Changes of woody plant interaction and spatial distribution between rocky and sandy soil areas in a semi-arid savanna, South Africa

We examined the impact of shrub-shrub interactions and soil type (rocky or sandy) on growth and spatial distribution of the two savanna shrub species Tarchonanthus camphoratus and Acacia mellifera. To explore plant interactions, we compared the size of juvenile and mature T. camphoratus shrubs between different locations (under the subcanopy of A. mellifera and in the open). Juvenile T. camphoratus shrubs had similar sizes regardless of location; however, in rocky soil, mature shrubs in the open were larger than those near A. mellifera, implying an inter-specific competitive effect of A. mellifera on T. camphoratus. Juvenile T. camphoratus shrubs grew faster in the sandy than in the rocky area. Furthermore, we used the Wiegand-Moloney O-ring statistics to explore the spatial distribution of T. camphoratus. T. camphoratus showed spatial aggregation, but in the rocky area T. camphoratus juveniles were positively associated with A. mellifera (indicating facilitation as the pattern-creating process), whereas in the sandy area they were positively associated with mature T. camphoratus shrubs (indicating seed dispersal as the pattern-creating process). T. camphoratus exhibited encroachment potential in the sandy area. We showed how spatial pattern analysis can help to explore processes determining woody plant spacing and recommend its further use.     

Parthenocarpy, attractiveness and seed predation by birds in Bursera morelensis

Parthenocarpy (production of seedless fruits) is a characteristic associated with a strategy to reduce seed predation by birds and to increase the attractiveness to frugivorous birds. Red cuajiote (Bursera morelensis) is an endemic tree species of Mexico that produces seeded and parthenocarpic fruits. The purpose of this work was to examine whether parthenocarpy is a factor that increases the number of bird visits to trees and if it is a strategy to diminish seeds predation by granivorous birds. We made field observations to determine the relation between the number of bird visits and fruit removal rate, to the total number of parthenocarpic fruits produced, and the number of birdpredated fruits to the number and proportion of parthenocarpic fruits in the trees. We found that in trees with a bigger amount of parthenocarpic fruits, both frugivoruos and granivorous birds made more visits; we then suggested that parthenocarpic fruits can act like a signal of attraction by means of a density and coloration effect. Also, we found that in trees with bigger proportions of parthenocarpic fruit, predation by granivorous birds is smaller. Parthenocarpy has an adaptative value for B.morelensis increasing bird visitation and reducing seed predation in the Tehuacan Valley.     

Dispersal of arbuscular mycorrhizal fungi and dark septate endophytes by Ctenomys cf. knighti (Rodentia) in the northern Monte Desert of Argentina

In arid and semi-arid environments, root-associated fungi may play a key role in plant communities (e.g., seedling establishment, nutrient acquisition, plant survival and heat tolerance). Several studies have shown the importance of small mammals as consumers and dispersal agents of mycorrhizal fungi spores in tropical and temperate ecosystems. However, little is known about the dispersal of infective propagules of endophytic fungi in arid and semi-arid ecosystems. This study analyzed the potential role of the subterranean rodent Ctenomys cf. knighti as a dispersal agent of root-associated fungi. In order to demonstrate this role, we analyzed: (1) the incidence of colonization of arbuscular mycorrhizae and dark septate endophytes on representative plant species of the Monte Desert; (2) the presence of fungal structures of arbuscular mycorrhizae and dark septate endophytes in C. cf. knighti fecal samples; and (3) the infectivity of the fungal propagules contained in the scat and their growth effects on nine native plant species. Data strongly suggest that this South American subterranean rodent may play a key role as a dispersal agent of arbuscular mycorrhizae and dark septate endophytes fungi in the arid environment of the northern Monte Desert of Argentina.     

