Spatial patterns of desert annuals in relation to shrub effects on soil moisture

Questions: What are the effects of a shrub (Haloxylon ammodendron) on spatial patterns of soil moisture in different seasons? How does productivity of understorey annuals respond to these effects? Are such effects always positive for annuals under shrubs? Location: South Gurbantunggut Desert, northwest China. Methods: Using geostatistics, we explored seasonal patterns of topsoil moisture in a 12 × 9‐m plot over the growing season. To determine spatial patterns of understorey annuals in response to H. ammodendron presence, biomass of annuals was recorded in four 0.2 × 5.0‐m transects from the centre of a shrub to the space between shrubs (interspace). We also investigated vertical distribution of root biomass for annuals and soil moisture dynamics across soil profiles in shrub‐canopied areas and interspaces. Results: Topsoil moisture changed from autocorrelation in the wet spring to random structure in the dry season, while soil moisture below 20 cm was higher in shrub‐canopied areas. Across all microhabitats, soil moisture in upper soil layers was higher than in deeper soil layers during the spring wet season, but lower during summer drought. Topsoil was close to air‐dry during the dry season and developed a ‘dry sand layer’ that reduced evaporative loss of soil water from deeper layers recharged by snowmelt in spring. Aboveground biomass of understorey annuals was lowest adjacent to shrub stems and peaked at the shrub margin, forming a ‘ring’ of high herbaceous productivity surrounding individual shrubs. To acclimate to drier conditions, annuals in interspaces invested more root biomass in deeper soil with a root/shoot ratio (R/S) twice that in canopied areas. Conclusions: Positive and negative effects of shrubs on understorey plants in arid ecosystems are commonly related to nature of the environmental stress and tested species. Our results suggest there is also microhabitat‐dependence in the Gurbantunggut Desert. Soil water under H. ammodendron is seasonally enriched in topsoil and deeper layers. Understorey annuals respond to the effect of shrubs on soil water availability with lower R/S and less root biomass in deeper soil layers and develop a ‘ring’ of high productivity at the shrub patch margin where positive and negative effects of shrubs are balanced.     

Impact of shrub canopies on understorey vegetation in western Eurasian tundra

Question: How does the composition and species richness of understorey vegetation associate with changing abundance of deciduous shrub canopies? What are the species‐specific associations between shrubs and understorey plants? Location: Tundra habitats along an over 1000‐km long range, spanning from NW Fennoscandia to the Yamal Peninsula in northwest Russia. Methods: The data from 758 vegetation sample plots from 12 sites comprised cover estimates of all plant species, including bryophytes and lichens, and canopy height of deciduous shrubs. The relationships between shrub volume and cover of plant groups and species richness of vegetation were investigated. In addition, species‐specific associations between understorey species and shrub volume were analysed. Results: Shrub abundance was shown to be associated with the composition of understorey vegetation, and the association patterns were consistent across the study sites. Increased forb cover was positively associated with shrub volume, whereas bryophyte, lichen, dwarf shrub and graminoid cover decreased in association with increasing volume of deciduous shrubs. The total species richness of vegetation declined with increasing shrub volume. Conclusions: The results suggest that an increase of shrubs – due to climatic warming or a decrease in grazing pressure – is likely to have strong effects on plant–plant interactions and lead to a decrease in the diversity of understorey vegetation.     

Remnant shrubs in Mediterranean semi-arid steppes: effects of shrub size, abiotic factors and species identity on understorey richness and occurrence

Isolated shrub patches are a key element for community structure and dynamics in semi-arid ecosystems, and may act as “hot spots” of understorey species diversity. Despite its importance, the relationships between shrub patch characteristics and understorey species richness, and the relative importance of these characteristics against other factors driving understorey species richness, are still poorly understood. We studied perennial species richness under the canopy of late-successional shrubs in semi-arid Stipa tenacissima steppes of SE Spain. The main aim of this study was to evaluate the relationship between shrub size and understorey richness, and to test if this relationship is modified by species identity. We also aimed to know the relative importance of shrub patch characteristics and abiotic factors as predictors of the distribution of understorey species under shrub canopies. Altitude and geographical co-ordinates were able to significantly explain the patterns of shrub occurrence and abundance in the study area. Understorey species richness was significantly related to the size of individual shrubs according to a simple power relationship for all evaluated species. Slope values of linear regressions with log-transformed data, ranging between 0.22 and 0.37, did not differ between species. Site characteristics, patch characteristics and patch species identity significantly explained the occurrence of species on patches. However, the relative importance of site characteristics was higher than that of patch characteristics and species identity in explaining these patterns. Our results agree with the general expectations of the theory of island biogeography and complement previous studies that emphasise the importance of late-successional shrubs in semi-arid Stipa steppes. Environmental management activities within these steppes should promote the conservation of remnant shrubs, as well as its introduction with restoration activities.     

