Hugo I. Martínez-Cabrera

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.

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.

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.

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 clades 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.

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

UNLABELLED 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.

Functional Traits in Parallel Evolutionary Radiations and Trait-Environment Associations in the Cape Floristic Region of South Africa

Evolutionary radiations with extreme levels of diversity present a unique opportunity to study the role of the environment in plant evolution. If environmental adaptation played an important role in such radiations, we expect to find associations between functional traits and key climatic variables. Similar trait-environment associations across clades may reflect common responses, while contradictory associations may suggest lineage-specific adaptations. Here, we explore trait-environment relationships in two evolutionary radiations in the fynbos biome of the highly biodiverse Cape Floristic Region (CFR) of South Africa. Protea and Pelargonium are morphologically and evolutionarily diverse genera that typify the CFR yet are substantially different in growth form and morphology. Our analytical approach employs a Bayesian multiple-response generalized linear mixed-effects model, taking into account covariation among traits and controlling for phylogenetic relationships. Of the pairwise trait-environment associations tested, 6 out of 24 were in the same direction and 2 out of 24 were in opposite directions, with the latter apparently reflecting alternative life-history strategies. These findings demonstrate that trait diversity within two plant lineages may reflect both parallel and idiosyncratic responses to the environment, rather than all taxa conforming to a global-scale pattern. Such insights are essential for understanding how trait-environment associations arise and how they influence species diversification.

A New Late Cretaceous (Coniacian-Maastrichtian) Javelinoxylon Wood From Chihuahua, Mexico

We describe a new fossil wood from the San Carlos Formation (Coniacian- Maastrichtian) in Chihuahua, northern Mexico. This Malvaceae s.l. wood is diffuse porous, vessels are solitary and in radial multiples, simple perforation plates, small alternate intervessel pits, vessel-ray parenchyma pits that are rounded with reduced borders, septate and nonseptate fibers, homocellular and heterocellular rays, and storied rays and vessel elements. These features support its inclusion within the genus Javelinoxylon, Malvaceae s.l., which occurs in other Upper Cretaceous localities in northern Mexico (Olmos Formation) and Texas (Aguja and Javelina Formations). This San Carlos fossil wood is the earliest occurrence of storied structure in the fossil record and the earliest angiosperm record for the State of Chihuahua, Mexico.

A NEW SPECIES OF TAPIRIRA (ANACARDIACEAE) FROM EARLY MIOCENE SEDIMENTS OF THE EL CIEN FORMATION, BAJA CALIFORNIA SUR, MEXICO

From the early Miocene El Cien Formation, Baja California Sur, Mexico, a new Tapirira species is described. Among the diagnostic features that relate T. peninsularis Martinez-Cabrera & Cevallos-Ferriz sp. nov. with extant species of the genus are libriform septate fibers, radial canals with 2–4 layers of epithelial cells, and scanty paratracheal to vasicentric axial parenchyma. In order to establish similarity between wood of T. peninsularis and fossil and extant species of the genus, cluster and principal component analyses were carried out. Numerical analyses support that Tapirira peninsularis is distinct from other extant and fossil species of the genus. Quantitative characters like frequency of radial canals, diameter of fibres, vessel element lumen, and height and width of the rays with radial canals are important in distinguishing between these taxa. Though these wood characters are quantitative, they also have been useful in the recognition of two subgenera within the genus. Recognition of this new species, along with the fossil record of the group, and a phytogeographic analysis suggest a low latitude North American origin for the genus, and a subsequent introduction to southern latitudes.

Ruprechtia in the Miocene El Cien Formation, Baja California Sur, Mexico

Fossil woods from the El Cien Formation have yielded important information on the taxonomic composition and climate of a flora established in the west coast of Mexico during the Miocene. This report of a new genus and species, Ruprechtioxylon multiseptatus Cevallos-Ferriz, Martinez Cabrera et Calvillo-Canadell, is based on woods with the following combination of features: vessels solitary and in radial multiples of 2–3; vestured, alternate, oval to polygonal intervessel pits; vessel-ray and vessel-parenchyma pits similar in size to intervessel pits, but with slightly reduced to reduced borders; 2–5 septa per fibre; scanty paratracheal, unilateral and vasicentric axial parenchyma; uniseriate homocellular rays, occasionally locally biseriate; crystals in fibres. The presence of Ruprechtioxylon (Polygonaceae) in the El Cien Formation confirms that plants of lineages growing today under contrasting climates lived together in the past. This record adds a new species to the growing list of Neotropical taxa that were present in Mexico prior to the great Plio-Pleistocene exchange of biota in the Americas.

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 Mexicos 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 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 , 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.