Pedro Luis Valverde

EVOLUTION OF MIXED STRATEGIES OF PLANT DEFENSE ALLOCATION AGAINST NATURAL ENEMIES

Abstract –In this study we present a simple optimization model for the evolution of defensive strategies (tolerance and resistance) of plants against their natural enemies. The model specifically evaluates the consequences of introducing variable costs and benefits of tolerance and resistance and nonlinear cost‐and‐benefit functions for tolerance and resistance. Incorporating these assumptions, the present model of plant defense predicts different evolutionary scenarios, not expected by previous work. Basically, the presence of an adaptive peak corresponding to intermediate levels of allocation to tolerance and resistance can arise when the shape parameter of the cost function is higher than the corresponding of the benefit function. The presence of two alternatives peaks of maximum tolerance and maximum resistance occurs only when benefits of tolerance and resistance interact less than additive. Finally, the presence of one peak of maximum resistance or maximum tolerance depends on the relative values of the magnitude of costs for tolerance and resistance. An important outcome of our model is that under a plausible set of conditions, variable costs of tolerance and resistance can represent an important aspect involved in the maintenance of intermediate levels of tolerance and resistance, and in favoring adaptive divergence in plant defensive strategies among populations. The model offers a framework for future theoretical and empirical work toward understanding spatial variation in levels of allocation to different defensive strategies.

POPULATION VARIATION IN THE COST AND BENEFIT OF TOLERANCE AND RESISTANCE AGAINST HERBIVORY IN DATURA STRAMONIUM

Abstract .–In this study we examine the hypothesis that divergent natural selection produces genetic differentiation among populations in plant defensive strategies (tolerance and resistance) generating adaptive variation in defensive traits against herbivory. Controlled genetic material (paternal half‐sib families) from two populations of the annual Datura stramonium genetically differentiated in tolerance and resistance to herbivory were used. This set of paternal half‐sib families was planted at both sites of origin and the pattern of genotypic selection acting on tolerance and resistance was determined, as well as the presence and variation in the magnitude of allocational costs of tolerance. Selection analyses support the adaptive differentiation hypothesis. Tolerance was favored at the site with higher average level of tolerance, and resistance was favored at the site with higher average level of resistance. The presence of significant environmentally dependent costs of tolerance was in agreement with site variation in the adaptive value of tolerance. Our results support the expectation that environmentally dependent costs of plant defensive strategies can generate differences among populations in the evolutionary trajectory of defensive traits and promote the existence of a selection mosaic. The pattern of contrasting selection on tolerance suggests that, in some populations of D. stramonium, tolerance may alter the strength of reciprocal coevolution between plant resistance and natural enemies.

Habitat restriction in Mammillaria pectinifera, a threatened endemic Mexican cactus

Abstract This study deals with the habitat restriction of Mammillaria pectinifera, a threatened cactus species, confined to a few low density localities of the Tehuacán valley in tropical Mexico. We analysed the patterns of presence/absence of M. pectinifera in relation to the presence/absence of 48 other plant species, and the variation of environmental factors in 120 sampling plots. A Principal Components Analysis revealed a clear segregation between plots with and without individuals of M. pectinifera. A classification analysis resulted in four groups: two with low prevalence and two with high prevalence of M. pectinifera. Paired comparisons between plots with and without M. pectinifera allowed the characterization of its patterns of occurrence related to the variation of environmental factors. M. pectinifera was found on deep alkaline soils with relatively high surface stoniness and high water retention capacity, showing low species richness compared with plots where it was absent. The limited distribution of M. pectinifera in the Tehuacán Valley seems to be related to particular requirements of this species, being restricted to certain suitable habitat patches. Nevertheless, it is likely that other aspects, such as poor dispersal and establishment abilities, or biotic interactions could be associated with the observed patterns. Nomenclature: Dávila-Aranda et al. (1993).

