Lonesome plants: How isolation affects seed set of a threatened dioecious shrub.

Plant reproductive failure is a critical concern for conserving rare and endangered species that typically have low-density and sparse populations. One important factor contributing to reproductive failure is the spatial arrangement of plants within a population, which can lead to isolation and negatively affect seed production, particularly in obligate outcrossers. Additionally, plant size can compound this effect, influencing seed production via multiple processes. Here, we investigate how spatial distribution and size influence the reproductive success of Vasconcellea chilensis, an endemic-threatened papaya species in Chile. We first examined whether V. chilensis can produce seeds via apomixis using pollinator exclusion experiments. We then used Spatial Point Pattern Analysis (SPPA) in three populations to explore the spatial arrangement of plants. Finally, we assessed whether plant size and neighbor distance influence the reproductive success V. chilensis is a dioecious shrub unable to produce fruits through apomixis. The SPPA revealed significant clustering of female and male plants at different spatial scales, indicating a non-random distribution. Moreover, a significant spatial association between the sexes was observed. In two populations, closer proximity to male plants was linked to higher seed production. Our study revealed that the reproductive system of V. chilensis is susceptible to distance-dependent reproductive failure due to pollen limitation. While the species' spatial structure may partially mitigate this risk, female plants isolated from male counterparts will likely experience reduced seed set.

Functional diversity and spatial association analyses at different spatial scales reveal no changes in community assembly processes along an aridity gradient in the Atacama Desert.

The structuring of plant assemblages along environmental gradients is typically explained by shifts from competition (limiting similarity) to environmental filtering as the environment becomes more stressful. However, facilitation, weaker-competitor exclusion, environmental heterogeneity, and the colonization-competition tradeoff can also structure plant assemblages along gradients. These assembly processes act on different plant traits and organs, and their prevalence varies with respect to the spatial scale. Using patterns of functional diversity, coupled with patterns of species association at two spatial scales, here we discern the assembly processes that structure shrub communities in four localities along an aridity gradient of the Atacama Desert. At each site, we calculated functional dispersion indexes for above- and below-ground traits, and patterns of species association at a patch and neighborhood scale. Our results revealed that at the patch scale in intermediate levels of aridity, the dominant assembly process was within-site environmental heterogeneity. At the neighborhood scale, communities are assembled mainly through random processes. Nonetheless, in some communities, the dominant assembly process was competition via limiting similarity or exclusion of the weaker competitor, and these did not change along the gradient. Together, these results reveal that environmental heterogeneity and competition are the main drivers of plant community assembly in a hyper-arid environment.

Dormancy-break and germination requirements for seeds of the threatened Austral papaya (Carica chilensis).

Seed dormancy is one of the most important adaptive mechanisms in plants, optimizing germination, seedling emergence, and establishment to ensure these processes occur when environmental conditions are favorable for plant survival and growth. Endemic to rocky environments of the southern Atacama Desert, the Austral papaya (Carica chilensis) is the papaya species with the southernmost distribution within the Caricaceae, thriving in the most extreme environmental conditions. This threatened plant exhibits low natural regeneration, primarily attributed to low germination, yet no information regarding seed dormancy release is available. In this study, we investigated the dormancy-break and germination requirements of C. chilensis. We hypothesized that if C. chilensis seeds exhibit physiological dormancy, then seeds with reduced moisture content and those treated with chemicals or growth hormones would exhibit higher germination percentages and faster germination than control seeds akin to other members of Caricacea. Our results confirmed this prediction and revealed that ultra-drying (< 3% moisture content) and treating seeds with sulfuric acid, gibberellic acid, or potassium nitrate are the most effective methods for germinating C. chilensis. Consequently, we suggest using these treatments to propagate this threatened papaya species.

Anachronic Fruit Traits and Natural History Suggest Extinct Megafauna Herbivores as the Dispersers of an Endangered Tree.

Megafaunal seed dispersal syndrome refers to a group of traits attributed to the evolution of plants in the presence of large mammals. Present-day plants that bear these traits in areas where megafauna are absent are presumed to represent anachronic dispersal systems. Gomortega keule is an endangered tree species from a monotypic family (Gomortegaceae), endemic to Chile. Its fruit traits suggest adaptation to seed dispersal by large vertebrates; however, none are present today along its area of distribution. Here, we conducted a detailed revision on the fruit morphology of G. keule to examine whether its fruit traits fit a megafaunal dispersal syndrome. Additionally, we examined the fruit processing behavior of large domestic and captive wild animals fed with G. keule fruits, and its effect on germination. G. keule fruits had traits consistent with those of a Type 1 megafaunal fruit. Compared to intact, whole stones, seed germination probabilities decreased when fruits were handled by animals, suggesting that the seed was damaged during mastication and/or ingestion. Moreover, results from our feeding trials with elephants may also imply low efficiency of extinct gomphotheres as seed dispersers of this species. Our results also suggest that although domestic animals may disperse G. keule, it is unlikely that at present they can substitute the services of its original dispersers. Further investigation on seedling survival, local livestock management and forest management practices may help reinstate sexual regeneration in G. keule. Finally, integrating observations on fruit ecology and local people's knowledge with experimental data enriches our species-centered approach and may help to address regeneration problems in other endangered plants.

