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.     

Reproducibility of species lists,visual cover estimates and frequency methods for recording high‐mountain vegetation

Question: When multiple observers record the same spatial units of alpine vegetation, how much variation is there in the records and what are the consequences of this variation for monitoring schemes to detect changes? Location: One test summit in Switzerland (Alps) and one test summit in Scotland (Cairngorm Mountains). Method: Eight observers used the GLORIA protocols for species composition and visual cover estimates in percentages on large summit sections (>100 m²) and species composition and frequency in nested quadrats (1 m²). Results: The multiple records from the same spatial unit for species composition and species cover showed considerable variation in the two countries. Estimates of pseudo‐turnover of composition and coefficients of variation of cover estimates for vascular plant species in 1 m × 1‐m quadrats showed less variation than in previously published reports, whereas our results in larger sections were broadly in line with previous reports. In Scotland, estimates for bryophytes and lichens were more variable than for vascular plants. Conclusions: Statistical power calculations indicated that unless large numbers of plots were used, changes in cover or frequency were only likely to be detected for abundant species (exceeding 10% cover) or if relative changes were large (50% or more). Lower variation could be reached with the point method and with larger numbers of small plots. However, as summits often strongly differ from each other, supplementary summits cannot be considered as a way of increasing statistical power without introducing a supplementary component of variance into the analysis and hence into the power calculations.     

Observer variation in field assessments of vegetation condition: Implications for biodiversity conservation

Summary   Uncertainty in assessments of vegetation condition that are used to inform land management and planning decisions for biodiversity conservation in Australia may lead to unexpected outcomes, including loss of biodiversity. This study investigates observer error in field estimates of vegetation attributes, one component of uncertainty in assessments of vegetation condition. Ten observers conducted vegetation condition assessments using two assessment protocols (BioMetric and Habitat Hectares) on 20 sites in a grassy woodland community. Observers' estimates varied substantially across multiple scoring categories for all vegetation attributes on almost all sites. Across all sites, the average coefficient of variation in total vegetation condition scores was 15–18% for both protocols, with a maximum of 60%. The primary cause of variation in total vegetation condition scores was random error in raw estimates of vegetation attributes, although sensitivity of some highly weighted attributes to error exacerbated variation in some cases. Observers generally agreed on the total scores and ranks of highly degraded (pasture) sites, but were less consistent on other sites. Rank correlations between pairs of observers were stronger for Habitat Hectares, suggesting BioMetric may be slightly more sensitive to observer error. It is recommended that: (i) research is undertaken into methods for reducing observer error; (ii) review is made of the sensitivity of index scoring structures to observer error; (iii) field observers estimate uncertainty around point estimates of vegetation condition; and, (iv) decision-makers explicitly incorporate uncertainty into the decision-making processes and aim for outcomes that are robust to this uncertainty.     

Observer error in vegetation surveys: a review

Aims Vegetation sampling employing observers is prone to both inter-observer and intra-observer error. Three types of errors are common: (i) overlooking error (i.e. not observing species actually present), (ii) misidentification error (i.e. not correctly identifying species) and (iii) estimation error (i.e. not accurately estimating abundance). I conducted a literature review of 59 articles that provided quantitative estimates or statistical inferences regarding observer error in vegetation studies.Important findings Almost all studies (92%) that tested for a statistically significant effect of observer error found at least one significant comparison. In surveys of species composition, mean pseudoturnover (the percentage of species overlooked by one observer but not another) was 10–30%. Species misidentification rates were on the order of 5–10%. The mean coefficient of variation (CV) among observers in surveys of vegetation cover was often several hundred % for species with low cover, although CVs of 25–50% were more representative of species with mean covers of>50%. A variety of metrics and indices (including commonly used diversity indices) and multivariate data analysis techniques (including ordinations and classifications) were found to be sensitive to observer error. Sources of error commonly include both characteristics of the vegetation (e.g. small size of populations, rarity, morphology, phenology) and attributes of the observers (e.g. mental fatigue, personal biases, differences in experience, physical stress). The use of multiple observers, additional training including active feedback approaches, and continual evaluation and calibration among observers are recommended as strategies to reduce observer error in vegetation surveys.     

