Phenological complementarity, species diversity, and ecosystem function

Increasing species diversity frequently enhances ecosystem function. Phenological complementarity, the asynchrony of species resource use and growth, may explain how species diversity influences ecosystem function but remains largely untested. We used an early successional plant community containing species with a variety of phenologies to test whether increasing species diversity enhances ecosystem function by increasing phenological complementarity. Over a two-year period, we increased environmental heterogeneity within an abandoned field with variation in disturbance, soil nutrients, water, light availability, and disturbance in 160 permanent plots, and measured percent cover of each plant species three times in each growing season. We did not manipulate species composition directly, and thus diversity and complementarity in each plot were the result of pre-existing conditions and responses of individuals to experimental treatments. Species diversity was measured in two ways, as the total number of species per plot and as the evenness of species abundances. Phenological complementarity was measured as the negative logarithm of the variance ratio. We tested whether the number of plant species per plot, species evenness, and their phenological complementarity in the first year predicted total annual cover in the second year. Total annual cover increased only moderately with number of species and evenness, consistent with studies that randomize species composition among replicate plots. Any effect that species number or evenness had on total annual cover, however, was not due to phenological complementarity. Rather, diversity was unrelated to phenological complementarity. These results indicate that naturally occurring variation in species diversity had little effect on whether phenological complementarity can enhance ecosystem function.     

Bat Diversity and Movement in an Agricultural Landscape in Matiguás, Nicaragua

Although agriculture dominates much of Central America, little is known about the bat assemblages that occur within agricultural landscapes and how bats use different types of tree cover within these landscapes. Using mist-nets and a mark-recapture protocol, we compared bat diversity and movement across six types of tree cover within an agricultural landscape in central Nicaragua. The tree cover types surveyed included secondary forests, riparian forests, forest fallows, live fences, pastures with high tree cover and pastures with low tree cover. We captured a total of 3084 bats of 39 species, including two new species records for the country ( Lonchorhina aurita and Molossops greenhalli ). Of these, 2970 bats and 27 species were in the Phyllostomidae family. There were significant differences in mean species density, abundance and evenness of phyllostomid bats across the different types of tree cover, but not in bat diversity. Riparian forests had the highest mean species density and bat abundance per plot. In contrast, mean bat abundance and species density were lowest in pastures with low tree cover. Of the 1947 phyllostomid bats marked, a total of 64 bats of eight species were recaptured. The average linear distance between extra-site recaptures was 2227 m (± 228 SE) and the maximum distance was 10.6 km. Bats were recorded moving between almost all types of tree cover, and especially to and from riparian forests. Our study suggests that agricultural landscapes retaining a heterogeneous tree cover may maintain a diverse bat assemblage, and that bats visit and use a variety of tree cover types within the agricultural matrix.     

How reliable is the monitoring of permanent vegetation plots? A test with multiple observers

Questions: A multiple plot design was developed for permanent vegetation plots. How reliable are the different methods used in this design and which changes can we measure? Location: Alpine meadows (2430 m a.s.l.) in the Swiss Alps. Methods: Four inventories were obtained from 40 m² plots: four subplots (0.4 m²) with a list of species, two 10m transects with the point method (50 points on each), one subplot (4m²) with a list of species and visual cover estimates as a percentage and the complete plot (40 m²) with a list of species and visual estimates in classes. This design was tested by five to seven experienced botanists in three plots. Results: Whatever the sampling size, only 45‐63% of the species were seen by all the observers. However, the majority of the overlooked species had cover < 0.1%. Pairs of observers overlooked 10‐20% less species than single observers. The point method was the best method for cover estimate, but it took much longer than visual cover estimates, and 100 points allowed for the monitoring of only a very limited number of species. The visual estimate as a percentage was more precise than classes. Working in pairs did not improve the estimates, but one botanist repeating the survey is more reliable than a succession of different observers. Conclusion: Lists of species are insufficient for monitoring. It is necessary to add cover estimates to allow for subsequent interpretations in spite of the overlooked species. The choice of the method depends on the available resources: the point method is time consuming but gives precise data for a limited number of species, while visual estimates are quick but allow for recording only large changes in cover. Constant pairs of observers improve the reliability of the records.     

