Using consecutive prescribed fires to reduce shrub encroachment in grassland by increasing shrub mortality

Woody plant encroachment into open grasslands occurs worldwide and causes multiple ecological and management impacts. Prescribed fire could be used to conserve grassland habitat but often has limited efficacy because many woody plants resprout after fire and rapidly reestablish abundance. If fire‐induced mortality could be increased, prescribed fire would be a more effective management tool. In California's central coast, shrub encroachment, especially of Baccharis pilularis (coyote brush), is converting coastal prairie into shrub‐dominated communities, with a consequent loss of native herbaceous species and open grassland habitat. B. pilularis has not been successfully controlled with single prescribed fire events because the shrub resprouts and reestablishes cover within a few years. We investigated whether two consecutive annual burns would control B. pilularis by killing resprouting shrubs, without reducing native herbaceous species or encouraging invasive plants. As expected, resprouting did occur; however, 2 years after the second burn, B. pilularis cover on burned plots was only 41% of the cover on unburned plots. Mortality of B. pilularis more than doubled following the second burn, likely maintaining a reduction in B. pilularis cover for longer than a single burn would have. Three native coastal prairie perennial grasses did not appear to be adversely affected by the two burns, nor did the burns result in increased cover of invasive species. Managers wanting to restore coastal prairie following B. pilularis encroachment should consider two consecutive annual burns, especially if moderate fire intensity is achievable.     

Facilitative and competitive effects of a large species with defensive traits on a grazing-adapted,small species in a long-term deer grazing habitat

Plants can adapt to grazing environments by developing defensive traits, such as spines and toxins, or having a small phenotype, such as short and prostrate growth forms. This study examined facilitative and competitive interactions between species with different types of grazing adaptation. We predicted that large species with defensive traits sometimes protect grazing-adapted species without defensive traits from herbivores, but competitively suppress them overall. We conducted an experiment using fences and removals of an unpalatable plant in the long-term deer grazing habitat of Nara Park in Nara, Japan. We evaluated the seasonal variations in the facilitative and competitive effects of a defensive perennial, Urtica thunbergiana, on the growth, survival, reproduction, and final fitness of a small palatable annual species, Persicaria longiseta, during a growing season. The populations of the two species in the park have adapted to the grazed habitat by increasing the density of stinging hairs (Urtica) and developing inherently short shoots and small leaves (Persicaria). We found that Urtica individuals had facilitative effects on the growth of Persicaria individuals under grazing during a few periods of the growing season, but had neutral effects on survival and plant fitness throughout the season. In the fenced plots, Urtica had negative effects on the growth, survival, and reproduction of Persicaria. These results suggest that the relative importance of the facilitative and competitive effects of Urtica on Persicaria fluctuated due to seasonal variations in grazing pressure and vegetative productivity. Although well-defended plants often facilitate less-defended species, we conclude that the facilitative effects of Urtica on Persicaria are limited in a plant community with a long history of intensive grazing.     

Environmental drivers of femaleness of an inter‐Andean monoecious shrub

Hetero‐and conspecific interactions, nutrient availability, climate, habitat heterogeneity, and disturbances can generate variation and spatial patterns of femaleness in plants. We assessed whether year, site, plant size, plant density, and canopy area of conspecific neighbors influenced the expression and spatial aggregation of femaleness in Croton aff. wagneri, a monoecious shrub from dry shrublands of the inter‐Andean valleys in Ecuador. We georeferenced in two sites (1,700 and 1,400 m.a.s.l) in five 10 × 10 m plots, within each site, the position of each Croton reproductive plant during first part of flowering season in two years, and measured their height, length, and width. The femaleness index of each plant was determined by the number of female and male buds and flowers. Plant density was determined for each plot, along with the number of neighbors and the summed canopy area of conspecific neighbors (at 1.0, 2.0, and 2.5 m radius, and the five closest plants) from each focal plant. Croton´s femaleness at the lower elevation site was greater than at the higher elevation site and increased with plant size and with canopy of the closest five neighbors. Soil at the lower elevation site had higher temperatures and lower water content. Aggregate patterns of femaleness were found in more plots at the lower elevation site. Our results indicate that location, plant size, and canopies of conspecific neighbors of Croton can affect femaleness and its aggregation and support the hypothesis that femaleness can be influenced by facilitative interactions. Abstract in Spanish is available with online material.     