Spatial segregation of plant species caused by Retama sphaerocarpa influence in a Mediterranean pasture: a perspective from the soil seed bank

Retama sphaerocarpa is a leguminous shrub whose important role in the semi-arid regions of south-eastern Spain has already been assessed: shrub canopies reduce light intensity, and thus evapotranspiration; also their extensive radical system take water and nutrients from great volumes of soil, concentrating them in the understorey. Consequently, subcanopy vegetation benefits from these facilitation processes, increasing its productivity. However, these shrublands have been rarely studied at a broader scale, i.e. as a savannah-like system composed of a variable number of shrubs scattered in a herbaceous matrix. As the microenvironmental conditions associated to the understorey are rather different from those of the open spaces among shrubs, species composition of the herbaceous matrix is expected to change accordingly. Thus, R. sphaerocarpa would be an important and still unknown source of spatial heterogeneity to the system. Our main purpose was to evaluate, through the soil seed bank, the heterogeneity in the herbaceous community induced by this shrub species. Soil samples were collected around adult shrubs from three positions relative to the canopy: near the centre of the shrubs, at the edge of the understorey, and completely outside the canopy. Floristic composition was evaluated by germination under greenhouse conditions. The results show that each position has a different floristic composition, characterised by a group of different species. The herbaceous species associated with the external position have functional traits which enable them to resist water stress and herbivore pressure, such as hairs, CAM metabolism, early flowering, horizontal growth or tiny stature. The species associated with the central position lack those traits, and are more competitive in more mesic environments, rich in nitrogen. The lowest number of seedlings and species was found in the internal position, suggesting that in our study the facilitation process may have less importance for community dynamics due to less stressful environmental conditions.     

Disentangling direct and indirect effects of a legume shrub on its understorey community

Direct and indirect interactions among plants contribute to shape community composition through above‐ and belowground processes. However, we have not disentangled yet the direct and indirect soil and canopy effects of dominants on understorey species. We addressed this issue in a semi‐arid system from southeast Spain dominated by the legume shrub Retama sphaerocarpa. During a year with an exceptionally dry spring, we removed the shrub canopy to quantify aboveground effects and compared removed‐canopy plots to open plots between shrubs to quantify soil effects, both with and without watering. We added a grass removal treatment in order to separate direct from indirect shrub effects and quantified biomass, abundance, richness and composition of the forb functional group. With watering, changes in forb biomass were primarily driven by indirect shrub effects, with contrasting negative soil and positive aboveground indirect effects; changes in forb abundance and composition were more influenced by direct shrub soil effects with contrasting species composition between open and Retama patches. As community composition was different between open and Retama patches the indirect effects of Retama on forb species did not concern forbs from the open community but forbs from Retama patches. Indirect effects are, thus, important at the functional group level rather than at the species level. Without watering, there were no significant interactions. Changes in species richness between treatments were weak and seldom significant. We conclude that shrub effects on understorey forbs are primarily due to their influence on soil properties, directly affecting forb species composition but indirectly affecting the biomass of the forbs of the Retama patches, and only with sufficient water.     

猕猴桃属植物狗枣子的修订

作者将海棠猕猴桃Actinidia maloides H.L.Li,心叶海棠猕猴桃A.maloides H.L.Lif.cordata C.F.Liang合并于狗枣子(Actinidia kollmikta (Maxim. and Rupr.)Maxim.)中作为异名,同时对狗枣子及其近缘种软枣猕猴桃A.arguta (Side. and Zucc.)Planch.ex Miq,木天蓼A.polygama(Sieb. and Zucc.)Maxim.和四数猕猴桃A.arguta (Sieb. and Zucc.)Planch.ex Miq,木天蓼A.polygama(Sied.and Zucc.)Maxim.和四数猕猴桃A.tetramera Maxim.的形态学特征进行了辨析。     

