Carlos Martorell

Testing the roles of competition, facilitation and stochasticity on community structure in a species-rich assemblage

Summary There is an ongoing debate about whether communities are closely integrated and bound together via interactions such as competition or facilitation, or are disintegrated and dominated by chance. We still lack community-wide data on the intensities of interactions and randomness, and measurements of their impacts on community structure. Using a long-term data set, we sought to measure the effects of interactions and stochasticity in structuring a highly diverse (>100 species) semi-arid grassland plant community, testing for positive and negative interactions at different stages of population growth of all species. During the colonization of new patches, most species were facilitated or inhibited by several others. These opposite effects can potentially have a large effect on species abundances, but they were correlated and cancelled out at the community level. Nevertheless, competition during colonization was strong enough to cause poor competitors to have small population sizes. The subsequent phase of population growth (increase and subsequent change in numbers within occupied patches) was mainly driven by intraspecific density dependence, and we found little evidence for interspecific interactions. Model results showed that stochasticity and recurrent colonization of transient, favourable patches maintained diversity, keeping poor competitors from becoming extinct. Synthesis: Our results, taken together with recent studies on tropical forests, suggest that weak interactions among established plants may be a general phenomenon, but that local interactions during colonization are important drivers of community composition. Most of the variance in species abundance in our community was explained by intraspecific competition and stochasticity, with interspecific interactions playing a minor role due to their overall weakness, interaction changes over ontogeny, and the cancellation of opposite-sign interactions when all the species in the community are considered. Despite this, some species were rare seemingly because they cannot withstand interspecific competition. Thus, to untangle the effects of interactions on community structure, future research should focus on interactions occurring at different phases of population growth and on whole communities.

The role of rustic coffee plantations in the conservation of wild tree diversity in the Chinantec region of Mexico

Rustic coffee plantations are characterised by the use of numerous wild and cultivated tree species for providing shade to the coffee shrubs. This paper analyses the role of these plantations in wild tree conservation through the examination of their patterns of floristic variation in southern Mexico. The studied plantations included a total of 45 plant species, most of which were wild tree species, including both mature forest and pioneer taxa. An extrapolation of the species accumulation curve among stands indicated that the whole system, composed of more than 100 coffee plantations, may harbour as many as 34 species of wild trees. The floristic structure of rustic coffee plantations was highly variable. This variation is a result of a combination of factors such as human management, original stand cover and the asynchrony in development stage of different plantations. This promotes a large β-diversity in the system. Thus, although a single plantation may have a limited potential to preserve wild tree species, it is the whole ensemble of floristically heterogeneous plantations which renders this agroforestry system valuable for plant diversity conservation, particularly in a region where native forest vegetation has almost disappeared.

The narrow-leaf syndrome: a functional and evolutionary approach to the form of fog-harvesting rosette plants

Plants that use fog as an important water-source frequently have a rosette growth habit. The performance of this morphology in relation to fog interception has not been studied. Some first-principles from physics predict that narrow leaves, together with other ancillary traits (large number and high flexibility of leaves, caudices, and/or epiphytism) which constitute the “narrow-leaf syndrome” should increase fog-interception efficiency. This was tested using aluminum models of rosettes that differed in leaf length, width and number and were exposed to artificial fog. The results were validated using seven species of Tillandsia and four species of xerophytic rosettes. The total amount of fog intercepted in rosette plants increased with total leaf area, while narrow leaves maximized interception efficiency (measured as interception per unit area). The number of leaves in the rosettes is physically constrained because wide-leafed plants can only have a few blades. At the limits of this constraint, net fog interception was independent of leaf form, but interception efficiency was maximized by large numbers of narrow leaves. Atmospheric Tillandsia species show the narrow-leaf syndrome. Their fog interception efficiencies were correlated to the ones predicted from aluminum-model data. In the larger xerophytic rosette species, the interception efficiency was greatest in plants showing the narrow-leaf syndrome. The adaptation to fog-harvesting in several narrow-leaved rosettes was tested for evolutionary convergence in 30 xerophytic rosette species using a comparative method. There was a significant evolutionary tendency towards the development of the narrow-leaf syndrome the closer the species grew to areas where fog is frequently available. This study establishes convergence in a very wide group of plants encompassing genera as contrasting as Tillandsia and Agave as a result of their dependence on fog.

