts (Kay et al., 2012; Vizcaíno et al., 2010) or globally along the prior to the Great American Biotic Interchange (GABI), when South entire temporal and geographic distribution of the formation based on America was mostly isolated from other Continents . Moreover, the SCF bibliography and historical collections (Croft, 2013). contains the best preserved high-latitude continental biotic record in the The approach by Vizcaíno et al. (2010) and Kay et al. (2012) was Southern Hemisphere providing further insights into mid-Miocene based on fossils collected from geographically and stratigraphically temperature and precipitation. It is well known that floras and faunas restricted sets of localities of the Atlantic Coast outcrops, which are often dependent on warm, wet conditions expanded to higher latitudes in in an excellent state of preservation, including partial or complete ar­ South America at that time (Frenguelli, 1953; Ortiz-Jaureguizar and ticulated skeletons, offering a unique opportunity to perform paleobio­ Cladera, 2006; Pascual and Odreman Rivas, 1971; Pascual and logical studies based on a form-function approach (Vizcaíno et al., Ortiz-Jaureguizar, 1990). The analysis of Hinojosa (2005) of early and 2012d). For these authors, the fact that many taxa come from certain ◦ middle Miocene floras at 33–34 S latitude between 21 and 13 Ma, levels deposited in a restricted time frame provides a narrow temporal suggests an increase in mean annual temperatures (MATs) from the late window that allows reliable paleoecological analysis (Kay et al., 2012; ◦ ◦ Oligocene-early Miocene from 16–17 C to MATs exceeding 22 C. Perkins et al., 2012). Vizcaíno et al. (2010) used the relationship be­ Likewise, at the beginning of the Miocene, the floras indicate an even tween population density and body size for estimating the on-crop more abrupt rise in rainfall, which culminated in values exceeding 1000 biomass (in kg/km2) of the species of these paleocommunities and mm. During the middle and late Miocene, rainfall subsided, reaching calculated their metabolic requirements. Kay et al. (2012) reconstructed minimum values of ~440 mm by around 10 Ma (Hinojosa, 2005). The the niche structure by identifying the number of species present, the expanded faunal samples and precise dating of our early-middle body size, locomotion, and diet of the mammalian genera at the suite of Miocene SCF localities (Trayler et al., 2020b) make them ideal candi­ localities FL 1–7 compared with similar kinds of data for extant faunas of ◦ dates for evaluating Patagonian climate and biota nearly 20 farther South America; using that data they interpreted the paleoclimate and south that Hinojosa’s localities, practically at the southern end of the paleoenvironment of FL 1–7. The FL 1–7 fauna was later studied by continent. Spradley et al. (2019) updating the approach of Kay et al. (2012) to Historically, the record from the Río Santa Cruz (RSC) represents the derive paleoecological predictive
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