Comparative study of seed germination and growth of Kochia prostrata and Kochia scoparia (Chenopodiaceae) under salinity

Soils of arid regions of Central Asia contain salts of different types that may differentially affect seed germination and plant development. We studied effect of NaCl, Na₂SO₄, 2NaCl + KCl + CaCl₂ and 2Na₂SO₄+K₂SO₄+MgSO₄ on germination of Kochia prostrata and Kochia scoparia seeds under a range of concentrations from 0.5 to 5% and at two constant temperature regimes +22 °C and +6 °C. The observed salt tolerance limit of germination at constant temperature +22 °C for both species was 5-6%, while at low temperature (+6 °C) this limit was 2%. The salt tolerance of young plants (before flowering) was 3% for NaCl. Low concentrations of sulfuric and mixed salts had a stimulating effect on seed germination in K. prostrata. Despite similarity of salt-tolerance limits the studied species showed a significant difference in seed recovery ability, i.e. the ability of ungerminated, salt-soaked seeds to germinate after transfer to fresh water. K. scoparia demonstrated a full germination recovery after seed transfer to distilled water while K. prostrata showed only a partial recovery.     

Neighbor competition modifies demographic responses of desert dune annuals to gradients in sand stability

A field experiment was conducted to test the hypothesis that competitive interactions affect demographic responses of desert dune annuals to gradients in sand stability. The experiment was based on a three-way factorial design with two types of neighbor treatments (complete neighbor removalvs. control), two types of habitats (stablevs. unstable sands) and two species which characterize sandy areas of the western Negev Desert (Neurada procumbensandBromus fasciculatus). Each of the eight treatment/habitat/species combinations was characterized by three demographic variables: seedling survival to reproduction, fruit number per reproductive plant and fruit number per seedling. All of these variables exhibited significant interactions between the effects of neighbor removal and habitat type, indicating that neighbor conditions were important in determining the observed patterns of demographic variation. Moreover, while plants ofNeurada procumbensgrowing without neighbors suffered more mortality on unstable sands, conspecific plants growing at natural density conditions suffered more mortality on stable sands. In the case ofBromus fasciculatus, individuals growing in the absence of competitive effects produced more fruits on stable sands, but conspecific plants growing at natural density conditions exhibited higher fruit production on unstable sands. These findings indicate that competitive effects not only modify, but may even reverse demographic responses of desert dune annuals to gradients in sand stability. Based on these results and evidence from two other studies it is suggested that seed dispersal from abiotically favorable into less favorable habitats may increase fitness of desert annuals by reducing the average effect of competition.     

Seed removal patterns in burned and unburned desert habitats: Implications for ecological restoration

In desert ecosystems, selective foraging by seed consumers affects distribution and composition of soil seed banks, influencing plant population dynamics. However, the roles of consumers in burned habitats where direct seeding is used during ecological restoration to replenish depleted seed banks have not been well established. We evaluated seed removal for nine seed species over 12 months in burned and unburned shrublands in the Mojave Desert, USA. Percentage of total seed removed was highest during spring (16% of offered seed) and summer (21%). Rodents removed much of the large-seeded Coleogyne ramosissima in both burned and unburned habitats, while seed removal of this species by ants was low in burned and moderate in unburned habitats. Ants removed the greatest amount of small-seeded species (Penstemon bicolor, Encelia farinosa, and Sphaeralcea ambigua) in unburned habitat, indicating that ants can exploit different seed masses. Seed removal imposes limitations on seed availability, particularly for large-seeded species, as both rodents and ants selected seeds of C. ramosissima. Successful restoration seeding projects in arid lands may require protecting seed from granivore pressure, and seed species selection and season of seeding warrant consideration to reduce seed loss.     

Factors controlling the regenerative cycle of Thymus funkii subsp. funkii in a semi-arid gypsum steppe: A seed bank dynamics perspective

