Atmospheric carbon dioxide is the raw material for the biosphere. Therefore, changes in the carbon isotopic composition of the atmosphere will influence the terrestrial δ13C signals we interpret. However, reconstructing the atmospheric δ13C value in the geologic past has proven challenging. Land plants sample the isotopic composition of CO2 during photosynthesis. We use a model of carbon isotopic fractionation during C3 photosynthesis, in combination with a meta–data set (519 measurements from 176 species), to show that the δ13C value of atmospheric CO2 can be reconstructed from the isotopic composition of plant tissue. Over a range of pCO2 (198–1300 ppmv), the δ13C value of plant tissue does not vary systematically with atmospheric carbon dioxide concentration. However, environmental factors, such as water stress, can influence the δ13C value of leaf tissue. These factors explained a relatively small portion of variation in the δ13C value of plant tissue in our data set and emerged strongly only when the carbon isotopic composition of the atmosphere was held constant. Members of the Poaceae differed in average δ13C value, but we observed no other differences correlated with plant life form (herbs, trees, shrubs). In contrast, over 90% of the variation the carbon isotopic composition of plant tissue was explained by variation in the δ13C value of the atmosphere under which it was fixed. We use a subset of our data spanning a geologically reasonable range of atmospheric δ13C values (−6.4‰ to −9.6‰) and excluding C3 Poaceae to develop an equation to reconstruct the δ13C value of atmospheric CO2 based on plant values. Reconstructing the δ13C value of atmospheric CO2 in geologic time will facilitate chemostratigraphic correlation in terrestrial sediments, calibrate pCO2 reconstructions based on soil carbonates offer a window into the physiology of ancient plants.

Better understanding of the functional biology of early angiosperms may clarify eco logical factors surrounding their origin and early radiation. Phylogenetic studies identify Amborella, Nymphaeales (water lilies), Austrobaileyales, and Chloranthaceae as extant lineages that branched before the radiation of core angiosperms. Among living plants, these lineages may rep resent the best models for the ecology and physiology of early angiosperms. Here we combine phylogenetic reconstruction with new data on the morphology and ecophysiology of these plants to infer early angiosperm function. With few exceptions, Amborella, Austrobaileyales, and Chlor anthaceae share ecophysiological traits associated with shady, disturbed, and wet habitats. These features include low and easily light-saturated photosynthetic rates, leaf anatomy related to the capture of understory light, small seed size, and clonal reproduction. Some Chloranthaceae, how ever, possess higher photosynthetic capacities and seedlings that recruit in canopy gaps and other sunny, disturbed habitats, which may have allowed Cretaceous Chloranthaceae to expand into more diverse environments. In contrast, water lilies possess ecophysiological features linked to aquatic, sunny habitats, such as absence of a vascular cambium, ventilating stems and roots, and floating leaves tuned for high photosynthetic rates in full sun. Nymphaeales may represent an early radiation into such aquatic environments. We hypothesize that the earliest angiosperms were woody plants that grew in dimly lit, disturbed forest understory habitats and/or shady streamside settings. This ecology may have restricted the diversity of pre-Aptian angiosperms and living basal lineages. The vegetative flexibility that evolved in the understory, however, may have been a key factor in their diversification in other habitats. Our inferences based on living plants are consistent with many aspects of the Early Cretaceous fossil record and can be tested with further study of the anatomy, chemistry, and sedimentological context of Early Cretaceous angiosperm fossils.

Reconstruction of changing C isotopic composition of Early Cretaceous atmospheric CO 2 from fossilized C3 vascular land-plant tissue revealed a brief and striking negative excursion (⌬ ഠ ؊5‰) in atmospheric ␦ 13 C, followed by a rapid positive compensation (⌬ ഠ ؉5‰) during the Aptian (ca. 117 Ma). Mass-balance calculations show that dissociation of a small amount of methane gas hydrate is the most tenable cause of the negative excursion; this would also result in an increased CO 2 :O 2 mixing ratio as O 2 is consumed during CH 4 oxidation to CO 2 , spurring the exponential phase of angiosperm biogeographic expansion.

Contents Summary 1 Introduction 2 Previous images of early angiosperms and their habitats 3 Progress in understanding angiosperm phylogeny: extant ‘basal’ relations 4 Early angiosperm ecology: inferences from extant plants and reflections from the fossil record 5 The ecology of angiosperm diversification: gaining a roothold and subsequent diversification 15 The environmental context of early angiosperm evolution 17 Conclusions 20 Acknowledgements 20 References 20 Summary The flowering plants – angiosperms – appeared during the Early Cretaceous period and within 10–30 Myr dominated the species composition of many floras worldwide. Emerging insights into the phylogenetics of development and discoveries of early angiosperm fossils are shedding increased light on the patterns and processes of early angiosperm evolution. However, we also need to integrate ecology, in particular how early angiosperms established a roothold in pre‐existing Mesozoic plant communities. These events were critical in guiding subsequent waves of angiosperm diversification during the Aptian–Albian. Previous pictures of the early flowering plant ecology have been diverse, ranging from large tropical rainforest trees, weedy drought‐adapted and colonizing shrubs, disturbance‐ and sun‐loving rhizomatous herbs, and, more recently, aquatic herbs; however, none of these images were tethered to a robust hypothesis of angiosperm phylogeny. Here, we synthesize our current understanding of early angiosperm ecology, focusing on patterns of functional ecology, by merging recent molecular phylogenetic studies and functional studies on extant ‘basal angiosperms’ with the picture of early angiosperm evolution drawn by the fossil record.