Selection Mosaic Exerted by Specialist and Generalist Herbivores on Chemical and Physical Defense of Datura stramonium

Selection exerted by herbivores is a major force driving the evolution of plant defensive characters such as leaf trichomes or secondary metabolites. However, plant defense expression is highly variable among populations and identifying the sources of this variation remains a major challenge. Plant populations are often distributed across broad geographic ranges and are exposed to different herbivore communities, ranging from generalists (that feed on diverse plant species) to specialists (that feed on a restricted group of plants). We studied eight populations of the plant Datura stramonium usually eaten by specialist or generalist herbivores, in order to examine whether the pattern of phenotypic selection on secondary compounds (atropine and scopolamine) and a physical defense (trichome density) can explain geographic variation in these traits. Following co-evolutionary theory, we evaluated whether a more derived alkaloid (scopolamine) confers higher fitness benefits than its precursor (atropine), and whether this effect differs between specialist and generalist herbivores. Our results showed consistent directional selection in almost all populations and herbivores to reduce the concentration of atropine. The most derived alkaloid (scopolamine) was favored in only one of the populations, which is dominated by a generalist herbivore. In general, the patterns of selection support the existence of a selection mosaic and accounts for the positive correlation observed between atropine concentration and plant damage by herbivores recorded in previous studies.

Does the microarchitecture of Mexican dry forest foliage influence spider distribution

Abstract Spider species diversity has been associated with vegetation structure and stratification but there are few studies comparing the spider distribution in different shrubs and trees. In this study we analyzed the species distribution of the spider community of 11 shrub and tree species in two different study sites in a Mexican tropical dry forest. We present results from multivariate analyses that explain their distribution. A classification analysis based on spider abundances separated one shrub, Croton ciliatoglanduliferus, from the rest of the plant species. This was explained by the presence of large numbers of the oxyopid Peucetia viridans (Hentz 1832) on this plant. A second cluster segregated broad-leaved from small-leaved, bipinnate species. This was mainly due to higher spider abundances in the latter type of plants. Four vegetation variables were estimated and their influence on the species distribution was assessed by means of a principal components and regression analysis. With the exception of P. viridans, all spiders were positively associated with number of leaves and number of branchlets per 50 cm branch and negatively with foliage area.

Geographic association and temporal variation of chemical and physical defense and leaf damage in Datura stramonium

The evolution of plant defense traits has traditionally been explained trough the “coevolutionary arms race” between plants and herbivores. According to this, specialist herbivores have evolved to cope effectively with the defensive traits of their host plants and may even use them as a cue for host location. We analyzed the geographic association between leaf trichomes, two tropane alkaloids (putative resistance traits), and leaf damage by herbivores in 28 populations of Datura stramonium in central Mexico. Since the specialist leaf beetles Epitrix parvula and Lema trilineata are the main herbivores of D. stramonium in central Mexico, we predicted a positive association between plant defense and leaf damage across populations. Also, if physical environmental conditions (temperature or precipitation) constrain the expression of plant defense, then the geographic variation in leaf damage should be explained partially by the interaction between defensive traits and environmental factors. Furthermore, we studied the temporal and spatial variation in leaf trichome density and leaf damage in five selected populations of D. stramonium sampled in two periods (1997 vs. 2007). We found a positive association between leaf trichomes density and atropine concentration with leaf damage across populations. The interaction between defensive traits and water availability in each locality had a significant effect on the geographic variation in leaf damage. Differences among populations in leaf trichome density are maintained over time. Our results indicate that local plant–herbivore interaction plays an important role in shaping the geographic and temporal variation in plant defense in D. stramonium.

Stem tilting, pseudocephalium orientation, and stem allometry in Cephalocereus columna-trajani along a short latitudinal gradient

Cephalocereus columna-trajani is a giant columnar cactus endemic of the Tehuacán-Cuicatlán Valley in Central Mexico. Stem tilting and northward pseudocephalium azimuth in C.␣columna-trajani have functional advantages in terms of interception of direct solar radiation at the northernmost portions of its range. Since the success of both characters strongly depends on the apparent position of the sun during the growing season, in this paper we test the hypothesis that the occurrence of such columnar morphology is restricted geographically and imposes mechanical restrictions that limit column height. Following a latitudinal gradient along the Tehuacán-Cuicatlán Valley, we selected five populations, recorded tilting angle and pseudocephalium azimuth, and carried out allometric and biomechanical analyses of height–diameter relationships. Northern populations showed higher tilting angles. Pseudocephalium azimuth significantly differed among populations, and pseudocephalium orientation was consistently North-Northwestern. Stem allometry showed that the stems of the southern populations increased in height at a far greater rate with respect to diameter than the northern populations. The southernmost population showed the lowest safety factor. These results support the hypothesis that stem tilting in C.␣columna-trajani is functionally advantageous in a restricted geographical range, and imposes mechanical restrictions to column height.

