Are Plant Censuses Carried Out on Small Quadrats More Reliable than on Larger Ones?

Plant censuses are known to be significantly affected by observers’ biases. In this study, we checked whether the magnitude of observer effects (defined as the % of total variance) varied with quadrat size: we expected the census repeatability (% of the total variance that is not due to measurement errors) to be higher for small quadrats than for larger ones. Variations according to quadrat size of the repeatability of species richness, Simpson equitability and reciprocal diversity indices, Ellenberg indicator values, plant cover and plant frequency were assessed using 359 censuses of vascular plants. These were carried out independently by four professional botanists during spring 2002 on the same 18 forest plots, each comprising one 400-m² quadrat, four 4-m² and four 2-m² quadrats. Time expenditure was controlled for. General Linear Models using random effects only were applied to the ecological indices to estimate variance components and magnitude of the following effects (if possible): plot, quadrat, observer, plant species and two-way interactions. High repeatability was obtained for species richness and Ellenberg indicator values. Species richness and Ellenberg indicator values were generally more accurate but also more biased in large quadrats. Simpson reciprocal diversity and equitability indices were poorly repeatable (especially equitability) probably because plant cover estimates varied widely among observers, irrespective of quadrat size. Grouping small quadrats usually increased the repeatability of the variable considered (e.g. species richness, Simpson diversity, plant cover) but the number of plant species found on those pooled 16 m² was much lower than if large plots were sampled. We therefore recommend to use large, single quadrats for forest vegetation monitoring.     

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.     

Latitudinal and altitudinal patterns of plant community diversity on mountain summits across the tropical Andes

The high tropical Andes host one of the richest alpine floras of the world, with exceptionally high levels of endemism and turnover rates. Yet, little is known about the patterns and processes that structure altitudinal and latitudinal variation in plant community diversity. Herein we present the first continental‐scale comparative study of plant community diversity on summits of the tropical Andes. Data were obtained from 792 permanent vegetation plots (1 m²) within 50 summits, distributed along a 4200 km transect; summit elevations ranged between 3220 and 5498 m a.s.l. We analyzed the plant community data to assess: 1) differences in species abundance patterns in summits across the region, 2) the role of geographic distance in explaining floristic similarity and 3) the importance of altitudinal and latitudinal environmental gradients in explaining plant community composition and richness. On the basis of species abundance patterns, our summit communities were separated into two major groups: Puna and Páramo. Floristic similarity declined with increasing geographic distance between study‐sites, the correlation being stronger in the more insular Páramo than in the Puna (corresponding to higher species turnover rates within the Páramo). Ordination analysis (CCA) showed that precipitation, maximum temperature and rock cover were the strongest predictors of community similarity across all summits. Generalized linear model (GLM) quasi‐Poisson regression indicated that across all summits species richness increased with maximum air temperature and above‐ground necromass and decreased on summits where scree was the dominant substrate. Our results point to different environmental variables as key factors for explaining vertical and latitudinal species turnover and species richness patterns on high Andean summits, offering a powerful tool to detect contrasting latitudinal and altitudinal effects of climate change across the tropical Andes.     

Vascular plant diversity and climate change in the alpine zone of the Snowy Mountains,Australia

This study examines vascular plant species richness along an altitudinal gradient in alpine Australia. Vascular plant composition and soil temperature records were obtained for five summits (from 1729 m to 2114 m a.s.l.) using sampling protocols from the Global Observation Research Initiative in Alpine Environments program. Species richness was examined against altitude, aspect and climatic variables at different spatial scales (10 × 10 cm quadrats, 1 m² quadrats, clusters of 4 * 1 m² quadrats, for the summit area above a line 5 m altitudinally below the summit (the −5 m isoline), for the extended summit down to the −10 m isoline). About 75 taxa (70 species, 5 graminoid genera) were recorded, 9 of which are endemic to the small alpine area of ∼100 km². There were significant linear relationships between species richness and altitude and climatic variables for the top to −5 isolines on the summits. However, there was no consistent pattern for species richness at other spatial scales, altitude, aspect or climatic variables. The proportion of species for the whole summits with localised distributions (local endemics) increased with altitude. Predicted increasing temperatures and reduced snowcover is likely to result in an increase in species richness as shrubs, herbs and introduced weeds become more common at higher altitude. Because Australian alpine areas occur in narrow altitudinal bands with no nival zone, there are no higher altitudinal refuges available for alpine species. Therefore many of these species are likely to be at risk of extinction from climate change.     

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.     

