Community development by forest understory plants after prolonged burial by tephra

Several processes bury plants, but sediment can also be subsequently removed, often by delayed erosion. Thus, the ability to survive multiple years of burial and to respond when released are important to vegetation changes and population dynamics. We experimentally evaluated the effects of delayed removal of tephra (aerially transported volcanic ejecta) in an old-growth forest understory near Mount St. Helens, using 1-m² plots assigned to three treatments: tephra removed 4 months after deposition (50 plots), tephra removed 28 months after deposition (the delayed erosion treatment, 50 plots), and undisturbed, natural tephra (100 plots). Prior to tephra removal, species density, cover, shoot density, and shoot size in the delayed erosion treatment were all similar to values in natural plots and significantly less than values in plots cleared initially, indicating that 24 months of additional burial adversely affected understory plants. However, all attributes eventually approached pre-eruption values for shrubs and herbs, indicating that erosion greatly facilitated vegetation recovery. Responses varied substantially among species and growth forms. Overall, our experimental results indicate that some plants of most species can respond effectively after release from burial of at least three growing seasons. In addition, the delay of erosion retards ecosystem recovery relative to early erosion, facilitates recovery relative to no erosion, and modifies the trajectory of post-disturbance vegetation change.

     

Causes of plant community divergence in the early stages of volcanic succession

Question: How do temporal changes in plant communities occur after volcanic eruptions? What characteristics determine successional divergence or convergence? Location: The summit area of Mount Usu, northern Japan, completely destroyed by 1‐3 m of thick ash and pumice during the 1977‐1978 eruptions. Habitats were classified into three types: gullies where the pre‐eruption topsoil was exposed due to the erosion of tephra (EG), gullies covered with tephra (CG), and outside of gullies covered with thick tephra (OG). Methods: Plant community structure was monitored for 15 years from 1983 to 1997 in 14 2 m × 5 m permanent plots. The data were summarized by species diversity, life form, and the detrended correspondence analysis. Results: The common species were perennial herbaceous plants, but habitat preferences differed between species. Seed bank species, including a nitrogen fixer Trifolium repens, were dominant in EG, and excluded the establishment of the later colonists. Pioneer trees slowly increased in cover. The detrended correspondence analysis indicated that species composition in the earlier stages did not differ greatly between plots. Thereafter, three patterns of temporal community changes were observed: seed bank species persisted in EG, and in OG and CG forest development proceeded or community structure did not change greatly. Conclusion: Pre‐eruption topsoil contributed to revegetation by the supply of seed bank and nutrients in the earliest stages, but resulted in the delay of forest development due to the persistence of seed bank species. Plant community divergence was driven by the persistence of earlier colonists.     

Recovery of forest understories buried by tephra from Mount St. Helens

To determine the effects of tephra (volcanic aerial ejecta) on forest understory plants, six sites were chosen along a tephra depth gradient (23 to 150 mm) northeast of Mount St. Helens, USA. All sites were in old forests beyond the limits of direct blast damage from the volcanic eruption. At each site, 150 one m² plots were permanently marked; all tephra was removed from 50 of these in 1980. Cover and density of plant species were recorded during 1980, 1981, and 1982.Tephra 23 mm deep had almost no effect on cover and density of vascular plants, and reduced bryophyte cover for only two years. Tephra 45 mm deep destroyed almost all bryophytes. Although damaged by 45 mm tephra, deciduous herbs recovered by 1982, but some evergreen species did not. Tephra 75 mm deep reduced herb cover in 1982 to 32% and density to 26% of that in cleared plots. At two sites with an average tephra depth of 150 mm, almost all herbs were eliminated except in microsites where tephra was thin, but shrub abundance was greatly reduced only where snow had been present during tephra deposition. Almost all cover was contributed by plants established previous to the eruption; seedling cover never exceeded 0.2%. Refugia with thin tephra, resulting from erosion, were vital to the survival of many species, especially bryophytes.Nomenclature of vascular plants follows Hitchcock & Cronquist (1973); moss nomenclature follows Lawton (1971).We thank T. Hill, C. Halpern and B. Smith for field assistance. The USDA Forest Service, and especially J. F. Franklin, facilitated entry into the restricted area around Mount St. Helens. This word was supported by the National Science Foundation, USDA Science and Education Administration, and the U.S. Forest Service Pacific Northwest Forest and Range Experiment Station.     

