Facilitation and herbivory during restoration of California coastal sage scrub

Using nurse plants to facilitate native plant recruitment in degraded habitats is a common restoration practice across various arid and semiarid environments. Living trees or shrubs are typically considered nurse plants, whereas dead shrubs left in the landscape from prolonged drought are understudied prospective facilitators for native plant recruitment. The interaction between nurse plants and biotic pressures, such as herbivory, on native recruitment is also not well understood in semiarid plant communities. We investigated the effects of facilitation and herbivory on native seedling germination, growth, and survival in the restoration of degraded coastal sage scrub (CSS) habitat. Native shrub seedlings (Artemisia californica and Salvia mellifera) were planted, and native annual species (Amsinckia intermedia, Deinandra fasciculata, Phacelia distans, and Pseudognaphalium californicum) were sown in three Shrub Type treatments (live shrub, dead shrub, and exposed areas), with a nested Cage treatment (no cage and cage) in each Shrub Type treatment. Annual species grew equally well in all Shrub Type treatments; shrub seedlings grew largest in exposed areas. While there was little evidence of facilitation for all species tested, there were strong positive effects of caging on growth and establishment of all species. Caging palatable native species or planting species with anti‐herbivory traits around target plants may be more strategic approaches compared to using nurse plants in restoring degraded CSS after extended drought.     

Positive effects of shrubs on plant species diversity do not change along a gradient in grazing pressure in an arid shrubland

Facilitative or positive interactions among species are driven mainly by the environmental amelioration or protection from grazing provided by nurse plants. Some studies have suggested that protection from grazing is inconsequential in water-limited environments because of low herbivore densities and their grazing effects. Others, however, argue that herbivores have a major effect on semi-arid plant communities, and that protection from grazing is a significant factor driving positive plant–plant interactions in such environments. We identified a gradient in grazing pressure in a semi-arid shrubland in south-eastern Australia along which we compared soil condition, incident radiation and plant composition beneath two nurse shrub species with open (shrub-free) interspaces. Our aim was to assess the degree of microclimatic amelioration provided by both shrubs, and changes in the interactions (intensity, importance and frequency) between both nurse shrubs and their understorey species, and their effects on species richness at the community level. Both the relative interaction intensity (RII) and interaction importance (I_imp) indices of plant–plant interactions were generally positive and independent of grazing pressure. Soil beneath both nurse plants had significantly greater indices of nutrient cycling and infiltration, and contained more C and N than soil in the open. Almost twice as many species occurred under the canopies of both shrubs (44 species) than in the open (23 species), and the composition of species differed significantly among microsites. Fifty-four percent of all perennial plant species occurred exclusively under shrubs. Our results suggest that environmental amelioration is a stronger driver of the facilitatory effect of shrubs on their understorey species than protection from grazing. Our conclusions are based on the fact that the substantial effect of plant–plant interactions on plant species richness was largely independent of grazing pressure. Irrespective of the underlying mechanism for this effect, our study illustrates the ecological role of shrubs as refugia for understorey plants in semi-arid environments and cautions against management practices aimed at reducing shrub populations.     

An unfortunate alliance: Native shrubs increase the abundance,performance, and apparent impacts of Bromus tectorum across a regional aridity gradient

Interspecific facilitation contributes to the assembly of desert plant communities. However, we know little of how desert communities invaded by exotic species respond to facilitation along regional-scale aridity gradients. These measures are essential for predicting how desert plant communities might respond to concomitant plant invasion and environmental change. Here, we evaluated the potential for Bromus tectorum (a dominant invasive plant species) and the broader herbaceous plant community to form positive associations with native shrubs along a substantial aridity gradient across the Great Basin, Mojave, and San Joaquin Deserts in North America. Along this gradient, we sampled metrics of abundance and performance for B. tectorum, all native herbaceous species combined, all exotic herbaceous species combined, and the total herbaceous community using 180 pairs of shrub and open microsites. Across the gradient, B. tectorum formed strong positive associations with native shrubs, achieving 1.6–2.2 times greater abundance, biomass, and reproductive output under native shrubs than away from shrubs, regardless of relative aridity. In contrast, the broader herbaceous community was not positively associated with native shrubs. Interestingly, increasing B. tectorum abundance corresponded to decreasing native abundance, native species richness, exotic species richness, and total species richness under but not away from shrubs. Taken together, these findings suggest that native shrubs have considerable potential to directly (by increasing abundance and performance) and indirectly (by increasing competitive effects on neighbors) facilitate B. tectorum invasion across a large portion of the non-native range.     