Seed Bank and Understorey Species Composition in a Semi-arid Environment: The Effect of Shrub Age and Rainfall

Understorey vegetation in patches of Retama sphaerocarpa shrubsin semi-arid environments is dependent on the overstorey shrublife history. Community structure changes with shrub age asa result of physical amelioration of environmental conditionsby the canopy and organic matter accumulation in the soil. Weinvestigated the effect of the canopy on understorey speciesdiversity in the field and its relationships with the soil seedbank under 50 shrubs from 5 to 25+ years old, and compared speciescomposition in the field in a wet and a dry year. Species compositionof the soil seed bank under R. sphaerocarpa shrubs did not differsignificantly with shrub age, but seed density increased asthe shrubs aged. In the field, community composition changedwith shrub age, increasing species richness in a process thatdepended on the amount of spring rainfall. Our results suggestthat the soil seed bank is rather uniform and that the shrubcanopy strongly selects which species appear in the understorey.There were seeds of many species present under both young andold shrubs but which only established under old shrubs. Thisshowed dispersal was not limiting species abundance and suggestedthat the canopy was an important sorting factor for speciespresent in the understorey. Less frequent species contributedthe most to patch diversity, and rainfall effectively controlledspecies emergence. Understorey community composition dependedon multiple interspecific interactions, such as facilitationby the shrub and competition from neighbours, as well as ondispersal processes. Facilitation in this environment is a keyfeature in the structuring of plant communities and in governingecosystem functioning. Copyright 2000 Annals of Botany Company Community structure, competition, dispersal, facilitation, species composition, rainfall variability, Retama sphaerocarpa, seed bank, semi-arid environments     

Responses of different herb life-history groups to a dominant shrub species along a dune stabilization gradient

Shrubs play a pivotal role in reversing desertification and in promoting ecological rehabilitation in severe environments. However, how the interactions between shrubs and their understorey species change during restoration remains unclear. Here, we report the results of an observational study conducted in the Mu Us Desert of northern China. This study explored how dune stabilization and the size of individual shrubs affect shrub–herb interactions. In particular, we aimed to determine how different life-history groups and performance indicators (e.g., biomass and richness) of subordinate species respond to shrub–herb interactions during dune stabilization. The shrub Artemisia ordosica had positive effects on understorey species in this dune system. The ability of the shrub to promote the growth of understorey species increased with dune stabilization, but decreased from small to large shrubs. This effect was due to an increase in the relative abundance and biomass of perennials and their higher sensitivity to the positive effects of the shrubs. In contrast, the ability of shrubs to improve the richness of understorey species decreased with dune stabilization, but increased from small to large shrubs. This effect occurred because perennials suffered high strain during recruitment in disturbed open patches of the semi-fixed dunes, particularly below small shrubs. Our results support the theory claiming that communities are sets of hidden interaction groups that have contrasting responses (neutral for annuals, but facilitative for perennials) to dominant neighbors in a single community, depending on their functional strategies. Furthermore, our study highlights the high efficiency of A. ordosica in facilitating ecological restoration of dry and disturbed sandy communities.     

Seedling and Sprout Response to Slash-and-Burn Agriculture in a Tropical Deciduous Forest1

Survival and growth of seedlings and sprouts were assessed in three plots for 16 mo following the slashing and burning of a tropical deciduous forest in Jalisco, Mexico. We encountered a total of 47 species: 21 seedling species and 35 sprout species. Calliandra formosa and Piptadenia flava were the most common seedling species; Bursera arborea, Cordia alliodora, and Piptadenia constricta were the most common sprouts. Colubrina triflora, Diphysa occidentalis, and Cnidoscolus spinosus had limited sprouting ability. Twenty-six species were represented by one seedling or one sprout. Thirty-eight percent of the seedlings were tree species, 59 percent were shrub species, and 2 percent were vines species. In contrast, 86 percent of the sprouts were from trees and 13 percent from shrubs. One year after the initial measurements, 29 percent of the seedlings and 13 percent of the sprouts were dead. Each of the seedling means (number of stems/individual, height and diameter of the tallest stem, and elliptical crown area) was significantly smaller (P < 0.05) than that of sprouts at all three measurement periods, but relative growth rates were similar. Total canopy area of seedlings had a larger relative increase than did the canopy area of sprouts. The presence of seedlings increased species diversity compared to calculated diversity excluding seedlings. Timing of fruit dispersal in relation to the date of burning and the high number of Leguminosae species in the forest appeared to favor seedling establishment for some species.     