Incorporating Restoration in Sustainable Forestry Management: Using Pine-Bark Mulch to Improve Native Species Establishment on Tephra Deposits

Native plant establishment is limited by harsh environmental conditions in areas affected by tephra deposition following volcanic eruptions. Late‐successional species might be lacking even decades after the disturbance. We assessed the effectiveness of pine‐bark mulch, a by‐product of sustainable timber production in the study area, in promoting the establishment and survival of a late‐successional species (Pinus pseudostrobus) and a nitrogen‐fixing legume (Lupinus elegans). We established a factorial experiment in areas covered with tephra during the eruption of the Paricutín volcano in the state of Michoacán, Mexico. After 1 year, P. pseudostrobus survival was significantly higher (p < 0.001) in plots with pine‐bark mulching (46.5%) than in plots without mulching (21.8%). After 2 years, surviving pines with mulching were significantly taller (p= 0.03) than pines without mulching (45.3 ± 3.8 cm and 31.2 ± 3.7 cm, respectively). Lupinus elegans plants survived longer when grown in plots with pine‐bark mulching than without mulching. Mulching reduced tephra temperatures during the dry season (when temperatures can reach up to 58°C 4 cm below the surface of bare tephra). Lupinus elegans plants were affected by herbivory by small rodents, run‐off, and frost at the end of the growing season. Our results suggest that mulching can ameliorate harsh environmental conditions on sites covered with tephra while incorporating a by‐product of sustainable forestry into restoration practice.     

Fine-scale dynamics of rhizomes in a grassland community

Spatial dynamics in grassland communities are constrained by the belowground spatial distribution of roots and rhizomes. Their dynamics are difficult to measure as underground data collection tends to be destructive and cannot be repeated at the same plot over time. We investigated rhizome dynamics indirectly by examining rhizome spatial structure on long‐term grassland study plots where aboveground shoot counts have been recorded using a fine‐scale grid over nine years. Number of rhizome apical ends, basal ends and total rhizome length of both live and dead rhizomes were obtained from the data by scanning rhizomes and processing them by GIS vectorization. These rhizome variables were correlated with the above‐ground shoot counts in grid cells over varying temporal lags. There was a general decrease in the intensity of correlation between live rhizomes and shoot counts with increasing time lags. Correlation of dead rhizomes increased with increasing time lag, reaching a maximum after several years, and then declined. Species differed strongly in the change of rhizome‐shoot counts correlation over varying time lags. These differences were used to infer rhizome growth dynamics, namely rhizome growth rate and lifespan, and rhizome mean decomposition time. The species involved differed in all these traits. Mean rhizome growth rate ranged from 0.2 (Polygonum bistorta) to 3.3 cm ur^?1 (Deschampsia flexuosa); mean rhizome lifespan ranged from 5 yr (Anthoxanthum alpinum) to over 8 yr (Nardus stricta) and mean decomposition time from one growing season (Anthoxanthum) to 7 yr (Polygonum). Presence of dead rhizomes below living rhizomes or aboveground shoots was taken as an indication of fine‐scale replacements between species. These were highly non‐random, with some species pairs replacing significantly more frequently. These differences in rhizome growth parameters underlie different strategies of horizontal growth and dieback between species. These can serve as one of the mechanisms of species replacements and contribute to the fine‐scale coexistence of species.     

SEEDLING ESTABLISHMENT IN FORESTS AFFECTED BY TEPHRA FROM MOUNT ST. HELENS

Seedling density on permanent plots at five sites was monitored for the first four summers following the deposition of 4.5 to 15 cm of tephra from the 1980 eruption of Mount St. Helens. Because the old-growth forests at the sites were not destroyed by the volcanic eruption, the plots were under a normal tree canopy. Almost no seedlings established in 1980. By 1983 tree seedling density exceeded 35 m^−-2 at all sites. Tsuga heterophylla seedlings were most common, with Abies amabilis seedlings next in abundance. A dense layer of small trees may develop on the tephra and long-term forest dynamics could be affected. There was no successful invasion by species not already present in the stands. Seedlings of some forest herbs, especially Tiarella trifoliata, were common, but many other common forest species produced none. It is still problematical if or when these species will return to areas of the forest from which they were extirpated.     

