Illecebrum verticillatum L. in the outer Hebrides

Summary

Scottish habitats are here reviewed in a European context. Examples are selected in order to identify those recognisable as distinctively Scottish as well as others related to Scandinavian or European counterparts. The paper begins with a resumé of the main environmental influences in Scotland, then describes examples of habitats almost unique to, or best represented in Scotland. This is followed by reference to some distinctively western versions of more widely distributed habitats, and others corresponding to related types elsewhere. The bearing of this on site selection for nature conservation is discussed. Hitherto, in the UK selection for National Nature Reserves and Sites of Special Scientific Interest has been based largely on the criteria listed in A Nature Conservation Review (Ratcliffe, 1977). It is important, however, also to review Scottish habitats in a European context, as attempted in this paper. In making proposals (now approaching completion) for Special Areas of Conservation, to be recommended by the UK Government for adoption by the European Commission, it is now our duty to include not only habitats for which we have special responsibility because they are unique to or best represented in Scotland, or have distinctively western features, but also sufficient examples of all the main European types occurring in our country.

Nomenclature of British flowering plants and ferns follows Stace (1991).     

Effects of local extinction of the plains vizcacha (Lagostomus maximus) on vegetation patterns in semi-arid scrub

We studied spatial and temporal effects of local extinction of the plains vizcacha (Lagostomus maximus) on plant communities following widespread, natural extinctions of vizcachas in semi-arid scrub of Argentina. Spatial patterns in vegetation were examined along transects extending outward from active and extinct vizcacha burrow systems. Responses of vegetation to removal of vizcachas were assessed experimentally with exclosures and by documenting vegetation dynamics for 6 years following extinctions. Transect data demonstrated clear spatial patterns in plant cover, particularly an increase in perennial grasses, outward from active vizcacha burrows. These patterns were consistent with predictions based on foraging theory and studies that document grasses as the preferred food of vizcachas. Removal of vizcachas, experimentally and with extinctions, resulted in an immediate increase in perennial and annual forbs indicating that intense herbivory can depress forb cover, as well as grasses. After a 1-year lag following cessation of herbivory, cover of grasses increased. Forbs declined as grasses increased. The long-term effect of extinction of vizcachas was a conversion of colony sites from open patches dominated by forbs to dense bunch grass characteristic of the matrix. Major changes in vegetation occurred within 2–3 years after extinction, resulting in a large pulse of landscape change. However, some species of grasses were uncommon until 5–6 years after the vizcacha extinction. With extinction and colonization, vizcachas generate a dynamic mosaic of patches on the landscape and create temporal, as well as spatial, heterogeneity in semi-arid scrub.     

Effects of Dry-Season N Input on the Productivity and N Storage of Mediterranean-Type Shrublands

Anthropogenic nitrogen (N) deposition is a globally important source of N that is expected to increase with population growth. In southern California, N input from dry deposition accumulates on vegetation and soil surfaces of chaparral and coastal sage scrub (CSS) ecosystems during the summer and fall and becomes available as a pulse following winter rainfall. Presumably, N input will act to stimulate the productivity and N storage of these Mediterranean-type, semi-arid shrublands because these ecosystems are thought to be N limited. To assess whether dry-season N inputs alter ecosystem productivity and N storage, a field experiment was conducted over a 4-year period where plots were exposed to either ambient N deposition (control) or ambient + 50 kgN ha⁻¹ y⁻¹ (added N) that was added as NH₄NO₃ during the fall dry-season of each year. Plots exposed to added N had significantly higher accumulation of NH₄ and NO₃ on ion exchange resins that was due in part to direct fertilization and N mineralization, and the increase in N availability lead to a significant increase in NO₃ leaching in chaparral but not CSS. Nitrogen addition also lead to an increase in litter and tissue N concentration and a decline in the C:N ratio, but failed to alter the ecosystem productivity and N storage of the chaparral and CSS shrublands over the 4-year study period. The reasons for the lack of a treatment response are unknown; however, it is possible that these semi-arid shrublands are not N limited, cannot respond rapidly enough to capture the ephemeral N pulse, are limited by other nutrients, or the N response is dependent on the amount and/or distribution of rainfall. These results have important implications for understanding the potential effects of anthropogenic N deposition on the C and N cycling and storage of Mediterranean-type, semi-arid shrublands. Author Contributions  GLV conceived or designed study, performed research, analyzed data, contributed new methods or models, and wrote the article. SCP performed research, analyzed data, and contributed to the writing of the article. RM performed research, analyzed data, and contributed to the writing of the article.     

