Late Campanian fossil of a legume fruit supports Mexico as a center of Fabaceae radiation.

Fabaceae is one of the most diverse angiosperm families and is distributed across the globe in a variety of environments. The earliest evidence of the family, previous to this work, was from Paleogene sediments where it was found to be diverse in many fossil assemblages around the world. Here, we describe a fossil legume fruit from the Olmos Formation (upper Campanian) in northern Mexico. We designated the fossil fruit as Leguminocarpum olmensis Centeno-González, Martínez-Cabrera, Porras-Múzquiz et Estrada-Ruiz sp. nov., and related it with the Fabaceae family based on the presence of a dehiscent pod with two valves, an apex bearing stylar base, short stipe, and reticulated veins in the pericarp. We propose a new fossil species of Leguminocarpum for this fossil fruit. This fossil provides critical information on the long geologic history of Leguminosae around the world, significantly extending the record into the Cretaceous of Mexico.

Vessel diameter is related to amount and spatial arrangement of axial parenchyma in woody angiosperms.

Parenchyma represents a critically important living tissue in the sapwood of the secondary xylem of woody angiosperms. Considering various interactions between parenchyma and water transporting vessels, we hypothesize a structure-function relationship between both cell types. Through a generalized additive mixed model approach based on 2,332 woody angiosperm species derived from the literature, we explored the relationship between the proportion and spatial distribution of ray and axial parenchyma and vessel size, while controlling for maximum plant height and a range of climatic factors. When factoring in maximum plant height, we found that with increasing mean annual temperatures, mean vessel diameter showed a positive correlation with axial parenchyma proportion and arrangement, but not for ray parenchyma. Species with a high axial parenchyma tissue fraction tend to have wide vessels, with most of the parenchyma packed around vessels, whereas species with small diameter vessels show a reduced amount of axial parenchyma that is not directly connected to vessels. This finding provides evidence for independent functions of axial parenchyma and ray parenchyma in large vesselled species and further supports a strong role for axial parenchyma in long-distance xylem water transport.

A global analysis of parenchyma tissue fractions in secondary xylem of seed plants.

Parenchyma is an important tissue in secondary xylem of seed plants, with functions ranging from storage to defence and with effects on the physical and mechanical properties of wood. Currently, we lack a large-scale quantitative analysis of ray parenchyma (RP) and axial parenchyma (AP) tissue fractions. Here, we use data from the literature on AP and RP fractions to investigate the potential relationships of climate and growth form with total ray and axial parenchyma fractions (RAP). We found a 29-fold variation in RAP fraction, which was more strongly related to temperature than with precipitation. Stem succulents had the highest RAP values (mean ± SD: 70.2 ± 22.0%), followed by lianas (50.1 ± 16.3%), angiosperm trees and shrubs (26.3 ± 12.4%), and conifers (7.6 ± 2.6%). Differences in RAP fraction between temperate and tropical angiosperm trees (21.1 ± 7.9% vs 36.2 ± 13.4%, respectively) are due to differences in the AP fraction, which is typically three times higher in tropical than in temperate trees, but not in RP fraction. Our results illustrate that both temperature and growth form are important drivers of RAP fractions. These findings should help pave the way to better understand the various functions of RAP in plants.

Wood anatomy reveals high theoretical hydraulic conductivity and low resistance to vessel implosion in a Cretaceous fossil forest from northern Mexico.

The Olmos Formation (upper Campanian), with over 60 angiosperm leaf morphotypes, is Mexico's richest Cretaceous flora. Paleoclimate leaf physiognomy estimates indicate that the Olmos paleoforest grew under wet and warm conditions, similar to those present in modern tropical rainforests. Leaf surface area, tree size and climate reconstructions suggest that this was a highly productive system. Efficient carbon fixation requires hydraulic efficiency to meet the evaporative demands of the photosynthetic surface, but it comes at the expense of increased risk of drought-induced cavitation. Here we tested the hypothesis that the Olmos paleoforest had high hydraulic efficiency, but was prone to cavitation. We characterized the hydraulic properties of the Olmos paleoforest using theoretical conductivity (Ks), vessel composition (S) and vessel fraction (F), and measured drought resistance using vessel implosion resistance (t/b)h(2) and the water potential at which there is 50% loss of hydraulic conductivity (P50). We found that the Olmos paleoforest had high hydraulic efficiency, similar to that present in several extant tropical-wet or semi-deciduous forest communities. Remarkably, the fossil flora had the lowest (t/b)h(2), which, together with low median P50 (-1.9 MPa), indicate that the Olmos paleoforest species were extremely vulnerable to drought-induced cavitation. Our findings support paleoclimate inferences from leaf physiognomy and paleoclimatic models suggesting it represented a highly productive wet tropical rainforest. Our results also indicate that the Olmos Formation plants had a large range of water conduction strategies, but more restricted variation in cavitation resistance. These straightforward methods for measuring hydraulic properties, used herein for the first time, can provide useful information on the ecological strategies of paleofloras and on temporal shifts in ecological function of fossil forests chronosequences.