Spatial patterns in a two-tiered semi-arid shrubland in southeastern Spain

Abstract. Single species and bivariate distribution patterns in a semi-arid shrubland in southeastern Spain, dominated by the tall leguminous shrub Retama sphaerocarpa, were investigated by second-order spatial analysis based on Ripley's K-function. Shrubs were significantly clumped because of a strong association of dwarf shrubs, mostly Artemisia barrelieri, under the canopy of Retama. Retama shrubs were randomly distributed, but when different size-classes were analysed separately, the pattern changed from significantly clumped to random and then to regular with increasing canopy diameter, suggesting increasing intraspecific competition with shrub size. Artemisia was significantly clumped at all scales because of aggregation under the canopy of large Retama shrubs. The association between the species became stronger with increasing canopy diameter of Retama shrubs, suggesting that facilitation prevailed over interspecific competition because of niche separation in different tiers, both above and below ground. Retama shrub size thus determined both the type of pattern for its own size class and tier, and the scale and intensity of the association with its understorey shrubs.     

Disentangling facilitation and seed dispersal from environmental heterogeneity as mechanisms generating associations between savanna plants

Question: How can we disentangle facilitation and seed dispersal from environmental heterogeneity as mechanisms causing spatial associations of plant species? Location: Semi‐arid savanna in the Kimberley Thorn Bushveld, South Africa. Methods: We developed a two‐step protocol for the statistical differentiation of association‐promoting mechanisms in plants based on the Acacia erioloba–Grewia flava association. Individuals of the savanna shrub G. flava and the tree A. erioloba were mapped on four study plots. Disentangling the mechanism causing the association of G. flava and A. erioloba involved tests of three spatial and one non‐spatial null model. The spatial null models include homogeneous and heterogeneous Poisson processes for spatial randomness based on the bivariate spatial point patterns of the four plots. With the non‐spatial analysis, we determined the relationship between the canopy diameter of A. erioloba trees and presence or absence of G. flava shrubs in the tree understorey to find whether shrub presence requires a minimum tree canopy diameter. Results: We first showed a significant positive spatial association of the two species. Thereafter, the non‐spatial analysis supported an exclusion of environmental heterogeneity as the sole cause of this positive association. We found a minimum tree size under which no G. flava shrubs occurred. Conclusions: Our two‐step analysis showed that it is unlikely that heterogeneous environmental conditions caused the spatial association of A. erioloba and G. flava. Instead, this association may have been caused by seed dispersal and/or facilitation (e.g. caused by hydraulic lift and/or nitrogen fixation by the host tree).     

Increase in nonnative understorey vegetation cover after nonnative conifer removal and passive restoration

Nonnative conifers are widespread in the southern hemisphere, where their use as plantation species has led to adverse ecosystem impacts sometimes intensified by invasion. Mechanical removal is a common strategy used to reduce or eliminate the negative impacts of nonnative conifers, and encourage native regeneration. However, a variety of factors may preclude active ecological restoration following removal. As a result, passive restoration – unassisted natural vegetation regeneration – is common following conifer removal. We asked, ‘what is the response of understorey cover to removal of nonnative conifer stands followed by passive restoration?' We sampled understorey cover in three site types: two‐ to ten‐year‐old clearcuts, native forest and current plantations. We then grouped understorey species by origin (native/nonnative) and growth form, and compared proportion and per cent cover of these groups as well as of bare ground and litter between the three site types. For clearcuts, we also analysed the effect of time since clearcut on the studied variables. We found that clearcuts had a significantly higher average proportion of nonnative understorey vegetation cover than native forest sites, where nonnative vegetation was nearly absent. The understorey of clearcut sites also averaged more overall vegetation cover and more nonnative vegetation cover (in particular nonnative shrubs and herbaceous species) than either plantation or native forest sites. Notably, 99% of nonnative shrub cover in clearcuts was the invasive nonnative species Scotch broom (Cytisus scoparius). After ten years of passive recovery since clearcutting, the proportion of understorey vegetation cover that is native has not increased and remains far below the proportion observed in native forest sites. Reduced natural regeneration capacity of the native ecosystem, presence of invasive species in the surrounding landscape and land‐use legacies from plantation forestry may inhibit native vegetation recovery and benefit opportunistic invasives, limiting the effectiveness of passive restoration in this context. Abstract in Spanish is available with online material.     