We researched the soil seed bank dynamics of Thymus funkii subsp. funkii, a gypsovag shrub endemic to the southeastern Iberian Peninsula, to determine possible reasons for low recruitment in isolated populations growing on crusted gypsum soils. Recruitment-restriction hypotheses involving main factors in the seed bank dynamics were analysed, and a diagrammatic dynamics model drafted from a population perspective. Viable seed production averaged 362 per plant. Seed shadows resulting from primary dispersal were concentrated under the mother plant. There were two thyme-seed predators: coleopteran larvae (pre-dispersal phase) and Messor bouvieri ants (pre- and post-dispersal phases), which predated 16.4% and 3.8% of population yield, respectively. However, ants may simultaneously contribute to the spatial secondary dispersal, since they lost a small fraction of seeds they harvested. Seeds showed innate physiological dormancy to high summer temperatures, although this disappeared rapidly. The response of germination to other temperature × light conditions was wide and rapid. No permanent seed bank was detected in the soil. Seeds buried in the upper soil layer (2 cm) germinated highly and fast. In contrast, a significant fraction of seeds buried deeply (8 cm) remained viable for over one year. Germination in the field accounted for only 1.0% of the seed output, and seed death due to pathogen attack and aging was also low (3.0%). Seedling emergence was confined to autumn and showed a high correlation with total plant cover, irrespective of plant species. Seedling survival was very low. The small spatial correspondence between primary dispersal and seedling emergence patterns may reflect the importance of secondary dispersal in T. funkii seed bank dynamics. Hypotheses and consequences of such horizontal seed movements for conservation management are discussed.     

Soil seed bank longevity of the exotic annual grass Bromus rubens in the Mojave Desert,USA

Although arid environments are often considered among the least invaded terrestrial biomes, the impacts of exotic plant species can be severe and long lasting. Bromus rubens (red brome) is an exotic annual grass species in the Mojave Desert known to outcompete native plant species, alter habitat, and promote accumulation of fuel that contributes to increasing fire frequency and severity. We assessed longevity of the exotic B. rubens seeds in the soil by burying seeds at four depths (0, 2, 5, and 10 cm) and recovering seeds 6, 12, 18, and 24 months after burial. Seed viability was reduced with greater burial depth and greater time since burial. A relatively small proportion of seeds retained viability for two years, suggesting that while the B. rubens seed bank can be large, it is relatively short-lived. Although B. rubens apparently relies more on the annual production, dispersal, and germination of seeds than on a long-lived seed bank for its annual recruitment, the numerous seeds produced by individual plants indicate that even a small proportion of seeds remaining viable for more than a year can aid recruitment from the seed bank and is an important factor in understanding population dynamics.     

Role of competition in restoring resource poor arid systems dominated by invasive grasses

Understanding the role competition intensity and importance play in directing vegetation dynamics is central to developing restoration strategies, especially in resource poor environments. We hypothesized 1) competition would be intense among invasive and native species, but 2) competition would be unimportant in explaining variation in target plant biomass and survivorship relative to other factors driving these variables. We performed a two year addition series field experiment to quantify competition intensity and importance. Densities of two invasive (cheatgrass and medusahead) and two native (Sandberg’s bluegrass and bluebunch wheatgrass) species were arranged in monocultures and mixtures of two, three and four species, producing varying total densities and species proportions. Multiple linear regression models predicting individual plant biomass and survivorship were developed. Based on biomass, competition intensity coefficients ranged from −0.38 to 0.63 with R² < 0.06. All survivorship data produced poor fitting regression models (R² < 0.05). Our results suggest neither competition intensity nor importance influenced plant dominance in resource poor environments during the two years of establishment. Land managers may be more successful at restoration of resource poor ecosystems by overcoming abiotic barriers to plant establishment rather than focusing on plant-plant interactions.     

Mechanical shaking and soil water affect the growth of Psammochloa villosa in the Mu Us Sandland

The Mu Us Sandland is basically characterized by water shortage and high wind. Thus, wind-induced mechanical perturbation (MP) and soil water availability are likely to interact to affect plant growth. Since high water availability and MP can induce responses that are in the opposite direction, we hypothesized that MP effects on perennial grasses might be mitigated by increased soil water availability in the Mu Us Sandland. We counducted an experiment in which seedlings of Psammochloa villosa were subjected to two levels of MP (non-MP vs. MP 1 min d⁻¹) and two levels of water availability (200 ml d⁻¹vs. 400 ml d⁻¹) and measured three plant traits. MP significantly decreased plant height, total biomass, and root/shoot ratio. There were significant interactions between MP and soil water availability on plant height and root/shoot ratio. These findings imply that MP alone is a stressful factor for P. villosa and MP effects on its growth can be partially mitigated by increased soil water availability, and also suggest that P. villosa may respond to MP in a way that allows plants to survive in the windy semiarid environments.     