Does size really matter? Species–area relationships in human settlements

Aim To describe species–area relationships in human settlements and compare them with those from a non‐urban habitat. Location West‐central Mexico. Methods We surveyed breeding birds in 13 human settlements and five shrubland patches. We estimated bird species richness using an abundance‐based coverage estimator with equal sample sizes to eliminate biases related to sampling effort differences. To assess species–area relationships, we performed log–log linear regressions between the size of the studied patches and their estimated bird richness. We also used a logarithmic approach to determine how the species–area relationship asymptoted and made use of the Michaelis–Menten model to identify the size at which the studied patches reached their maximum species richness. We also investigated (1) possible relationships among the estimated bird richness and other variables known to affect urban‐dwelling birds (built cover, plant species richness, tree cover or human population density) and (2) changes in bird community composition related to the size of the studied human settlements. Results Species–area relationships exhibited different patterns among the studied habitats. The log–log regression slope was steeper in human settlements, while the intercept was higher in shrublands. The maximum number of species was more than twofold higher in shrublands. Human settlement patch size was the only variable significantly related to bird richness. Our community composition results show that two main bird groups are related to human settlement size, and that as the size of human settlements increases, bird community similarity in relation to the largest city increases. Main conclusions Human settlements act as ecological islands, with pronounced species–area relationships. Our results suggest that an important threshold for bird species richness and community composition is reached in human settlements > 10.2 km². This threshold is unlikely to be generalizable among bio‐regions, and thus should be quantified and considered when studying, managing and/or planning urban systems.     

Seed Rain in High-Altitude Restoration Plots in Switzerland

Seed rain was studied in restoration plots installed in 1985 and 1987, respectively, on an alpine downhill ski run at circa 2,500 m above sea level. The study was initiated in late autumn 1996 and completed in autumn 1998; it included temporal and spatial variation in density per m², as well as alpha diversity (species richness), and species composition of the seed rain versus that of the resident vegetation. This is the first report on post-restoration monitoring of seed rain above the timberline. Seed rain density and alpha diversity varied seasonally, with the first peak occurring immediately after spring snowmelt and the second in early autumn. The density of seed rain varied between plots and years (1,528–1,778 seeds per m² in one plot [RPF] versus 1,096–3,557 seeds per m² in another plot [RPG]). Total species number per plot was nearly twice as high in RPF as in RPG in both study years. Seed rain totaled 18 species; all but one represented either transplants introduced in restoration or colonizers established in the plots soon after restoration. Distribution of species in seed rain was largely asymmetric and only a few species provided substantial contributions. Composition of species and their respective contribution to seed rain differed between plots and was clearly influenced by performance of some species used in restoration as transplants; together they provided as much as 51% of the total seed rain. The results of the study demonstrate that restoration enhanced increase of species richness as well as seed rain in situ.     

The influence of scale and species pool on the relationship between vascular plant species richness and cover in an alpine area in Norway

Recent studies emphasise the potential importance of scale and species pool on the humped-back or unimodal relationship between species richness and productivity. We use a classic phytosociological data-set from Rondane, central south Norway, to evaluate the relative importance of these factors in an alpine area. The effect of species pool is assessed using plot scores from a Correspondence Analysis (CA) of the data. Generalised Additive Models (GAM) are used to relate vascular plant species richness to cover of vascular plants, CA plot scores, and plot area in different combinations. Species richness of vascular plants is unimodally related to total vascular plant cover. Plot scores of the first three CA axes (representing the effect of species pool) have a complex relationship with species richness, but explain a large fraction of the total deviance in richness. A humped relationship between richness and cover remains after accounting for CA plot scores in the model, i.e. the relationship is independent of species pool. The results suggest that the relationship between richness and cover changes from one vegetation type to another, as evaluated statistically through the importance of the interaction between cover and CA scores in explaining variation in richness. Plot area also influences the relationship. A unimodal relationship is only evident when small plot sizes are used, whereas a monotonically increasing relationship is found at large plot sizes. Plot area has the strongest effect on the unimodal relationship between richness and cover, whereas vegetation type has only a minor effect on this relationship. This revised version was published online in June 2006 with corrections to the Cover Date.     