The impact of human trampling on a threatened coastal Mediterranean plant: The case of Anchusa littorea Moris (Boraginaceae)

In a 5-period study (2007–2011) we examined the effects of human trampling on the last remaining population of Anchusa littorea on its sandy coastal habitat in Sardinia. This species, considered extinct in the wild for several years, was casually rediscovered in a small population at Is Arenas (SW Sardinia). We monthly monitored six trampled and six un-trampled permanent plots, mapping individuals and recording their size and reproductive variables. Trampled and un-trampled plots showed significant differences with respect to plant density, plant size and reproductive performance (flowers and fruits production) of A. littorea. This study demonstrated that human trampling represents a severe threat to the conservation of this species that can be appreciated as a keystone species with concern to human trampling effects on coastal dune plants. In case of A. littorea urgent measures should be undertaken to protect this unique remnant population in the dune system of Is Arenas. In particular, tourist paths should be redirected and confined to others areas in order to promote the natural expansion of A. littorea in its original habitat. A possible integrated strategy for the conservation and management of the species consists in a combination of in situ and ex situ measures.     

Sexual dimorphism and resource allocation in male and female shrubs of Simmondsia chinensis

Summary Desert populations of the evergreen dioecious shrub Simmondsia chinensis exhibit sex-related leaf and canopy dimorphisms not present in populations from more mesic coastal environments. Leaves on female shrubs have characteristically larger sizes, greater specific weights, and greater water-holding capacity than male leaves in desert habitats. In coastal scrub environments no significant difference is present, with leaf characteristics of both sexes similar to those of desert male shrubs. Desert female shrub canopies are typically relatively open with little mutual branch shading. In male shrubs canopies are more densely branched with considerable mutual shading of branches. Female plants allocate a greater proportion of their vegetative resources to leaves than do male plants. Considering total biomass, male plants allocate 10–15% of their resources (biomass, calories, glucose-equivalents, nitrogen, phosphorus) to reproductive tissues. Female allocation is dependent on seed set. At 100% seed set females would allocate 30–40% of their resources to reproduction, while female reproductive investment would equal that of males at approximately 30% seed set. Sexual dimorphism and the associated physiological characteristics in Simmondsia act as an alternative to differential habitat selection by male and female plants. Female plants respond to limited water resources in desert areas by increasing their efficiency in allocating limited resources to reproductive structures.     

Multiple effects of shrubs on annual plant communities in arid lands of South Australia

Abstract The presence of shrubs in arid lands creates spatial heterogeneity that affects the distribution and performance of annual plants; several possible mechanisms have been implicated. A preliminary survey in a chenopod shrubland in South Australia showed differences in the distribution of annual plants under canopies of Atriplex vesicaria and Maireana sedifolia (the two dominant shrub species) and open spaces. A series of experiments were conducted to test the potential contribution to these patterns of nutrient enrichment under shrubs, differential seed accumulation, stress reduction by the canopy, competition by shrub roots, and protection against grazing. The germinable soil seed‐bank under A. vesicaria and M. sedifolia was different from that in open spaces, but these differences can only explain a fraction of the differences observed in the growing annual plant community in different microsites. The soil under A. vesicaria had higher total nitrogen content than soil in open spaces, whereas soil under M. sedifolia had lower available phosphorus than open spaces. Although annual plant densities under A. vesicaria were higher than in open spaces, experimental removal of shrubs increased their density, suggesting that shrub canopies inhibit annual plants in this system. Surprisingly, trenching of open areas close to shrubs (severing lateral shrub roots) decreased annual plant density. We suggest that water moves laterally through shrub roots, in a process akin to a hydraulic lift, increasing water availability for the annual plants. Exclusion of vertebrate grazers had a stronger effect on annual plant biomass in open spaces than under M. sedifolia, suggesting that this shrub provides shelter against herbivory. Overall our results show that shrubs can have simultaneously facilitative and inhibitory effects on the annual plant community through different mechanisms, but more importantly that different shrub species have different effects. This is a potential mechanism allowing for species coexistence of annual plants.     