Restoring habitat for riparian birds in the lower Colorado River watershed: An example from the Las Vegas Wash, Nevada

The success of riparian restoration projects in the arid southwestern U.S. is often measured in terms of vegetation characteristics such as growth, cover, and structure. Among low-elevation riverine environments within the Colorado River watershed, restoration is typically conducted to improve degraded habitats for birds of conservation concern by replacing the exotic tamarisk (Tamarix ramosissima) with native cottonwoods (Populus spp.) and willows (Salix spp.). The working assumption for many restoration practitioners is that replacing exotic plants with native plants will improve habitat quality and will, therefore, benefit birds. Based on data collected at exotic and restored (i.e., native) sites along the Las Vegas Wash, Nevada, not all birds benefit from restoration. Broad measures of community benefit, including benefits to birds of conservation concern and riparian obligate/dependent birds, were not detected. There were, however, some species-specific benefits. Some environmental variables that were associated with exotic and native sites were significant in explaining the composition of the bird community. For example, the richness of forbs and grass-like plants (a proxy of soil moisture), invertebrate mass, and percent shade (a proxy for canopy characteristics) were important. Considering our results and depending on restoration goals, tamarisk replacement projects may not inherently provide benefits to birds.     

Patterns of seed occurrence in corsac and red fox diets in Mongolia

Corsac fox (Vulpes corsac) and red fox (Vulpes vulpes) diets often consist of small quantities of vegetation, including fruit and seeds that may represent important sources of energy and nutrition. However, details of the species consumed are few, but may provide insight into the role of foxes as seed dispersers. We analyzed scats of corsac (n = 408) and red foxes (n = 533) collected in an arid region of Mongolia from June 2006 to June 2007. We found 13 seed species in corsac scats, the most common being Allium polyrhizum, Asparagus gobicus, and Tribulus terrestris, and 12 species in red fox scats, the most common being Amygdalus pedunculata, Corispermum mongolicum, and T. terrestris. The proportion of scats with seeds varied significantly by season for both foxes. Seeds occurred most frequently during the breeding period, which corresponded with winter, and least frequently during the dispersal period in autumn for both species. We detected no differences in the proportion of scats with seeds between species seasonally. Our results suggest that fruit represents an important component of diet, especially during winter when availability of other prey items is low, and that both foxes may facilitate seed dispersal of some plant species.     

Speciation in gall-inducing thrips on Acacia in arid and non-arid areas of Australia

Habitat modification mediated by abiotic processes imposes consequences for the diversification of plants and plant specialists. Host plant diversification is expected to be dependent on environmental and ecological constraints that are fundamentally connected to physiography (physical geography). A phylogenetic approach is used to reconstruct ancestral host plant affiliation of gall-inducing thrips that specialize on Australian Acacia. Diversification rate models are applied to gall-thrips clades that are affiliated with different Acacia host lineages. Gall-thrips diversification is reconciled with contemporary patterns of species diversity relating to the different host lineages. Results indicate that diversification on host lineages that are mostly distributed in non-arid areas are best explained by a rate-constant model. However, this model disguised additive effects of rate-variable lineage accumulation that indicated diversification decreases in sub-clades within this larger group. Gall-thrips affiliated with hosts distributed mostly in arid areas were best characterized by rate-variable decreasing net diversification. The work infers parent gall-thrips lineages are less species diverse compared to daughter lineages that tend to be distributed away from the arid interior of Australia. Contrasting ecological and environmental interactions unique to parent and daughter lineages are suggested to influence the mode of speciation and phenotypic diversity represented by each.     