Extreme drought stress shifts net facilitation to neutral interactions between shrubs and sub‐canopy plants in an arid desert

The stress gradient hypothesis (SGH) predicts that the importance or intensity of competition and facilitation will change inversely along abiotic stress gradients. It was originally postulated that increasing environmental stress can induce a monotonic increase in facilitation. However, more recent models predicted that the relationship between severity and interaction exhibits a hump‐shaped pattern, in which positive interactions prevail under moderate stress but decline at the extreme ends of stress gradients. In the present study, we conducted a field experiment along a temporal rainfall gradient for five consecutive years, in order to investigate interactions in a shrub‐herbaceous plant community at the southern edge of the Badain Jaran Desert, and, more specifically, investigated the effects of Calligonum mongolicum, a dominant shrub species, on both abiotic environmental variables and the performance of sub‐canopy plant species. We found that shrubs can improve sub‐canopy water regimes, soil properties, plant biomass, density, cover, and richness and, more importantly, that the positive effect of shrubs on sub‐canopy soil moisture during the summer diminishes as rainfall decreases, a pattern that partly explains the collapse of the positive interaction between shrubs and their understory plants. These results provide empirical evidence that the positive effect of shrubs on understory plant communities in extreme arid environments may decline and become neutral with increasing drought stress.     

The role of nurse plants in the establishment of shrub seedlings in the semi-arid subtropical Andes

In the nurse plant syndrome, or nurse association, seedlings (beneficiaries) are associated with adult shrubs/trees (benefactors). This phenomenon has been documented in several regions of the planet. Abiotic stress amelioration (one mechanism of facilitation) is one of the causes of this association. Most of the studies addressing the nurse syndrome have been conducted on spatial scales of a few hectares and have focused on only one or a few species. Moreover, there is an almost complete lack of studies addressing the incidence and characteristics of the nurse phenomenon in the arid Andes of South America. We undertook a first approximation to the study of facilitation in these ecosystems. The study was conducted at local and regional scales and involved the assessment of the spatial distribution of juveniles (seedlings and saplings) of 51 populations of 16 shrub and 12 cactus species in relation to shrub cover at 20 localities of the Prepuna (subtropical Andes of Bolivia and Argentina, 20–26°S). In terms of spatial distribution, the juveniles of most of the populations of shrubs studied were distributed both under the shrubs and in open spaces, thereby showing an apparent indifference to microhabitat. Globose and opuntioid cacti were preferentially distributed below the canopies of shrubs and were usually more associated with the dominant shrub species, which stood out as better potential nurses. The pattern was consistent throughout the region, including the more mesic and arid localities. The fact that Prepuna woody species are capable of establishing in open spaces would confer this region a greater resilience. Our findings further suggest that community dynamics in arid and semi-arid environments are more variable than previously thought.     

生态恢复中灌丛保育效应研究进展

灌木作为干旱半干旱地区植被的主要组成部分,对于维持生态系统稳定性和发挥生态服务功能具有重要作用。灌丛保育效应可以减轻限制性因子对目标植物生长和繁殖的不利影响,促进退化生境植物物种自然更新和植被恢复。本文综述了近20年有关灌丛保育效应研究和应用的主要成果,从微生境改善、肥岛形成、防御和抵抗作用发挥、有益微生物引入和协助繁殖体传播或保存以及种间竞争网络格局改变等方面阐述灌丛保育效应的促进机制,并从非生物环境、生物干扰、生活史阶段和生长繁殖策略等方面分析影响灌丛保育效应发挥的关键因素。从完善灌丛保育效应的理论机制和开展多因素共同作用下多植物物种相互作用关系模型构建角度,对灌丛保育效应的研究进行了展望。     

Facilitation by a dwarf shrub enhances plant diversity of human-valued species at high elevations in the Himalayas of Nepal