Water relations of understorey shrubs in a Banksia woodland,Swan Coastal Plain,Western Australia

Abstract Diurnal and seasonal water relations were measured in understorey species from a Banksia woodland. The shrubs exhibited various responses to summer drought. Stirlingia latifolia had high xylem pressure potential and transpiration in late summer. Adenanthos cygnorum maintained high xylem pressure potential year round with dawn values around ? 0.3 MPa and minimum values around ?1.3 MPa, but showed severe restriction of transpiration in late summer. Eremaea pauciflora and Jacksonia floribunda had high transpiration and xylem pressure potential levels in early summer, but exhibited water stress in late summer when transpiration rates were low and minimum xylem pressure potentials were as low as ? 5.5 MPa. Late summer xylem pressure potentials in 27 other shrub species were, in general, inversely related to root system depth with minimum values below ? 5.0 MPa in some species. The water relations of S. latifolia, E. pauciflora and J. floribunda indicated a phreatophytic habit: all possessed deep roots of sufficient size to reach groundwater that was located 6–7 m deep at the study site. Stirlingia latifolia functioned phreatophytically year round, while E. pauciflora and J. floribunda were phreatophytes until the falling water table carried ground-water beyond the reach of their roots in late summer. However, most understorey species depended on soil-stored water. Water use by the understorey was greatest in early summer.     

Shrubs as a source of spatial heterogeneity—the case of Retama sphaerocarpa in Mediterranean pastures of central Spain

The processes underlying the positive effect exerted by woody species on other plant species can be grouped into a physical effect due to the canopy itself (mainly amelioration of temperature and light extremes), and an edaphic effect (nutrient enrichment). Both groups of effects are present in the savanna-like systems originated by Retama sphaerocarpa in the Iberian Peninsula. The consequences of these shrubs for the spatial heterogeneity induced in the herbaceous community are not well known. The herbaceous community was sampled by means of radial transects around 20 adult shrubs, from the canopy centre to the open areas among the shrubs, both northwards and southwards. Floristic composition, diversity and pasture mean height change both along the inside–outside gradient, and from north to south. As a nutrient enriched environment, the centre of the understorey showed the lowest species richness and evenness, and the highest standing crop. In contrast, the environmentally stressful open areas showed the highest number of species and evenness, and the lowest standing crop. This general effect is only partially due to nutrient availability, and is not homogeneous around the shrub, as open area conditions penetrate into the understorey driving the herbaceous community towards that of open areas. An individual shrub effect is also suggested by our results. Different conditions are thus provided by R. sphaerocarpa understoreys, favouring different subsets of species from the general pool, and therefore defining shrub canopies as sources of spatial heterogeneity in the whole savanna-like system.     

The effect of fire interval on post‐fire understorey communities in Yellowstone National Park

Questions: How does the time interval between subsequent stand‐replacing fire events affect post‐fire understorey cover and composition following the recent event? How important is fire interval relative to broad‐ or local‐scale environmental variability in structuring post‐fire understorey communities? Location: Subalpine plateaus of Yellowstone National Park (USA) that burned in 1988. Methods: In 2000, we sampled understorey cover and Pinus contorta density in pairs of 12–yr old stands at 25 locations. In each pair, the previous fire interval was either short (7–100 yr) or long (100–395 yr). We analysed variation in understorey species richness, total cover, and cover of functional groups both between site pairs (using paired t‐tests) and across sites that experienced the short fire intervals (using regression and ordination). We regressed three principal components to assess the relative importance of disturbance and broad or local environmental variability on post‐fire understorey cover and richness. Results: Between paired plots, annuals were less abundant and fire‐intolerant species (mostly slow‐growing shrubs) were more abundant following long intervals between prior fires. However, mean total cover and richness did not vary between paired interval classes. Across a gradient of fire intervals ranging from 7–100 yr, total cover, species richness, and the cover of annuals and nitrogen‐fixing species all declined while the abundance of shrubs and fire‐intolerant species increased. The few exotics showed no response to fire interval. Across all sites, broad‐scale variability related to elevation influenced total cover and richness more than fire interval. Conclusions: Significant variation in fire intervals had only minor effects on post‐fire understorey communities following the 1988 fires in Yellowstone National Park.     

