The inselberg flora of Atlantic Central Africa. I. Determinants of species assemblages

Aims To identify the relative contributions of environmental determinism, dispersal limitation and historical factors in the spatial structure of the floristic data of inselbergs at the local and regional scales, and to test if the extent of species spatial aggregation is related to dispersal abilities. Location Rain forest inselbergs of Equatorial Guinea, northern Gabon and southern Cameroon (western central Africa). Methods We use phytosociological relevés and herbarium collections obtained from 27 inselbergs using a stratified sampling scheme considering six plant formations. Data analysis focused on Rubiaceae, Orchidaceae, Melastomataceae, Poaceae, Commelinaceae, Acanthaceae, Begoniaceae and Pteridophytes. Data were investigated using ordination methods (detrended correspondence analysis, DCA; canonical correspondence analysis, CCA), Sørensen's coefficient of similarity and spatial autocorrelation statistics. Comparisons were made at the local and regional scales using ordinations of life‐form spectra and ordinations of species data. Results At the local scale, the forest‐inselberg ecotone is the main gradient structuring the floristic data. At the regional scale, this is still the main gradient in the ordination of life‐form spectra, but other factors become predominant in analyses of species assemblages. CCA identified three environmental variables explaining a significant part of the variation in floristic data. Spatial autocorrelation analyses showed that both the flora and the environmental factors are spatially autocorrelated: the similarity of species compositions within plant formations decreasing approximately linearly with the logarithm of the spatial distance. The extent of species distribution was correlated with their a priori dispersal abilities as assessed by their diaspore types. Main conclusions At a local scale, species composition is best explained by a continuous cline of edaphic conditions along the forest‐inselberg ecotone, generating a wide array of ecological niches. At a regional scale, these ecological niches are occupied by different species depending on the available local species pool. These subregional species pools probably result from varying environmental conditions, dispersal limitation and the history of past vegetation changes due to climatic fluctuations.     

Environmental gradients in a southern Europe estuarine system: Ria de Aveiro,Portugal implications for soft bottom macrofauna colonization

Four seasonal sampling surveys were carried out between December 1985 and September 1986 in Canal de Mira (Ria de Aveiro, Portugal). A total of 40 sampling stations, distributed over 13 transects, was used. Salinity, temperature, dissolved oxygen and pH of the water mass were measured. Sediment temperature, and salinity and pH of interstitial water were determined. Sediment variables also included granulometric composition and organic matter contents. Bottom macrofauna samples were collected at each station. Ordination (PCA and MDS) and classification of the sampling stations were performed, using the physicochemical and the biological data sets separately. Average linkage cluster analysis using the unweighted paired-group method, arithmetic averages, was used for both sets of data. With a salinity range from 35.1‰ to 0.0‰, Canal de Mira behaves like a tidally and seasonally poikilohaline estuary. Water temperature (8.5–24.7°C) decreased along the channel towards its inner part during the cold season; an inverse and more pronounced trend was observed during the hot season. Dissolved oxygen contents was generally high during the day (50% to 240% saturation). Oversaturation was observed throughout the growing season, with peaks in areas with large amounts of rooted vegetation. The pH values, largely correlated with dissolved oxygen, ranged from 6.8 to 8.9. Four types of sediment were present in Canal de Mira, medium and muddy sands being dominant. Two major gradients were identified: (i) a typical longitudinal estuarine gradient, associated with distance from the mouth, representing physicochemical variables such as tidal amplitude, salinity and temperature; this gradient was accompanied by an upstream increase in dominance; the community composition changes were mainly related to salinity; (ii) a lateral gradient, related to current velocity, depth and sediment composition; the subtidal community had a comparatively low species richness and abundance. Groups of stations could be recognized along the environmental gradients. Benthic community changes, however, appeared to be gradual rather than marked by abrupt transitions.     