Facilitation is a global phenomenon that occurs when one species promotes the growth, survival, or reproduction of another species, mostly in stressful environments. However, the importance of facilitation by shrubs in maintaining plant community diversity is not well evaluated in the Himalayas, especially for the richness and conservation of medicinal and human-valued species. Therefore, we aimed to explore the facilitative role of a dwarf shrub species, Berberis angulosa, in maintaining plant composition and richness of human-valued species in the Langtang valley of Nepal's Himalayas. We censused plant species in open patches and beneath Berberis during monsoon and post-monsoon (dry) seasons at three elevations.Total species richness and richness of human-valued species were significantly higher inside the Berberis canopy than in gaps; the former being 39% and the latter 46% greater under shrubs than in open sites. Facilitation by Berberis shrubs promoted plant community diversity irrespective of season and elevation; however, higher differences in mean species richness for both total plant species and human-valued species during the dry season and at high elevation indicated increased facilitation intensity under more stressful conditions. The facilitative effect of Berberis shrubs increased, combining both seasons, overall plant diversity by 19% (total=105), and human-valued species by 16% (total=56). Our results show the importance of facilitation by nurse shrubs in structuring plant communities and protecting medicinal and socio-ecologically important plants, thus enriching ecosystem services in the Himalayas. These results suggest nurse plant species should be incorporated into conservation policies and management strategies for effective biodiversity conservation and sustainability, especially in the face of climate change.     

Effects of plant-plant interactions and herbivory on the plant community structure in an arid environment of Saudi Arabia

There is currently considerable evidence support that plant community structures are driven by plant-plant interactions (e.g., competition and facilitation). In contrast, there is also evidence demonstrating that plant community structure is affected by the impact of consumer pressure (e.g., grazing). In this study, 15 and 10 Acacia gerrardii nurse plants were selected inside and outside Sudyrah natural reserve (protected) area in western Saudi Arabia, respectively. The understory vegetation abundance (e.g. cover and density) was measured among quadrats around the nurse trees in both protected and unprotected areas to examine the impact of grazing and the positive interaction on the understory species. I found that understory vegetation associated with nurse trees (A. gerrardii) has been driven by both the positive impact of nurse plant and the grazing. Although the understory vegetation was positively affected by the impact of facilitation, the composition of such vegetation has been changed due to the impact of herbivory.     

Changes in Spatial Patterns of Caragana stenophylla along a Climatic Drought Gradient on the Inner Mongolian Plateau

Few studies have investigated the influence of water availability on plant population spatial patterns. We studied changes in the spatial patterns of Caragana stenophylla along a climatic drought gradient within the Inner Mongolian Plateau, China. We examined spatial patterns, seed density, “nurse effects” of shrubs on seedlings, transpiration rates and water use efficiency (WUE) of C. stenophylla across semi-arid, arid, and intensively arid zones. Our results showed that patches of C. stenophylla populations shifted from a random to a clumped spatial pattern towards drier environments. Seed density and seedling survival rate of C. stenophylla decreased from the semi-arid zone to the intensively arid zone. Across the three zones, there were more C. stenophylla seeds and seedlings underneath shrub canopies than outside shrub canopies; and in the intensively arid zone, there were almost no seeds or seedlings outside shrub canopies. Transpiration rates of outer-canopy leaves and WUE of both outer-canopy and inner-canopy leaves increased from the semi-arid zone to the intensively arid zone. In the intensively arid zone, transpiration rates and WUE of inner-canopy leaves were significantly lower and higher, respectively, than those of outer-canopy leaves. We conclude that, as drought stress increased, seed density decreased, seed proportions inside shrubs increased, and “nurse effects” of shrubs on seedlings became more important. These factors, combined with water-saving characteristics associated with clumped spatial patterns, are likely driving the changes in C. stenophylla spatial patterns.     

Shrubs, granivores and annual plant community stability in an arid ecosystem

Compensatory population dynamics among species stabilise aggregate community variables. Inter-specific competition is thought to be stabilising as it promotes asynchrony among populations. However, we know little about other inter-specific interactions, such as facilitation and granivory. Such interactions are also likely to influence population synchrony and community stability, especially in harsh environments where they are thought to have relatively strong effects in plant communities. We use a manipulative experiment to test the effects of granivores (harvester ants) and nurse plants (dwarf shrubs) on annual plant community dynamics in the Negev desert, Israel. We present evidence for weak and inconsistent effects of harvester ants on plant abundance and on population and community stability. By contrast, we show that annual communities under shrubs were more species rich, had higher plant density and were temporally less variable than communities in the inter-shrub matrix. Species richness and plant abundance were also more resistant to drought in the shrub under-storey compared with the inter-shrub matrix, although population dynamics in both patch types were synchronised. Hence, we show that inter-specific interactions other than competition affect community stability, and that hypothesised mechanisms linking compensatory dynamics and community stability may not operate to the same extent in arid plant communities.     

