Oldest fossil flowers of hamamelidaceous affinity, from the Late Cretaceous of New Jersey.

Exceptionally well-preserved staminate inflorescences, pistillate inflorescences, and detached stamens with important phylogenetic and paleoecological implications have been discovered from the Turonian (ca. 88.5-90.4 million years B.P.) Raritan Formation of New Jersey. The fossils have a combination of floral and pollen characters found in various genera of modern entomophilous and anemophilous Hamamelidaceae and anemophilous Platanus (Platanaceae). The floral characters of the fossils, including a sepal cup, staminal tube, and apparently nectariferous staminodes, indicate that this taxon was probably insect pollinated. The juxtaposition of character complexes in an extinct taxon from disparate modern taxa provides an interesting phylogenetic perspective on the origins of Hamamelidaceae and is a striking example of a fossil that is a mosaic of familial level characters relative to modern taxa. Of even broader interest, however, is the occurrence of staminodal nectaries that have structural characters intermediate between the fossil's functional stamens and modern hamamelidaceous petals. This transitional staminode morphology in the context of the other fossil characters suggests a staminodal origin of petals in the hamamelid-rosid lineage. This hypothesis is supported by the apparent staminode position within the fossil flowers where petals are found in modern genera. The character complex of morphologically transitional staminodes, a staminal tube, and sepal cup can be viewed as prehypanthial, lacking only fusion of the staminal tube to the sepal cup. The appearance of the character complex embodied in these flowers during the late mid-Cretaceous may signal the early stages of the relationship between specialized pollinators, such as bees, and the hamamelid-rosid-asterid lineage of angiosperms, arguably one of the most important events in angiosperm radiation.