Mónica R. Carvalho

Insect leaf-chewing damage tracks herbivore richness in modern and ancient forests.

The fossil record demonstrates that past climate changes and extinctions significantly affected the diversity of insect leaf-feeding damage, implying that the richness of damage types reflects that of the unsampled damage makers, and that the two are correlated through time. However, this relationship has not been quantified for living leaf-chewing insects, whose richness and mouthpart convergence have obscured their value for understanding past and present herbivore diversity. We hypothesized that the correlation of leaf-chewing damage types (DTs) and damage maker richness is directly observable in living forests. Using canopy access cranes at two lowland tropical rainforest sites in Panamá to survey 24 host-plant species, we found significant correlations between the numbers of leaf chewing insect species collected and the numbers of DTs observed to be made by the same species in feeding experiments, strongly supporting our hypothesis. Damage type richness was largely driven by insect species that make multiple DTs. Also, the rank-order abundances of DTs recorded at the Panamá sites and across a set of latest Cretaceous to middle Eocene fossil floras were highly correlated, indicating remarkable consistency of feeding-mode distributions through time. Most fossil and modern host-plant pairs displayed high similarity indices for their leaf-chewing DTs, but informative differences and trends in fossil damage composition became apparent when endophytic damage was included. Our results greatly expand the potential of insect-mediated leaf damage for interpreting insect herbivore richness and compositional heterogeneity from fossil floras and, equally promisingly, in living forests.

Phytogeographic implications of fossil endocarps of Menispermaceae from the Paleocene of Colombia

PREMISE OF THE STUDY Fossil leaves of Menispermaceae were previously described from the Paleocene of Colombia. Because of strong homoplasy of leaf characters, the fossils could not be placed more specifically within recognized clades, and additional data were needed to specify intrafamilial and paleogeographic relationships during the Paleocene. METHODS Fossil endocarps of Menispermaceae were collected from the Cerrejón Formation, the recently discovered Bogotá flora, and Wyoming (∼60 Ma). We surveyed the endocarp morphology of almost all extant genera, conducted character optimization, a molecular scaffold analysis, and critically reviewed the related fossil genera. KEY RESULTS Parallel syndromes of fruit characters have appeared in unrelated clades of the family according to current phylogenetic reconstructions. However, mapping selected endocarp characters across those clades that contain horseshoe-shaped endocarps facilitates identification and phylogenetic assessment of the fossils. Three fossil species are recognized. One of them belongs to the extant genus Stephania, which today grows only in Africa and Australasia. Palaeoluna gen. nov. is placed within the pantropical clade composed of extant Stephania, Cissampelos, and Cyclea; this morphogenus is also recognized from the Paleocene of Wyoming. Menispina gen. nov. shows similarity with several unrelated clades. CONCLUSIONS The new fossils from Colombia reveal a complex paleobiogeographic history of the recognized clades within Menispermaceae, suggesting a more active exchange among neotropical, paleotropical, North American, and European paleoforests than previously recognized. In addition, the new fossils indicate that neotropical forests were an important biome for the radiation and dispersal of derived lineages in Menispermaceae after the Cretaceous-Paleogene boundary.

First record of Todea (Osmundaceae) in South America, from the early Eocene paleorainforests of Laguna del Hunco (Patagonia, Argentina)

UNLABELLED PREMISE OF THE STUDY The early Eocene Laguna del Hunco caldera-lake paleoflora (ca. 52 Ma) from Chubut Province, Argentina, is notably diverse and includes many conifer and angiosperm lineages that are extinct in South America but extant in Australasian rainforests. No ferns have been previously described from Laguna del Hunco. We describe and interpret a new species of fossil Osmundaceae based on fertile and sterile pinnae. • METHODS The fossil specimens were compared with other extant and fossil Osmundaceae based on living and herbarium material and published descriptions. A morphological matrix based on 29 characters was constructed for 17 living species in Osmundaceae, four species assigned to the fossil genus Todites, and the new fossil species. Phylogenetic analyses were conducted under parsimony using morphology and total evidence matrices. • KEY RESULTS Both the new fossil and the Todites species were consistently resolved within the leptopteroid clade of Osmundaceae, and the new species resolved in a clade with the two living Todea species, which are now restricted to Australia, New Guinea, New Zealand, and southern Africa. • CONCLUSIONS Todea amissa sp. nov. is the first record of Todea, living or fossil, in South America and only the second fossil record worldwide. The distribution of extant Todea on Gondwanan continents other than South America is broadly shared with other taxa from Laguna del Hunco, further indicating that a large component of this flora represents a Gondwanic biome that is no longer found on the South American continent.

Eocene lantern fruits from Gondwanan Patagonia and the early origins of Solanaceae

Shedding light on fossil lantern fruit The Solanaceae (or nightshades) are one of the best-studied plant families, yet their evolutionary origins have thus far been relatively obscure. Corroborative fossil evidence of molecular phylogenetic divergence dates has been lacking. Wilf et al. present 52-million-year-old fossils of lantern fruits from Argentina, which they ascribe to the modern genus Physalis. These fossil finds suggest a much earlier origin of the lantern fruit lineage and indicate that the Solanaceae may have diversified before the final breakup of the Gondwanan supercontinent. Science, this issue p. 71 52-million-year-old fossils of lantern fruit demonstrate ancient Gondwanan history for the diverse nightshade family. The nightshade family Solanaceae holds exceptional economic and cultural importance. The early diversification of Solanaceae is thought to have occurred in South America during its separation from Gondwana, but the family’s sparse fossil record provides few insights. We report 52.2-million-year-old lantern fruits from terminal-Gondwanan Patagonia, featuring highly inflated, five-lobed calyces, as a newly identified species of the derived, diverse New World genus Physalis (e.g., groundcherries and tomatillos). The fossils are considerably older than corresponding molecular divergence dates and demonstrate an ancient history for the inflated calyx syndrome. The derived position of these early Eocene fossils shows that Solanaceae were well diversified long before final Gondwanan breakup.