Effects of experimental rainfall manipulations on Chihuahuan Desert grassland and shrubland plant communities

Aridland ecosystems are predicted to be responsive to both increases and decreases in precipitation. In addition, chronic droughts may contribute to encroachment of native C₃ shrubs into C₄-dominated grasslands. We conducted a long-term rainfall manipulation experiment in native grassland, shrubland and the grass–shrub ecotone in the northern Chihuahuan Desert, USA. We evaluated the effects of 5 years of experimental drought and 4 years of water addition on plant community structure and dynamics. We assessed the effects of altered rainfall regimes on the abundance of dominant species as well as on species richness and subdominant grasses, forbs and shrubs. Nonmetric multidimensional scaling and MANOVA were used to quantify changes in species composition in response to chronic addition or reduction of rainfall. We found that drought consistently and strongly decreased cover of Bouteloua eriopoda, the dominant C₄ grass in this system, whereas water addition slightly increased cover, with little variation between years. In contrast, neither chronic drought nor increased rainfall had consistent effects on the cover of Larrea tridentata, the dominant C₃ shrub. Species richness declined in shrub-dominated vegetation in response to drought whereas richness increased or was unaffected by water addition or drought in mixed- and grass-dominated vegetation. Cover of subdominant shrubs, grasses and forbs changed significantly over time, primarily in response to interannual rainfall variability more so than to our experimental rainfall treatments. Nevertheless, drought and water addition shifted the species composition of plant communities in all three vegetation types. Overall, we found that B. eriopoda responded strongly to drought and less so to irrigation, whereas L. tridentata showed limited response to either treatment. The strong decline in grass cover and the resistance of shrub cover to rainfall reduction suggest that chronic drought may be a key factor promoting shrub dominance during encroachment into desert grassland.     

Interactive effects of grazing and shrubs on the annual plant community in semi‐arid Mediterranean shrublands

Question: We studied the interactive effects of grazing and dwarf shrub cover on the structure of a highly diverse annual plant community. Location: Mediterranean, semi‐arid shrubland in the Northern Negev desert, Israel. Methods: Variation in the biomass and plant density of annual species in the shrub and open patches was monitored during four years, inside and outside exclosures protected from sheep grazing, in two contrasting topographic sites: north and south‐facing slopes that differed in their dominant dwarf shrubs species: Sarcopoterium spinosus and Corydothymus capitatus, respectively. Results: Above‐ground biomass, density and richness of annual species were lower under the canopy of both shrub species compared to the adjacent open patches in the absence of grazing. Grazing reduced the biomass of annuals in open patches of both topographic sites, but not in the shrub patches. On the north‐facing slope, grazing also reduced plant density and richness in the open patches, but increased plant density in the shrub patches. At the species level, various response patterns to the combined effects of grazing and patch type were exhibited by different annuals. Protection against the direct impacts of grazing by shrub cover as well as species‐specific interactions between shrubs and annuals were observed. A conceptual mechanistic model explaining these interactions is proposed. Conclusion: In semi‐arid Mediterranean shrublands grazing and dwarf shrub cover interact in shaping the structure of the annual plant community through (1) direct impacts of grazing restricted to the open patches, (2) species‐specific facilitation/ interference occurring in the shrub patches and (3) subsequent further processes occurring among the interconnected shrub and open patches mediated through variation in seed flows between patches.     

Ecological stoichiometry explains larger-scale facilitation processes by shrubs on species coexistence among understory plants

Plant facilitation (positive plant–plant interactions) strongly influences biodiversity, structure, and dynamics in plant communities, and the topic has received considerable attention among ecologists. Most studies of facilitation processes by shrubs have been conducted at small spatial scales between shrubs and their neighboring species. Yet, we know little about whether facilitation processes by shrubs at a small scale (i.e., a patch scale) also work at a larger scale (i.e., a site scale) in terms of the maintenance of biodiversity. Here, we report that the facilitative effects of shrubs on plant diversity at a larger scale can be explained by changing ecological stoichiometry. The soil fertility showed unimodal shape along shrub cover gradient, suggesting that the facilitative effects of a shrub do not necessarily increase as the shrub develops. The unimodal shape of dependence of plant species richness on shrub cover probably was generated by the unimodal dependence of soil fertility on shrub cover. Soil nutrient enrichment by shrubs shifted low N:P ratios of plant communities with low levels of shrub cover to more balanced N:P ratios at intermediate levels of shrub cover. At the peak N:P ratio along the gradient in shrub cover, the maximum species richness and functional richness were observed, which was consistent with the unimodal relationship predicted by the resource balance hypothesis. Thus, our findings showed that facilitation processes by shrubs at a patch scale also work at a larger scale in terms of the maintenance of biodiversity. Because observed larger-scale facilitation processes are enhanced at some intermediate levels of shrub cover, this study offers practical insight into the need for management practices that allow some intermediate levels of grazing by livestock for optimizing the role of larger-scale facilitation processes in the maintenance of biodiversity and ecosystem functioning in arid and semi-arid rangelands.     

