Climbing since the early Miocene: The fossil record of Paullinieae (Sapindaceae).

Paullinieae are a diverse group of tropical and subtropical climbing plants that belong to the soapberry family (Sapindaceae). The six genera in this tribe make up approximately one-quarter of the species in the family, but a sparse fossil record limits our understanding of their diversification. Here, we provide the first description of anatomically preserved fossils of Paullinieae and we re-evaluate other macrofossils that have been attributed to the tribe. We identified permineralized fossil roots in collections from the lower Miocene Cucaracha Formation where it was exposed along the Culebra Cut of the Panama Canal. We prepared the fossils using the cellulose acetate peel technique and compared the anatomy with that of extant Paullinieae. The fossil roots preserve a combination of characters found only in Paullinieae, including peripheral secondary vascular strands, vessel dimorphism, alternate intervessel pitting with coalescent apertures, heterocellular rays, and axial parenchyma strands of 2-4 cells, often with prismatic crystals. We also searched the paleontological literature for other occurrences of the tribe. We re-evaluated leaf fossils from western North America that have been assigned to extant genera in the tribe by comparing their morphology to herbarium specimens and cleared leaves. The fossil leaves that were assigned to Cardiospermum and Serjania from the Paleogene of western North America are likely Sapindaceae; however, they lack diagnostic characters necessary for inclusion in Paullinieae and should be excluded from those genera. Therefore, the fossils described here as Ampelorhiza heteroxylon gen. et sp. nov. are the oldest macrofossil evidence of Paullinieae. They provide direct evidence of the development of a vascular cambial variant associated with the climbing habit in Sapindaceae and provide strong evidence of the diversification of crown-group Paullinieae in the tropics by 18.5-19 million years ago.

Effects of dry-season irrigation on leaf physiology and biomass allocation in tropical lianas and trees.

Lianas are more abundant in seasonal forests than in wetter forests and are thought to perform better than trees when light is abundant and water is limited. We tested the hypothesis that lianas perform better than trees during seasonal drought using a common garden experiment with 12 taxonomically diverse species (six liana and six tree species) in 12 replicated plots. We irrigated six of the plots during the dry season for four years, while the remaining six control plots received only ambient rainfall. In year 5, we measured stem diameters for all individuals and harvested above- and belowground biomass for a subset of individuals to quantify absolute growth and biomass allocation to roots, stems, and leaves, as well as total root length and maximum rooting depth. We also measured rate of photosynthesis, intrinsic water use efficiency (iWUE), pre-dawn and midday water potential, and a set of functional and hydraulic traits. During the peak of the dry season, lianas in control plots had 54% higher predawn leaf water potentials (ΨPD ), and 45% higher photosynthetic rates than trees in control plots. By contrast, during the peak of the wet season, these physiological differences between lianas and trees become less pronounced and, in some cases, even disappeared. Trees had higher specific leaf area (SLA) than lianas; however, no other functional trait differed between growth forms. Trees responded to the irrigation treatment with 15% larger diameters and 119% greater biomass than trees in control plots. Liana growth, however, did not respond to irrigation; liana diameter and biomass were similar in control and irrigation plots, suggesting that lianas were far less limited by soil moisture than were trees. Contrary to previous hypotheses, lianas did not have deeper roots than trees; however, lianas had longer roots per stem diameter than did trees. Our results support the hypothesis that lianas perform better and experience less physiological stress than trees during seasonal drought, suggesting clear differences between growth forms in response to altered rainfall regimes. Ultimately, better dry-season performance may explain why liana abundance peaks in seasonal forests compared to trees, which peak in abundance in less seasonal, wetter forests.