Structure and dynamics in seasonal dry evergreen forest in northeastern Thailand

Abstract. Studies of tropical forest dynamics have often been based on one large-scale permanent plot, representative of a given forest type. Broad classifications of tropical forest types are expected to include a wide range of stand structures, dynamics patterns and species compositions – a range which cannot be represented in a single plot. To demonstrate this problem two 1-ha permanent plots, dominated by Hopea ferrea and Shorea henryana (both Dipterocarpaceae), respectively, were established in 1987 in seasonal dry evergreen forest at the Sakaerat Environmental Research Station in northeastern Thailand. In 1997 the plots were remeasured as to patterns of recruitment, mortality and growth. The Hopea plot was relatively static with low mortality, recruitment and growth. The Shorea plot was very dynamic with high rates of growth, mortality and recruitment. If the current trends continue, the plots are likely to further diverge. Even if the study of a large forest plot provides a good insight into tropical forest dynamics, it is necessary to consider the entire local pattern of variation.     

The effect of fluctuations in tidal inundation frequency on a salt-marsh vegetation

Over a period of 15 years recordings were made of the species cover in permanent plots on the salt marsh of one of the West Frisian Islands, Schiermonnikoog (The Netherlands). Correlations between annual changes in the cover of the major species, and fluctuations in the monthly frequency of inundation by seawater were studied. First, a spectral analysis was carried out on the inundation frequency data to look for predictable patterns. Subsequently, fluctuations were defined as deviations from these predictable patterns. In a repeated multiple regression model, the effects of the season in which the fluctuations occurred, and the elevational position of the plots on the salt marsh were studied as factors influencing the correlation patterns. The behaviour of various species is discussed in relation to their seed bank characteristics and their salt tolerance.     

Fine‐scale five‐year vegetation change in boreal bog vegetation

Abstract. Within an ombrogenous part of N. Kisselbergmosen, Rødenes, SE Norway, fine‐scale changes in species abundance, successional trends relative to the main gradients (as represented by DCA axes), and co‐ordinated change within pairs of the bottom layer species are studied. Data sets were sampled twice with a five‐year interval, and included species abundance and cover of mud bottom, naked peat and litter in 436 sample plots (16 cm× 16 cm), and species abundance in 6976 subplots (4 cm× 4 cm). Depth from the surface of subplots to the water table was estimated in 1991. Most summers and growing seasons were somewhat drier than normal in the 5‐yr period. The area covered by mud‐bottom, naked peat and litter increased significantly, as did the frequencies of the dwarf shrubs Calluna vulgaris and Andromeda polifolia in hummocks and upper lawn. Sample plots were significantly displaced downward the peat productivity gradient (DCA 2), reflecting the reduced cover of many bottom layer species, including all Sphagnum spp. Significant coordinated changes in cover of bottom layer species are described. The changes observed in hummocks support the existence of a local regeneration cycle, as suggested by other researchers. Some of the vegetation changes seem parallel to those reported from areas with a higher nitrogen deposition, but it is not likely that nitrogen deposition alone is the major cause of the observed changes. Between‐year variation in population size and climatic fluctuations may as well explain the observed changes.     

Ordination of vegetation change in Guadalupe Mountains,New Mexico,USA

Vegetation change over a nine-year period was studied in Guadalupe Mountains, New Mexico. Permanent transects in desert shrub vegetation were sampled in 1972 and 1980. Emphasis was given to shrubs because of their importance to big game diets. Univariate paired t-tests and reciprocal averaging ordination were used to detect and display coordinated changes in species composition over time. Despite apparently less browsing pressure in desert shrub vegetation in 1980 there were few significant changes in species composition. In addition, preferred forage species showed reduced reproduction while species of intermediate and poor forage value dis-played increased reproduction during this time. These data do not support traditional rangeland succession theory which states that enhanced reproduction of preferred species should follow grazing or browsing pressure reduction.     

