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The spread of grasslands in the Miocene and of C4 grasses in the late Miocene-Pliocene represents a major development in terrestrial plant evolution that affected the climate system and faunal evolution. The macrofossil record of grasses is sparse, likely due to the limited preservation potential of grasses. Diagnosis of the C3 or C4 photosynthetic pathway depends on preservation of both cellular structures and organic carbon for isotope analysis. Here we analyze the anatomical and isotopic composition of newly-collected grass fossils from the Dove Spring Formation, Red Rock Canyon State Park, California, USA, located in the El Paso Basin on the western side of the Basin and Range Province, a site previously identified as one of the earliest known C4 grass fossil bearing localities. We analyzed the anatomical and geochemical characteristics of these new grass fossils dated to 12.01–12.15 Ma. The fossils analyzed in this study include grass shoots and in cross-section display anatomy indicative of the C3 photosynthetic pathway. We isolated organic carbon from the stem fossils and determined the carbon isotopic composition to be − 24.8 ± 0.5‰. Together, the anatomical and geochemical analyses confirm that these plants used the C3 photosynthetic pathway. Our findings are consistent with dietary evidence based on tooth enamel from grazing mammals of available C3 resources in the same sections. These newly reported Miocene-age C3 grass fossils contribute to a sparse macrofossil record of grass evolution. Overall, paleoecological reconstructions at this site indicate more humid conditions during the Miocene compared to the modern Mojave Desert with C3 grasses and diverse grazing mammals.

The stable carbon isotope ratios of tooth enamel from 89 herbivores from the Dove Spring Formation (DSF) of the El Paso Basin, western Mojave Desert, California, were analyzed to determine if C4plants may have been present in the late Miocene of southern California. Taxa analyzed include antilocaprids, castorids, camelids, equids, gomphotheriids, merycoidodontids, and rhinocerotids. The atmospheric CO2-δ13C value has varied significantly in the geologic past, leading to changes in the δ13C values of plants and tooth enamel. The estimated δ13C value of atmospheric CO2in the middle to late Miocene is −5.8±0.2‰, making the pure C3diet cutoff value higher than the modern by ~2‰. Given this, much of the evidence of an early spread of C4, which inappropriately uses the modern C3diet cutoff value, may be a false signal from water-stressed C3plants and the amount of C4biomass in the geologic past may have been over-estimated. The enamel-δ13C values of all taxa sampled are −10.1±1.1‰ (n=89). Accounting for the changes in the δ13C values of atmospheric CO2, enamel-δ13C values from the DSF indicate that herbivores from that area had C3diets and lived in an environment dominated by C3plants. Serial enamel isotope data from the DSF revealed negligible intra-tooth variations in the δ13C values, confirming that these herbivores consumed C3plants year-round. If the C4photosynthetic pathway did spread to southern California prior to the rapid, global rise in C4in the late Miocene, the taxa sampled did not incorporate C4biomass into their diets. The results of the stable carbon isotope analysis, along with the evidence from the rock record and the life habits of the fossil taxa present, indicate that the El Paso Basin consisted of diverse habitats including C3grasslands, wooded C3grasslands, and riparian forests.