Rarefaction does not eliminate the species richness‐biomass relationship in calcareous blackland prairies

Abstract. Relationships between species richness and biomass are common, but the causes remain controversial. It has been suggested that a hump‐shaped relationship may be an artifact of few individuals in both low and high biomass samples. We used rarefaction to standardize species richness to a constant number of individuals in 99 quadrat samples from relict calcareous prairies. Animal ecologists have long used rarefaction to standardize species richness to a constant number of individuals. The expected species richness of each quadrat was determined for 10, 25, 50, 100 and 150 individual plant ramets using the EcoSim program which resampled the data in each quadrat 1000 times. Because we had few samples with biomass over 300 g/m², our data showed a positive relationship between species richness and biomass. The relationship was largely unaffected by rarefaction. Even at the most extreme case of rarefying to 10 individual ramets, the relationship was clear. Monte Carlo covariance tests did not change the shape of the richness‐biomass relationship and significant correlations between observed species richness and rarified species richness suggest that the number of individuals had negligible effects on richness patterns. Because the data had no strong declining phase, these results may not apply to the declining phase of a humped relationship. The results are discussed with reference to other methods of standardizing species richness. Most importantly, we suggest that expressing species richness per number of individuals is inappropriate and may lead to incorrect conclusions.     

Temporal and spatial patterns in fire occurrence during the establishment of mixed‐oak forests in eastern North America

Question: What was the role of fire during the establishment of the current overstory (ca. 1870–1940) in mixed‐oak forests of eastern North America? Location: Nine sites representing a 240‐km latitudinal gradient on the Allegheny and Cumberland Plateaus of eastern North America. Methods: Basal cross‐sections were collected from 225 trees. Samples were surfaced, and fire scars were dated. Fire history diagrams were constructed and fire return intervals were calculated for each site. Geographic patterns of fire occurrence, and fire‐climate relationships were assessed. Results: Fire was a frequent and widespread occurrence during the formation of mixed‐oak forests, which initiated after large‐scale land clearing in the region ca. 1870. Fire return ranged from 1.7 to 11.1 years during a period of frequent burning from 1875 to 1936. Fires were widespread during this period, sometimes occurring across the study region in the same year. Fires occurred in a variety of climate conditions, including both drought and non‐drought years. Fires were rare from 1936 to the present. Conclusions: A variety of fire regime characteristics were discerned. First, a period of frequent fire lasted approximately 60 years during the establishment of the current oak overstory. Second, fire occurred during a variety of climate conditions, including wet climates and extreme drought. Finally, there was within‐site temporal variability in fire occurrence. These reference conditions could be mimicked in ongoing oak restoration activities, improving the likelihood of restoration success.     

Scale‐dependent determinants of plant species richness in a semi‐arid fragmented agro‐ecosystem

Aims: (1) Understanding how the relationship between species richness and its determinants depends on the interaction between scales at which the response and explanatory variables are measured. (2) Quantifying the relative contributions of local, intermediate and large‐scale determinants of species richness in a fragmented agro‐ecosystem. (3) Testing the hypothesis that the relative contribution of these determinants varies with the grain size at which species richness is measured. Location: A fragmented agro‐ecosystem in the Southern Judea Lowland, Israel, within a desert–Mediterranean transition zone. Methods: Plant species richness was estimated using hierarchical nested sampling in 81 plots, positioned in 38 natural vegetation patches within an agricultural matrix (mainly wheat fields) among three land units along a sharp precipitation gradient. Explanatory variables included position along that gradient, patch area, patch isolation, habitat heterogeneity and overall plant density. We used general linear models and hierarchical partitioning of variance to test and quantify the effect of each explanatory variable on species richness at four grain sizes (0.0625, 1, 25 and 225 m²). Results: Species richness was mainly affected by position along a precipitation gradient and overall plant density, and to a lesser extent by habitat heterogeneity. It was also significantly affected by patch area and patch isolation, but only for small grain sizes. The contribution of each explanatory variable to explained variance in species richness varied with grain size, i.e. scale‐dependent. The influence of geographic position and habitat heterogeneity on species richness increased with grain size, while the influence of plant density decreased with grain size. Main conclusions: Species richness is determined by the combined effect of several scale‐dependent determinants. Ability to detect an effect and effect size of each determinant varies with the scale (grain size) at which it is measured. The combination of a multi‐factorial approach and multi‐scale sampling reveals that conclusions drawn from studies that ignore these dimensions are restricted and potentially misleading.     

