Degradation of the upper pulvinus in modern and fossil leaves of Cercis (Fabaceae)

Abstract
Identification of fossil leaf impressions as Cercis has been questioned based upon the presence or absence of a pulvinus at the base of the lamina (upper pulvinus). In the present study, leaves of Cercis canadensis were examined before and after abscission to explore the degradation processes that could occur prior to fossilization, and the North American record for fossil foliage of Cercis was revised accordingly. Results for C. canadensis indicate that: (1) the pulvinus consists largely of tissues with nonlignified cells (a wide cortex, a nonlignified fiber sheath, phloem, and pith) that degrade rapidly after leaf abscission, (2) the lignified xylem tissue that remains in the pulvinus after degradation is in brittle strands, (3) the pulvinus degrades at a faster rate than the lamina or the petiole, and (4) the degraded pulvinus cushion leaves a semicircular pattern on the lamina. From examination of fossils as well as extant species, we: (1) demonstrated that in fossils, the upper pulvinus can show a greater degree of degradation than the adjoining petiole or lamina tissue, suggesting the degradation of upper pulvinus tissue is similar in modern vs. fossil specimens, (2) defined numerous other laminar characters that can be used in conjunction with, or in the absence of, an upper pulvinus to confirm the presence of Cercis in the fossil record, and (3) showed from those criteria that the earliest known North American fossil leaf record for Cercis, from a specimen newly reported in the present study, is from the middle Miocene Succor Creek flora of Oregon.