Marine influence controls plant phenological dynamics in Mediterranean Mexico

Aims To assess the role of moisture in phenological timing in the mediterranean coastal flora of Baja California, and specifically to assess the role of coastal fog and ocean-derived moisture in plant phenology. Moisture seems to be the primary driver of flowering times and durations at the arid end of the mediterranean-climate region, where rainfall is often sporadic (temperature and day length can be expected to play a much lesser role as they are not growth limiting). We aimed to understand: What factors drive climatic variation between sites? Are there general flowering patterns allowing us to identify phenological categories? Do flowering patterns vary in relation to site-specific weather? and most importantly, does maritime influence on weather affect flowering dynamics in coastal mediterranean ecosystems?Methods The southernmost extent of the California Floristic Province (in Baja California, Mexico) is a biological diversity hotspot of high endemism and conservation value, with two steep moisture gradients: rainfall (N–S) and coastal fogs (W–E), providing an ideal study system. We installed five weather stations across the moisture gradients, recording data hourly. We monitored flowering phenology in the square kilometer surrounding each weather station from 2010 to 2013. About 86 plant taxa were monitored across the five sites, every 6–8 weeks. Averaged climatic data is presented with general trends in flowering, and specific flowering syndromes were observed. Data for flowering intensity across the sites was analyzed using a principal components analysis.Important findings Data analysis demonstrates a general seasonal pattern in flowering times, but distinct differences in local weather and phenology between the five study sites. Three flowering syndromes are revealed in the flora: (i) water responders or spring bloomers, (ii) day-length responders or fall-blooming taxa and (iii) aseasonal bloomers with no seasonal affinity. The two moisture gradients are the strongest drivers of flowering times. Inland sites showed higher phenological variation than coastal sites where seasonality is dampened by ocean-derived moisture, which extends and buffers perennial plant phenology and is a probable driver of local endemism. Phenological controls vary globally with climate and geography; moisture is the primary driver of phenology in mediterranean climates and fog is an important climatic variable in coastal Mexico.     

Bolete Productivity of Cistaceous Scrublands in Northwestern Spain<Superscript>1</Superscript>

Bolete Productivity of Cistaceous Scrublands in Northwestern Spain. Cistaceous scrublands dominated by Cistus ladanifer and Halimium lasianthum are widely distributed in northwestern Spain and are known to harbor the popular edible mushrooms Boletus edulis and B. aereus. As commercial harvest of these species is an important activity in Spain’s forests, in some cases generating greater revenue than timber production, we tried to determine whether there is commercial potential for bolete harvest in young cistaceous scrublands. Transects of 50 m × 2 m were randomly established in cistaceous scrublands and bolete yield was measured. High production of these valuable mushrooms was associated with 3-year-old plants, a much earlier onset than observed for Pinus and Quercus stands. Management recommendations for the studied scrub ecosystems are proposed that take into account the economic value of the mushrooms.     

Predicting drought tolerance from slope aspect preference in restored plant communities

Plants employ strategies of tolerance, endurance, and avoidance to cope with aridity in space and time, yet understanding the differential importance of such strategies in determining patterns of abundance across a heterogeneous landscape is a challenge. Are the species abundant in drier microhabitats also better able to survive drought? Are there relationships among occupied sites and temporal dynamics that derive from physiological capacities to cope with stress or dormancy during unfavorable periods? We used a restoration project conducted on two slope aspects in a subwatershed to test whether species that were more abundant on more water‐limited S‐facing slopes were also better able to survive an extreme drought. The attempt to place many species uniformly on different slope aspects provided an excellent opportunity to test questions of growth strategy, niche preference, and temporal dynamics. Perennial species that established and grew best on S‐facing slopes also had greater increases in cover during years of drought, presumably by employing drought tolerance and endurance techniques. The opposite pattern emerged for annual species that employed drought‐escape strategies, such that annuals that occupied S‐facing slopes were less abundant during the drought than those that were more abundant on N‐facing slopes. Our results clarify how different functional strategies interact with spatial and temporal heterogeneity to influence population and community dynamics and demonstrate how large restoration projects provide opportunities to test fundamental ecological questions.     

Acclimation of photosynthesis and respiration to elevated atmospheric CO2 in two Scrub Oaks

Abstract For two species of oak, we determined whether increasing atmospheric CO₂ concentration (C_a) would decrease leaf mitochondrial respiration (R) directly, or indirectly owing to their growth in elevated C_a, or both. In particular, we tested whether acclimatory decreases in leaf‐Rubisco content in elevated C_a would decrease R associated with its maintenance. This hypothesis was tested in summer 2000 on sun and shade leaves of Quercus myrtifolia Willd. and Quercus geminata Small. We also measured R on five occasions between summer 1999 and 2000 on leaves of Q. myrtifolia. The oaks were grown in the field for 4 years, in either current ambient or elevated (current ambient + 350 µmol mol^?1) C_a, in open‐top chambers (OTCs). For Q. myrtifolia, an increase in C_a from 360 to 710 µmol mol^?1 had no direct effect on R at any time during the year. In April 1999, R in young Q. myrtifolia leaves was significantly higher in elevated C_a—the only evidence for an indirect effect of growth in elevated C_a. Leaf R was significantly correlated with leaf nitrogen (N) concentration for the sun and shade leaves of both the species of oak. Acclimation of photosynthesis in elevated C_a significantly reduced maximum RuBP‐saturated carboxylation capacity (V_{c max}) for both the sun and shade leaves of only Q. geminata. However, we estimated that only 11–12% of total leaf N was invested in Rubisco; consequently, acclimation in this plant resulted in a small effect on N and an insignificant effect on R. In this study measurements of respiration and photosynthesis were made on material removed from the field; this procedure had no effect on gas exchange properties. The findings of this study were applicable to R expressed either per unit leaf area or unit dry weight, and did not support the hypothesis that elevated C_a decreases R directly, or indirectly owing to acclimatory decreases in Rubisco content.     

