Santiago Arizaga

Architecture, Light Interception, and Distribution of Larrea Species in the Monte Desert, Argentina

This paper analyzes the correlation between leaf orientation and the environmental conditions that prevail within the biogeographic range of each of the four South American Larrea species (L. ameghinoi, L. cuneifolia, L. divaricata, and L. nitida; Zygophyllaceae). Data on the distribution of Larrea species were gathered from herbarium specimens. Measurements of leaf orientation were made throughout the Monte Desert, and in more detail in northern Patagonia, where the four species coexist. The direct solar radiation intercepted by the unshaded leaves of each species was estimated through a computer model and plotted as a function of the hourly time for the summer and winter solstices. L. ameghinoi presents horizontal leaves and prostrate growth, characters which allow its development on sites that are exposed to the Patagonian westerlies. The species, however, is an inefficient light interceptor in winter and early spring, when moisture conditions are adequate in Patagonia. Its architecture is the result of selection for cushion—type, wind resistant forms, at the expense of light interception. It is restricted to windy, open areas of the Patagonian steppe. L. cuneifolia shows erect, east—facing leaves and branches, which maximize interception in the early morning and late afternoon, keeping noon interception at a minimum. It can tolerate very hot environments by physically evading the midday sun and intercepting more early morning and late afternoon light. It colonizes the hotter and drier parts of the Monte Desert. L. divaricata has divaricate leaves with folioles uniformly distributed in all azimuthal directions, and showing an inclination of around 70°. Although it never shows maximum light interception efficiencies, it preforms relatively well in all seasons and at all hours of the day. Its distribution is wide, not only in the arid Monte, but reaching also the Chaco woodlands and the Pacific coastal deserts. L. nitida shows erect, north—facing leaves and branches. Its leaf orientation distribution allows the gradual warming of the leaf surfaces during the morning, with a maximum light interception near winter noons. The species grow in the Patagonian Monte and on the slopes of the Andes. Its general affinity with the colder west side of the Patagonian and Monte Deserts links its distribution with winter—type rains of Pacific origin. The results suggest that the contrasting leaf orientations of the four South American Larrea species reflect the prevalent selective conditions endured under long periods in isolation, and that leaf orientation is an adaptive character that influences the habitat specificity of the different species.

Propagation mechanisms in Agave macroacantha (Agavaceae), a tropical arid-land succulent rosette.

Agave macroacantha can sexually reproduce by seeds and propagate vegetatively by aerial bulbils and ground-level basal shoots and rhizomes. It forms compact patches apparently generated by the multiplication of ground-level offshoots. We experimentally evaluated the establishment and survival of bulbils and seedlings of A. macroacantha in the Tehuacán Valley, Mexico, between 1991 and 1994 and studied comparatively the effectiveness of sexual reproduction against vegetative propagation.Seedlings showed low survival rates. Cohorts placed outside nurse plants died in less than 1 yr, while 1-10% of cohorts under nurse plants survived for more than 2 yr. Herbivores negatively affected seedling survival in non-nursed plots. In rainy years, survival rates increased. Bulbils showed higher survival rates than seedlings.The excavation of rosettes showed that most are derived from vegetative shoots, as indicated by remains of rhizomes in their base. Most rosettes had ground-level vegetative offspring totaling almost three shoots per rosette.In A. macroacantha, the establishment of seedlings and bulbils is a rare event that possibly only occurs under nurse plants in rainy years, while ground-level cloning is highly effective as a propagation mechanism. These results are consistent with the aggregated spatial pattern of the species.

Pollination ecology of Agave macroacantha (Agavaceae) in a Mexican tropical desert. I. Floral biology and pollination mechanisms

In a study of sexual reproduction in long-lived semelparous plants, we observed Agave macroacantha in the tropical desert of Tehuacán-Cuicatlán, Mexico, describing duration of flowering, flower phenology, and nectar production patterns. We also performed two manipulative experiments evaluating (a) the seed production efficiency of different crossing systems (selfing, cross-pollination, apomixis, and control), and (b) the effect of different pollinators (diurnal exposure to pollinators, nocturnal exposure, exclusion, and control) on the seeds produced. Flowering occurred from early May to late July and had a mean duration of 29 days in the individual rosettes. The flowers were protandrous; anthesis occurred in the afternoon of the third day after floral opening, and the pistils matured in the afternoon of the fifth day. The stigmas remained receptive from dusk to the following morning. Pollination was mostly allogamous. Nectar was produced principally during the night, from the first stages of floral aperture until the stigmas wilted and flowering ceased. The flowers were visited during the day by hymenoptera, butterflies, and hummingbirds and during the night by bats and moths. Only the nocturnal visitors, however, were successful pollinators. Agave macroacantha is extremely dependent on nocturnal pollinators for its reproductive success.