Modern vegetation–pollen–climate relationships for the Pampa grasslands of Argentina

Aim To analyse the relationships between potential natural vegetation, pollen and climate in order to improve the interpretation of fossil pollen records and provide the background for future quantitative palaeoclimatic reconstructions. Location Pampa grasslands of Argentina, between 33–41° S and 56–67° W. Methods Modern pollen data were obtained from a pollen data base developed by the Grupo de Investigación de Paleoecología y Palinología, Universidad Nacional de Mar del Plata, Argentina (143 surface samples and 17 pollen types). Analysis of pollen and climate data involved multivariate statistics (cluster analysis and principal components analysis), scatter diagrams, Pearson’s correlation and isopoll mapping. Results Vegetation patterns at regional scales (grasslands and xerophytic woodlands) and local scales (edaphic communities) were identified by cluster analysis of pollen surface samples. The main climatic variables that appear to constrain the vegetation distribution and abundance of taxa are mean annual precipitation, annual effective precipitation and summer temperature. Individual pollen types such as Chenopodiaceae, Apiaceae, Cyperaceae, Prosopis, Schinus, Condalia microphylla and other xerophytic taxa are good indicators of moisture regime. Many pollen types are significantly correlated with summer temperature. The modern vegetation–pollen–climate relationships vary in a broadly predictable manner, supporting the contention that fossil pollen assemblages can be related to particular climatic characteristics. Main conclusions An expanded suite of modern analogues facilitated new insights into vegetation–pollen–climate relationships at the regional scale in Pampa grasslands. Relationships between individual pollen types and climate are appraised at a regional scale and new modern analogues are presented. The results provide the basis for improved vegetation and climate reconstruction from fossil records of the study area.     

Numerical Characteristics of Pollen Assemblages of Surface Samples from the West Kunlun Mountains

A total of 31 suface sediment samples were collected from West Kunlun Mountain in south Xinjiang Autonomous Region in northwest China. These samples are from seven types of vegetation: Picea schrenkiana Fisch. et Mey. forest, Sabina Spach. woodland, sub-alpine steppe, alpine meadow, desert vegetion, cushion-vegetation and vegetation adjancent to glaciers. Pollen percentages and pollen concentrations were calculated in all samples. The dominant pollen types in the region are Chenopodiaceae, Artemisia, Picea, Ephedra, Gramineae, Cyperaceae, Rosaceae, Leguminosae, Compositae etc. In order to reveal the relationship between pollen composition and the vegetation type from which the soil sample was collected, principal component analysis and group average cluster analysis were employed on the pollen data. The results revealed that the major vegetation types in this region could be distinguished by pollen composition: a. Samples from desert vegetation were dominated by pollen of Chenopodiaceae (about 60195%). The percentages of all other pollen types were low. b. Picea forest samples were rich in Picea pollen (about 20%) Sabina forest had more Sabina pollen grains than other vegetation types (about 5%, others <1%). Pollen percentages of Artemisia, Chenopodiaceae and Ephedra were comparatively higher (each about 20%) in these samples from the two types of vegetations. C. Pollen percentages of Artemisia, Cyperaceae, Gramineae and Chenopodiaceae were high in both sub-alpine steppe and alpine meadow. But steppe containal more Artemisia and Chenopodiaceae (steppe 33.75% and 32.30%, meadow 15.57% and 19.48% in average), less Cyperaceae and Gramineae (steppe 2.58% and 7.60%, meadow 22.35% and 12.93% in average) than meadow. d. Samples from cushion-vegetation and vegetation adjacent to glaciers were mainly composed of pollen grains transported from other sites. It was not easy to distinguish them from other vegetation types. Principal component analysis and cluster analysis distinguish samples from Picea forest, Sabina woodland, sub-alpine steppe, alpine meadow and desert vegetation. Therefore we think it will be possible to apply the module to reconstruct past vegetation in this region and other similar regions. Regression analysis was also applied to reveal the relationships between pollen and plant percentages of Artemisia, Chenopodiaceae, Cyperaceae and Gramineae. The results indicated that a linear relationship existed between pollen and plant percentages for Artemisia, Chenopodiaceae and Cyperaeeae.     