Dynamics of submersed aquatic vegetation on shallow soft bottoms in the Baltic Sea

Abstract. Persistence and colonization of submersed aquatic plant species were studied in permanent plots (20 cm × 20 cm) at three shallow sites adjacent to Askö island, in the northern Baltic Sea. The study started in 1991 at two of the sites, in 1992 at a third site and continued until 1997. Two major weather‐induced disturbances occurred during the study: a long, stormy period during a mild winter and a cold winter with extreme low water levels. The stormy period caused a large loss of species from plots (95 ± 5% SE) at the most wave‐exposed site, resulting in a low species number per plot (0.8 ± 0.2) the following summer. During the three following years the mean species number increased to 3.6 ± 0.2. The cold winter caused high species turnover and increased species number per plot at the two most wave‐exposed sites. The species turnover at the sheltered site was highest in the two years with low water level in May, possibly due to increased waterfowl herbivory. Annuals, loosely anchored and highly reproductive species had significantly higher plot colonization rates and lower persistence than perennials, well‐anchored species and species with none or low reproduction. The extent of lateral growth had no significant effect on colonization or persistence. Although these disturbances have a large effect on the dynamics, species mobility was also high in other years. Relative to other, similar, studies in terrestrial vegetation mean persistence in plots was remarkably low and mean mobility and species turnover rates were very high.     

Resilience and alternative equilibria in a mire plant community after experimental disturbance by volcanic ash

Following disturbance events vegetation can either be resilient and return to its original state, or there can be shifts in vegetation composition and abundance patterns that may indicate alternative equilibiria. We conducted a long‐term field experiment that simulated impact by aerially transported volcanic ejecta (tephra) in a Sphagnum‐dominated plant community in order to test the effects of this type of large‐scale disturbance. Sixty plots of 1.4 × 1.4 m were established at Sarobetsu mire in northern Hokkaido, Japan and subjected to seven treatments (including the control) with natural tephra or ground glass shards differing in grain size, layer thickness and season of application. Water chemistry and vegetation were surveyed before tephra application and during five and eight years after the perturbation, respectively. Leaching of ions from fine‐grained glass shards caused a sustained increase of soil water pH and electric conductivity. Under coarser materials water quality differed little from the control, but a short‐lived peak of potassium suggested that mechanisms like nutrient release from decomposing plant material may influence water chemistry after volcanic disturbance. The perturbation initially reduced the cover of the dominant functional group (Sphagnum mosses) in all treatments; vascular plants were less affected. All species were able to recover by growing through the tephra, and open tephra surfaces were colonized by ubiquitous cryptogams, but not by spermatophytes. In contrast to the overall resilient behaviour of the vegetation, in some plots that received natural tephra an alternative state with high cover of the dwarf shrub Myrica gale developed. The patterns indicated that physical and chemical properties of the tephra determined the initial effects on plants, but that stochastic processes contributed to subsequent succession. These are likely to have effects on ecosystem functioning, e.g. hydrological processes and carbon sequestration.     

Plant traits and functional types in response to reduced disturbance in a semi‐natural grassland

Abstract. Question: How do functional types respond to contrasting levels of herbage use in temperate and fertile grasslands? Location: Central France (3°1’E, 45°43’N), 870 m a.s.l. Methods: Community structure and the traits of dominant plant species were evaluated after 12 years of contrasted grazing and mowing regimes in a grazing trial, comparing three levels of herbage use (high, medium and low). Results and Conclusions: Of 22 measured traits (including leaf traits, shoot morphology and composition, phenology), seven were significantly affected by the herbage use treatment. A decline in herbage use reduced individual leaf mass, specific leaf area and shoot digestibility, but increased leaf C and dry matter contents. Plants were taller, produced larger seeds and flowered later under low than high herbage use. Nine plant functional response types were identified by multivariate optimization analysis; they were based on four optimal traits: leaf dry matter content, individual leaf area, mature plant height and time of flowering. In the high‐use plots, two short and early flowering types were co‐dominant, one competitive, grazing‐tolerant and moderately grazing‐avoiding, and one grazing‐avoiding but not ‐tolerant. Low‐use plots were dominated by one type, neither hardly grazing‐avoiding nor grazing‐tolerant, but strongly competitive for light.     