Shifts in climate foster exceptional opportunities for species radiation: the case of South african geraniums.

Climate change is often assumed to be a major driver of biodiversity loss. However, it can also set the stage for novel diversification in lineages with the evolutionary ability to colonize new environments. Here we tested if the extraordinary evolutionary success of the genus Pelargonium was related to the ability of its species to capitalize on the climate niche variation produced by the historical changes in southern Africa. We evaluated the relationship between rates of climate niche evolution and diversification rates in the main Pelargonium lineages and disentangled the roles of deep and recent historical events in the modification of species niches. Pelargonium clades exhibiting higher ecological differentiation along summer precipitation (SPP) gradients also experienced higher diversification rates. Faster rates of niche differentiation in spatially structured variables, along with lower levels of niche overlap among closely related species, suggest recent modification in species niches (e.g. dispersal or range shift) and niche lability. We suggest that highly structured SPP gradients established during the aridification process within southern Africa, in concert with niche lability and low niche overlap, contributed to species divergence. These factors are likely to be responsible for the extensive diversification of other lineages in this diversity hot spot.

Wood anatomical correlates with theoretical conductivity and wood density across China: evolutionary evidence of the functional differentiation of axial and radial parenchyma.

BACKGROUND AND AIMS: In recent years considerable effort has focused on linking wood anatomy and key ecological traits. Studies analysing large databases have described how these ecological traits vary as a function of wood anatomical traits related to conduction and support, but have not considered how these functions interact with cells involved in storage of water and carbohydrates (i.e. parenchyma cells). METHODS: We analyzed, in a phylogenetic context, the functional relationship between cell types performing each of the three xylem functions (conduction, support and storage) and wood density and theoretical conductivity using a sample of approx. 800 tree species from China. KEY RESULTS: Axial parenchyma and rays had distinct evolutionary correlation patterns. An evolutionary link was found between high conduction capacity and larger amounts of axial parenchyma that is probably related to water storage capacity and embolism repair, while larger amounts of ray tissue have evolved with increased mechanical support and reduced hydraulic capacity. In a phylogenetic principal component analysis this association of axial parenchyma with increased conduction capacity and rays with wood density represented orthogonal axes of variation. In multivariate space, however, the proportion of rays might be positively associated with conductance and negatively with wood density, indicating flexibility in these axes in species with wide rays. CONCLUSIONS: The findings suggest that parenchyma types may differ in function. The functional axes represented by different cell types were conserved across lineages, suggesting a significant role in the ecological strategies of the angiosperms.

Phylogenetic influences on leaf trait integration in Pelargonium (Geraniaceae): convergence, divergence, and historical adaptation to a rapidly changing climate.

PREMISE OF THE STUDY: Trait integration may improve prediction of species and lineage responses to future climate change more than individual traits alone, particularly when analyses incorporate effects of phylogenetic relationships. The South African genus Pelargonium contains divergent major clades that have radiated along the same seasonal aridity gradient, presenting the opportunity to ask whether patterns of evolution in mean leaf trait values are achieved through the same set of coordinated changes among traits in each clade. METHODS: Seven leaf traits were measured on field-collected leaves from one-third of the species (98) of the genus. Trait relationships were examined using phylogenetic regression within major clades. Disparity analysis determined whether the course of trait evolution paralleled historical climate change events. KEY RESULTS: Divergence in mean trait values between sister clades A1 and A2 was consistent with expectations for leaves differing in longevity, despite strong similarity between clades in trait interactions. No traits in either clade exhibited significant relationships with multivariate climate axes, with one exception. Species in clades C and A2 included in this study occupied similar environments. These clades had similar values of individual trait means, except for δ(13)C, but they exhibited distinctive patterns of trait integration. CONCLUSIONS: Differing present-day patterns of trait integration are consistent with interpretations of adaptive responses to the prevailing climate at the time of each clade's origin. These differing patterns of integration are likely to exert strong effects on clade-level responses to future climate change in the winter rainfall region of South Africa.

Low levels of climate niche conservatism may explain clade diversity patterns in the South African genus Pelargonium (Geraniaceae).

