Joel Flores

Are nurse-protégé interactions more common among plants from arid environments?

Abstract Harsh conditions in arid and semi-arid environments make seedling establishment rare. Plant recruitment in arid environments often occurs only in years with above average rainfall or in safe sites under the canopy of nurse plants that provide shelter from high temperatures and low moisture. Associations of establishing seedlings with adult plants are referred to as nurse-protégé interactions and are thought to be commensalisms in which seedlings benefit from the micro-environment created by adult plants with no effect for the latter. This phenomenon is thought to be more frequent in harsh than in mild environments and appears to occur frequently in deserts and arid and semi-arid biomes. Here, we investigate whether nurse-protégé interactions are more common in arid environments by searching the published literature from the previous 92 years using the terms nurse plants, protégé plants, facilitation, nucleation and facultative mutualism. We then quantitatively compared these reports from arid zones to other environments. A total of 296 papers were found which referred to nurse-protégé interactions. More than half (158) focused on arid and semi-arid zones. This information was also used to explore hypotheses of potential causative forces that might have selected for such interactions in the arid zones such as seed trapping, nutrient, moisture, protection from browsing or trampling and support availability. Because of the large number of different nurse species (147, from 98 genera and 40 families) and protégé species (429, from 273 genera and 84 families), described across a diversity of environments, we suggest that there may be more than one causative factor selecting for nurse-protégé interactions in arid and semi-arid environments. Nomenclature: See references in (App. 1).

Effect of predation and solar exposure on the emergence and survival of desert seedlings of contrasting life-forms

Abstract We evaluated the effect of solar exposure and predation on seedlings of six species of three life-forms. Columnar succulents and arborescent semi-succulents emerged faster under shrubs (24 and 55 days) than in open places (44 and 60 days). Succulents had a longer survival time (138 days) without predators and under shrubs. Predator exclusion led to a longer survival in semi-succulents under shrubs (237 days) and in open places (198 days). Predation did not affect the survival time of shrubs. Our results extend to the seedling stage, the hypothesis that in desert environments, each life-form adopts different strategies for survival and growth.

Are fungi important for breaking seed dormancy in desert species? Experimental evidence in Opuntia streptacantha (Cactaceae)

Seeds of Opuntia spp. have physiological dormancy; they need a period of after-ripening to break dormancy, and the embryos have low growth potential. We evaluated the combined effects of seed age and presence of fungi on the testa on germination of Opuntia streptacantha, an abundant species in the Chihuahuan Desert (Mexico), assuming that older seeds have broken seed dormancy and fungi can reduce mechanical resistance to germination. In a preliminary experiment, we found no germination of 9-year-old (1998) and freshly collected (2007) seeds. However, we obtained 67% and 27% germination from 9-year-old and fresh non-sterilized seeds, respectively, and found fungi growing on the testa of all germinated seeds. Two fungal strains were isolated and identified using ribosomal internal transcribed spacer (ITS) sequence analysis: Penicillium chrysogenum and Phoma sp. In a second experiment, we inoculated seeds with strains of P. chrysogenum and Phoma sp., as well as Trichoderma koningii and binucleate Rhizoctonia (Gto17S2), to evaluate their ability to break seed dormancy. Seeds inoculated with P. chrysogenum, Phoma sp. and T. koningii had higher germination than controls for both seed ages, but germination was higher in older seeds. Scanning electron microscopy showed that these fungi eroded the funiculus, reducing its resistance. Binucleate Rhizoctonia did not lead to germination and controls had almost no germination. Our results strongly indicate that fungi are involved in breaking seed dormancy of O. streptacantha, and that the effect of fungi on seeds is species-specific.