Modern pollen–vegetation relationships along transects on the Whangapoua Estuary, Great Barrier Island, northern New Zealand

Aims To quantify pollen–vegetation relationships from saline to freshwater in an estuarine gradient from surface samples of the modern pollen rain, to allow more accurate interpretations of the stratigraphic palynological record. Location Whangapoua Estuary, Great Barrier Island, northern New Zealand. Methods Six transects were laid out along a vegetation sequence running from estuarine mud to freshwater swamp. Along these transect lines, 108 plots were sampled for vegetation and surface sediments from wet sand, mud, plant litter or moss (sand and mud sites are inundated by most tides, other sites less frequently). All sediment samples were analysed for pollen. The relationships between plant species frequency and pollen representation were examined at a community scale using twinspan and ordination analyses, and for individual species using fidelity and dispersibility indices, regression and box‐plot analyses. Results The quantitative relationships between source taxon vegetation frequency and its pollen representation varied between species due to differential pollen production and dispersal. twinspan of the surface pollen samples suggests five vegetation types: (A) mangrove (Avicennia marina); (C) Leptocarpus similis salt meadow; (D) Baumea sedges; (E) Leptospermum shrubland; and (F) Typha/Cordyline swamp forest. The (B) Juncus kraussii community is not represented palynologically owing to the destruction of its delicate pollen grains during acetolysis of samples. Detrended correspondence analysis places these communities on an estuarine‐to‐freshwater gradient. However, pollen assemblages at the seaward end of the salinity gradient are less clearly representative of the associated vegetation than those at the landward end, probably because the open vegetation at the former allows the influx of wind‐ and water‐dispersed pollen from surrounding vegetation. Main conclusions The vegetation pattern (zonation) at Whangapoua is reflected in the pollen rain. When the long‐distance and over‐represented pollen types are excluded, five out of six of the broad vegetation communities can be identified by their pollen spectra. Species with high fidelity and low‐to‐moderate dispersibility indices can be used to identify the vegetation types in the sedimentary sequences. The more open vegetation types at the ‘marine end’ of the sequence tend to be ‘overwhelmed’ by regional pollen, but the nature of the sediments and the presence of discriminatory species (e.g. A. marina, Plagianthus divaricatus, Cordyline australis), even in small amounts, will allow correct identification of the local vegetation represented in sedimentary palynological sequences. A box‐plot analysis indicates that the pollen and spore types A. marina (mangroves), Sarcocornia quinqueflora (salt meadow), P. divaricatus (sedges), Gleichenia (shrubland) and C. australis (swamp forest) are highly discriminatory in relation to vegetation type. These discriminatory palynomorphs help with the interpretation of stratigraphic pollen studies. However, salt marsh vegetation communities in the sediments must be interpreted with caution as the marine sediments are easily affected by erosion, bioturbation and tidal inundation effects.     

A survey of modern pollen and vegetation along a south–north transect in Mongolia