Pollination ecology of Agave macroacantha (Agavaceae) in a Mexican tropical desert. II. The role of pollinators

We did a series of observational studies and manipulative experiments on the guild of nocturnal visitors of Agave macroacantha, including (1) a description of the hourly patterns of visits by moths and bats, (2) an evaluation of the relative contribution of bats and moths to flowering success, and (3) an evaluation of the pollination efficiency of the different bat species. Scapes exposed to moths but excluded to bats yielded ∼50% fewer fruits than those exposed to both pollinator groups. Flowers exposed to the bat species Leptonycteris curasoae showed similar fruiting success to those exposed to Choeronycteris mexicana and to those exposed to the whole nocturnal visitor guild. However, the fruits originated from flowers pollinated by Leptonycteris curasoae yielded significantly more seed than those exposed to Choeronycteris mexicana or to the whole pollinator guild. It is concluded that Agave macroacantha is extremely dependent on nocturnal pollinators for its reproductive success and that bats are especially important for successful pollination. Some of these pollinators are migratory and have been reported to be steadily declining. A continuing decline in the populations of pollinators may impede the successful sexual reproduction of the plant host and may put the long-term survival of this agave species under risk.

Insurance against reproductive failure in a semelparous plant : bulbil formation in Agave macroacantha flowering stalks

Bulbils are small aerial rosettes that occur on the flowering stalks of semelparous Agave plants and in related families, and that are capable of acting as clones of the parent plant. We hypothesized that bulbil formation was inversely related to fruiting success in the flowering stalk, and we explored this hypothesis in A. macroacantha, a species from South-Central Mexico. Forty randomly chosen plants were divided amongst three treatments: (a) elimination of all floral buds, (b) exclusion of pollinators, and (c) control. We also studied 22 plants in which the flowering stalk had been felled by goat grazing. Between September and November 1991 we kept a record of the numbers of bulbils and capsules produced in each flowering stalk. Significant (P<0.0001) differences between treatments were found in the proportion of plants hearing capsules and bearing bulbils. The control treatment had the highest proportion of plants producing capsules, treatment a had the highest proportion of individuals bearing bulbils, while treatment b showed an intermediate response. In the goat-grazed group, 45% of the plants failed to produce any propagative structure after the stalk was cut, and half of all plants produced bulbils on the remaining stump. A significant inverse relationship between the numbers of capsules and the numbers of bulbils per plant was found for the three randomly assigned treatments. Our results suggest that once the production of the flowering stalk has been triggered and the death of the rosette is irreversible, bulbils may act as an insurance mechanism that increases the probability of successful reproduction of the genet.

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.

Landscape genetics reveals inbreeding and genetic bottlenecks in the extremely rare short-globose cacti Mammillaria pectinifera (Cactaceae) as a result of habitat fragmentation

Mammillaria pectinifera is an endemic, short-globose cactus species, included in the IUCN list as a threatened species with only 18 remaining populations in the Tehuacán-Cuicatlán Valley in central Mexico. We evaluated the population genetic diversity and structure, connectivity, recent bottlenecks and population size, using nuclear microsatellites. M. pectinifera showed high genetic diversity but some evidence of heterozygote deficiency (FIS), recent bottlenecks in some populations and reductions in population size. Also, we found low population genetic differentiation and high values of connectivity for M. pectinifera, as the result of historical events of gene flow through pollen and seed dispersal. M. pectinifera occurs in sites with some degree of disturbance leading to the isolation of its populations and decreasing the levels of gene flow among them. Excessive deforestation also changes the original vegetation damaging the natural habitats. This species will become extinct if it is not properly preserved. Furthermore, this species has some ecological features that make them more vulnerable to disturbance such as a very low growth rates and long life cycles. We suggest in situ conservation to prevent the decrease of population sizes and loss of genetic diversity in the natural protected areas such as the Tehuacán-Cuicatlán Biosphere Reserve. In addition, a long-term ex situ conservation program is need to construct seed banks, and optimize seed germination and plant establishment protocols that restore disturbed habitats. Furthermore, creating a supply of living plants for trade is critical to avoid further extraction of plants from nature.