Soil seed bank and species turnover in a limestone grassland

Abstract. The vegetation and soil seed bank in a limestone grassland in southern Sweden were studied in permanent 1-m² subplots which had been either grazed or not grazed for 17 yr. Of the 92 species recorded, 18 were present only in the seed bank, 28 were more frequent and 24 were less frequent in the seed bank than in the vegetation and 22 were not detected in the seed bank. Among the species present in the seed bank, therophytes were over-represented. Species turnover in the vegetation was estimated from presence/absence data collected in 1980, 1986 and 1990. Turnover was high, but there were no differences between grazed and ungrazed subplots. The turnover for individual species was also high in many cases. There was no clear relationship between the turnover of a species and its presence in the persistent seed bank.     

Short-term and long-term effects of mowing on the vegetation of two calcareous fens

Abstract. Short-term field experiments are often used to predict and evaluate long-term management effects. Based on a mowing experiment in two calcareous fens near Lake Neuchâtel, Switzerland, we investigated whether short-term treatment effects (i.e. during the first four years) were confirmed by long-term results (13 - 14 yr). Plots were mown in summer or in winter or left unmown. The main long-term trends in overall species composition (based on percentage cover estimates) were already observable in the first four years: mown and unmown plots diverged, whereas summer-cut and winter-cut plots remained similar. At the individual species level, however, short-term and long-term treatment effects differed considerably: many species whose abundance seemed affected by treatments during the first four years showed no response in the long term, and vice versa. These discrepancies were similar when based on cover estimates or on counts of shoots. Species responses did actually depend on the time scale considered. Short-term and long-term treatment effects on species richness were similar (i.e. a decrease in unmown plots), although only long-term effects were significant. Treatment effects on the above-ground biomass varied considerably, and short-term trends (lower biomass in unmown plots) differed from long-term trends (higher biomass in unmown plots). Our sites showed little overall change in species composition during the period investigated, and treatment effects were low compared with other similar experiments. If study sites are less stable or treatment effects more drastic, a short-term evaluation is expected to be even less reliable. Knowledge on species dynamics at a site may help to choose the adequate spatial and temporal scale of investigation, and thus increase the efficiency of management experiments.     

Implications from the Buell‐Small Succession Study for vegetation restoration

Abstract. Succession is relevant to restoration because managers have to prevent, enhance or replace natural vegetation dynamics. Features of a permanent plot study of post‐agricultural succession in central New Jersey, USA, illustrate important implications of vegetation dynamics for restoration. In the past, such implications had to be drawn from chronosequences and coarse resolution studies, neither of which exposes the local contingencies relevant to site specific restoration. However, the fine scale and continuous nature of the current study reveal that succession is highly contingent on historical and local spatial heterogeneity. For example, the absence of one generally expected dominant stage, the demise of shrubs without replacement by later successional tree species, and the long and multimodal persistence of individual species suggest that neither the relay floristic or initial floristic models of succession is adequate to guide restoration. At the local scale, volleys of species appear through the succession, and reflect spatial contingencies such as neighboring vegetation and edge relationships, and patchy behavior of different functional groups. The role of introduced species and of plant consumers are additional sources of local contingency. These local and time‐specific behaviors in the vegetation are the patterns that restorationists must understand either to choose appropriate reference states, to prevent unexpected local dynamics, or to design interventions that are appropriate to the specific site of interests.     

Fifty years vegetation development of a xerothermic calcareous grassland in Central Europe after heavy disturbance