Aim This modern pollen‐rain study documents the spatial and quantitative relationships between modern pollen and vegetation in Mongolia, and explores the potential for using this relationship in palaeoclimatic reconstructions. Location East‐central Mongolia. Methods We collected 104 pollen surface samples along a south–north transect across five vegetation zones in Mongolia. Discriminant analysis was used to classify the modern pollen spectra into five pollen assemblages corresponding to the five vegetation zones. Hierarchical cluster analysis was used to divide the main pollen taxa into two major groups and seven subgroups representing the dry and moist vegetation types and the main vegetation communities within them. Results Each vegetation zone along the transect can be characterized by a distinctive modern pollen assemblage as follows: (1) desert zone: Chenopodiaceae–Zygophyllaceae–Nitraria–Poaceae pollen assemblage; (2) desert‐steppe zone: Poaceae–Chenopodiaceae pollen assemblage; (3) steppe zone: Artemisia–Aster‐type–Poaceae–Pinus Haploxylon‐type pollen assemblage; (4) forest‐steppe zone: Pinus Haploxylon‐type–Picea–Artemisia–Betula, montane forb/shrub and pteridophyte pollen assemblage; and (5) mountain taiga zone: Pinus Haploxylon‐type–Picea–Poaceae–Cyperaceae, montane forb/shrub and Pteridophyte pollen assemblage. Main conclusions Based on the ratio between the major pollen taxon groups and subgroups, we propose two pollen–climate indices that represent the precipitation and temperature conditions in the study region. When plotted along our south–north transect, the moisture indices (M) and temperature indices (T) mimic the regional gradients of precipitation and temperature across Mongolia very closely. These pollen–climate indices can be used for palaeoclimatic reconstruction based on fossil pollen data.     

Pollen grain morphology of Nolana L. (Solanaceae: Nolanoideae: Nolaneae) and related genera of southern South American Solanaceae

Nolana comprises 89 species almost entirely restricted to the Atacama and Peruvian deserts. These species are adapted to the arid lomas environment of northern Chile and southern Peru, where they depend on marine fog locally known as garúa or camanchaca for their development. Recent molecular phylogenetic studies provide support for the placement of Nolana in the Solanaceae rather than Nolanaceae. Additionally, a sister relationship with the tribe Lycieae has been suggested. The pollen morphology of 111 species from 18 genera of southern South American Solanaceae was investigated under light and scanning electron microscopes in order to establish the palynological affinities of Nolana within the Solanaceae and to investigate if pollen morphology is congruent with the phylogenetic relationships and former taxonomic classification of the species of Nolana. The following genera were studied: Nolana, Cestrum, Combera, Dunalia, Fabiana, Jaborosa, Latua, Lycium (incl. Grabowskia, Phrodus), Nicandra, Nicotiana, Nierenbergia, Petunia, Physalis, Reyesia, Salpiglossis, Schizanthus, Solanum and Vestia. Two pollen types that differ in exine structure were distinguished within Nolana; these pollen types are congruent with recent phylogenetic studies, thus, we propose to reinstate Nolana sect. Sorema.     

Floristic composition in relation to environmental gradients across KwaZulu‐Natal,South Africa

Conservation planning in the face of global change is still in its infancy. A suggested approach is to incorporate environmental gradients into conservation planning as they reflect the ecological and evolutionary processes generating and maintaining diversity. Our study provides a framework to identify the dominant environmental gradients determining floristic composition and pattern. Nonmetric multidimensional scaling was used on 2155 sampling plots in savanna and grassland habitat located across the province of KwaZulu‐Natal, South Africa (94 697 km²), a floristically rich region having steep environmental gradients, to determine the dominant gradients. Hierarchical cluster analysis was used to group similar plots which were then used in a Classification and Regression Tree analysis to determine the environmental delimiters of the identified vegetation clusters. Temperature‐related variables were the strongest delimiters of floristic composition across the province, in particular mean annual temperature. Frost duration was the primary variable in the Classification and Regression Tree analysis with important implications for savanna/grassland dynamics. Soil properties (base, pH status) and moisture variables accounted for most of the variation for the second and third axes of floristic variation. Given that climatic and edaphic variables were well correlated with floristic composition, it is anticipated that a changing climate will have a marked influence on floristic composition. We predict warmer temperatures may facilitate the spread of frost sensitive savanna species into previously cooler, grassland areas. Species associated with specific soil types will not easily be able to move up the altitudinal gradient to cooler climes because geology is aligned in an approximately north‐south direction compared with increasing altitude from east‐west. Future conservation planning should take cognisance of these gradients which are surrogates for ecological and evolutionary processes promoting persistence.     