Climatic and biotic extreme events moderate long‐term responses of above‐ and belowground sub‐Arctic heathland communities to climate change

Climate change impacts are not uniform across the Arctic region because interacting factors causes large variations in local ecosystem change. Extreme climatic events and population cycles of herbivores occur simultaneously against a background of gradual climate warming trends and can redirect ecosystem change along routes that are difficult to predict. Here, we present the results from sub‐Arctic heath vegetation and its belowground micro‐arthropod community in response to the two main drivers of vegetation damage in this region: extreme winter warming events and subsequent outbreaks of the defoliating autumnal moth caterpillar (Epirrita autumnata). Evergreen dwarf shrub biomass decreased (30%) following extreme winter warming events and again by moth caterpillar grazing. Deciduous shrubs that were previously exposed to an extreme winter warming event were not affected by the moth caterpillar grazing, while those that were not exposed to warming events (control plots) showed reduced (23%) biomass from grazing. Cryptogam cover increased irrespective of grazing or winter warming events. Micro‐arthropods declined (46%) following winter warming but did not respond to changes in plant community. Extreme winter warming and caterpillar grazing suppressed the CO₂ fluxes of the ecosystem. Evergreen dwarf shrubs are disadvantaged in a future sub‐Arctic with more stochastic climatic and biotic events. Given that summer warming may further benefit deciduous over evergreen shrubs, event and trend climate change may both act against evergreen shrubs and the ecosystem functions they provide. This is of particular concern given that Arctic heath vegetation is typically dominated by evergreen shrubs. Other components of the vegetation showed variable responses to abiotic and biotic events, and their interaction indicates that sub‐Arctic vegetation response to multiple pressures is not easy to predict from single‐factor responses. Therefore, while biotic and climatic events may have clear impacts, more work is needed to understand their net effect on Arctic ecosystems.     

Facilitation by nurse shrubs of resprouting behavior in a post-fire shrubland in northern Patagonia,Argentina

Abstract. The nurse-plant syndrome is a widely recognized example of positive (facilitative) influences of plant species on the establishment and growth of other species. Most studies of the nurse-plant syndrome have been on species that reproduce mainly from seed rather than vegetatively. In this study, we experimentally compared the influences of two species of nurse shrubs, Schinus patagonicus and Berberis buxifolia, on the survival and growth of vegetatively reproducing herbaceous and woody plants in a post-fire shrubland in northern Patagonia, Argentina. The vegetation beneath shrubs was removed by clipping and, in a paired-sample design, one half of the canopy of each shrub was removed. We determined species richness, counted number of resprouts, and measured photon flux density and soil moisture beneath cut and uncut halves of each shrubs. Abundances of resprouts were several times greater beneath the uncut vs. the cut shrubs, as was the mean number of species. Thus, shrubs have a strong facilitating influence as measured by resprout densities and the number of species. Numbers of resprouts and of species were twice as high beneath Schinus as beneath Berberis, implying important differences in the facilitative effects of the two shrubs species. Microsites beneath Schinus were characterized by lower and more heterogeneous light levels but by greater soil moisture. Even though the reproductive mode in this post-fire shrubland is overwhelmingly vegetative, facilitation by nurse shrubs is important and differentially effective for different species of nurse shrubs.     

The influence of rice plant age on susceptibility to the rice water weevil,Lissorhoptrus oryzophilus

Plant age‐ and plant stage‐related changes in the resistance of rice, Oryza sativa, to its most important insect pest in the US, the rice water weevil (Lissorhoptrus oryzophilus), were investigated in a series of field and greenhouse choice and no‐choice studies. Rice plants were susceptible to infestation by rice water weevils over a broad range of plant ontogenetic stages, from at least the early vegetative stage to well into the reproductive stage. There was, however, a clear preference expressed by rice water weevils in both choice and no‐choice experiments for plants in (or nearly in) the tillering stage of development, with pre‐tillering and reproductive stage plants less preferred. The relationship between rice plant age and susceptibility to weevils is thus nonlinear. This study constitutes one of the most thorough studies to date of the relationship in a grass species between plant age and susceptibility to herbivores. The results provide a biological explanation for observed patterns of weevil infestations and a rationale for the cultural practice of delayed flooding.     