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     

Soil nutrients trump intraspecific effects on understory plant communities

Understanding the links between intraspecific genetic variation and patterns of diversity in associated communities has been the primary focus of community genetics or ‘genes-to-ecosystem’ research in ecology. While other ecological factors, such as the abiotic environment, have well-documented influences on communities, the relative contributions of genetic variation versus the environment to species interactions remains poorly explored. In this study, we use a common garden experiment to study a coastal dune plant community dominated by the shrub, Baccharis pilularis, which displays a morphological dimorphism in plant architecture. We found the differences in the understory plant community between erect and prostrate morphs of Baccharis to be statistically significant, but small relative to the impacts of nutrient additions (NPK and C additions), for the richness, cover, and biomass of the understory plant community. There were no significant interactions between Baccharis morphology and nutrient-addition treatments, suggesting the influence of nutrient addition was consistent between erect and prostrate morphs. Moreover, we found no difference in overall plant community composition between Baccharis morphs, while NPK additions led to shifts in understory community composition compared to unfertilized shrubs. In sum, our results indicate that nutrients are the more important factor governing understory plant community structure in a coastal dunes ecosystem followed by intraspecific variation in dominant shrub architecture. Our results address a growing call to understand the extended consequences of intraspecific variation across heterogeneous environments in terrestrial ecosystems.     

锦鸡儿属灌丛化对草原化荒漠区土壤线虫群落组成和代谢足迹的影响

调查了内蒙古草原化荒漠区3种锦鸡儿[垫状锦鸡儿(Caragana tibetica)、荒漠锦鸡儿(C.roborovskyi)和狭叶锦鸡儿(C.stenophylla)]灌丛内、外土壤线虫群落多样性、组成和代谢足迹,以及相关的土壤理化性质和植物群落特征,旨在探讨锦鸡儿属灌丛对土壤线虫群落的影响,并分析这种影响是否具有灌木种间差异,同时从线虫功能团水平上探究灌丛如何通过非生物因素和生物因素影响线虫群落组成。结果表明：锦鸡儿属灌丛对土壤线虫多度、丰富度和多样性无显著影响,但却显著影响土壤线虫群落组成。原因是灌丛对土壤线虫群落具有物种选择性;灌丛可能主要是通过根系分泌物、凋落物质量等因素,而不是通过土壤理化性质和林下植物影响线虫群落组成。土壤线虫组成的变化引起线虫代谢足迹发生明显变化。灌丛内土壤线虫功能代谢足迹大于灌丛外(除狭叶锦鸡儿外),说明灌丛内土壤线虫群落对碳的利用率更高。3种锦鸡儿属灌木中,狭叶锦鸡儿的结构代谢足迹最大,表明狭叶锦鸡儿对捕食杂食类线虫代谢活性的促进作用更强。土壤线虫组成的变化通过线虫代谢足迹导致土壤食物网结构发生显著性变化。狭叶锦鸡儿灌丛土壤干扰程度低,营养富集状况好...     