A dendro‐ecological study of forest overstorey productivity following the invasion of the non‐indigenous shrub Lonicera maackii

Question: Will a non‐indigenous, invasive, understorey shrub, such as Lonicera maackii (Amur honeysuckle) have an impact on the productivity of overstorey trees in hardwood forests? Location: Trees from 12 invaded and four non‐invaded sites were sampled in hardwood forests of southwestern Ohio, US. Methods: Changes in radial and basal area tree growth in the ten years prior to L. maackii invasion vs. ten years after invasion were examined using dendrochronological techniques. Intervention analysis was used to detect growth changes 25 years prior to and 25 years following invasion, and estimates of load impacts for L. maackii population and biomass were also calculated. Results: We found that the rate of radial and basal area growth of overstorey trees was reduced significantly in eleven out of twelve invaded sites. Non‐invaded sites did not exhibit this consistent pattern of reduced growth. For invaded vs. non‐invaded sites, the mean basal area growth was reduced by 15.8%, and the overall rate of basal area growth was reduced by 53.1%. Intervention analysis revealed that the first significant growth reductions were 6.25 ± 1.24 (mean ± SE) years after invasion with the greatest frequency of negative growth changes occurring 20 years after invasion. In invaded stands, 41% of trees experienced negative growth changes. In terms of invasive load estimates per 1000 L. maackii individuals, radial tree growth was reduced by 0.56 mm.a^?1, and basal area growth was reduced by 0.74 cm².a^?1, Given these findings, significant economic losses could occur in hardwood forests of Ohio. Conclusions: To our knowledge, this is the first study using dendrochronological techniques to investigate the impact of a non‐indigenous, understorey plant on overstorey tree growth. Active management will likely be needed to maintain forest productivity in L. maackii impacted landscapes.     

Three centuries of fire in montane pine‐oak stands on a temperate forest landscape

Question: What was the role of fire in montane pine‐oak (Pinus‐Quercus) stands under changing human land uses on a temperate forest landscape in eastern North America? Location: Mill Mountain in the central Appalachian Mountains, Virginia, US. Methods: A dendroecological reconstruction of fire history was generated for four stands dominated by xerophytic pine and oak species. The fire chronology began under presettlement conditions following aboriginal depopulation. Subsequent land uses included European settlement, iron mining, logging, and US Forest Service acquisition and fire protection. Results: Fires occurred approximately every 5 years until 1930 without any evidence of a temporal trend in fire frequency. Burning ceased after 1930. Area‐wide fires affecting multiple pine stands were common, occurring at intervals of approximately 16 years. Most living pines became established during the late 1800s and early 1900s. Dead pines indicated that an older cohort established ca. 1730. Most hardwoods were established between the 1920s and 1940s. Conclusions: Except for fire protection, changes in land use had no discernible influence on fire frequency. Lightning ignitions and/or large fire extent may have been important for maintaining frequent burning in the 1700s, while fuel recovery may have constrained fire frequency during later periods. The disturbance regime appears to be characterized by frequent surface fires and occasional severe fires, insect outbreaks or other disturbances followed by pine recruitment episodes. Industrial disturbances appear to have had little influence on the pine stands. The greatest impact of industrial society is fire exclusion, which permitted hardwood establishment.     