Putative local adaptations modulate the interactions of the carnivorous plant Drosera uniflora Willd (1809) (Droseraceae) with cushion and shrub nurse plants

Cushion and shrub plants are typical high mountain nurse plants. In Magellanic tundras of the Cordillera del Sarao on the coast of Chile, the carnivorous plant Drosera uniflora grows in association with cushions of Donatia fascicularis and the shrubs Chusquea montana var. nigricans and Lepidothamnus fonkii. The different microhabitats for recruitment, in addition to the limited gene flow of D. uniflora, enable us to hypothesise that this plant manifests putative local adaptations, expressed in seed germination and population abundance. Our aim was to evaluate the local adaptation of D. uniflora to cushion and shrub microhabitats by estimating germination and abundance. Local adaptation of seed germination was determined by means of reciprocal transplant experiments. Abundance was determined in small plots located inside, on the edge and outside the nurse plants. Seed origin and growth substrate play a decisive role in D. uniflora germination. Seeds that originate from cushions plant habitat and germinate in the same substrate do so in greater numbers than when those which originate from shrubby habitats. Conversely, seeds that originate from shrubby habitat and germinate in the same substrate do not exhibit significant differences. Abundance was always greater inside than outside of the nurse plants. We concluded that availability and quality of different microhabitats for seed germination and recruitment, together with a limited gene flow, would trigger putative local adaptations in D. uniflora, modulating a long history of interactions with the plants that act as nurse plants.     

Effects of local and landscape factors on the abundance of an endangered multivoltine butterfly at riverbanks

Plebejus argyrognomon is one of the grassland‐dwelling butterflies undergoing rapid decline in recent decades. Grassland habitats for butterflies are generally threatened by fragmentation and invasive species, hence are among the most vulnerable ecosystems. We studied the seasonal abundance of P. argyrognomon at habitat patches along the banks of the Kinugawa River in eastern Japan, to identify environmental factors suitable for population persistence of this species, including habitat patch connectivity. Results showed that the patch's host plant cover had a positive effect on abundance in all three seasons, while the shading of the host plants by surrounding non‐host plants and nearby forested area showed negative effects. Additionally, habitat patch connectivity and nectar richness could be considered as positive factors in autumn and summer, respectively. Analysis of habitat connectivity also showed that the Kinugawa River did not appear to act as a dispersal barrier for P. argyrognomon. Our findings emphasize the importance of understanding environmental factors that may vary among seasons, and such understanding could contribute to habitat management of multivoltine butterflies in fragmented landscapes.     

Effects of temperature and elevation on habitat use by a rare mountain butterfly: implications for species responses to climate change

Abstract 1. The present study used the mountain specialist butterfly Parnassius apollo as a model system to investigate how climate change may alter habitat requirements for species at their warm range margins. 2. Larval habitat use was recorded in six P. apollo populations over a 700 m elevation gradient in the Sierra de Guadarrama (central Spain). Larvae used four potential host species (Sedum spp.) growing in open areas amongst shrubs. 3. Parnassius apollo host‐plant and habitat use changed as elevation increased: the primary host shifted from Sedum amplexicaule to Sedum brevifolium, and larvae selected more open microhabitats (increased bare ground and dead vegetation, reduced vegetation height and shrub cover), suggesting that hotter microhabitats are used in cooler environments. 4. Larval microhabitat selection was significantly related to ambient temperature. At temperatures lower than 27 °C, larvae occupied open microhabitats that were warmer than ambient temperature, versus more shaded microhabitats that were cooler than ambient conditions when temperature was higher than 27 °C. 5. Elevational changes in phenology influenced the temperatures experienced by larvae, and could affect local host‐plant favourability. 6. Habitat heterogeneity appears to play an important role in P. apollo larval thermoregulation, and may become increasingly important in buffering populations of this and other insect species against climatic variation.     

Population dynamics of the climate‐sensitive endangered perennial Ferula sadleriana Ledeb. (Apiaceae)