Influence of an overstorey tree (Prosopis glandulosa) on associated shrubs in a savanna parkland: implications for patch dynamics

The arborescent legume, honey mesquite (Prosopis glandulosa), appears to play a central role in patch dynamics of southern Texas savannas by modifying soils and microclimate and by facilitating the ingress, establishment and/or growth of shrubs in its understorey. As an indirect test for the occurrence and persistence of facilitation in mature shrub clusters (patches), we examined the gas exchange, water relations and production of associated shrubs growing in patches where a Prosopis overstorey was present and in patches where Prosopis had succumbed to natural mortality. Surface (0–10 cm) soils associated with shrub patches were enriched in total [N] and [C] compared to soils of neighboring herbaceous zones. However, there were no detectable differences in soil [N] or [C] in patches with and without Prosopis. Foliar [N] and biomass of various shrub species were also statistically comparable for patches with and without Prosopis. These results are in accordance with other studies that indicate the nutrient legacy associated with Prosopis occupation of a patch may persist for decades after its demise. In comparison to plants growing in the absence of Prosopis, leaf water potentials (predawn and midday), and net photosynthesis and water vapor conductance (morning and midday) of outer-canopy sunlit leaves over an annual growth cycle were comparable for two common evergreen shrubs, Zanthoxylum fagara and Berberis trifoliolata, growing in patches with a live Prosopis. These findings indicate that the presence of Prosopis was not enhancing the growth or activity of mature understorey shrubs; facilitation may, therefore, be important only during early stages of cluster development. In addition, we found no indication that the loss of Prosopis has initiated a downward phase in a cyclic succession of patch initiation, growth and death. Rather, the understorey shrubs appear to be able to maintain growth and productivity in the absence of a Prosopis overstorey, and may, therefore, represent persistent components of woody patches on these savanna landscapes.     

Severe dry winter affects plant phenology and carbon balance of a cork oak woodland understorey

Mediterranean climates are prone to a great variation in yearly precipitation. The effects on ecosystem will depend on the severity and timing of droughts. In this study we questioned how an extreme dry winter affects the carbon flux in the understorey of a cork oak woodland? What is the seasonal contribution of understorey vegetation to ecosystem productivity?We used closed-system portable chambers to measure CO₂ exchange of the dominant shrub species (Cistus salviifolius, Cistus crispus and Ulex airensis), of the herbaceous layer and on bare soil in a cork oak woodland in central Portugal during the dry winter year of 2012. Shoot growth, leaf shedding, flower and fruit setting, above and belowground plant biomass were measured as well as seasonal leaf water potential. Eddy-covariance and micrometeorological data together with CO₂ exchange measurements were used to access the understorey species contribution to ecosystem gross primary productivity (GPP).The herbaceous layer productivity was severely affected by the dry winter, with half of the yearly maximum aboveground biomass in comparison with the 6 years site average. The semi-deciduous and evergreen shrubs showed desynchronized phenophases and lagged carbon uptake maxima. Whereas shallow-root shrubs exhibited opportunistic characteristics in exploiting the understorey light and water resources, deep rooted shrubs showed better water status but considerably lower assimilation rates. The contribution of understorey vegetation to ecosystem GPP was lower during summer with 14% and maximum during late spring, concomitantly with the lowest tree productivity due to tree canopy renewal. The herbaceous vegetation contribution to ecosystem GPP never exceeded 6% during this dry year stressing its sensitivity to winter and spring precipitation.Although shrubs are more resilient to precipitation variability when compared with the herbaceous vegetation, the contribution of the understorey vegetation to ecosystem GPP can be quite variable and will ultimately depend of tree density and canopy cover.     

Shrub facilitation drives tree establishment in a semiarid fog‐dependent ecosystem

Questions

The exceptional occurrence of tall rain forest patches on foggy coastal mountaintops, surrounded by extensive xerophytic shrublands, suggests an important role of plant–plant interactions in the origin and persistence of these patches in semi‐arid Chile. We asked whether facilitation by shrubs can explain the growth and survival of rain forest tree species, and whether shrub effects depend on the identity of the shrub species itself, the drought tolerance of the tree species and the position of shrubs in regard to wind direction.