Past and current trends of change in a dune prairie/oak savanna reconstructed through a multiple-scale history

Abstract. The history of a rapidly changing mosaic of prairie and oak savanna in northern Indiana was reconstructed using several methods emphasizing different time scales ranging from annual to millennial. Vegetation change was monitored for 8 yr using plots and for 30 yr using aerial photographs. A 20th century fire history was reconstructed from the stand structure of multiple-stemmed trees and fire scars. General Land Office Survey data were used to reconstruct the forest of A.D. 1834. Fossil pollen and charcoal records were used to reconstruct the last 4000 yr of vegetation and fire history. Since its deposition along the shore of Lake Michigan about 4000 yr ago, the area has followed a classical primary dune successional sequence, gradually changing from pine forest to prairie/oak savanna between A.D. 264 and 1007. This successional trend, predicted in the models of Henry Cowles, occurred even though the climate cooled and prairies elsewhere in the region retreated. Severe fires in the 19th century reduced most tree species but led to a temporary increase in Populus tremuloides. During the last few decades, the prairie has been invaded by oaks and other woody species, primarily because of fire suppression since A.D. 1972. The rapid and complex changes now occurring are a response to the compounded effects of plant succession, intense burning and logging in the 19th century, recent fire suppression, and possibly increased airborne deposition of nitrates. The compilation of several historical research techniques emphasizing different time scales allows this study of the interactions between multiple disturbance variables.     

Influence of nests of leaf‐cutting ants on plant species diversity in road verges of northern Patagonia

Abstract. It has been suggested that ant nests are the most frequent small‐scale disturbance that affect vegetation patterns. However, their effects on plant diversity are little studied. We document effects of nests of the leaf‐cutting ant Acromyrmex lobicornis on physical‐chemical soil properties and their influence on plant diversity near road verges in a desert steppe in NW Patagonia, Argentina. We analysed nest soils and controls for nitrogen, phosphorus, organic matter, moisture retention capacity and texture. We also analysed the vegetation on 42 nests (30 active and 12 abandoned or without life) and 42 areas without nests. Soil around nests had a greater nutrient content and capacity to retain moisture than control soils, which is mainly due to the presence of organic waste that the ants deposit on the soil surface. We found no association between the occurrence of nests and specific groups of plants, but plant diversity was higher at nest‐sites than at nearby non‐nest sites. This increased diversity – which is also found on abandoned nests – is mainly due to the occurrence of a larger number of native and exotic plant species on nest‐sites that are uncommon elsewhere in the study area. The most abundant plant species showed similar cover values at nest and non‐nest sites. This suggests that changes in diversity are associated to edaphic changes caused by nests rather than by changes in competitive balance caused by dominant species exclusion. We propose that the nests of Acromyrmex lobicornis, through increasing the availability of resources, generate favourable microsites that can function both as ‘refuges’ for less frequent native species, and as‘stepping stones’ for less frequent exotic plant species.     

Species richness in deciduous forests: Effects of species pools and environmental variables

Abstract. The study was conducted in deciduous forests of two Swedish regions, Öland and Uppland. It had two objectives: to (1) test the species pool hypothesis by examining if differences in small‐scale species richness are related to differences in large‐scale species richness and the size of the regional species pool, and (2) to examine the relationship between species richness and productivity and its scale‐dependence. The first data set comprised 36 sites of moderate to high productivity. In each site, we recorded the presence of vascular plant species in nested plots ranging from 0.001 to 1000 m² and measured several environmental variables. Soil pH and Ellenberg site indicator scores for nitrogen were used as estimators of productivity. The second data set included 24 transects (each with 20 1‐m² plots) on Öland in sites with low to high productivity. Species number, soil pH and relative light intensity were determined in each plot. The forest sites on Öland were more species‐rich than the Uppland sites on all spatial scales, although environmental conditions were similar. Small‐scale and large‐scale species richness were positively correlated. The results present evidence in favour of the species pool hypothesis. In the nested‐plots data set, species number was negatively correlated with pH and nitrogen indicator scores, whereas a unimodal relationship between species number and pH was found for the transect data set. These results, as well as previously published data, support the hump‐shaped relationship between species richness and productivity in Swedish deciduous forests. Two explanations for the higher species richness of the sites with moderate productivity are given: first, these sites have a higher environmental heterogeneity and second, they have a larger ‘habitat‐specific’ species pool.     

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.     