Predators and dispersers: Context-dependent outcomes of the interactions between rodents and a megafaunal fruit plant.

Many plant species bear fruits that suggest adaptation to seed dispersal by extinct megafauna. Present-day seed dispersal of these megafaunal plants is carried out by rodents, which can act as predators or dispersers; whether this interaction is primarily positive or negative can depend on the context. Here, we parameterized a stochastic model using data from the field and experimental arenas to estimate the effect of rodents on the recruitment of Myrcianthes coquimbensis -an Atacama Desert shrub with megafaunal fruits- and examine whether environmental conditions can alter the sign and strength of these rodent-plant interactions. We show that the outcome of these interactions is context-dependent: in wet conditions seed removal by rodents negatively impacts the recruitment probability of M. coquimbensis; in contrast, in dry conditions, the interaction with rodents increases recruitment success. In all cases, the strength of the effect of rodents on the recruitment success was determined mainly by their role as dispersers, which could be positive or negative. This study demonstrates that by caching seeds, rodents can be effective dispersers of a megafaunal fruit plant, but that the sign and magnitude of their effect on recruitment changes as a function of the environmental context in which the interaction occurs.

Conspecific plants are better 'nurses' than rocks: consistent results revealing intraspecific facilitation as a process that promotes establishment in a hyper-arid environment.

Harsh environmental conditions in arid ecosystems limit seedling recruitment to microhabitats under nurse structures, such as shrubs or rocks. These structures, however, do not necessarily afford the same benefits to plants because nurse rocks provide only physical nurse effects, whereas nurse plants can provide both physical and biological nurse effects. Nevertheless, if the nurse plant is a conspecific, the benefits it provides may be outweighed by higher mortality due to negative density-dependent processes; consequently, negative density-dependence is expected to limit plants from acting as nurses to their own seedlings. The degree to which an abiotic nurse may be more beneficial than a conspecific one remains largely unexplored. Here, we examine the role and elucidate the mechanisms by which conspecific plants and rocks promote plant establishment in a hyper-arid desert. For 4 years, we examined establishment patterns of Myrcianthes coquimbensis (Myrtaceae), a threatened desert shrub that recruits solely in rock cavities and under conspecific shrubs. Specifically, we characterized these microhabitats, as well as open interspaces for comparison, and conducted germination, seed removal and seedling survival experiments. Our results revealed that conspecific shrubs and nurse rocks modified environmental conditions in similar ways; soil and air temperatures were lower, and water availability was higher than in open interspaces. We found no evidence on negative density-dependent recruitment: seed removal was lowest and seedling emergence highest under conspecific plants, moreover seedling survival probabilities were similar in rock cavities and under conspecific plants. We conclude that the probability of establishment was highest under conspecific plants than in other microhabitats, contrasting what is expected under the Janzen-Connell recruitment model. We suggest that for species living in stressful environments, population regulation may be a function of positive density-dependence and intraspecific facilitation may be a process that promotes the persistence of some plant species within a community.

Fruit Size Determines the Role of Three Scatter-Hoarding Rodents as Dispersers or Seed Predators of a Fleshy-Fruited Atacama Desert Shrub.

Scatter-hoarding rodents can act as both predators and dispersers for many large-seeded plants because they cache seeds for future use, but occasionally forget them in sites with high survival and establishment probabilities. The most important fruit or seed trait influencing rodent foraging behavior is seed size; rodents prefer large seeds because they have higher nutritional content, but this preference can be counterbalanced by the higher costs of handling larger seeds. We designed a cafeteria experiment to assess whether fruit and seed size of Myrcianthes coquimbensis, an endangered desert shrub, influence the decision-making process during foraging by three species of scatter-hoarding rodents differing in body size: Abrothrix olivaceus, Phyllotis darwini and Octodon degus. We found that the size of fruits and seeds influenced foraging behavior in the three rodent species; the probability of a fruit being harvested and hoarded was higher for larger fruits than for smaller ones. Patterns of fruit size preference were not affected by rodent size; all species were able to hoard fruits within the entire range of sizes offered. Finally, fruit and seed size had no effect on the probability of seed predation, rodents typically ate only the fleshy pulp of the fruits offered and discarded whole, intact seeds. In conclusion, our results reveal that larger M. coquimbensis fruits have higher probabilities of being harvested, and ultimately of its seeds being hoarded and dispersed by scatter-hoarding rodents. As this plant has no other dispersers, rodents play an important role in its recruitment dynamics.