Palaeoecological perspectives on pattern and process in plant diversity and distribution adjustments: a comment on recent developments*

Two developments in the understanding of the relationship between sedimentary pollen assemblages, vegetation and plant diversity are discussed. The Prentice model of vegetation–pollen relationships has improved our understanding of how and at what scale sedimentary pollen records vegetation. The modelling framework allows improved palaeoecological study designs that may, potentially, give important new insights into processes of plant migrations in response to climate change. Also, reconstructions of plant communities and landscape openness may be improved. Competing hypotheses for the relationship between vegetation and palynological diversity are discussed and it is concluded that more attention should be focused on evenness aspects of palynological diversity and on hypothesis testing.     

An example of Holocene vegetation stability from Camerons Lagoon,a near treeline site on the Central Plateau,Tasmania

Abstract A discontinuous record of vegetation over the past 7500 years was obtained through pollen analysis of pond sediments from an extensive treeless plain on the relatively dry Central Plateau of Tasmania. The results demonstrate continuity of treeless conditions, which probably persisted from the Pleistocene throughout the Holocene, up to the present day. Some changes to the structure of the grassland were observed, especially over the past 200 years. Analysis of carbonized particles showed that fires had been infrequent over the period examined. It seems that this area, close to the altitudinal tree limit, has remained a natural grassland, and the hypothesis of Jackson (1973), that unreliable summer conditions may be a major factor in maintaining open conditions, is supported. Parts of the Central Plateau may thus preserve plant communities with some of the floristic elements and structure of widespread Bassian grasslands of the Late Pleistocene.     

Species richness–environment relationships within coastal sclerophyll and mesophyll vegetation in Ku‐ring‐gai Chase National Park,New South Wales,Australia

Abstract Patterns in species richness from a wide range of plant communities in Ku‐ring‐gai Chase National Park, New South Wales, Australia, were examined in relation to a number of environmental variables, including soil physical and chemical characteristics. Total species richness and richness of three growth‐form types (trees, shrubs and ground cover) were determined in duplicate 500‐m² quadrats from 50 sites on two geological substrata: Hawkesbury Sandstone and Narrabeen shales and sandstones. Generalized linear models (GLM) were used to determine the amount of variation in species richness that could be significantly explained by the measured environmental variables. Seventy‐three per cent of the variation in total species richness was explained by a combination of soil physical and chemical variables and site attributes. The environmental variables explained 24% of the variation in tree species richness, 67% of the variation in shrub species richness and 62% of the variation in ground cover species richness. These results generally support the hypothesis of an environmental influence on patterns in total species richness and richness of shrubs and ground cover species. However, tree species richness was not adequately predicted by any of the measured environmental variables; the present environment exerts little influence on the richness of this growth‐form type. Historical factors, such as fire or climatic/environmental conditions at time of germination or seedling establishment, may be important in determining patterns in tree species richness at the local scale.     

Fish Communities of the Sacramento River Basin: Implications for Conservation of Native Fishes in the Central Valley, California

The associations of resident fish communities with environmental variables and stream condition were evaluated at representative sites within the Sacramento River Basin, California between 1996 and 1998 using multivariate ordination techniques and by calculating six fish community metrics. In addition, the results of the current study were compared with recent studies in the San Joaquin River drainage to provide a wider perspective of the condition of resident fish communities in the Central Valley of California as a whole. Within the Sacramento drainage, species distributions were correlated with elevational and substrate size gradients; however, the elevation of a sampling site was correlated with a suite of water-quality and habitat variables that are indicative of land use effects on physio-chemical stream parameters. Four fish community metrics – percentage of native fish, percentage of intolerant fish, number of tolerant species, and percentage of fish with external anomalies – were responsive to environmental quality. Comparisons between the current study and recent studies in the San Joaquin River drainage suggested that differences in water-management practices may have significant effects on native species fish community structure. Additionally, the results of the current study suggest that index of biotic integrity-type indices can be developed for the Sacramento River Basin and possibly the entire Central Valley, California. The protection of native fish communities in the Central Valley and other arid environments continues to be a conflict between human needs for water resources and the requirements of aquatic ecosystems; preservation of these ecosystems will require innovative management strategies.     