Limits to convergence of vegetation during early primary succession

Questions: Primary succession, measured by changes in species composition, is slow, usually forcing a chronose‐quence approach. A unique data set is used to explore spatial and temporal changes in vegetation structure after a 1980 volcanic eruption. On the basis of data from a transect of 20 permanent plots with an altitudinal range of 250 m sampled through 2005, two questions are asked: Do changes along the transect recapitulate succession? Do plots converge to similar composition over time? Location: A ridge between 1218 and 1468 m on Mount St. Helens, Washington, USA. Methods: Repeat sampling of plots for species cover along a 1‐km transect. Floristic changes were characterized by techniques including DCA, clustering and similarity. Results: Species richness and cover increased with time at rates that decreased with increasing elevation. The establishment of Lupinus lepidus accelerated the rate of succession and may control its trajectory. Diversity (H) at first increased with richness, then declined as dominance hierarchies developed. Primary succession was characterized by overlapping phases of species assembly (richness), vegetation maturation (diversity peaks, cover expands) and inhibition (diversity declines). Each plot passed through several community classes, but by 2005, only four classes persisted. Succession trajectories (measured by DCA) became shorter with elevation. Similarity between groups of plots defined by their classification in 2005 did not increase with time. Similarity within plot groups converged slightly at the lower elevations. Despite similarities between temporal and spatial trends in composition, trajectories of higher plots do not recapitulate those of lower plots, apparently because Lupinus was not an early colonist. Any vegetation convergence has been limited to plots that are in close proximity.     

Use of sheep grazing in the restoration of semi‐natural meadows in northern Finland

Abstract. The biodiversity of species‐rich semi‐natural meadows is declining across Europe due to ceased management. In this study we aimed to find out how successfully the local species richness of an overgrown semi‐natural mesic meadow could be restored by sheep grazing after a long period of abandonment. The cover of vascular plant species in grazed plots and ungrazed exclosures was studied for five years and the responses of different functional plant groups were followed (herbs vs grasses, tall vs short species, species differing in flowering time, species representing different Grime's CSR strategies and species indicative of rich vs poor soil). Grazing increased species number by nearly 30%. On grazed plots the litter cover practically disappeared, favouring small herbs such as Rhinanthus minor, Ranunculus acris, Trifolium pratense and the grass Agrostis capillaris. Grazing decreased the cover of the late flowering tall herb Epilobium angustifolium but had no effect on the abundance of the early flowering tall herbs Anthriscus sylvestris or Geranium sylvaticum. We suggest that to succeed in restoration it is useful to determine the responses of different functional plant groups to grazing. Grassland managers need this information to optimize the methods and timing of management used in restoration. Additional management practices, such as mowing, may be needed in mesic meadows to decrease the dominance of tall species. The availability of propagules seemed to restrict further increase of species richness in our study area.     

Relationship between topographic heterogeneity and vegetation patterns in a Californian salt marsh

Questions: Are species richness and species abundances higher in the presence of tidal creeks? Do species richness and species abundances vary with plot size? Location: Intertidal plain of Volcano Marsh, Bahia de San Quintin, Mexico. Methods: We analysed vegetation patterns in large areas (cells) with tidal creeks (+creek) and without (‐creek). We surveyed vegetation cover, microtopography, habitat type, and distance to creeks in nested plots of five sizes, 0.1, 0.25, 1, 2.5, and 10 m². Results: Species richness, frequency, cover, and assemblages differed between ±creek cells. Richness tended to be higher in +creek cells, and cover and frequency of individual species differed significantly between ±creek cells. We found consistent patterns in vegetation structure across plot sizes. We encountered 13 species that occurred in 188 unique assemblages. The most common assemblage had six species: Batis maritima, Frankenia salina, Salicornia bigelovii, S. virginica, Salicornia spec. and Triglochin concinna. This assemblage occurred in ±creek cells and at all spatial scales. Of the most common assemblages all but one were composed of multiple species (3–9 species/plot). Conclusions: The persistence of vegetation patterns across a 100‐fold range in spatial scale suggests that similar environmental factors operate broadly to determine species establishment and persistence. Differences in assemblage composition result from variation of frequency and cover of marsh plain species, particularly Suaeda esteroa and Monanthochloe littoralis. The recommendation for restoration of Californian salt marshes is to target (and plant) multi‐species assemblages, not monocultures.     