Contrasting responses to water-deficit among Encelia canescens populations distributed along an aridity gradient.

PREMISE OF THE STUDY: Drought is the most limiting factor for plant growth and recruitment in arid environments. For widespread species, however, plant responses to drought can vary across populations because environmental conditions can vary along the range of the species. Here, we assessed whether plants of Encelia canescens from different populations along an aridity gradient in the Atacama Desert respond differently to water-deficit conditions. METHODS: We conducted a common-garden experiment using plants grown from seeds from three populations distributed along an aridity gradient to test for differences in relative growth rate (RGR), biomass, root to shoot ratios, and photosynthesis between watered and water-deficit plants. Additionally, we examined the relationship between root to shoot ratios with RGR and total plant biomass along the gradient. KEY RESULTS: Water deficit affected root to shoot ratios, biomass, and RGR, but not photosynthesis. Populations varied in RGR and biomass; plants from the most arid population had higher RGRs, but lower biomass than those from the least arid population. In watered conditions, root to shoot ratios did not vary with RGR or biomass. Conversely, with the water deficit, root to shoot ratios were negatively and positively related to biomass and RGR, respectively. CONCLUSIONS: Response to water deficit differed among E. canescens populations; plants from the lowest rainfall environment adjusted root to shoot ratios, which may have allowed for equal biomass production across treatments. In contrast, plants from the wettest population did not adjust root to shoot ratios, but were reduced in biomass. These morphological and physiological changes to water availability showed that populations can use different strategies to cope with water deficit.

Context-dependent recruitment of Guettarda viburnoides in a Neotropical forest-savanna mosaic.

PREMISE OF THE STUDY: Seed dispersal is a key process for plant regeneration in tropical areas. Differences in disperser behavior result in a nonrandom distribution of seeds among habitats. Patterns of seed dispersal may cascade through the entire recruitment phase or uncoupling between developmental stages may occur such that patterns of seed dispersal are discordant with patterns of recruitment. Here, we analyzed how habitat and interannual variability affect the recruitment dynamics of a bird-dispersed tree, Guettarda viburnoides (RUB.). METHODS: Over 3 years, we determined the habitats where seeds of G. viburnoides are dispersed, and we experimentally quantified postdispersal seed predation, seedling emergence, and survival in four habitats of a forest-savanna mosaic in Bolivia. KEY RESULTS: Habitat affected seed dispersal, seed predation, and seedling emergence. The strength of postdispersal processes, however, varied between years and no consistent within-habitat pattern emerged. Uncoupling among different life-stages was observed across habitats, and spatial concordance was found between seed rain and sapling recruitment patterns. CONCLUSIONS: Habitat can affect seed dispersal, postdispersal processes and the recruitment dynamics of a Neotropical tree in a heterogeneous landscape. Additionally, our results show interannual variability in the strength of postdispersal processes, which leads to shifts in habitat suitability between years. Therefore, to better understand the role of site suitability for recruitment, we need to explicitly consider not only variation in habitat-specificity for dispersal and postdispersal processes, but also how this variation can shift under different environmental conditions-that is, the context dependence of suitability.

Seed dispersal by pulp consumers, not "legitimate" seed dispersers, increases Guettarda viburnoides population growth.

We examined the effect of seed dispersal by Purplish Jays (Cyanocorax cyanomelas; pulp consumers) and the Chestnut-eared Araçari (Pteroglossus castanotis; "legitimate" seed dispersers) on population growth of the small tree Guettarda viburnoides (Rubiaceae) in northeastern Bolivian savannas. Because each bird species differs with respect to feeding and post-feeding behavior, we hypothesized that seed dispersal by each species will contribute differently to the rate of increase of G. viburnoides, but that seed dispersal by either species will increase population growth when compared to a scenario with no seed dispersal. To examine the effects of individual dispersers on the future population size of G. viburnoides, we projected population growth rate using demographic models for G. viburnoides that explicitly incorporate data on quantitative and qualitative aspects of seed dispersal by each frugivore species. Our model suggests that seed dispersal by C. cyanomelas leads to positive population growth of G. viburnoides, whereas seed dispersal by P. castanotis has a detrimental effect on the population growth of this species. To our knowledge, this is the first study to report negative effects of a "legitimate" seed disperser on the population dynamics of the plant it consumes. Our results stress the importance of incorporating frugivore effects into population projection matrices, to allow a comprehensive analysis of the effectiveness of different dispersers for plant population dynamics.