Ancient plant species surviving in isolated small populations are particularly vulnerable to extinction, therefore understanding their population dynamics is necessary for conservation. The iteroparous perennial relic endemic Ferula sadleriana Ledeb. (Apiaceae) is restricted to seven distant localities in the Carpathian Basin, where it inhabits rocky hills. We monitored the species' largest population on the Pilis Hill, Hungary, over 14–19 years (depending on trait) between 1979 and 2010, and relationships were sought between climatic properties and population attributes. The population of 4000 ± 1509 emergent individuals underwent large interannual fluctuations, with the vegetative stage displaying sevenfold and the reproductive stage twenty‐eight‐fold differences. Spring and early summer precipitation had a marked influence on abundances and seed set. Alternating years of high and low counts of reproductive plants suggest costs of reproduction that most probably incur prolonged dormancy and retrogression to the vegetative stage. Seed set was positively influenced by number of reproductive plants over years and by plant size within a year. Ungulates nullify yearly reproductive output by grazing on reproductive individuals. This is particularly intense in dry summers, when reproductive output is already low. The strong precipitation response of abundance, absence of clonal propagation and soil seed bank, and geographical isolation of the populations place F. sadleriana at considerable risk under an increasingly variable and extreme climate. Management should seek to maintain the species' original habitat mosaic (potentially compensating for climate variation), minimize grazing damage and anthropogenic disturbance, and establish ex situ conservation programs to provide propagules for eventual reintroduction.     

Conservation and restoration of Nardus grasslands in the Swiss northern Alps

Aim: Species‐rich Nardus grasslands are high nature‐value habitats. In Switzerland, many of these grasslands are degraded even though they have been under protection since the 1980s. Degradation shows two divergent trends: Nardus grasslands are either dominated by Nardus stricta or by eutrophic plants, both trends leading to the disappearance of typical Nardus grassland species. With this study, we aim to identify the factors that could be adjusted to conserve the integrity of this habitat. Location: Bernese Alps, Switzerland. Methods: In 2016, we investigated the underlying causes of this degradation process by assessing vegetation composition in 48 Nardus grasslands located in the Swiss northern Alps of canton Bern and linking it to soil, management and environmental variables. To explore the effect of the degradation on higher trophic levels, orthopteran species richness and densities were assessed. Results: Results show that Nardus meadows (mown) are rarely degraded compared to Nardus pastures (grazed). Within pastures, eutrophic plants are most abundant on small pastures with low soil carbon/nitrogen ratio, indicating high nutrient availability. Nardus stricta dominance is most problematic on north‐exposed slopes and in summer pastures. A plausible driver of both degradation trends is the grazing management regime: within small pastures at low elevation where the grazing periods are short but intense, soil carbon/nitrogen ratio is low because of high dung deposition, thus the eutrophic species become dominant. Contrastingly, on large summer pastures with low‐intensity and long‐term grazing, N. stricta becomes dominant due to selective grazing. Both degradation trends show a negative impact on the orthopteran density. Conclusion: Species‐rich Nardus grasslands are a precious alpine habitat for specialised plant species and orthopterans. With an extensive mowing regime or a more controlled grazing regime that homogenises intensity in time and space, species‐rich Nardus grasslands can be conserved in Switzerland.     

Frugivorous birds mediate sex‐biased facilitation in a dioecious nurse plant

Abstract. Facilitation by dispersal occurs if the nurse plant acts as a focus which is actively selected by seed dispersers and enhances the fitness of the facilitated plant. Sex‐biased facilitation may be produced if seed dispersers tend to concentrate the seeds under female, fruit‐bearing plants of dioecious species more often than under conspecific males. Juniperus sabina is a dioecious shrub with a prostrate growth form from Mediterranean high mountains that modifies many microhabitat characteristics related to seedling establishment and survival. Soil water availability, maximum soil temperature in summer, organic matter and total nitrogen content, were different on open ground as compared with beneath J. sabina shrubs, irrespective of its sex. Other studied characteristics such as soil bulk density and soil compaction after rain did not differ between the microhabitats considered. Some species, such as Juniperus communis, Pinus nigra, Helleborus foetidus and Euphorbia nicaeensis, are spatially associated to J. sabina shrubs, strongly suggesting a facilitative role. The anemochorous P. nigra and myrme‐chorous H. foetidus and E. nicaeensis did not associate preferentially to any sex of J. sabina. Only J. communis, an endozoochorous species sharing the same bird dispersers as J. sabina, presented a female‐biased spatial association with the nurse plant. Seed dispersal mediated by birds attracted by the fruit‐rewarding females of J. sabina explains the sex‐biased spatial pattern of Juniperus communis.     

Suppression of OsRAD51D results in defects in reproductive development in rice (Oryza sativa L.)