Short-term signals of climate change along an altitudinal gradient in the South Alps

Short-term changes in plant species number, frequency and composition were studied along an altitudinal gradient crossing four summits from the treeline ecotone to the subnival zone in the South Alps (Dolomites, Italy). Large-scale (summit areas) and small-scale patterns (16 plots of 1 m2/summit) were monitored. After 5 years, a re-visitation of the summit areas revealed a considerable increase of species richness at the upper alpine and subnival zone (10% and 9%, respectively) and relatively modest increases at the lower alpine zone and the treeline ecotone (3% and 1%, respectively). At the small scale, the results were partly different, with species richness decreasing at the lower summits and increasing at the higher summits. The changes can most likely be attributed to climate warming effects and to competitive interactions. The main newcomers at the lower three summits were species from the treeline and the lower altitudinal zones. Only at the highest summit, the newcomers came from the alpine species pool. At the treeline ecotone, the abundance of Pinus cembra, of dwarf shrubs and clonal graminoid species increased. Here, displacements of alpine species may be predicted for the near future. At the higher summits, expansions of the established alpine species and further invasions of species from lower altitudes are forecasted.     

Reproducibility of vegetation cover estimates in south‐central Alaska forests

Abstract. Reproducibility of vegetation measurements is critical for large‐scale or long‐term studies, where numerous observers collect data, but past studies have questioned repro‐ducibility of some techniques. Five methods of evaluating understory composition were appraised for reproducibility among six observers in two forest types in south‐central Alaska: ocular estimates in quadrats, overall community species rank and cover estimates, nested rooted frequency, horizontal‐vertical profiles, and pin drop (systematic points). One forest type was selected to represent structure of coastal communities, another to represent structure of interior Alaska communities. Three general methods of evaluating reproducibility were considered: standard deviations (precision among observers), components of variance (percentage of total variance attributable to observers), and analysis of variance (significance of observer variance). Observer variances were generally similar among techniques and significant in most cases. No technique stood out as being more reproducible than others. Features of techniques other than reproducibility may be more important when selecting a technique. Management decisions based on vegetation cover data should consider the observer errors involved as well as biological significance.     

Evaluating outcomes of restoration ecology projects on limited budgets: assessment of variation in sampling intensity and sampling frequency for four habitat types

While best practices for evaluating restoration ecology projects are emerging rapidly, budget constraints often limit postrestoration monitoring, which emphasizes the need for practical and efficient monitoring strategies. We examined the postrestoration outcome for an ENGO (Nature Conservancy of Canada) project, to assess retroactively how variation in intensity and frequency of sampling would have affected estimates of plant species composition, diversity, and richness over time. The project restored four habitat types (mesic forest, oak woodland, wet meadow, and sand barren) using sculptured seeding of tallgrass prairie and woody species. Species‐level plant cover was monitored annually for 10 years in 168 2 × 2–m quadrats. We performed randomization tests to examine estimates of species diversity and richness as a function of the number of quadrats sampled, and assessed the necessity of annual sampling for describing changes in species composition and successional trajectories. The randomization tests revealed that sampling 10–17 quadrats, depending on habitat type, was sufficient to obtain estimates of species diversity that were at least 95% of values obtained from the whole dataset. Species richness as a function of number of quadrats sampled did not plateau, which suggests that rather than increasing the number of sampling quadrats, richness could be estimated more efficiently using nonquadrat based sampling techniques. Nonmetric multidimensional scaling analysis revealed that plant species composition largely stabilized by 3–5 years postrestoration depending on habitat type. By that time, native, seeded species dominated the restoration, and the benefits of annual sampling for tracking changes in species composition diminished.     

The relationship between psyllid leaf galls and redbay (Persea borbonia) fitness traits in sun and shade

Spatial autocorrelation in vegetation has been discussed extensively, but little is yet known about how standard plant sampling methods perform when confronted with varying levels of patchiness. Simulated species maps with a range of total abundance and spatial autocorrelation (patchiness) were sampled using four methods: strip transect, randomly located quadrats, the non-nested multiscale modified Whittaker plot and the nested multiscale North Carolina Vegetation Survey (NCVS) plot. Cover and frequency estimates varied widely within and between methods, especially in the presence of high patchiness and for species with moderate abundances. Transect sampling showed the highest variability, returning estimates of 19–94% cover for a species with an actual cover of 50%. Transect and random methods were likely to miss rare species entirely unless large numbers of quadrats were sampled. NCVS plots produced the most accurate cover estimates because they sampled the largest area. Total species richness calculated using semilog species-area curves was overestimated by transect and random sampling. Both multiscale methods, the modified Whittaker and the NCVS plots, overestimated species richness when patchiness was low, and underestimated it when patchiness was high. There was no clear distinction between the nested NCVS or the non-nested modified Whittaker plot for any of the measures assessed. For all sampling methods, cover and especially frequency estimates were highly variable, and depended on both the level of autocorrelation and the sampling method used. The spatial structure of the vegetation must be considered when choosing field sampling protocols or comparing results between studies that used different methods.     