Gap disturbances in northern old‐growth forests of British Columbia,Canada

Abstract. We characterized the abundance, size and spatial patterning of canopy gaps, as well as gap‐forming processes and light availability in boreal, sub‐boreal, northern temperate and subalpine old‐growth forests of northwestern British Columbia. The proportion of area in canopy gaps ranged from 32% in northern temperate forests to 73% in subalpine forests. Evenly distributed developmental gaps were dominant but permanent openings created by edaphic components and by shrub communities were also common, particularly in subboreal forests. Abundant gaps, large gap sizes, high numbers of gap makers per gap and frequent gap expansion events suggest that gaps have long tenure in these forests. Snapped stems and standing dead mortality were the most common modes of mortality in all forest types resulting in little forest floor disturbance, creating few germination sites for seedling establishment. We found high mean light levels (16–27% full sun) and little difference between non‐gap and gap light environments. Our results suggest that gap dynamics in these forests differ fundamentally from those in temperate and tropical forest ecosystems.     

A regional‐scale consideration of the effects of species richness on above‐ground biomass in temperate natural grasslands of China

Question: Can species richness be a predictor for above‐ground biomass in natural grasslands at a regional scale? Location: A total of 647 sites across temperate natural grasslands of northern China. Methods: Structural equation modelling (SEM) was used to examine the effect of species richness on above‐ground biomass. Asymptotically distribution‐free estimation was selected for parameter estimation. The SEM process was performed at five sample sizes (n=50, 100, 200, 300 and 647). Spatial structure in the original data was examined by calculating Moran's I. Results: SEM run at n=647 revealed a positive effect of species richness on above‐ground biomass after controlling for the influences of bioclimatic factors and grazing. At the four reduced sample size levels, the positive effect held true for most cases in 400 observations (>92%). Conclusions: Contrary to observations in previous studies in natural grasslands, our data showed a positive effect of species richness on above‐ground biomass. This suggests that, as a short‐term effect of diversity on productivity, niche complementarity among coexisting species tends to be an important process in arid and semi‐arid natural grasslands. We hold that biodiversity conservation is among the fundamental approaches required to maintain productivity of grasslands in arid and semi‐arid areas.     

Distribution and abundance of trees in floodplain forests of the Wisconsin River: Environmental influences at different scales

Questions: 1. How do physiography, flooding regime, landscape pattern, land‐cover history, and local soil conditions influence the presence, community structure and abundance of overstorey trees? 2. Can broad‐scale factors explain variation in the floodplain forest community, or are locally measured soil conditions necessary? Location: Floodplain of the lower 370 km of the Wisconsin River, Wisconsin, USA. Methods: Floodplain forest was sampled in 10 m × 20 m plots [n= 405) during summers of 1999 and 2000 in six 12‐ to 15‐km reaches. Results: Species observed most frequently were Fraxinus pennsylvanica, Acer saccharinum and Ulmus americana. Physiography (e.g. geographic province) and indicators of flooding regime (e.g. relative elevation and distance from main channel) were consistently important in predicting occurrence, community composition, and abundance of trees. Correspondence analysis revealed that flood‐tolerant and intolerant species segregated along the primary axis, and late‐successional species segregated from flood‐tolerant species along the secondary axis. Current landscape configuration only influenced species presence or abundance in forests that developed during recent decades. Land‐cover history was important for tree species presence and for the abundance of late‐successional species. Comparison of statistical models developed with and without soils data suggested that broad‐scale factors such as geographic province generally performed well. Conclusions: Physiography and indicators of flood regime are particularly useful for explaining floodplain forest structure and composition in floodplains with a relatively high proportion of natural cover types.     