Resource availability and plant diversity explain patterns of invasion of an exotic grass

Aims In this study, we examine two common invasion biology hypotheses—biotic resistance and fluctuating resource availability—to explain the patterns of invasion of an invasive grass, Microstegium vimineum.Methods We used 13-year-old deer exclosures in Great Smoky Mountains National Park, USA, to examine how chronic disturbance by deer browsing affects available resources, plant diversity, and invasion in an understory plant community. Using two replicate 1 m 2 plots in each deer browsed and unbrowsed area, we recorded each plant species present, the abundance per species, and the fractional percent cover of vegetation by the cover classes: herbaceous, woody, and graminoid. For each sample plot, we also estimated overstory canopy cover, soil moisture, total soil carbon and nitrogen, and soil pH as a measure of abiotic differences between plots.Important findings We found that plant community composition between chronically browsed and unbrowsed plots differed markedly. Plant diversity was 40% lower in browsed than in unbrowsed plots. At our sites, diversity explained 48% and woody plant cover 35% of the variation in M. vimineum abundance. In addition, we found 3.3 times less M. vimineum in the unbrowsed plots due to higher woody plant cover and plant diversity than in the browsed plots. A parsimonious explanation of these results indicate that disturbances such as herbivory may elicit multiple conditions, namely releasing available resources such as open space, light, and decreasing plant diversity, which may facilitate the proliferation of an invasive species. Finally, by testing two different hypotheses, this study addresses more recent calls to incorporate multiple hypotheses into research attempting to explain plant invasion.     

Does the energy equivalence rule apply to intertidal macrobenthic communities?

Energy equivalence assumes equal contribution of large and small species to production and energy flow in communities. As in a double logarithmic plot, physiological rates decline with body weight by –0.25, log biomass should increase by 0.25 and log abundance decline by –0.75 with log species weight, when this concept is valid. This was tested with annual data sets of the macrobenthos of 4 intertidal sites in the German Wadden Sea (Königshafen) and 3 sites in a south Portuguese lagoon (Ria Formosa). Only abundance data from two of these sites displayed significantly negative slopes with mean body size of the species. Biomass and secondary production data were significantly positively correlated with mean body size for all Ria Formosa sites and also for the biomass of a mussel bed in Königshafen. However, high variation in body size of the individuals of a species limits interpretation of these plots.It is preferable to test this concept by body weight classes regardless of its species composition. At Königshafen, biomass and production displayed two distinct peaks. One peak at small body size was caused by browsing species. The other peak at larger body size was caused by animals which potentially extract their food from the water column. This bimodality was only vaguely reflected at one station in the Ria Formosa, possibly because of a dominance of detritus feeding species. In a normalized form (log biomass or production / width of size classvs. log size class), these spectra imply a dominance of small individuals in biomass and production at all sites (except for a mussel bank at Königshafen). This is interpreted as a consequence of permanent disturbances.     

Canopy cover and tree regeneration in old-growth cove forests of the Appalachian Mountains