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.     

An examination of observer bias as a source of error in surveys of seagrass shoots

Abstract Mensurative experiments investigated the effects of different observers on estimates of the density of shoots of two species of seagrass: Posidonia australis Hook and Zostera capricorni Aschers. Balanced programmes of sampling were used to examine variation in counts of seagrass shoots attributable to different observers, sizes of quadrats, depths and locations within large beds of each species of seagrass. A separate experiment examined differences between novice observers and a more experienced observer, when an ‘optimal’ size of sampling unit was used. Estimated densities of Zostera shoots varied inconsistently among observers, quadrats, depths and locations. Differences between observers were not affected by the size of quadrat used to count Posidonia shoots, but varied between locations in the seagrass bed. Experience had only a minor impact on biases. Only two of 12 novices produced counts that were different from the experienced observer. These results emphasize the importance of considering both accuracy and precision in the design of field studies of seagrasses.     

Monitoring vegetation change caused by trampling: a study in the Cairngorms,Scotland

Summary

A study was made in the Cairngorms, Scotland to make recommendations for a monitoring scheme capable of detecting changes in the vegetation caused by recreational pressure following the development of a funicular railway. Four methods were used in field trials to assess percentage cover of plant species and gravel, rock and bare ground, where appropriate, in two vegetation types (open and closed). The methods used were visual estimates in 50 × 40 cm quadrats (Q), the mean of visual estimates in twenty 10 × 10 cm sub-quadrats of the 50 × 40 cm quadrats (Q₂₀), a modified point intercept method (RL) and photography. Variances between observers and between-quadrats were estimated for the different methods. The sampling design for detecting change was based on a model of variance, constructed from field trial data.

Between-observer and between-quadrat variances were related to mean percentage cover and approximated to a binomial distribution. The between-quadrat variance was larger than observer variance. The Q₂₀ method achieved appreciably better precision than the other methods. Analysis of half of the 10 × 10 cmsub-quadrats (1/2Q₂₀) selected in a checker board design achieved a relative efficiency of 78% compared with the Q₂₀. This result suggests that comparable precision to the Q₂₀ method could be achieved by choosing about 14 sub-quadrats in a larger quadrat, thus saving some time. Variation between quadrats also suggested that the Q₂₀ method was the one of choice for maximising precision. The precision of the photographic method was based on fewer data points, so is less accurate than other estimates.

Minimum sample sizes were estimated for detecting a 10% relative change of a species in open vegetation with 30% cover (i.e. a change from 30% to <27 or to >33% cover). With a 10 % Type II error rate and 5 % Type I error rate the minimum sample sizes were 47 quadrats for Q, 18 for Q ₂₀, 43 for RL, and 23 for the means of ten 10 × 10 cm sub-quadrats in open vegetation.

The most time-efficient field recording appeared to be the use of Q despite the required sample size being 2.6 times higher than that of Q₂₀. The far lower time requirement per quadrat, however, compensated for the higher numbers. The number of quadrats would depend on the specified change in percentage cover and on the statistical significance level used. For example, to detect a 10% absolute change in cover (i.e. from 30% to either <20 % or >40 % cover) at 95 % probability the net effective recording time is estimated at 5 h per vegetation type while to detect a 5 % change at 99 % probability would require c. 25 h. Larger samples may be required for other species or for species with a low initial cover.     

Effects of sampling time,species richness and observer on the exhaustiveness of plant censuses

Question: How may sampling time affect exhaustiveness of vegetation censuses in interaction with observer effect and quadrat species richness? Location: French lowland forests. Methods: Two data sets comprised of 75 timed, one‐hour censuses of vascular plants carried out by five observers on 24 400‐m² forest quadrats were analysed using mixed‐effect models. Results: The level of exhaustiveness increased in a semi‐logarithmic way with sampling time and decreased with quadrat species richness. After one hour, 20 to 30% of the species remained undetected by single observers. This proportion varied among observers and the discrepancy increased with increasing sampling time. Fixing the sampling time may make richness estimates vary less between observers but the time limit should be at least 30 min to reduce the bias in exhaustiveness between rich and poor quadrats. Conclusions We advocate the use of sampling methods based on spatially or temporally‐replicated censuses and statistical analyses that correct for the lack of census exhaustiveness in vegetation studies.     