Modern surface pollen assemblages from the Middle and High Atlas,Morocco: insights into pollen representation and transport

Thirty-three modern surface samples were collected in the environmentally and climatologically contrasting regions of the Middle and High Atlas Mountains, Morocco. Samples representing forest and steppe montane environments (1935–2760 m above sea level) are clustered around study sites at Lake Tislit (High Atlas, semi-arid oro-Mediterranean bioclime) and Lake Sidi Ali and Michliffen (Middle Atlas, sub-humid montane Mediterranean bioclime). Good discrimination between regional pollen spectra is evident, with Middle Atlas samples reflecting higher arboreal cover (Cedrus and evergreen Quercus) and High Atlas samples with high abundances of non-arboreal taxa, including Artemisia and Fabaceae. These four taxa (Cedrus, evergreen Quercus, Artemisia and Fabaceae) are furthermore shown to be reliable indicators of local source vegetation within a 100 m² quadrat, taking into account threshold abundances of 7%, 20%, 4% and 10%, respectively. Deciduous Quercus, Olea and Phillyrea show long-distance pollen dispersal across both regions, contributing to non-trivial arboreal pollen (AP) values of up to 35% (typically 20–30%) in the High Atlas spectra. In the Middle Atlas, AP values of 40 to 50% occur in open sampling locations and > 60% under forest canopy cover. These insights should be taken into account when interpreting ancient pollen spectra from regional lakes and bogs for palaeoenvironmental reconstruction.     

Modern pollen/vegetation/land-use relationships in mountain environments: an example from the Champsaur valley (French Alps)

This study aims at elucidating modern pollen spectra/environmental data relationships from both natural and human-induced vegetation types as an aid for palaeo-ecological reconstructions. A set of 51 surface moss polsters was sampled from different vegetation and land-use types in the Champsaur area (French Alps) and analysed to obtain modern pollen analogues of ancient cultural landscapes in mountain ecosystems. Samples were selected from grazed areas, mown meadows, cultivated fields, fallow land and deciduous and coniferous forests. Vegetation composition around the sampling points and seventeen types of environmental variables (e.g., management type, soil and topography) were collected all for these 51 sites. Patterns of modern local pollen variation in relation to the environmental variables were explored by means of canonical correspondence analysis (CCA) and associated statistical tests. This correlative model allows us to determine the major explanatory variables and to identify taxa indicative of particular anthropogenic activities, and thus may help to calibrate fossil pollen assemblages.The indicator pollen types are evaluated in the light of comparable material from lowland and mid-elevation areas of western Europe. The results of the French data-set confirm some of the conclusions drawn from the North European data-sets, but also show some site specific features. Pollen markers with a broader global significance common to other regions include Rhinanthus type, Apiaceae and Dipsacaceae for mown meadows, Urtica type, Plantago media/major, Trifolium type and Potentilla type for grazed areas, and Cerealia type, Centaurea cyanus, and Polygonum aviculare for cultivated fields. New pollen anthropogenic indicators typical of our study area are Sanguisorba officinalis, Vicia type (mowing), Lotus type, Onobrychis type, Centaurea nigra type, Serratula type (grazing), Sinapis type and Papaver rhoeas (cultivation). This study provides potentially valuable analogues for human-induced vegetation types, and it may then become possible to interpret more objectively local pollen diagrams from Alpine mountain environments in terms of past cultural landscape development.     

Observations on desmids from Scotland,Shetland, Cumbria and the English Midlands

Summary

Samples from bogs and lochs collected in the parish of Assynt, south — west Sutherland; loch and pool samples from Shetland; tarn samples from Cumbria and a small pool in Oadby, Leicestershire, have been examined. New desmid varieties and formae, some rare desmids and interesting observations on common desmids can be reported.     

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.