Relationships between canopy cover and tree regeneration were determined for various species in cove forests of the Great Smoky Mountains. Old-growth stands were sampled with six plots covering a total area of 4.8 ha. Each plot was subdivided into contiguous 10×10 m quadrats. Canopy cover overlying each of the 480 quadrats was characterized with three different indices based on visual estimates of cover. Influences of: (1) overlying cover, (2) proximate openings, and (3) total area of proximate openings on quadrat regeneration densities were determined. Most species reproducing by seed and some species reproducing by vegetative means had higher densities in quadrats with openings, but only the intolerants were highly dependent on gaps. Tsuga canadensis, a very shade-tolerant species, was one of the few species with abundant regeneration beneath dense canopy cover. In general, understory areas near gaps had somewhat higher regeneration densities than other areas with overlying cover. Several shade-tolerant species showed a positive regeneration density response to canopy openings and an ability to regenerate in gaps 0.01–0.03 ha in area. These openings were too small for intolerant species. Many species exhibited a positive response to total size of the proximate opening(s). A sharp increase in regeneration density with area of the opening(s) was evident at approximately 0.04 ha for the shade-intolerant species.     

Population structure and environmental relationships of the tropical tree Nectandra rudis (Lauraceae), a rare species in western Mexico

Population structure and environmental relationships of the tropical tree Nectandra rudis (Lauraceae), a rare species in western Mexico. The tree N. rudis is a rare species from western Mexico of which community and population features are unknown. We studied a population in an altitudinal gradient, from 550-1,850 m above sea level in the Sierra de Manantlan, Jalisco, Mexico. We established four 60x48 m sample sites at vertical distances of 100 m along this altitudinal gradient. Within each plot, ten 100 m2 circular sub-sampling units were randomly located. At each unit, we recorded diameter at breast height (dbh) and tree height for all woody vegetation > or =2.5 cm dbh. Basal area, tree density, frequency, species richness and importance values per species and plot. We estimated the vertical structure (total tree height) and diameter( as M=5log(10)N) for all N. rudis individuals. A direct ordination through Canonical Correspondence Analysis was done, involving amongst other species, edaphic and environmental data matrices. The record of 44 N. rudis individuals, in seven out the 56 plots sampled, represents the most septentrional record for the species and the first in Western Mexico. Its density and basal area represented 4.5 % and 8.7 % respectively of the total estimated for the community. The greatest importance values were observed at 1 650 m above sea level. The population structure of N. rudis is structured into five diameter categories in an inverse "J" shaped distribution. This is a typical behavior observed to occur in the Lauraceae, which produces big seeds of short viability that germinate when there is high soil moisture content. The species tend to form dense seedling banks although only a reduced number of them are able to survive. Species richness varies from 27 to 39 at plot level; the greatest importance values for the plots on which N. rudis was found, corresponds to Urera verrucosa (Liebm.) V.W. Steinm., N. rudis, Ficus sp., Beilschmiedia manantlanensis Cuevas y Cochrane, amongst others. Canonical Correspondence Analysis suggests that environmental variables such as rooted trees, crown cover, litter depth and soluble magnesium are the more significant explanatory variables for the distribution and abundance of N. rudis.     

Nonlogarithmic death rate calculations for Byssochlamys fulva and other microorganisms.

Survivor curves for heat-resistant ascospores of Byssochlamys fulva exposed to lethal heat were nonlogarithmic. At lower heating temperatures, the log survivor curves were characterized by a shoulder plus an accelerating death rate; with increased temperatures, the rate approached logarithmic death. The formula (log No -- log N)a = kt + C was adapted to linearize these data. No and N are the initial and surviving numbers of organisms at the time t. The death rate is given by k, and C is a constant for a set of data. The a value is derived from the least-squares slope of a plot of log (log No -- log N) against log time and is used to linearize the thermal death rate curves. This formula permitted calculations of parameters analogous to those for logarithmic death (D and z). Use of formula is illustrated for selected nonlinear microbial death rate curves from the literature.     

Nonlogarithmic death rate calculations for Byssochlamys fulva and other microorganisms.

Survivor curves for heat-resistant ascospores of Byssochlamys fulva exposed to lethal heat were nonlogarithmic. At lower heating temperatures, the log survivor curves were characterized by a shoulder plus an accelerating death rate; with increased temperatures, the rate approached logarithmic death. The formula (log No -- log N)a = kt + C was adapted to linearize these data. No and N are the initial and surviving numbers of organisms at the time t. The death rate is given by k, and C is a constant for a set of data. The a value is derived from the least-squares slope of a plot of log (log No -- log N) against log time and is used to linearize the thermal death rate curves. This formula permitted calculations of parameters analogous to those for logarithmic death (D and z). Use of formula is illustrated for selected nonlinear microbial death rate curves from the literature.     