The hydraulic architecture of Ginkgo leaves

PREMISE OF THE STUDY The hydraulics of xylem has been widely studied in numerous species and organ types. However, comparatively little is known about how phloem and xylem are hydraulically coupled or about many of the basic structural properties of phloem (such as conducting cell numbers and conductive areas), which nevertheless have direct bearing on understanding phloem loading and unloading. METHODS Using a combination of light, epifluorescence, confocal, and transmission electron microscopy, we quantified the hydraulic architecture of Ginkgo biloba leaf laminae and examined the scaling relationships between phloem and xylem in five fully mature leaves. KEY RESULTS The conductive areas and lengths of sieve cells and tracheids increase basipetally toward the petiole in a manner that is consistent with Münchs pressure flow hypothesis for phloem transport. This trend holds true for individual veins, the sum of conductive areas across all veins at any distance from the petiole, and for individual sieve cells and tracheids. Further, the conductive areas of phloem and xylem are isometrically correlated across the entire vasculature of the leaf lamina. The data for conducting cell areas do not conform with the predictions of the hydraulic models of da Vinci and Murray. CONCLUSIONS The scaling of Ginkgo lamina hydraulics complies with that observed in leaves of other gymnosperms and most angiosperms and is inconsistent with theoretical models that assume that the volume of transported incompressible fluids is conserved.

Spatiotemporal distribution of essential elements through Populus leaf ontogeny

Highlight Calcium, potassium, and zinc have a preferential spatiotemporal distribution throughout leaf development, and compartmentalization of calcium in poplar leaves enhances the evapotranspiration stream and phloem maturation.

A Late Cretaceous Piper (Piperaceae) from Colombia and diversification patterns for the genus

PREMISE OF THE STUDY Documented fossil floras in the neotropics are sparse, yet their records provide evidence on the spatial and temporal occurrence of taxa, allowing for testing of biogeographical and diversification scenarios on individual lineages. A new fossil Piper from the Late Cretaceous of Colombia is described here, and its importance for assessing diversification patterns in the genus is addressed. METHODS Leaf architecture of 32 fossil leaf compressions from the Guaduas Formation was compared with that of 294 extant angiosperm species. The phylogenetic position of the fossil named Piper margaritae sp. nov. was established based on leaf traits and a molecular scaffold of Piper. The age of the fossil was independently used as a calibration point for divergence time estimations. KEY RESULTS Natural affinities of P. margaritae to the Schilleria clade of Piper indicate that the genus occurred in tropical America by the Late Cretaceous. Estimates of age divergence and lineage accumulation reveal that most of the extant diversity of the genus accrued during the last ∼30 Myr. CONCLUSIONS The recent radiation of Piper is coeval with both the Andean uplift and the emergence of Central America, which have been proposed as important drivers of diversity. This pattern could exemplify a recurrent theme among many neotropical plant lineages.

Paleocene wind-dispersed fruits and seeds from Colombia and their implications for early Neotropical rainforests

Abstract Extant Neotropical rainforests are well known for their remarkable diversity of fruit and seed types. Biotic agents disperse most of these disseminules, whereas wind dispersal is less common. Although wind-dispersed fruits and seeds are greatly overshadowed in closed rainforests, many important families in the Neotropics (e.g., Bignoniaceae, Fabaceae, Malvaceae, Orchidaceae, Sapindaceae) show numerous morphological adaptations for anemochory (i.e. wings, accessory hairs). Most of these living groups have high to moderate levels of plant diversity in the upper levels of the canopy. Little is known about the fossil record of wind-dispersed fruits and seeds in the Neotropics. Six new species of disseminules with varied adaptations for wind dispersal are documented here. These fossils, representing extinct genera of Ulmaceae, Malvaceae, and some uncertain families, indicate that wind-dispersed fruit and seed syndromes were already common in the Neotropics by the Paleocene, coinciding with the early development of multistratal rainforests. Although the major families known to include most of the wind-dispersed disseminules in extant rainforests are still missing from the Paleogene fossil record of South and Central America, the new fossils imply that anemochory was a relatively important product and/or mechanism of plant evolution and diversification in early Neotropical rainforests.

Phloem networks in leaves

The survival of all vascular plants depends on phloem and xylem, which comprise a hydraulically coupled tissue system that transports photosynthates, water, and a variety of other molecules and ions. Although xylem hydraulics has been extensively studied, until recently, comparatively little is known quantitatively about the phloem hydraulic network and how it is functionally coupled to the xylem network, particularly in photosynthetic leaves. Here, we summarize recent advances in quantifying phloem hydraulics in fully expanded mature leaves with different vascular architectures and show that (1) the size of phloem conducting cells across phylogenetically different taxa scales isometrically with respect to xylem conducting cell size, (2) cell transport areas and lengths increase along phloem transport pathways in a manner that can be used to model Münchs pressure-flow hypothesis, and (3) report observations that invalidate da Vincis and Murrays hydraulic models as plausible constructs for understanding photosynthate transport in the leaf lamina.