Predictability of plant and fungal species richness of old‐growth boreal forest islands

Abstract. The fragmentation and deterioration of old‐growth forest habitat by modern forestry have become a major threat to species diversity in Fennoscandia. In order to develop a conservation strategy for the remaining diversity it is essential to identify the existing diversity and to develop appropriate conservation and monitoring programs. For these purposes indicators of conservation value for administrative prioritization are required. This study examines the predictability of plant and fungal species richness on two spatial scales on 46 isolated old‐growth forest islands (0.17 ‐ 12 ha) in a forest‐wetland mosaic. We explore (1) to what extent area, isolation and stand structure variables can explain the variation in species richness and (2) if richness patterns of individual species groups correlate. Isolation showed no relation to species richness. Area explained 50 ‐ 70% of the variation in total species richness and was positively related to the density of crustose lichens and Red‐list species in island interiors. Stand structure variables explained 28 ‐ 66% of the residual variation in total species richness after controlling for island size, and 15 ‐ 73% of the variation in density of species in island interiors. The highest predictability of species richness was found among substrate‐specific fungi and Red‐list species. Different stand structure variables were found to explain richness in the different species groups, and only among a few species groups species richness correlated. Thus, species richness of one single species group is unlikely to be a good indicator for total biodiversity. The results show that measurements of stand size and stand structure variables may be a strong complementary tool, and sometimes a substitute to extensive species inventories when one aims to estimate and monitor plant and fungal species diversity in old‐growth Picea abies forests.     

Causes of pattern in plant communities where environmental change is rapid and species longevity is short

Questions : To what extent can spatial structure and its causes be determined in a highly disturbed environment? What are the main determinants of pattern and are these species‐specific? How much do spatial patterns change over generations? Location : Wimmera region of southern Australia. Methods : Broad‐leaved weeds were counted in 225 000 contiguous 20‐cm square quadrats. A substantial number of these quadrats were recorded again after two and four years. An hierarchical ‘adaptive analysis’ approach was used to select spatial analytical methods to examine specific aspects of pattern and variation in pattern from year to year. Results : Patterns varied among species and included both dense and sparse patches surrounded by areas of zero density, diffuse gradations of density and clear anisotropy. Patterns in Erodium botrys and Oxalis pes‐caprae persisted over years, whereas patterns in Arctotheca calendula were less pronounced and varied over time. Edaphic factors appeared to have only a minor influence over the spatial distribution of the weed community as a whole. In Oxalis pes‐caprae, whose patches were hypothesized to have been shaped by cultivation, there was no spread in four years, despite further tillage. Outlying plants of O. pes‐caprae failed to establish new patches, even in the year of greatest population increase. Little evidence of localised recruitment events was found. Conclusions : Despite repeated annual disturbances by natural and anthropogenic mechanisms, clear and interpretable spatial structure develops in annual weeds over a range of spatial resolutions. Adaptive analysis is a useful approach to the characterization of such patterns.     

Effects of the expansion by Hippophaë rhamnoides on plant species richness in coastal dunes

Question: Is the expansion of Hippophaë rhamnoides in coastal dunes associated with a decline in plant species richness, and is this decline best described by a hump‐backed relationship between species number and shrub cover? Location: Grey and yellow dunes on the East Frisian islands Spiekeroog and Norderney. Methods: Total plant species richness as well as the number of herbaceous and cryptogam species were determined in 2001 using plots of 16 m² size. We compared shrubland plots with varying cover of Hippophaë with neighbouring dune grassland plots without shrubs as reference sites. Soil samples were collected to determine the values of some important edaphic variables (pH, organic matter, nitrogen). Results: The shrubland plots with Hippophaë had or tended to have lower soil pH and C/N ratios and higher contents of organic matter and nitrogen than the grassland plots. Total species richness was marginally significantly related to the cover of Hippophaë in a hump‐backed manner on both islands. The pattern was more pronounced for mosses and lichens than for herbaceous species. For all species groups on Spiekeroog and for the herbaceous species on Norderney, the hump‐backed relationship was much improved when using the difference in species number between shrubland and grassland plot as a dependent variable. Relationships could be improved by including the soil parameters as co‐variables. Species richness was highest at moderate levels of shrub expansion, while it was much reduced in very dense shrubland. The decrease in species number is caused by the decline in grassland species typical of the open dunes, including some rare taxa. Conclusions: The expansion of Hippophaë rhamnoides is a serious threat to the plant species richness of open coastal dunes, and needs to be counteracted by management measures.