Changes in species diversity and canopy cover in steppe vegetation in Inner Mongolia under protection from grazing

This study describes changes in species diversity and canopy cover in relation to variation in livestock grazing in a semi-arid area in Inner Mongolia, China. Canopy cover for each species was recorded 2 and 3 years after cessation of livestock grazing, as well as in an area with continued grazing. Total species richness, alpha diversity, beta diversity and canopy cover were analysed. Sixty species were recorded during the study; 25 of them were annuals. The total number of species was the same, 52, in the grazed and the protected area, but species richness and alpha diversity per plot were lower in the area protected from grazing. The beta diversity showed little difference between the protected area and the grazed control. The total canopy cover was highest in the protected area, but the cover of annuals was higher in the grazed area. In CA ordination, the difference between treatments increased with time of protection. However, in the short period covered by this study it was difficult to separate the effects of protection from grazing and fluctuation in weather conditions, particularly of precipitation.     

Composition and structure of a species rich Amazonian rain forest obtained by two different sample methods

Inventory data for trees ≥ 10 cm DBH from a hectare plot are compared to data obtained by the Point-Centered Quarter Method along a line transect from the same locality in Anangu, Amazonian Ecuador. The one-hectare quadrat plot of 100 × 100 m had 734 individuals, 153 species, 46 families, a total basal area of 22.2 m², and an estimated above ground tree volume of 240.5 m³. The line transect had a calculated density of 728 individuals per hectare, which included 239 species, 51 families, a total basal area of 34.1 m², and an estimated above ground tree volume of 409.6 m³. Of the 20 species with the highest IVI, only four were shared by the two samples. The most important species were Quararibea ochrocalyx on the hectare plot and Iriartea deltoidea on the line transect, constituting 26.6 and 13.3% of the individuals, respectively. The five families with the highest FIV on the hectare plot (Bombacaceae, Arecaceae, Moraceae, Caesalpinaceae, and Lauraceae) and on the line transect (Arecaceae, Moraceae, Meliaceae, Mimosaceae and Caesalpinaceae) constitute 40.4% and 35.4% of the Family Importance Values of the samples, respectively. The Point-Centered Quarter Method used along a line transect reflects maximum diversity and provides average values of density and tree size in the area. The quadrat plot reflects the local structure and composition of the forest within the plot.     

Exotic plant invasions over 40 years of old field successions: community patterns and associations

While exotic plant species often come to dominate disturbed communities, long-term patterns of invasion are poorly known. Here we present data from 40 yr of continuous vegetation sampling, documenting the temporal distribution of exotic plant species in old field succession. The relative cover of exotic species decreased with time since abandonment, with significant declines occurring ≥20 yr post-abandonment. The number of exotic species per plot also declined with time since abandonment while field-scale richness of exotics did not change. This suggests displacement occurring at small spatial scales. Life history types changed from short-lived herbaceous species to long-lived woody species for both native and exotic plant species. However, shrubs and lianas dominated woody cover of exotic plants while trees dominated native woody cover. The species richness of exotic and native species was positively correlated at most times. In abandoned hay fields, however, the proportion of exotic plant cover per plot was inversely related to total species richness. This relationship suggests that it is not the presence, but the abundance of exotic species that may cause a reduction in community diversity. While the development of closed-canopy forest appears to limit most introduced plant species, several shade-adapted exotic species are increasing within the fields. These invasions may cause a reversal of the patterns seen in the first 40 yr of succession and may result in further impacts on community structure.     