PREMISE OF THE STUDY: Sharp climatic gradients in South Africa and in particular in the Cape Floristic Region (CFR) provide a diversity of niches over short distances that may have promoted ecological diversification in local clades. Here we measured the extent to which closely related species occupy divergent climates and test whether niche lability is correlated with higher species diversity in the genus. METHOD: We integrated phylogenetic information and environmental niche models (ENM) to assess the levels of climate niche conservatism. ENMs for 113 species of Pelargonium were calculated using maximum entropy. We used two tests, one assessing climate niche equivalency and the other testing niche similarity between sister species and within sections. We also examined whether niche similarity was correlated with phylogenetic relatedness across the genus. KEY RESULTS: Niche similarity was mostly independent of phylogenetic relationships. Compared to random expectations, 23% of closely related species pairs had climate niches that were more similar, and only 6.5% were more disparate; the remaining 70% of comparisons had similarities that fell within random expectations. Similar trends were observed when analyses were restricted to only sister species pairs. Although the overall proportion of niche divergence was low, this was significantly related to sectional diversity. We also found a negative relationship between diversity and the proportion of random niches. CONCLUSIONS: Lack of widespread niche conservatism in a highly heterogeneous landscape and few instances of significant climate niche lability suggest that an adaptive divergence process was implicated in the Pelargonium radiation.

Integration of vessel traits, wood density, and height in angiosperm shrubs and trees.

PREMISE OF THE STUDY: Trees and shrubs tend to occupy different niches within and across ecosystems; therefore, traits related to their resource use and life history are expected to differ. Here we analyzed how growth form is related to variation in integration among vessel traits, wood density, and height. We also considered the ecological and evolutionary consequences of such differences. METHOD: In a sample of 200 woody plant species (65 shrubs and 135 trees) from Argentina, Mexico, and the United States, standardized major axis (SMA) regression, correlation analyses, and ANOVA were used to determine whether relationships among traits differed between growth forms. The influence of phylogenetic relationships was examined with a phylogenetic ANOVA and phylogenetically independent contrasts (PICs). A principal component analysis was conducted to determine whether trees and shrubs occupy different portions of multivariate trait space. KEY RESULTS: Wood density did not differ between shrubs and trees, but there were significant differences in vessel diameter, vessel density, theoretical conductivity, and as expected, height. In addition, relationships between vessel traits and wood density differed between growth forms. Trees showed coordination among vessel traits, wood density, and height, but in shrubs, wood density and vessel traits were independent. These results hold when phylogenetic relationships were considered. In the multivariate analyses, these differences translated as significantly different positions in multivariate trait space occupied by shrubs and trees. CONCLUSIONS: Differences in trait integration between growth forms suggest that evolution of growth form in some lineages might be associated with the degree of trait interrelation.

Upper Cretaceous woods from the Olmos Formation (late Campanian-early Maastrichtian), Coahuila, Mexico.

PREMISE OF THE STUDY: The Olmos Formation was part of a system of deltas that existed in the southern portion of the Western Interior of North America during the Campanian-Maastrichtian. The paleofloristic composition from the northern portions of the Epicontinental Sea is relatively well known, but less intensive exploration in the south has precluded more detailed floristic comparison across the entire latitudinal span of the Sea. The Olmos Formation flora, with more than 100 different leaf morphotypes so far recognized and several wood types, has the most diverse Cretaceous fossil plant assemblage in Mexico and represents a valuable opportunity for comparative studies. • METHODS: The fossil woods here described were collected in the Coahuila State, Mexico. The samples were studied using standard thin section technique and identified by comparison with fossil and extant material. • KEY RESULTS: We described four new genera (Olmosoxylon, cf. Lauraceae; Coahuiloxylon, ?Anacardiaceae, ?Burseraceae; Muzquizoxylon, Cornaceae; and Wheeleroxylon, Malvaceae s.l.) and three xylotypes of angiosperms. • CONCLUSIONS: Some of the genera present in the Olmos Formation such as Javelinoxylon and Metcalfeoxylon have been described from geologic units in the USA (San Juan Basin, New Mexico and Big Bend National Park, Texas), suggesting similarity in the taxonomic composition of the floras that inhabited southern portions of the western margin of the Campanian-Maastrichtian Epicontinental Sea. Other species, however, have only been reported for the Olmos Formation, indicating some degree of local floristic differentiation among the assemblages that inhabited the southern portion of the Western Interior.

Wood anatomy and wood density in shrubs: Responses to varying aridity along transcontinental transects.

Wood density plays a key role in ecological strategies and life history variation in woody plants, but little is known about its anatomical basis in shrubs. We quantified the relationships between wood density, anatomy, and climate in 61 shrub species from eight field sites along latitudinal belts between 31° and 35° in North and South America. Measurements included cell dimensions, transverse areas of each xylem cell type and percentage contact between different cell types and vessels. Wood density was more significantly correlated with precipitation and aridity than with temperature. High wood density was achieved through reductions in cell size and increases in the proportion of wall relative to lumen. Wood density was independent of vessel traits, suggesting that this trait does not impose conduction limitations in shrubs. The proportion of fibers in direct contact with vessels decreased with and was independent of wood density, indicating that the number of fiber-vessel contacts does not explain the previously observed correlation between wood density and implosion resistance. Axial and radial parenchyma each had a significant but opposite association with wood density. Fiber size and wall thickness link wood density, life history, and ecological strategies by controlling the proportion of carbon invested per unit stem volume.