Clear‐felling effects on colonization rates of shade‐tolerant forest herbs into a post‐agricultural forest adjacent to ancient forest

Question: Does clear‐felling influence forest herb colonization into post‐agricultural forest? Location: A stand of poplar cultivars with a dense understorey of Acer pseudoplatanus in Muizen forest (northern Belgium), planted in 1952 on farmland adjacent to ancient forest and clear‐felled in 1997. Methods: Shade‐tolerant forest herbs were surveyed in 112 grid‐based sample plots: just before clear‐felling, and 5 and 10 yr afterwards. Shade‐tolerant herbs were subdivided into ancient forest species (AFS) and other shade‐tolerant species (OSS). Effects of clear‐felling on species number per plot, total cover per plot and colonization rate of species groups were compared using non‐parametrical tests. Species number per plot was modelled by means of generalized linear mixed models (GLMMs), with inventory time, distance to the nearest parcel edge, and cover of light‐loving species (LS) as explanatory variables. The C‐S‐R signature (competitive, stress‐tolerant and ruderal strategies, respectively) shift of sample plots was calculated on the selected shade‐tolerant species. Results: Frequency of most species increased during the 10‐yr period. Number of OSS increased more and faster than that of AFS. OSS increased to the level of the adjacent forest, but was lower where LS cover remained high. There was a positive correlation between the change of the colonization rate and the competitive plant strategy. Conclusions: We assume that clear‐felling stimulated generative reproduction of shade‐tolerant herbs, whereas quickly emerging woody species controlled competitive exclusion by LS. Succession of dark and light phases, such as provided by an understorey managed as a coppice, could promote colonization of shade‐tolerant herbs into post‐agricultural forest.     

Effects of plot size on the ordination of vegetation samples

Questions: Do ordination patterns differ when based on vegetation samples recorded in plots of different size? If so, how large is the effect of plot size relative to the effects of data set heterogeneity and of using presence/absence or cover‐abundance data? Can we combine plots of different size in a single ordination? Methods: Two homogeneous and two heterogeneous data sets were sampled in Czech forests and grasslands. Cover‐abundances of plant species were recorded in series of five or six nested quadrats of increasing size (forest 49‐961 m2; grassland 1‐49 m²). Separate ordinations were computed for plots of each size for each data set, using either species presences/absences or cover‐abundances recorded on an ordinal scale. Ordination patterns were compared with Procrustean analysis. Also, ordinations of data sets jointly containing plots of different size were calculated; effects of plot size were evaluated using a Monte Carlo test in constrained ordination. Results: The results were consistent between forest and grassland data sets. In homogeneous data sets, the effect of presence/absence vs. cover‐abundance was similar to, or larger than, the effect of plot size; for presence/absence data the differences between ordinations of differently sized plots were smaller than for cover‐abundance data. In heterogeneous data sets, the effect of plot size was larger than the effect of presence‐absence vs. cover‐abundance. The plots of smaller size (= 100 m2 in forests, = 4 m² in grasslands) yielded the most deviating ordination patterns. Joint ordinations of differently sized plots mostly did not yield patterns that would be artifacts of different plot size, except for plots from the homogeneous data sets that differed in size by a factor of four or higher. Conclusions: Variation in plot size does influence ordination patterns. Smaller plots tend to produce less stable ordination patterns, especially in data sets with low ß‐diversity and species cover‐abundances. Data sets containing samples from plots of different sizes can be used for ordination if they represent vegetation with large ß‐diversity. However, if data sets are homogeneous, i.e. with low ß‐diversity, the differences in plot sizes should not be very large, in order to avoid the danger of plot size differences distorting the real vegetation differentiation in ordination patterns.     

Morphological and phenological shoot plasticity in a Mediterranean evergreen oak facing long-term increased drought

Mediterranean trees must adjust their canopy leaf area to the unpredictable timing and severity of summer drought. The impact of increased drought on the canopy dynamics of the evergreen Quercus ilex was studied by measuring shoot growth, leaf production, litterfall, leafing phenology and leaf demography in a mature forest stand submitted to partial throughfall exclusion for 7 years. The leaf area index rapidly declined in the throughfall-exclusion plot and was 19% lower than in the control plot after 7 years of treatment. Consequently, leaf litterfall was significantly lower in the dry treatment. Such a decline in leaf area occurred through a change in branch allometry with a decreased number of ramifications produced and a reduction of the leaf area supported per unit sapwood area of the shoot (LA/SA). The leafing phenology was slightly delayed and the median leaf life span was slightly longer in the dry treatment. The canopy dynamics in both treatments were driven by water availability with a 1-year lag: leaf shedding and production were reduced following dry years; in contrast, leaf turnover was increased following wet years. The drought-induced decrease in leaf area, resulting from both plasticity in shoot development and slower leaf turnover, appeared to be a hydraulic adjustment to limit canopy transpiration and maintain leaf-specific hydraulic conductivity under drier conditions.     