Fifty years ago on the south-facing slope of a limestone hill 4 km NE of Göttingen (Germany) a small plot of 4 m² size was chosen in a xerothermic calcareous grassland (Mesobromion alliance) for investigation. The formerly grazed slope was abandoned a few years prior to the study, but in 1987 intermittent grazing started again. The mean annual temperature in Göttingen (1953–2003) was 8.8 °C on the average; the annual mean increased by 1.1 °C during the study period. Mean annual precipitation amounted to 642 mm (393–1093 mm). The mean monthly precipitation during summer (April–August) varied in the range between 29 and 101 mm. In the years 1953–2003 the presence of species was recorded annually in 400 subplots (100 cm² each). In part of this time also the amount of bare soil within the plot was recorded, and standing crop and the nitrogen contents in leaves of Bromus erectus were measured in the immediate surrounding of the plot.In fall 1953 the vegetation was removed completely in half of the plot, in the other half the two dominant grasses Bromus erectus and Brachypodium pinnatum were left. The differences between the two halves vanished already after 2 years (starting phase 1953–1955). Thus, only results concerning the plot as a whole are reported here. During the investigation three periods could be distinguished. After the starting phase in period I (1955–1986) total presence, presence of the most important species, species richness, number of endangered species, diversity and evenness increased. In this way the grassland regenerated after the heavy disturbance, and a dense, species-rich vegetation formed. Only a few species showed maximum values already within the first 10 years and faded out later.Period II (1987–1997) spanned 10 years of grazing. During this time total presence, presence of many important species, species richness, number of endangered species, diversity and evenness decreased, whereas areas with bare soil increased. Only two species showed maximum values during this phase, and just one new species invaded (Lolium perenne). During this period a more open type of grassland was formed. Period III (1998–2003) spanned 5 years without grazing and only one grazing year (2002). Here, a partial regeneration of a more closed grassland took place with traits similar to period I.Several phenomena occurred continuously through all three periods. Bromus erectus always was the dominating species. Nevertheless, proportions of graminoids (in % total presence) were lowered, and the proportion of herbs increased. The ratio Bromus erectus/Brachypodium pinnatum was determined mainly by Brachypodium. Seedlings of woody plants were always present, but in most cases survived only one summer. Shrubs did not settle inside the plot, although small bushes developed in the immediate surrounding. In summary the 50-year process can be described as a slow recovery of a calcareous grassland after very heavy disturbance, modified by intermittent grazing, without formation of shrub vegetation.     

Long-term vegetation change in the Juncus squarrosus grassland at Moor House,northern England

Data from the long-term experiment on sheep grazing versus zero sheep grazing (i.e. in an enclosure) on the composition of the Juncus squarrosus grassland at the Moor House NNR are presented. The data have been analysed to assess change in (1) the abundance of individual species, (2) higher plant and bryophyte communities, and (3) vegetation structure. The higher plant data suggested that both the enclosed-and grazed plots were changing in the same direction, but the enclosed plot was moving much faster. The general tendency was towards a blanket bog vegetation dominated by Eriophorum vaginatum and Calluna vulgaris. Juncus squarrosus has declined very quickly in the enclosed plot, but there has also been a slower decline in the grazed one. A significant increase in Calluna vulgaris has occurred only in the enclosed plot. In contrast, the bryophyte data showed completely different successional trajectories in the two treatment plots.The vegetation structure changed markedly after release from grazing, with a decrease in phytomass, but increase in necromass in the lowest height strata. There was no noticeable change in structure over the 18 year period in the grazed plot.Particular problems found in this study were that some species either fluctuated widely in response, or changed in a curvilinear manner.     

Permanent plot research in wet meadows and cutting experiment

Vegetation mowing is usually recommended for preserving wet meadows, in particular for the prevention of land warping and brushwood encroachment. The concept and the methods for the study of the influence of different cutting rhythms on the vegetation of two wet meadows, as well as some short term results are presented. Cutting experiments with permanent plots have been undertaken in vegetation communities such as Orchio-Schoenetum nigricantis and Caricetum elatae, abandoned for many years. The strategy of the study was to have diverse but complementary methodological approaches in order to assess changes in the vegetation and species abundance, especially concerning the botanical composition, the vegetation structure and the species vitality. The methods include counting of plants or part of plants, morphometry of Phragmites communis, biomass measurements, phenological records, structure relevés with the point-intercept technique as well as classical vegetation relevés with coverage estimation.On a short term base we can conclude, on the whole, that the yearly winter cut treatment induces a better vitality (number of individuals, flowering, biomass) of the studied species, while in the control plots the vitality is low, due notably to the important layer of accumulating litter and the lack of light on the ground, which hamper soil warming. The yearly summer cut treatment and the winter cut made every three years show in general intermediate characteristics. Furthermore, the vegetation structure is noticeably different according to whether the vegetation is cut (with hay removal) or not. Concerning the methodology, it is concluded that a diverse approach is a good way to assess changes in the vegetation with a minimum loss of information. If the rather subjective method of vegetation relevés with cover estimation may be suitable for long term survey on permanent plots, more quantitative and objective techniques have to be set if short term or precise changes in the botanical composition, the species vitality and the biomass, as well as structural changes, are sought.This paper was presented at the meeting of the working Group for Succession on Permanent Plots, held in Bern (Switzerland), 4th–7th September 1991.     