Recruitment of fleshy-fruited species under different shrub species: Control by under-canopy environment

Seedling densities and the mortality of fleshy-fruited species under 10 European tall-shrub species and the tree Sorbus aria were analyzed in 10-year-old experimental scrub for one growing season. The mean total densities of seedlings of 18 fleshy-fruited species differed under the different hosts, being highest under Crataegus monogyna, Viburnum opulus and Rosa canina (24–20 m^–2); intermediate under Sorbus aria, Rhamnus catharticus, Cornus sanguinea, Ligustrum vulgare and Juniperus communis (17–12 m^–2); and lowest under Euonymus europaeus, Viburnum lantana and Prunus spinosa (10–9 m^–2). This ranking was similar for the 11 planted species and for nine bird-dispersed, non-planted species which invaded the site, notably Hedera helix and Solanum dulcamara. Within the experimental scrub as a whole mortality of seedlings differed significantly among species, being highest in those with the smallest number of recruits. Mortality ranged from 94% and 89% for Solanum dulcamara and Viburnum opulus to 26% for Hedera helix and Ligustrum vulgare. The mean mortality across all species of seedlings did not differ significantly between host species, although it was slightly higher under Juniperus and Cornus (56%, 53%) and lower under Viburnum lantana and Rhamnus (42%, 41%). In four species, the mortality of seedlings was significantly higher under conspecific adults. Mean topsoil water content after a dry spell was negatively correlated with the diffuse site factor (DSF). The mean density of accumulated seedlings of the most abundant seedling species (Hedera) was positively correlated with the topsoil water content. Densities of other species showed no strong correlations with either topsoil water content or DSF. The probable factors controlling recruitment are discussed.     

The narrow-leaf syndrome: a functional and evolutionary approach to the form of fog-harvesting rosette plants

Plants that use fog as an important water-source frequently have a rosette growth habit. The performance of this morphology in relation to fog interception has not been studied. Some first-principles from physics predict that narrow leaves, together with other ancillary traits (large number and high flexibility of leaves, caudices, and/or epiphytism) which constitute the “narrow-leaf syndrome” should increase fog-interception efficiency. This was tested using aluminum models of rosettes that differed in leaf length, width and number and were exposed to artificial fog. The results were validated using seven species of Tillandsia and four species of xerophytic rosettes. The total amount of fog intercepted in rosette plants increased with total leaf area, while narrow leaves maximized interception efficiency (measured as interception per unit area). The number of leaves in the rosettes is physically constrained because wide-leafed plants can only have a few blades. At the limits of this constraint, net fog interception was independent of leaf form, but interception efficiency was maximized by large numbers of narrow leaves. Atmospheric Tillandsia species show the narrow-leaf syndrome. Their fog interception efficiencies were correlated to the ones predicted from aluminum-model data. In the larger xerophytic rosette species, the interception efficiency was greatest in plants showing the narrow-leaf syndrome. The adaptation to fog-harvesting in several narrow-leaved rosettes was tested for evolutionary convergence in 30 xerophytic rosette species using a comparative method. There was a significant evolutionary tendency towards the development of the narrow-leaf syndrome the closer the species grew to areas where fog is frequently available. This study establishes convergence in a very wide group of plants encompassing genera as contrasting as Tillandsia and Agave as a result of their dependence on fog. Electronic Supplementary Material The online version of this article () contains supplementary material, which is available to authorized users.     

Vegetation-type conversion of evergreen chaparral shrublands to savannahs dominated by exotic annual herbs: causes and consequences for ecosystem function

Woody, evergreen shrublands are the archetypal community in mediterranean-type ecosystems, and these communities are profoundly changed when they undergo vegetation-type conversion (VTC) to become annual, herb-dominated communities. Recently, VTC has occurred throughout southern California chaparral shrublands, likely with changes in important ecosystem functions. The mechanisms that lead to VTC and subsequent changes to ecosystem processes are important to understand as they have regional and global implications for ecosystem services, climate change, land management, and policy. The main drivers of VTC are altered fire regimes, aridity, and anthropogenic disturbance. Some changes to ecosystem function are certain to occur with VTC, but their magnitudes are unclear, whereas other changes are unpredictable. I present two hypotheses: (1) VTC leads to warming that creates a positive feedback promoting additional VTC, and (2) altered nitrogen dynamics create negative feedbacks and promote an alternative stable state in which communities are dominated by herbs. The patterns described for California are mostly relevant to the other mediterranean-type shrublands of the globe, which are biodiversity hotspots and threatened by VTC. This review examines the extent and causes of VTC, ecosystem effects, and future research priorities.