A field test of the stress‐gradient hypothesis along an aridity gradient

Aims: The stress‐gradient hypothesis (SGH) predicts how plant interactions change along environmental stress gradients. We tested the SGH in an aridity gradient, where support for the hypothesis and the specific shape of its response curve is controversial. Location: Almería, Cáceres and Coimbra, three sites in the Iberian Peninsula that encompass the most arid and wet habitats in the distribution range of a nurse shrub species –Retama sphaerocarpa L. (Boiss) – in Europe. Methods: We analysed the effect of Retama on its understorey plant community and measured the biomass and species richness beneath Retama and in gaps. We estimated the frequency (changes in species richness), importance and intensity of the Retama effects, and derived the severity–interaction relationship pattern, analysing how these interaction indices changed along this aridity gradient. Results and conclusions: The intensity and frequency of facilitation by Retama increased monotonically with increasing environmental severity, and the importance tended to have a similar pattern, overall supporting the SGH. Our data did not support predictions from recent revisions of the SGH, which may not apply to whole plant communities like those studied here or when interactions are highly asymmetrical. Facilitation by Retama influenced community composition and species richness to the point that a significant fraction of species found at the most arid end of the gradient were only able to survive beneath the nurse shrub, whereas some of these species were able to thrive in gaps at more mesic sites, highlighting the ecological relevance of facilitation by nurse species in mediterranean environments, especially in the driest sites.     

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.     

Non-trophic interactions in deserts: Facilitation,interference, and an endangered lizard species

Research on plant–animal interactions has been focused on direct consumer interactions (i.e. plants as resources), but non-trophic interactions including providing shelter or interference with movement can also affect the fine-scale distribution of animals. In particular, non-trophic interactions that are positive could support threatened animal populations. Positive interactions have been used in the restoration of plant communities, but have not yet been extended to the management of animal habitat. In this study, we tested the hypothesis that non-trophic interactions influence the occurrence of an endangered lizard species in an arid shrub-annual system. At a location known to have a population of blunt-nosed leopard lizards (Gambelia sila), we geotagged 700 shrubs, measured shrub morphometric traits, collected biomass samples, and surveyed for lizard presence using scat detection dogs over two years. Relative to 2014, in 2013 plant productivity was high and lizard scats were found more frequently in areas with low invasive grass cover (i.e. residual dry matter, RDM). In 2014, plant productivity was low because of an extreme drought year, and lizard scats were more frequently observed under shrub canopies, particularly those with relatively dense cover. These findings support the novel theory that positive non-trophic interactions are a critical form of plant–animal interactions in addition to consumption. Dominant shrubs can act as a foundation species by functioning as a basal node in structuring both plant and animal communities through a network of interactions. Managing dominant plants, in addition to habitat, is therefore important for conserving animal species in arid ecosystems.     

Establishing Native Grasses in a Big Sagebrush–Dominated Site: An Intermediate Restoration Step

Many semiarid rangelands in the Great Basin, U.S.A., are shifting dominance to woody species as a consequence of land degradation including intense livestock grazing and fire suppression. Whereas past rehabilitation efforts in Big sagebrush (Artemisia tridentata) steppes removed the shrub and added introduced forage grasses to successfully shift communities from shrublands to grasslands, current consensus is that native species should be included in restoration projects and that retention of some woody plants is desirable. We examined the potential for interseeding grasses into dense shrub communities as a precursor to thinning shrubs and releasing grasses from shrub interference. We compared seedling establishment of the native grass, Bluebunch wheatgrass (Pseudoroegneria spicata), with that of the Eurasia grass, Crested wheatgrass (Agropyron desertorum), in dense Ar. tridentata stands. Shrubs may play an important role as nurse plants for seedling establishment (reduced solar radiation, “island of fertility” effect) but result in highly contrasting light environments and root interference for seedlings. In experimental plots, we examined effects of Ar. tridentata shade levels (0, 40, 70, and 90% reduction of solar radiation) and initial root exclusion (present/absent) on the establishment and growth of P. spicata and Ag. desertorum seedlings. With this design we evaluated the interference effects of Ar. tridentata on the two grasses and identified the most beneficial microsites for grass restoration in Ar. tridentata–dominated communities. We predicted seedling survival and growth to be greater under moderate shade (40% reduction) and limited root competition than under no or strong shade conditions (0 and 90%) and unrestricted root interactions. Fifty to 85% of the P. spicata and Ag. desertorum seedlings survived the dry summer months of 1995 and 1996 and the intervening winter. Neither shading nor root exclusion from Ar. tridentata affected final seedling survival of either species. Seedling biomass of both grass species was negatively affected by initial root interactions with Ar. tridentata. However, the analysis of seedling biomass variability (coefficient of variation) indicated that in all shade and root‐exclusion treatments, some seedlings of both species developed to large individuals to survive in Ar. tridentata–dominated rangelands. Thus, the use of interseeding techniques shows promise for restoring herbaceous species in dense Ar. tridentata stands and should be given further consideration when shrub retention is an important consideration.