Seedling establishment under native tamaulipan thornscrub and Leucaena leucocephala plantation

Abstract In order to investigate the influence that plantations and clearing of native vegetation might have on plant regeneration in northeastern Mexico, seedling survival and germination (i.e., seedling emergence from the soil surface) were measured monthly in native fragments and an exotic plantation over a one-year period. Seedlings were measured in 1-m 2 plots, using 10 replicates under each of three different treatments: (i) dense thornscrub, (ii) semi-cleared thornscrub, and (iii) a dense L. leucocephala plantation. Seedling number peaked during summer and early autumn, lowest number of seedlings occurred during early spring. Excluding grasses, 36 species were present during the study, and only 14 were present in L. leucocephala plantation. Thornscrub and open thornscrub had similar number of grass seedlings, whereas more forb seedlings were present in open thornscrub. Seedlings of woody species were similar for all treatments. Under L. leucocephala germination was low for most species other than itself, most seedlings, however, died during the winter months.

Further evidence from the effect of fungi on breaking Opuntia seed dormancy.

Recently, we found that fungi are involved in breaking seed dormancy of Opuntia streptacantha, and that the effect of fungi on seeds is species-specific. However, the effect of fungi on seed germination from other Opuntia spp has not been evaluated. Thus, we evaluated the effect of four fungal species (Penicillium chrysogenum, Phoma sp., Trichoderma harzianum, Trichoderma koningii) on the germination of Opuntia leucotricha, an abundant species in the Chihuahuan Desert, Mexico. We found that seeds inoculated with the four fungal species had higher germination than control seeds. Trichoderma spp. were the most effective. Our results strongly indicate that fungi are involved in breaking seed dormancy of O. leucotricha. Thus, we suggest that these fungi could promote seed germination from other Opuntia species.

Effect of fungi and light on seed germination of three Opuntia species from semiarid lands of central Mexico

Fungal attack under light reduces mechanical resistance of the testa of Opuntia seeds, making it easier for the embryo to emerge. However, the effect of fungi on Opuntia seed germination in darkness is unknown. We evaluated the combined effects of light and inoculation with Phoma medicaginis, Trichoderma harzianum, Trichoderma koningii, and Penicillium chrysogenum on germination of O. streptacantha, O. leucotricha, and O. robusta seeds, from central Mexico. We also evaluated the combined effects of seed age (2-, 3-, and 12-year-old seeds) and presence of fungi on the testa on O. streptacantha germination. All fungal species eroded the funicular envelope and promoted seed germination for O. leucotricha and O. streptacantha, but did more so in light than in darkness. For the latter species, younger seeds inoculated with fungi had lower germination than older ones. For O. robusta, we found that seeds inoculated with P. medicaginis and T. harzianum had similar germination in light and in darkness. Our results strongly indicate that deterioration of the testa by fungi is higher in light than in darkness.

Ecophysiological and anatomical mechanisms behind the nurse effect: which are more important? A multivariate approach for cactus seedlings.

Background Cacti establish mostly occurs under the canopy of nurse plants which provide a less stressful micro-environment, although mechanisms underlying this process are unknown. The impact of the combination of light and watering treatments on Opuntia streptacantha (Cactaceae) seedlings was examined. Methods/Principal Findings Ecophysiological [titratable acidity, osmotic potential (‘solute potential’, Ψs), relative growth rate (RGR) and their components (NAR, SLA, and LWR)], anatomical (chloroplast density, chloroplast frequency, and cell area), and environmental [photosynthetic photon flux density (PPFD) and air temperature] sets of variables were analyzed, assessing relationships between them and measuring the intensity of the relationships. Three harvests were carried out at days 15, 30, and 45. Ψs and acidity content were the most important responses for seedling establishment. The main anatomical and environmental variables were chloroplast density and water availability, respectively. Opuntia streptacantha seedlings establish better in the shade-watering treatment, due to higher Ψs and acidity, unaffected chloroplasts, and lower PPFD. In addition, the chloroplasts of cells under high-light and non-watering treatment were clumped closer to the center of the cytosol than those under shade-drought, to avoid photoinhibition and/or to better distribute or utilize the penetrating light in the green plant tissue. Conclusions Opuntia seedlings grow better under the shade, although they can tolerate drought in open spaces by increasing and moving chloroplasts and avoiding drastic decreases in their Ψs. This tolerance could have important implications for predicting the impact of climate change on natural desert regeneration, as well as for planning reforestation-afforestation practices, and rural land uses.