Disturbance-Response Analysis: a Method for Rapid Assessment of the Threat to Species in Disturbed Areas

We developed a method with which to analyze a species response to chronic anthropogenic disturbance (CAD). We regressed density of individuals on the intensity of 3 disturbance agents (human activities, raising livestock, and land degradation) along CAD gradients to determine how much CAD is experienced by a species; whether species are ruderal (achieving maximum densities in disturbed sites); whether density declines as a result of CAD; which disturbance agents are responsible for this reduction; and the number of populations that decline as a result of CAD. We tested the method on 9 threatened Mammillaria species (Cactaceae). Seven species were ruderal. In 4 species, actual disturbance surpassed the CAD intensity in which plants achieved their maximum densities; thus, the density of those 4 species appeared to be declining. For 7 species, some populations were threatened by human activities or livestock, whereas others were favored by them. Land degradation negatively affected all species. Our results allowed us to distinguish 4 groups of species that had similar responses and thus may require different forms of management. Our method provided an estimation of the growth rate (lambda) of the studied populations that was significantly correlated with lambdas obtained from demographic studies. The size structures of populations were consistent with the predictions of the analysis, which suggests our results are consistent and reliable. Disturbance-response analysis provides a basis for management in heavily populated areas, where conservation must be achieved along with development activities that cause CAD. The method provides readily interpretable information, which facilitates participative decision making; the data are rapidly generated, which makes it appropriate when results are required promptly or for assessment of large numbers of species; and it provides a comprehensive perception of how threatened species behave in the real world.

Predicting Tropical Dry Forest Successional Attributes from Space: Is the Key Hidden in Image Texture?

Biodiversity conservation and ecosystem-service provision will increasingly depend on the existence of secondary vegetation. Our success in achieving these goals will be determined by our ability to accurately estimate the structure and diversity of such communities at broad geographic scales. We examined whether the texture (the spatial variation of the image elements) of very high-resolution satellite imagery can be used for this purpose. In 14 fallows of different ages and one mature forest stand in a seasonally dry tropical forest landscape, we estimated basal area, canopy cover, stem density, species richness, Shannon index, Simpson index, and canopy height. The first six attributes were also estimated for a subset comprising the tallest plants. We calculated 40 texture variables based on the red and the near infrared bands, and EVI and NDVI, and selected the best-fit linear models describing each vegetation attribute based on them. Basal area (R 2 = 0.93), vegetation height and cover (0.89), species richness (0.87), and stand age (0.85) were the best-described attributes by two-variable models. Cross validation showed that these models had a high predictive power, and most estimated vegetation attributes were highly accurate. The success of this simple method (a single image was used and the models were linear and included very few variables) rests on the principle that image texture reflects the internal heterogeneity of successional vegetation at the proper scale. The vegetation attributes best predicted by texture are relevant in the face of two of the gravest threats to biosphere integrity: climate change and biodiversity loss. By providing reliable basal area and fallow-age estimates, image-texture analysis allows for the assessment of carbon sequestration and diversity loss rates. New and exciting research avenues open by simplifying the analysis of the extent and complexity of successional vegetation through the spatial variation of its spectral information.

Cultural or Ecological Sustainability? The Effect of Cultural Change on Sabal Palm Management Among the Lowland Maya of Mexico

Sabal palm has been used for thatching the traditional Maya house for over 3000 yr. The great importance of this resource has promoted its management within home gardens. Although traditionally managed populations in home gardens are capable of ecological long-term persistence, the impact of cultural change on sustainable resource management is poorly understood. By means of interviews in 108 households, we obtained information about Sabal management practices, leaf demand, and sociocultural data. Density and size structure of the palm populations in the respective home gardens were also measured. By means of principal components analysis, the sociocultural data were summarized into a cultural change index, which was then statistically related to palm density, size structure, leaf demand, and management practices. Leaf demand along the cultural change gradient was estimated. Sabal populations were affected by the cultural change index. Palm density and the proportion of harvestable individuals were higher in the more traditional households. The number of management practices decreased, and the probability of felling adult palms increased with cultural change. As a result, the percentage of the total leaf demand satisfied by home garden production diminished from 118.2-69.4% as cultural change increased. Traditional practices seem oriented to increasing the palm availability. Seed sowing and the protection of seedlings and adults affect the life stages with the largest impact on the population growth rate, as measured through sensitivity analysis. This means that abandoning traditional practices and felling adults more frequently should reduce rapidly, which is consistent with the low palm density observed in less traditional households. The application of demographic models to Sabal tells us that traditional management warrants the persistence of the resource as long as the current conditions remain unchanged. In contrast, our data show that Sabal management may not be sustainable from a cultural perspective, since the cultural attitudes that affect palm management and demand change over time. Both approaches assess the same problem from different viewpoints reaching different but complementary conclusions. In this study, we propose new methods that integrate the ecological and cultural processes, which affect the sustainable management of the natural resources.