Species turnover and diversity during early stages of vegetation recovery on the volcano Usu,northern Japan

To study the rate of revegetation during succession on the volcano Usu, northern Japan, vegetation structure and species composition were monitored from 1984 to 1988 in permanent quadrats near the summit of the volcano, which had been almost completely deforested by 1 to 3 m thick volcanic deposits in 1977–78. Analyses of vegetation structure included species richness, species diversity (IT), evenness (J'); year-to-year changes were quantified using Community Coefficient (CC), and Percentage Similarity (PS). While species turnover rates fluctuated and vegetation cover gradually increased, species richness, diversity and evenness did not fluctuate much. The diversity parameters showed slightly different levels for three habitat types distinguished. CC values comparing subsequent years with the starting year 1984 suggest only minor changes in qualitative species composition; PS values decreased more rapidly, indicating larger changes in species cover. A few well-rooted perennial plants were predominant, therefore, PS decline resulted from dominance-controlled community structure. PS-value detected not only habitat difference, but also the rate of the volcanic succession, suggesting that PS is most effective to evaluate successional pace.     

Succession and fluctuation in a wet dune slack in relation to hydrological changes

Vegetation changes in a wet dune slack complex have been studied over a period of 23 yr. The vegetation was recorded in 40 permanent plots: all plots in 1964, 1977, 1983 and 1987 and some also in five other years. There were large fluctuations in annual precipitation during this period, including some extremely dry (1976) and wet (1985) years. Many species of wet calcareous and pioneer habitats declined in cover between 1964 and 1977, when there were prolonged dry summer periods, though few species were lost from the sampled quadrats. Practically all calciphilous and pioneer species disappeared between 1977 and 1983, when there were relatively many wet summers. It is suggested that extremely dry conditions temporarily obscure the impacts of hydrological changes in local hydrological systems, by retarding the vegetation succession. After a period of excessive precipitation the (acidifying) effects of ground-water withdrawal rapidly became evident in the species composition of the dune slack.     

Effects of sampling teams and estimation methods on the assessment of plant cover

Abstract. We evaluated variability in cover estimation data obtained by (1) two sampling teams who double sampled plots and (2) one team that used two methods (line intercepts and visual estimation of cover classes) to characterize vegetation of herbaceous wetlands. Species richness and cover estimates were similar among teams and among methods, but one sampling team scored cover higher than the other. The line intercept technique yielded higher cover estimates but lower species richness estimates than the cover class method. Cluster analyses of plots revealed that 36% and 11% of plots sampled consecutively by two teams or using two methods, respectively, were similar enough in species composition and abundance to be paired together in the resulting clustering tree. Simplifying cover estimate data to presence/absence increased the similarity among both teams and methods at the plot scale. Teams were very similar in their overall characterization of sites when cover estimation data were used, as assessed by cluster analysis, but methods agreed best on their overall characterization of sites when only presence/absence data were considered. Differences in abundance estimates as well as pseudoturnover contribute to variability. For double sampled plots, pseudoturnover was 19.1%, but 57.7% of pseudo‐turnover cases involved taxa with ≤ 0.5% cover while only 3.4% involved taxa with > 8% cover. We suggest that vegetation scientists incorporate quality control, calibrate observers and publish their results.     

In the eye of the beholder: bias and stochastic variation in cover estimates

Cover estimates by eye is a prevailing method to assess abundance. We examined cover estimates with regard to bias and random variation. Ten observers working with a national forest vegetation survey estimated sixteen 100 m²-plots, placed in two different vegetation types. These had similar species composition but were clearly distinguishable in the field. In species-wise analyses, observer bias varied greatly, with Dicranum spp., Vaccinium vitis-idaea and Vaccinium myrtillus having the largest bias. Experience had a surprisingly small impact on variation. Power analysis revealed only small differences between observers in the ability to distinguish the two vegetation types, and little value in averaging the assessments from two, three or four observers. Cover estimates did better than presence/absence data in separating the two vegetation types and multivariate analyses were more powerful than univariate ones.     

A six year study of a secondary succession after deforestation in North Hungary

Succession in a herbaceous vegetation developed on a site formerly occupied by aQuercetum petraeae-cerris forest is described and discussed. The deforestation took place in 1979. In the clearing 15 sample plots of 1 m² were established in three strata. Every year the floristic composition and the cover of each species was determined in permanent quadrats. For calculation of diversity values based on relative cover the formula given byHill (1973) was used.     

Habitat preferences of sub-montane spiders in northem England

Thirteen study sites were selected to cover the dominant vegetation types on the plateaux of three summits in the North Pennines, England The spider community of these sites were sampled by pitfall trapping between April and October 1991 Several environmental variables were collected for each site, including local topography, soil characteristics and vegetation structure Multivariate methods were used to classify the sites based on their spider species composition Canonical correspondence analysis was used to assess the influence of the external factors on the distribution of the species The results suggest that vegetation density is the major factor influencing spider distribution on the summit plateaux with slope and soil depth also contributing to the variation Three spider associations for the plateaux are proposed based on their common species composition Two short Festuca grassland assemblages and a Nardus/ Ertophorum spp assemblage associated with higher vegetation density The habitat preferences of some species common to the plateaux are discussed in light of the results, and compatisons are made with studies from other upland areas