Foliole movement and canopy architecture of Larrea tridentata (DC.) Cov. in Mexican deserts

SummaryThe creosote bush (Larrea tridentata) is a common desert perennial with bifoliate, amphistomatic, divaricate leaves. The leaves can vertically close their folioles and vary their profile with respect to direct solar radiation. Field data from different Mexican deserts showed a significant correlation between foliole aperture and mean foliole inclination: in plants in which folioles were more open, the foliole surfaces were less vertical. In a series of field experiments in the Chihuahuan Desert, foliole aperture varied significantly with the water-status of the plant and the hour of the day. In moist plants, folioles opened in the early morning and closed in the afternoon. Water-stressed plants showed significantly lower foliole apertures. A simulation of the light interception patterns of the plants showed that foliole closure in water-stressed individuals reduces direct radiation interception by around 24%. Most (64%) of the reduction in interception was due to the vertical inclination of the photosynthetic surfaces induced by foliole closure in the water-stressed plants. The rest (36%) of the reduction in interception was due to differential self-shading between foliole pairs, which was higher in the closed folioles of the water-stressed plants, but operated more towards the early hours of the day.

Ground Spider Guilds and Functional Diversity in Native Pine Woodlands and Eucalyptus Plantations

Abstract Vegetation structure and floristics have a strong influence on the relative abundance of spider guilds and functional diversity of terrestrial arthropods. Human activities have transformed much of the temperate woodlands. The aim of this study was to test five predictions related to the guild distribution and functional diversity of the ground spider communities of Eucalyptus plantations and native pine woodlands in western Mexico. Spiders were collected every fortnight from September to November from 15 pitfalls positioned in each of the eight sites. We also assessed the cover of grasses, herbs, shrubs, and leaf litter in each site. We found that the abundances of ground hunters and sheet weavers between plantations and pine woodlands were different. Nevertheless, there was not a consistent difference between sites of each of the vegetation types. Most species of ground hunters, sheet web weavers, and many other hunters were associated with litter and the grass cover. Nonetheless, in some cases, species of different families belonging to the same guild responded to different variables. Wolf spiders were related to the grass Aristida stricta Micheaux, 1803, while the species of the other families of ground hunters were associated with leaf litter. One Eucalyptus plantation and one pine woodland had the highest functional diversity of all sites. These sites have a well developed litter and grass cover. Our study suggests that the abundance of litter and a high cover of grasses explain the occurrence of species with different traits, and these habitat components results in a high functional diversity.

Alternative glacial-interglacial refugia demographic hypotheses tested on Cephalocereus columna-trajani (Cactaceae) in the intertropical Mexican drylands

Historic demography changes of plant species adapted to New World arid environments could be consistent with either the Glacial Refugium Hypothesis (GRH), which posits that populations contracted to refuges during the cold-dry glacial and expanded in warm-humid interglacial periods, or with the Interglacial Refugium Hypothesis (IRH), which suggests that populations contracted during interglacials and expanded in glacial times. These contrasting hypotheses are developed in the present study for the giant columnar cactus Cephalocereus columna-trajani in the intertropical Mexican drylands where the effects of Late Quaternary climatic changes on phylogeography of cacti remain largely unknown. In order to determine if the historic demography and phylogeographic structure of the species are consistent with either hypothesis, sequences of the chloroplast regions psbA-trnH and trnT-trnL from 110 individuals from 10 populations comprising the full distribution range of this species were analysed. Standard estimators of genetic diversity and structure were calculated. The historic demography was analysed using a Bayesian approach and the palaeodistribution was derived from ecological niche modelling to determine if, in the arid environments of south-central Mexico, glacial-interglacial cycles drove the genetic divergence and diversification of this species. Results reveal low but statistically significant population differentiation (FST = 0.124, P < 0.001), although very clear geographic clusters are not formed. Genetic diversity, haplotype network and Approximate Bayesian Computation (ABC) demographic analyses suggest a population expansion estimated to have taken place in the Last Interglacial (123.04 kya, 95% CI 115.3–130.03). The species palaeodistribution is consistent with the ABC analyses and indicates that the potential area of palaedistribution and climatic suitability were larger during the Last Interglacial and Holocene than in the Last Glacial Maximum. Overall, these results suggest that C. columna-trajani experienced an expansion following the warm conditions of interglacials, in accordance with the GRH.