Seed dispersal and seed fate in Joshua tree (Yucca brevifolia)

Joshua tree (Yucca brevifolia) is a charismatic symbol of the Mojave Desert. Despite its familiarity, we know little about the reproduction of this species, including mechanisms of seed dispersal. Here we examine mechanisms of seed dispersal and resulting seed fate. We experimentally tracked fruit and seed removal and followed the fates of Joshua tree seeds using radioactive tracers. The majority of Joshua tree fruits monitored were taken directly from the tree canopy by white-tailed antelope squirrels, and seeds and fruits on the soil surface were quickly removed by animals. Rodents given seeds labeled with scandium-46 cached them between 0.1 cm and 4.1 cm deep. Seedling emergence was most common for seeds planted 1 cm deep, whereas seeds placed on the soil surface seldom germinated. Wind dispersal is unlikely because fruits and seeds lack adaptations for wind dispersal; wind speeds required to move Joshua tree seeds and fruits across the soil surface were higher than those typically found in the Mojave Desert. Further, rodents removed most seeds before abiotic burial was possible. We conclude that most Joshua tree seeds are dispersed by scatter hoarding by rodents, and that caches made by rodents are suitable sites for seedling emergence.     

Effects of overgrazing and rainfall variability on the dynamics of semiarid banded vegetation patterns: A simulation study with cellular automata

The effects of factors acting at small scale (grass/grass and grass/shrub biological interactions) and landscape scale (overgrazing and rainfall changes) on the development and permanence of banded vegetation patterns (VB) are assessed with a spatially explicit cellular automaton model. In particular, the influence of two environmental factors that are changing in many VB arid lands (rainfall and grazing pressures) is studied. Three rainfall regimes (representing 0.5, 1.0, and 2.0 times the long-term rainfall average), five grazing intensities, two types (reversible and irreversible) of grazing disturbance, two grazing periodicities (chronic and pulse), two levels of grass colonization ability, positive and negative interactions between shrubs and grasses, and the efficiency of endozoochorous seed dispersal are simulated.The results show that the permanence of undisturbed VB depends on the interaction of two factors, rainfall regime and grass colonization ability. Type and intensity of grazing also modify VB cover and permanence; furthermore, long-term overgrazing may convert mixed grass/shrub plant communities to pure scrublands dominated by endozoochorous-dispersed shrubs due to competitive interactions between shrubs and grasses.Besides providing an adequate representation of the system’s dynamics, the model is a useful tool that may be used to explore the consequences of climate change on management scenarios.     

Xylem efficiency vs. safety: Acclimation to drought of seedling root anatomy for six Patagonian shrub species

With the aim to seek evidences on the possible trade off between hydraulic efficiency and hydraulic safety, from both an evolutionary and an acclimation point of view, we compared root xylem anatomy and derived functional properties for seedlings of six Patagonian shrub species grown in a greenhouse under two levels of water availability (control and drought). Root central cylinder area, vessel diameter (b) and double-wall thickness (t) were measured; from these data, the sum of vessel radii to the fourth power and wall strength [(t/b)²] were calculated as indicators of hydraulic efficiency and safety respectively. Across species, we observed only a weak negative correlation between hydraulic efficiency and hydraulic safety. Within species, Lycium chilense, the species with the most mesic leaves of the group, showed significant acclimation to drought for both functional attributes, lowering efficiency and increasing safety by developing a higher proportion of small and more resistant xylem vessels.     

Predispersal reproductive ecology of an arid land crucifer,Moricandia moricandioides: effect of mammal herbivory on seed production

In this paper the effect of herbivory on the seed production ofMoricandia moricandioides(Cruciferae), an annual species living in arid habitats of south-eastern Spain, is analysed. Firstly, the flowering and fruiting phenology and the composition and structure of the pollinator guild is described, in order to compare the overall effect of mutualistic speciesvs. that of antagonistic species onM. moricandioidesseed production. The loss of female reproductive success due to each herbivore species attacking the plant is then quantified, in an attempt to differentiate the effects on plant seed production of each herbivorous species. The results indicate that only antagonistic organisms interacting withM. moricandioidesinfluenced the female fitness of this plant species. In fact, pollinator abundance or activity did not correlate with the number of seeds dispersed by the plants. In addition, of all the herbivore species, only sheep activity significantly influenced seed production ofM. moricandioidesindividuals, the effect of the remaining crucivorous herbivores being cancelled by the stronger herbivory of sheep. The results of this study suggest that herbivory by sheep, in addition to decreasing the seed production ofM. moricandioides, could be an important determinant of the current distribution pattern of this crucifer species on the ramblas of the Baza basin.     