Location

Open area–shrubland–forest matrix, Fray Jorge Forest National Park, Chile.

Methods

We recorded survival after 12 years of a ~3600 tree saplings plantation (originally ~30‐cm tall individuals) of Aextoxicon punctatum, Myrceugenia correifolia and Drimys winteri placed outside forests, beneath the shrub Baccharis vernalis, and in open (shrub‐free) areas. We assessed the effects of neighbouring shrubs and soil humidity on survival and growth along a gradient related to the direction of fog movement.

Results

B. vernalis had a clear facilitative effect on tree establishment and survival since, after ~12 years, saplings only survived beneath the shrub canopy. Long‐term survival strongly depended on tree species identity, drought tolerance and position along the soil moisture gradient, with higher survival of A. punctatum (>35%) and M. correifolia (>14%) at sites on wind‐ and fog‐exposed shrubland areas. Sites occupied by the shrub Aristeguietia salvia were unsuitable for trees, presumably due to drier conditions than under B. vernalis.

Conclusions

Interactions between shrubs and fog‐dependent tree species in dry areas revealed a strong, long‐lasting facilitation effect on planted tree's survival and growth. Shrubs acted as benefactors, providing sites suitable for tree growth. Sapling mortality in the shrubland interior was caused by lower soil moisture, the consequence of lower fog loads in the air and thus insufficient facilitation. While B. vernalis was a key ecosystem engineer (nurse) and intercepted fog water that dripped to trees planted underneath, drier sites with A. salvia were unsuitable for trees. Consequently, nurse effects related to water input are strongly site and species specific, with facilitation by shrubs providing a plausible explanation for the initiation of forest patches in this semi‐arid landscape.     

Greater shrub dominance alters breeding habitat and food resources for migratory songbirds in Alaskan arctic tundra

Climate warming is affecting the Arctic in multiple ways, including via increased dominance of deciduous shrubs. Although many studies have focused on how this vegetation shift is altering nutrient cycling and energy balance, few have explicitly considered effects on tundra fauna, such as the millions of migratory songbirds that breed in northern regions every year. To understand how increasing deciduous shrub dominance may alter breeding songbird habitat, we quantified vegetation and arthropod community characteristics in both graminoid and shrub dominated tundra. We combined measurements of preferred nest site characteristics for Lapland longspurs (Calcarius lapponicus) and Gambel's White‐crowned sparrows (Zonotrichia leucophrys gambelii) with modeled predictions for the distribution of plant community types in the Alaskan arctic foothills region for the year 2050. Lapland longspur nests were found in sedge‐dominated tussock tundra where shrub height does not exceed 20 cm, whereas White‐crowned sparrows nested only under shrubs between 20 cm and 1 m in height, with no preference for shrub species. Shrub canopies had higher canopy‐dwelling arthropod availability (i.e. small flies and spiders) but lower ground‐dwelling arthropod availability (i.e. large spiders and beetles). Since flies are the birds' preferred prey, increasing shrubs may result in a net enhancement in preferred prey availability. Acknowledging the coarse resolution of existing tundra vegetation models, we predict that by 2050 there will be a northward shift in current White‐crowned sparrow habitat range and a 20–60% increase in their preferred habitat extent, while Lapland longspur habitat extent will be equivalently reduced. Our findings can be used to make first approximations of future habitat change for species with similar nesting requirements. However, we contend that as exemplified by this study's findings, existing tundra modeling tools cannot yet simulate the fine‐scale habitat characteristics that are critical to accurately predicting future habitat extent for many wildlife species.     

766. CERCIDIPHYLLUM MAGNIFICUM – Systematic placement and fossil history of Cercidiphyllum Siebold & Zuccarini

Cercidiphyllum Sieb. & Zucc. comprises two very similar living species, Cercidiphyllum japonicum Sieb. & Zucc. and Cercidiphyllum magnificum (Nakai) Nakai, both commonly referred to as the katsura tree. The systematic placement and fossil history of Cercidiphyllum are described, and aspects of its cultivation and conservation are discussed. Cercidiphyllum magnificum is illustrated.