Effect of tephra depth on vegetation development in areas affected by volcanism

Plant establishment on sites affected by major volcanic disturbances is limited by several factors, such as lack of suitable microsites for germination and establishment in sites affected by tephra from volcanic eruptions. Even after long periods of time, tephra deposited over un-vegetated areas (agricultural fields and other barren areas) lack closed vegetation cover and in many cases late successional species. To assess limiting factors for plant establishment, a field survey in a tephra deposit from the Paricutin volcano eruption (19°30′42.4′′ N, 102°12′03.0′′) and greenhouse experiments were carried out. The field survey determined the relationship between tephra depth and vegetation distribution. Greenhouse experiments determined the effect of tephra depth on establishment and growth of two dominant species in the tephra deposit surveyed, Eupatorium glabratum and Lupinus elegans. Our results suggest that size and spatial distribution of vegetation patches is related to tephra depth in the field (77% of the vegetation patches were on tephra 38.8 cm deep or less and only 2% on tephra of more than 46.8 cm). Under greenhouse conditions, Eupatorium glabratum and Lupinus elegans height sharply decreased as depth of the tephra layer increased. Lupinus elegans plants growing in tephra less than 30 cm deep had a mean weight of 10.56 g (±0.53 g) compared with 3.11 g (±0.46 g) for plants growing in tephra more than 30 cm deep. Our results suggest that tephra depth is a limiting factor for canopy development in barren areas affected by tephra deposition.     

Mowing Reduces Exotic Annual Grasses but Increases Exotic Forbs in a Semiarid Grassland

Cheatgrass (Bromus tectorum) and other exotic winter‐active plants can be persistent invaders in native grasslands, growing earlier in the spring than native plants and pre‐empting soil resources. Effective management strategies are needed to reduce their abundance while encouraging the reestablishment of desirable native plants. In this 4‐year study, we investigated whether mowing and seeding with native perennial grasses could limit growth of exotic winter‐actives, and benefit growth of native plants in an invaded grassland in Colorado, United States. We established a split‐plot experiment in October 2008 with 3 mowing treatments: control, spring‐mowed, and spring/summer‐mowed (late spring, mid‐summer, and late summer), and 3 within‐plot seeding treatments: control, added B. tectorum seeds, and added native grass seeds. Cover of plant species and aboveground biomass were measured for 3 years. In March and June of 2010, 2011, and March of 2012, B. tectorum and other winter‐annual grasses were half as abundant in both mowing treatments as in control plots; however, cover of non‐native winter‐active forbs increased 2‐fold in spring‐mowed plots and almost 3‐fold in spring/summer‐mowed plots relative to controls. These patterns remained consistent 1 year after termination of treatments. Native cool‐season grasses were most abundant in spring‐mowed plots, and least abundant in control plots. There was higher cover of native warm‐season grasses in spring/summer‐mowed plots than in control plots in July 2011 and 2012. The timing of management can have strong effects on plant community dynamics in grasslands, and this experiment indicates that adaptive management can target the temporal niche of undesirable invasive species.     

Spatial scale of observation affects α, β and γ diversity of cavity-nesting bees and wasps across a tropical land-use gradient