Distribution patterns of acorns after primary dispersion in a fragmented oak forest and their consequences on predators and dispersers

Forest fragmentation affects seed production, but little is known about how seeds are distributed in fragmented forests after primary dispersion and how this affects their interactions with other organisms. This study addressed these issues in a fragmented oak forest of Quercus laeta where four habitat types were recognized: interior and edge of patches, deforested matrix around patches, and an abandoned field. Areas and distances to the abandoned field were measured for 40 patches, and acorns were sampled in all habitats. Only samples from the interior and edge of patches had acorns, which were classified as aborted, viable, and parasitized by insects or fungi. Acorns in all these classes were regressed against patch area and distance to the abandoned field. Field experiments were also conducted to determine whether acorn removal rates varied across habitats. Aborted acorns within patches were not related to area or distance, but they decreased with increasing values of both variables at patch edges. Within patches, viable acorns were positively related to increases in area and distance, while the converse occurred for insect-damaged acorns; fungal-infested acorns were not related to patch variables in this habitat. Complex interactions between area and distance explained how acorns in these classes were distributed across patch edges. Acorn removal was higher within patches and their edges when compared to human-disturbed habitats; both mice and birds were identified as potential acorn dispersers. These results suggest that forest fragmentation affects acorn-mediated processes and that this may influence the recovery of patch connectivity in the landscape.

Seedling survival of three endemic and threatened Mexican cacti under induced climate change

Human-induced warming may increase the risk of local extinction for plant species with low tolerance of elevated temperatures. The Chihuahuan desert harbors the highest diversity of globose cacti in the world and most of them are at risk of extinction. Predictive models of climate change indicate an increase in summer temperature of 1–2°C by 2030 for this desert. Nevertheless, studies on the vulnerability of cacti species in early development phases to future climate change are scarce. We assessed the survival of three threatened cacti species from the Chihuahuan desert under induced warming. Open-top chambers (OTCs) were used to simulate the effect of global warming on 2-year seedlings of Echinocactus platyacanthus f. visnaga, Ferocactus histrix and Stenocactus coptonogonus. OTCs had higher temperature and lower humidity than control plots, and these elevated temperatures reduced seedling survival. Within the OTCs, no living individuals of any species were found after 105 days. Conversely, in the control plots, the three cacti species showed variable numbers of survivors after this period. Therefore the predicted global warming scenarios will greatly limit plant recruitment and the long-term persistence of natural populations of Mexican endemic cacti species.

Shade shelters increase survival and photosynthetic performance of oak transplants at abandoned fields in semi-arid climates

Forest restorations conducted in semiarid, seasonally dry climates must deal with the intense drought stress that affects tree seedlings during the dry season. Although this water deficit is the most commonly invoked source of mortality for seedlings, several other environmental factors may also preclude survival of transplants. For instance, it has been widely reported that excessive light reduces the efficiency of the photosynthetic apparatus, hence decreasing plant survival, but most seedling transplants in deforested areas are conducted under these light conditions. This study is focused in determining whether excessive light affects the photosynthetic performance and survival of Quercus coccolobifolia, a Mexican oak species, when their seedlings are transplanted in semiarid deforested areas. Further, this study tests the possibility of using artificial shade shelters to improve the ecophysiological performance and survival of seedlings. Oak seedlings were transplanted under full sunlight conditions and beneath artificial shade shelters of two different colors: white and black. To reduce water stress, and hence isolate the effects of light treatments, a drip irrigation system was implemented at each experimental plot. Seedling survival was monitored weekly for 128 days and photosynthetic performance was assessed by measuring chlorophyll fluorescence at three opportunities during the experiment. Sun-exposed seedlings showed lower photosynthetic performance and survival rates than those beneath shelters of both colors. These results suggest that sunlight damage can reduce seedling survival when they are transplanted in exposed sites, and that shade shelters can improve the success of forest restoration programs in semiarid climates.