Seed size in relation to phylogeny,growth form and longevity in a subalpine meadow on the east of the Tibetan plateau

This study examined seed size distribution, and seed size in relation to phylogeny, growth form and longevity, where seed size is expected to be approximately of a log-normal frequency distribution and correlate dwith phylogeny, growth form and longevity. We made use of a data set of 229 species from alpine meadow on the east of the Tibetan plateau. Species spanned 10⁴ range of seed size, and the frequency of seed mass classes on a logarithmic scale produced an approximately normal distribution, but largely shifted towards smaller-sized seeds compared to those of the temperate zone and the tropics. It was evident that seed size was strongly related to phylogeny, where order, family, genus and species accounted for 2%, 32.9%, 48.9% and 16.2% of total variation in log seed mass. However, both growth form and longevity did not account for variation in log seed mass, which was opposite to the previously reported patterns of trait correlation. The implications of our results are discussed.     

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.     

Scale‐dependent variation in visual estimates of grassland plant cover

Abstract. Plant cover was visually estimated by five observers, independent of each other, in a species‐rich grassland in the Bílé Karpaty Mts., southeastern Czech Republic, in seven plots ranging from 0.001 to 4 m². Variation of total plant cover among the observers was high at small scales: 0.001–0.016 m²; coefficient of variation, CV = 35 to 45%, but much lower at larger scales: 0.06–4 m²; CV = 7 to 15%. Differences between visual estimates of plant cover of individual species made by different observers were affected by plot size, total cover and morphology of particular plants. CV of the cover of individual species ranged from 0 to 225% and decreased with increasing plot size. For abundant plants the CV attained ca. 50%, independent of plot size. In spite of a very high number of sterile plants with similar leaf morphology and colour, the observed variation in cover estimates in the studied grassland was comparable with results reported from other vegetation types. Differences between estimates by individual observers were often larger than usual year to year changes in undisturbed grasslands. Therefore, I suggest that to avoid difficulties in the interpretation of results based on plant cover data obtained from visual estimates, several observers should always work together, adjusting their extreme estimates.     

Restoration of a species‐rich fen‐meadow after abandonment: response of 64 plant species to management

Abstract. Eleven years of abandonment of a species‐rich fen‐meadow under undisturbed environmental conditions resulted in transformation into areas with tall herb‐, sedge‐ and rush‐dominated communities and areas with Alnus thicket. Species cover was measured in permanent plots in both community types and succession was monitored during 14 yr of restoration following reintroduction of management. The annual increase in accumulated species number followed a log‐log‐time linear regression during 10 yr of grazing management. The expected number of years taken before this annual rate was equal to annual extinction, i.e. a stable situation according to species density, was up to six. The response of 64 species to management was evaluated through paired statistical tests of changes in cover and frequency over time. In total, 55 species could each be allocated to one unique response model (monotone or non‐monotone, concave models) independently of the importance value used (cover or frequency) and type of management (grazing following felling or mowing and mowing without grazing). Species which increased in response to grazing had the most persistent seed banks and CR‐strategies, while species decreasing in response to grazing had less persistent seed banks and CS‐strategies. Some of the species which increased due to grazing followed a model with a local maximum in cover and frequency. The results are discussed in relation to management of species with high cover value during restoration succession.     

Effect of Differentiation and Cell Density on Glycosphingolipid Class and Molecular Species Composition of Mouse Neuroblastoma NB2a Cells

The effects of cell density and retinoic acid-induced differentiation on the class and molecular species composition of mouse neuroblastoma NB2a cell glycosphingolipids were examined under conditions where the period of culture was controlled. The total amount of neutral glycosphingolipids per cell decreased both with differentiation and as the cells became confluent. The relative amount of the neutral glycosphingolipid classes was not affected by differentiation, whereas there were small but significant changes in the relative amount of the neutral glycosphingolipid classes as the cells became confluent. The total amount of the gangliosides was unaffected by either differentiation or cell density, but there were significant changes in the ganglioside class composition as a result of both cell density and differentiation, and the effects were additive. The molecular species of all the major neutral glycosphingolipid and ganglioside classes were essentially identical, and were altered only slightly by either differentiation or cell density.