Early and Mid-Cretaceous Aristolochiaceous Seeds from Portugal and Eastern North America

Premise of research. The fossil record of Aristolochiaceae (Piperales, magnoliids) is sparse, particularly from Cretaceous strata. Fossil seeds from the Early and mid-Cretaceous of Portugal and North America provide the earliest unequivocal documentation of the group. Methodology. Detailed morphological and anatomical investigations of the fossil aristolochiaceous seeds were carried out using SEM and synchrotron radiation X-ray tomographic microscopy (SRXTM). Comparison with other seeds, extinct as well as extant, was based on published data and SRXTM analyses. Pivotal results. Two new genera and two new species, Aristospermum huberi and Siratospermum mauldinense, are described based on fossil seeds from the Early and mid-Cretaceous of Portugal and North America. The seeds are anatropous and bitegmic, with the micropyle formed by the inner integument. The testa consists of an exotesta of varied thickness and an endotesta of crystalliferous cells. The tegmen is three cell layers thick and consists of an outermost layer of longitudinally aligned fibers, a middle layer of transversely aligned fibers perpendicular to the longitudinal fibers, and an inner layer of thin-walled cuboidal cells. In most seeds the exotesta is abraded, exposing the crystalliferous cells of the endotesta. Among extant plants, a similar seed coat with a crystalliferous endotesta and crossing fibers in the tegmen is unique to Aristolochiaceae. Conclusion. The unique seed coat allows Aristospermum and Siratospermum to be assigned confidently to the lineage that today includes extant Aristolochiaceae. Aristospermum and Siratospermum provide the first unequivocal documentation of the Aristolochiaceae lineage of the Piperales during the Early Cretaceous diversification of angiosperms.We thank the Paul Scherrer Institut, Villigen, Switzerland, for provision of synchrotron radiation beam time at the TOMCAT Beamline X02DA of the Swiss Light Source. We are also grateful to Federica Marone for her help at the Beamline (projects 20141047, 20190071 to P. C. J. Donoghue, S. Bengtson, andE. M. Friis) and Anna Lindström for help with the SRXTM analyses. Support for this researchwas also obtained from the Swedish Research Council (2014-5228 to E.M. Friis) and from the Oak SpringGarden Foundation.