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Abstract Premise The spurge family Euphorbiaceae is prominent in tropical rainforests worldwide, particularly in Asia. There is little consensus on the biogeographic origins of the family or its principal lineages. No confirmed spurge macrofossils have come from Gondwana. Methods We describe the first Gondwanan macrofossils of Euphorbiaceae, represented by two infructescences and associated peltate leaves from the early Eocene (52 Myr ago [Ma]) Laguna del Hunco site in Chubut, Argentina. Results The infructescences are panicles bearing tiny, pedicellate, spineless capsular fruits with two locules, two axile lenticular seeds, and two unbranched, plumose stigmas. The fossils' character combination only occurs today in some species of the Macaranga-Mallotus clade (MMC; Euphorbiaceae), a widespread Old-World understory group often thought to have tropical Asian origins. The associated leaves are consistent with extant Macaranga. Conclusions The new fossils are the oldest known for the MMC, demonstrating its Gondwanan history and marking its divergence by at least 52 Ma. This discovery makes an Asian origin of the MMC unlikely because immense oceanic distances separated Asia and South America 52 Ma. The only other MMC reproductive fossils so far known are also from the southern hemisphere (early Miocene, southern New Zealand), far from the Asian tropics. The MMC, along with many other Gondwanan survivors, most likely entered Asia during the Neogene Sahul-Sunda collision. Our discovery adds to a substantial series of well-dated, well-preserved fossils from one undersampled region, Patagonia, that have changed our understanding of plant biogeographic history. 

Abstract The Cretaceous–Paleogene (K/Pg) extinction appears to have been geographically heterogeneous for some organismal groups. Southern Hemisphere K/Pg palynological records have shown lower extinction and faster recovery than in the Northern Hemisphere, but no comparable, well-constrained Southern Hemisphere macrofloras spanning this interval had been available. Here, macrofloral turnover patterns are addressed for the first time in the Southern Hemisphere, using more than 3500 dicot leaves from the latest Cretaceous (Maastrichtian) and the earliest Paleocene (Danian) of Argentine Patagonia. A maximum ca. 90% macrofloral extinction and ca. 45% drop in rarefied species richness is estimated across the K/Pg, consistent with substantial species-level extinction and previously observed extirpation of host-specialized leaf mines. However, prior palynological and taxonomic studies indicate low turnover of higher taxa and persistence of general floral composition in the same sections. High species extinction, decreased species richness, and homogeneous Danian macrofloras across time and facies resemble patterns often observed in North America, but there are several notable differences. When compared with boundary-spanning macrofloras at similar absolute paleolatitudes (ca. 50°S or 50°N) from the Williston Basin (WB) in the Dakotas, both Maastrichtian and Danian Patagonian species richnesses are higher, extending a history of elevated South American diversity into the Maastrichtian. Despite high species turnover, our analyses also reveal continuity and expansion of leaf morphospace, including an increase in lobed and toothed species unlike the Danian WB. Thus, both Patagonian and WB K/Pg macrofloras support a significant extinction event, but they may also reflect geographically heterogeneous diversity, extinction, and recovery patterns warranting future study. 

Denk et al . agree that we reported the first fossil Fagaceae from the Southern Hemisphere. We appreciate their general enthusiasm for our findings, but we reject their critiques, which we find misleading and biased. The new fossils unequivocally belong to Castanopsis , and substantial evidence supports our Southern Route to Asia hypothesis.