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.

A liana from the lower Miocene of Panama and the fossil record of Connaraceae.

PREMISE OF THE STUDY: Permineralized wood is common in the Miocene beds exposed during the expansion of the Panama Canal. We describe a stem with the distinctive anatomy of a liana and evaluate the evolutionary, biogeographic, and ecological significance of this discovery. METHODS: The object of the study was obtained from a collection of fossil woods and fruits from a locality in the lower Miocene Cucaracha Formation, where the formation is exposed by the Culebra Cut of the Panama Canal. Thin sections were prepared using the cellulose acetate peel technique and examined using transmitted light microscopy. We described the anatomy and compared it with that of extant and fossil species. We also reviewed and evaluated published reports of fossils attributed to Connaraceae. KEY RESULTS: The anatomy of this fossil wood matches the genus Rourea (Connaraceae). The stem is only 1 cm in diameter, but vessels >200 µm in diameter also occur, indicating the perennial climbing habit. We evaluated 12 other pre-Quaternary occurrences attributed to Connaraceae. Four are accepted, three are rejected, and we consider five unknown or uncertain. CONCLUSIONS: The discovery of this Rourea stem confirms the presence of Connaraceae in the Neotropics by the early Miocene, provides the oldest evidence of the climbing habit in the family, and contributes to our understanding of the flora of Panama 19 mya. Although the fossil record of Connaraceae is sparse, reliable occurrences span three continents and indicate that the family originated as early as the Late Cretaceous-Paleocene and was widespread by the early Miocene.

Fruits and wood of Parinari from the early Miocene of Panama and the fossil record of Chrysobalanaceae.

PREMISE OF THE STUDY: Chrysobalanaceae are woody plants with over 500 species in 20 genera. They are among the most common trees in tropical forests, but a sparse fossil record has limited our ability to test evolutionary and biogeographic hypotheses, and several previous reports of Chrysobalanaceae megafossils are doubtful. METHODS: We prepared fossil endocarps and wood collected from the lower Miocene beds along the Panama Canal using the cellulose acetate peel technique and examined them using light microscopy. We compared the fossil endocarps with previously published fossils and with fruits from herbarium specimens. We compared the fossil wood with photographs and descriptions of extant species. KEY RESULTS: Parinari endocarps can be distinguished from other genera within Chrysobalanaceae by a suite of features, i.e., thick wall, a secondary septum, seminal cavities lined with dense, woolly trichomes, and two ovate to lingulate basal germination plugs. Fossil endocarps from the Cucaracha, Culebra, and La Boca Formations confirm that Parinari was present in the neotropics by the early Miocene. CONCLUSIONS: The earliest unequivocal evidence of crown-group Chrysobalanaceae is late Oligocene-early Miocene, and the genus Parinari was distinct by at least 19 million years ago. Parinari and other Chrysobalanaceae likely reached the neotropics via long-distance dispersal rather than vicariance. The presence of Parinari in the Cucaracha flora supports the interpretation of a riparian, moist tropical forest environment. Parinari was probably a canopy-dominant tree in the Cucaracha forest and took advantage of the local megafauna for seed dispersal.