Aim Anthropogenic changes in land use may have major consequences for global biodiversity. However, species diversity is determined by a suite of factors that may affect species differently at different spatial scales. We tested the combined effects of land use and spatial scale on α, β and γ diversity in the tropics using experimental communities of cavity‐nesting bees and waSPS (Hymenoptera: Aculeata). We aimed to determine whether: (1) land‐use intensity negatively affects species richness of cavity‐nesting Hymenoptera, (2) β diversity, both within and between plots, is higher in more natural systems, (3) species richness of flowering herbs correlates positively with species richness of Hymenoptera within and across habitats, (4) richness of cavity‐nesting Hymenoptera in highly modified habitats declines with increasing distance from natural or semi‐natural habitats, (5) the effects of land use, herb diversity and forest distance on Hymenoptera α and β diversity vary at different spatial scales, and (6) bees and waSPS respond to land use in a similar way. Location Manabi, south‐west Ecuador. Methods We examined diversity (species richness) within 48 plots of five habitat types that comprised a gradient of decreasing agricultural intensity from rice and pasture to coffee agroforests, unmanaged abandoned agroforests and forest fragments, using standardized nesting resources for reproducing communities of cavity‐nesting bees and waSPS. Results (1) Land use significantly affected α diversity of trap‐nesting bees and waSPS at the subplot (per trap) scale, but not subplot β diversity or plot‐scale species richness (γ diversity). (2) Beta diversity was surprisingly higher between plots within a land‐use type than between land‐use types. (3) Species richness of bees and waSPS increased with diversity of flowering herbs at the subplot (trap) scale only. (4) Forest distance correlated positively with bee species richness at the plot scale only. (5) Land use, herb diversity and forest distance each showed significant correlations with bee and wasp diversity at only one spatial scale. (6) Despite differences in life history, bees and waSPS responded to land‐use intensity in a similar way. Main conclusions The effects of land use on species richness were highly dependent on spatial scale. Subplot‐scale analyses showed that rice and pasture contained the highest species diversity, whereas plot‐scale analyses showed no significant difference in the diversity of different land‐use types. We emphasize caution in the estimation of biodiversity at only one spatial scale, and highlight the surprisingly large contribution of managed land to the regional biodiversity of these species.     

Plant diversity,species turnover and shifts in functional traits in coastal dune vegetation: Results from permanent plots over a 52‐year period

Question: What is the relationship between plant diversity and species turnover in coastal dune vegetation plots? How is the long‐term change in species composition of vegetation plots related to shifts in functional traits, and what does it tell us about the dominant processes? Location: Coastal dunes, the Netherlands. Methods: Our data set comprised 52 years of vegetation data from 35 permanent plots in grassland/scrub/woodland vegetation. Vegetation dynamics were described in terms of changes in species composition and abundance, and shifts in 13 functional traits related to resources capture and forage quality, regeneration and dispersal. Results: Species turnover in the plots was high, because of local extinction and colonization. Species‐rich plots were more stable in terms of species abundance and composition compared with species‐poor plots. Over time, the plots converged with respect to their abiotic conditions, as reflected by Ellenberg indicator values – indicating that the prevailing process was succession. The high species turnover reflected high invasibility: accordingly, the relative importance of annuals increased. Most newcomer annuals, however, were competitive generalists of little conservation value. The functional trait analysis allowed us to unravel the complexity of effects of disturbances and succession, and yielded information on the processes driving the observed vegetation dynamics. Conclusions: In this study, small‐scale species turnover was negatively related to species diversity, indicating more stability in species‐rich communities. Regarding shifts in trait diversity, unifying filters appeared to be more dominant than diversifying filters. Counteracting this homogenization process poses a challenge for nature management.     

Direct effects of tephra fallout from the Puyehue–Cordón Caulle Volcanic Complex on Nothofagus pumilio ring widths in northern Patagonia

We evaluated the radial growth response of adult Nothofagus pumilio (Poepp. et Endl) Krasser trees affected by tephra deposition following historical volcanic eruptions of the Puyehue–Cordón Caulle Volcanic Complex (PCCVC) in northern Patagonia. Standard tree–ring width chronologies were developed for trees from two sites that were affected by up to 55 cm of tephra during the 2011 eruption, which allowed us to detect the general tree–growth response to eruptions VEI ≥ 3 and VEI ≤ 2. The tree growth trend satisfactorily followed the mean temperature record (r = 0.42); however, the analysis of studentized residuals identified outliers (≥ ± 2 SD) directly related to the volcanic eruptions of the years 1921–1922 and 2011 and the respective post–eruption years, while for the 1960 eruption and following year, they largely exceeded the mean value of the residuals. The large amount of tephra deposited during the 1921–22 and 2011 eruptions caused physical damage to the tree canopy leading to the appearance of white rings and to locally absent rings. The rate of change in radial growth of trees during these eruptions presented significant declines in relation to the growth of five years before the eruption and to the following year. The low amount of tephra deposited during the 1960 eruption did not cause damage to the stands and trees increased their radial growth. In general, trees that had reduced radial growth experienced a remarkable recovery starting in the second or third post–eruption year. The amount of tephra deposited and the time of year of the volcanic eruptions had an important influence on tree rings. Some ecophysiological causes that could explain the growth responses of N. pumilio to tephra fall are discussed herein. Our study may provide useful insights to clarify the uncertain characteristics of some eruptions in the past or to detect the occurrence of large, undocumented volcanic eruptions throughout the Andes.