The cellular roles of RAD51 paralogs in somatic and reproductive growth have been extensively described in a wide range of animal systems and, to a lesser extent, in Arabidopsis, a dicot model plant. Here, the OsRAD51D gene was identified and characterized in rice (Oryza sativa L.), a monocot model crop. In the rice genome, three alternative OsRAD51D mRNA splicing variants, OsRAD51D.1, OsRAD51D.2, and OsRAD51D.3, were predicted. Yeast two‐hybrid studies, however, showed that only OsRAD51D.1 interacted with OsRAD51B and OsRAD51C paralogs, suggesting that OsRAD51D.1 is a functional OsRAD51D protein in rice. Loss‐of‐function osrad51d mutant rice plants displayed normal vegetative growth. However, the mutant plants were defective in reproductive growth, resulting in sterile flowers. Homozygous osrad51d mutant flowers exhibited impaired development of lemma and palea and contained unusual numbers of stamens and stigmas. During early meiosis, osrad51d pollen mother cells (PMCs) failed to form normal homologous chromosome pairings. In subsequent meiotic progression, mutant PMCs represented fragmented chromosomes. The osrad51d pollen cells contained numerous abnormal micro‐nuclei that resulted in malfunctioning pollen. The abnormalities of heterozygous mutant and T2 Ubi:RNAi‐OsRAD51D RNAi‐knock‐down transgenic plants were intermediate between those of wild type and homozygous mutant plants. The osrad51d and Ubi:RNAi‐OsRAD51D plants contained longer telomeres compared with wild type plants, indicating that OsRAD51D is a negative factor for telomere lengthening. Overall, these results suggest that OsRAD51D plays a critical role in reproductive growth in rice. This essential function of OsRAD51D is distinct from Arabidopsis, in which AtRAD51D is not an essential factor for meiosis or reproductive development.     

The Role of Habitat and Herbivory on the Restoration of Tidal Freshwater Submerged Aquatic Vegetation Populations

Submerged aquatic vegetation (SAV) has declined precipitously throughout coastal areas and its reestablishment has long been an important objective of coastal management. We investigated restoration success of Vallisneria americana (wild celery) using seeds, seed pods, and whole shoot transplants at sites in the Chesapeake Bay in the United States where historical aerial photography has indicated that the species once grew. In addition, we evaluated habitat conditions and established herbivore exclosures to assess the impacts of water quality, sediment conditions, and grazers on planting success. Whole shoot transplants resulted in the most rapid cover of the bottom, but required greater planting effort. Direct dispersal of individual seeds was generally more successful than dispersal of intact seed pods, resulting in more rapid initial seedling growth. Overall, 100% bottom cover of whole shoot transplant plots could be reached in approximately 3 years, despite light attenuation coefficients (K_d) of 3.0 to 4.0. Transplants at shallow depths (<0.5 m) were able to rapidly grow and elongate to the surface at mid‐to‐low tidal heights. Transplants were successful in both muddy (8% organic) and sandy (<2%) substrates. Using mesh exclosures to protect the plants from herbivory was critical to restoration success. Although water quality and other habitat conditions are important for SAV growth and survival, restoration in the unvegetated areas studied here was limited by grazing of initial recruits. The establishment of protected founder colonies of sufficient size to withstand initial grazing pressures may be required to reestablish SAV in similar areas.     

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.     

Reindeer reduce biomass of preferred plant species

Abstract. Reduced weights in reindeer that graze in pastures with high reindeer densities have raised the question if coastal summer pastures are modified by grazing. To evaluate this, the impact of reindeer grazing on standing crop was measured by the plant intercept method inside and outside grazing exclosures in the understorey of a coastal mountain birch forest in northern Norway. The understories of coastal birch forests are dominated by vascular plants and are important summer pastures to reindeer. Based on the literature, we made a priori categorization of the vascular plant species into functional groups of preferred forage, less preferred forage and forage of unknown value to reindeer. Intercept frequency was measured within the same plots on three occasions in the summer of 1996. At the end of the grazing season, total standing crop was 33% lower in open plots compared to plots protected by exclosures. However, the reduction varied between the functional groups, with only preferred forage plants being significantly reduced in standing crop (by 49%). Results suggest that reindeer have a strong annual impact on most of the preferred forage species. However, some of the preferred graminoids are tolerant of grazing and dominate the understorey despite decades of high grazing pressure. We suggest that current grazing pressure is favouring the establishment of a few grazing tolerant graminoids, and that this reduces the forage plant variability. The results are discussed in relation to the grazing optimization hypothesis and the potential importance of plant variability for pasture quality.