Changes in the flora and vegetation of the coastal dunes of Voorne (The Netherlands) in relation to environmental changes

Ecologically the former isle of Voorne is very varied. This is reflected by an exceptionally high number of plant species and vegetation types. Many data regarding flora and vegetation have been collected during more than 25 years. A comparison of the early and the later data show that in this period many changes have taken place in flora and vegetation. The changes are partly the result of a natural development of the vegetation, but to a considerable extent they are due to human activities in the coastal area itself and in its surroundings. Examples of changes produced by natural developments and by human activities are given. The effects on flora and vegetation of the area are illustrated by comparing flora inventories, vegetation maps and permanent plot studies. In addition, the future development in the region which will strongly affect its natural value, is discussed.I would like to thank Michiel Boeken, Joke van der Linden and Elly van Oosterhoud who carried out the second flora inventory and Jeanette van Beuzekom and Inge Boelens who mapped parts of the Voorne dune area. I am particularly indebted to Wiecher Smant who offered assistance in many ways.     

Quantitative studies of phenology and early effects of non-grazing in Cretan phrygana vegetation

Abstract. The phenology of five types of phrygana vegetation situated at various altitudes in SW Kríti (Crete), Greece, was studied over a period of 17 months. Two treatments were compared - traditional pasturing and non-grazing since spring 1993. The numbers of species in blossom and the numbers of flowering units were recorded. For each site, management type and year were displayed in symphenological diagrams. Differences in phenology were mainly related to altered management or the weather regime, or both. The numbers of flowering units were more evident than those of species in blossom for describing the early response of phrygana vegetation to non-grazing. Increases in numbers of flowering units in the ungrazed areas are mainly due to certain abundant therophytes. Unfavourable weather conditions in late winter or early spring may cause a temporary decline in annual species. Rainfall in May at low altitudes promotes resprouting and reflowering of annuals.     

Temporal variation in the woody understory of an old-growth Fagus-Acer forest and implications for overstory recruitment

Abstract. Changes in woody vegetation were examined over eight years, using a 1.05-ha permanent plot in which the location of every shrub and tree > 1m height was mapped. There was little change in the overstory vegetation, as expected for an old-growth forest. Much greater change occurred in the understory, primarily related to a 40 % increase in density. Differences occurred among species in the under-story, as Acer saccharum and Prunus serotina increased and Fraxinus americana and Fagus grandifolia decreased. Canopy gap dynamics are implicated in differences among species in the establishment and growth of individuals in the understory and their recruitment into the overstory. It is concluded that because understory is temporally variable, overstory recruitment from the understory may take different courses at different times in the same forest.     

Observer aging and long‐term avian survey data quality

Long‐term wildlife monitoring involves collecting time series data, often using the same observers over multiple years. Aging‐related changes to these observers may be an important, under‐recognized source of error that can bias management decisions. In this study, we used data from two large, independent bird surveys, the Atlas of the Breeding Birds of Ontario (“OBBA”) and the North American Breeding Bird Survey (“BBS”), to test for age‐related observer effects in long‐term time series of avian presence and abundance. We then considered the effect of such aging phenomena on current population trend estimates. We found significantly fewer detections among older versus younger observers for 13 of 43 OBBA species, and declines in detection as an observer ages for 4 of 6 vocalization groups comprising 59 of 64 BBS species. Consistent with hearing loss influencing this pattern, we also found evidence for increasingly severe detection declines with increasing call frequency among nine high‐pitched bird species (OBBA); however, there were also detection declines at other frequencies, suggesting important additional effects of aging, independent of hearing loss. We lastly found subtle, significant relationships between some species' published population trend estimates and (1) their corresponding vocalization frequency (n ≥ 22 species) and (2) their estimated declines in detectability among older observers (n = 9 high‐frequency, monotone species), suggesting that observer aging can negatively bias long‐term monitoring data for some species in part through hearing loss effects. We recommend that survey designers and modelers account for observer age where possible.