Factors that influence the intensity of non-agricultural management of plant resources

We investigated the relationships of land tenure, biological, cultural and spatial variables and their effect on the intensity of management of 20 edible plants used by the Santa Maria Tecomavaca community in Oaxaca State, Mexico. We developed a non-linear generalized model showing that land ownership, cultural importance and biological characteristics of a plant are the most significant factors influencing farmers’ decisions to intensify management of plant resources. On common land, species with high cultural importance and long life cycles are managed intensively, while, regardless of cultural importance, species with short life cycles are simply gathered. On land where particular people have an acknowledged right of use, species with long life cycles are always managed at an intermediate level of intensity regardless of their cultural importance, while for species with short life cycles, management intensity tends to slightly decrease as cultural importance increases. Even though management is generally more intensive in lands where individuals have use rights, as expected from the “supervised collective action model”, culturally important resources are more intensively managed in communal areas, probably as a result of a long tradition of consumption and of their economic importance.

The adaptive value of cued seed dispersal in desert plants: Seed retention and release in Mammillaria pectinifera (Cactaceae), a small globose cactus

Serotiny, or delayed seed dispersal, is common in fluctuating environments because it hedges the risks of establishment. Mammillaria pectinifera (Cactaceae) facultatively expels fruits in the year they are produced or retains them to disperse the seed over several years. We tested whether M. pectinifera increased fruit expulsion as a response to increased rainfall. While no fruit expulsion was observed in 1997, a dry year, in the wetter 1998 around 20% of all fruits formed were expelled from the maternal plant. A greenhouse experiment showed that high moisture results in the plants expelling all their fruits. Because in 1998 establishment was five times higher than in 1997, this response seems to be highly adaptive: Active fruit expulsion and consequent seed release increases the probability of establishment during pulses of high precipitation.

Ruderality in extreme-desert cacti? Population effects of chronic anthropogenic disturbance on Echinocereus lindsayi

Ruderal species, i.e., those that increase their numbers in the presence of disturbance, are not expected to occur in extreme environments. We test whether Echinocereus lindsayi, a cactus from an extreme desert, follows the ruderal trend observed in similar species from mild deserts, or, as theory suggests, it is a non ruderal. Contrary to expectations, its density and fraction of small individuals in the population increased with disturbance. This seemingly results from increased establishment, as it is nursed by rocks exposed by disturbance. A demographic model for two populations, one nearly pristine and another highly disturbed, showed that at the latter site recruitment was more frequent and likely. At the disturbed site the performance of E. lindsayi individuals was usually poor, except on favorable years. Then, competition release caused by disturbance apparently allowed for a better performance compared to the less disturbed site. Despite that this opportunistic behavior would suffice to maintain the population size stable, the large mortality produced by an insect outbreak in two of the four study years caused the population to diminish. In contrast, the population at the less disturbed site was near equilibrium. If the insect outbreak is associated to disturbance, E. lindsayi at the disturbed site would be already experiencing more disturbance than it tolerates. This agrees with the fact that no populations were found at greater disturbance intensities. While, contrary to our hypothesis, E. lindsayi is ruderal, this extreme desert species appears to tolerate far less disturbance than its counterparts from milder areas.

Does retained-seed priming drive the evolution of serotiny in drylands? An assessment using the cactus Mammillaria hernandezii

PREMISE OF THE STUDY Serotinous plants retain their seeds for a long time. In deserts, retained seeds undergo hydration-dehydration cycles and thus may become primed. Priming enhances germination and seedling vigor. We test the hypothesis that serotiny evolves because it provides a site protected from predators in which seeds can become primed. Rainfall-cued dispersal of primed seeds may enhance this effect. METHODS We tested this hypothesis with Mammillaria hernandezii through protein-content analyses; field and laboratory germination experiments with primed, unprimed, and retained seeds; and fitness estimations from demographic models. KEY RESULTS Hydration-dehydration cycles induced priming, enhancing germination. Artificial priming and retention in the parent plant for 1 yr induced similar changes in seed protein patterns, suggesting that priming occurs naturally while seeds are retained. Under field conditions, germination of seeds retained for 1 yr more than doubled that of seeds of the same cohort that were not primed or that remained buried for 1 yr. The first seeds to germinate died rapidly. Serotinous plants whose seeds underwent priming had higher fitness than those whose seeds were in the soil seed bank or that did not experience priming. CONCLUSIONS Priming in soil seed banks may be costly because of high predation, so seed protection during priming is sufficient to promote the evolution of serotiny. Bet hedging contributes to this process. Rapid germination of primed seeds that respond to brief rainfall events is disadvantageous because such rainfall is insufficient for seedling survival. Serotinous species counteract this cost by cueing dispersal with heavy precipitation.