Seed germination of five mountain steppe species of Central Asia

This paper presents data on the germination traits of five perennials (Allium polyrrhizum, Agropyron cristatum, Arenaria meyeri, Artemisia frigida and Artemisia santolinifolia) widespread in the mountain steppes of southern Mongolia. Germination and seed viability were assessed at three alternate temperatures (8/4 °C, 20/10 °C, 32/20 °C), three levels of osmotic stress (deionized water; −0.5 MPa, −1 MPa Mannitol solution), and under conditions of alternate light/darkness versus complete darkness. The results of a factorial ANOVA with treatments and species as main effects showed that all five species germinated best at higher temperatures, with only Agropyron cristatum showing some seed mortality. Osmotic stress reduced seed viability and total germination in all five species. Darkness had no influence on viability, but positively affected seed germination of Allium polyrrhizum and Agropyron cristatum. We therefore conclude that, in the field, germination of all five species is mainly controlled by ambient temperatures and water availability, both of which drop towards the end of summer when dispersal takes place and effectively delay seedling recruitment until the next vegetation period.     

Passage time, viability, and germination of seeds ingested by foxes

Seed passage through the gut of vertebrates is critical for endozoochorus seed dispersal because it may influence seed viability and germination capacity. Foxes are important fruit-eaters and presumed seed dispersers in the Chaco ecoregion. We experimentally assessed the seed passage time (SPT) and its effect on seed viability and germination in two fox species (Pseudalopex gymnocercus and Cerdocyon thous) that occur in the Chaco. Passage time was measured using both artificial seeds of three sizes and wild seeds of four native fleshy-fruit species commonly consumed by foxes (Ziziphus mistol, Acacia aroma, Celtis tala, and Syagrus romanzoffiana). The average SPT was 6.8 h for the Pampa fox and 7.7 h for the crab-eating fox. Passage time did not differ either among seeds of different sizes or between fox species. Passage through digestive tract of both fox species did not affect seed survival, whereas germination rate was slightly improved. Our data suggest that both fox species are legitimate seed dispersers of the four species studied, having a positive effect on both germination rate and dispersal distance from the parent plant.     

Dieback affects forest structure in a dry Afromontane forest in northern Ethiopia

Forests are highly susceptible to dieback under ongoing climate warming. In degraded forests, dead standing trees, or snags, have become such prominent features that they should be taken into account when setting management interventions. This study investigated (1) the extent and spatial pattern of standing dead stems of Juniperus procera and Olea europaea subsp. cuspidata along an elevational gradient, and (2) the effect of dieback on forest stand structure. We quantified abundance, size, and spatial pattern of tree dieback in 57 plots (50 m × 50 m) established at 100 m intervals along five transects. The snag density and basal area (mean ± SE) of the two species combined were 147 ± 23 stems ha⁻¹ and 5.35 ± 0.81 m² ha⁻¹, respectively. The percentages of snags were extremely high for both J. procera (57 ± 7%) and O. europaea subsp. cuspidata (60 ± 5%), but showed a decreasing trend with increasing elevation suggesting that restoration is even more urgent at the lower elevations. Snags of the two species accounted for 31 and 45% of total stand density and basal area, respectively. Living stems exhibited truncated inverse-J-shaped diameter and height class distributions, indicating serious regeneration problems of these foundation species in the study area. In addition to direct interventions to assist recruitment of climax tree species, sites with high dieback would probably benefit from snag reduction to prevent fire incidents in the remaining dry Afromontane forests in northern Ethiopia.