Habitat and management preference of Bromus racemosus L., a rare species in mesic meadows of Northwest Europe, MR Simmelink, JAM Janssen, JHJ Schaminée

Tags: Netherlands, Bromus racemosus, grasslands, grass species, grassland management, nutrient availability, winter annual, rare species, Nature Conservation, Abstract Bromus, LUTZ, moist conditions, Wageningen University, Introduction Bromus, plant communities
Content: Tuexenia 37: 289­312. Gцttingen 2017. doi: 10.14471/2017.37.002, available online at www.tuexenia.de Habitat and management preference of Bromus racemosus L., a rare species in mesic meadows of Northwest Europe Habitat- und Managementprдferenz von Bromus racemosus L., einer seltenen Art in mдЯig nдhrstoffreichen Feuchtwiesen Nordwesteuropas Max R. Simmelink1, *, John A.M. Janssen2, Joop H.J. Schaminйe2, 3, 4 & Eddy J. Weeda5 1Graduate MSc Forest and Nature Conservation, Wageningen University, Grote Koppel 9, 3813AA Amersfoort, The Netherlands; 2Alterra-Wageningen University and Research, P.O. Box 47, 6700 AA Wageningen, The Netherlands; 3Nature Conservation and Plant Ecology Group, Wageningen University, The Netherlands; 4Radboud University Nijmegen, The Netherlands; 5Veerallee 28, 8019 AC Zwolle, The Netherlands *Corresponding author, e-mail: [email protected] Abstract Bromus racemosus L. is a rather rare grass species of moist meadows. It has strongly decreased in the course of the 20th century due to intensification of agricultural grassland management, and is therefore included in Red Lists of several European countries. Its winter annual life-cycle is remarkable for a species of permanent grasslands. The aim of this study is to determine the habitat preference and optimal management of B. racemosus in the Netherlands and surrounding countries. Vegetation, soil and hydrological data from 28 sites in the Netherlands have been compared with B. racemosus cover, and with vegetation data from surrounding countries. The results indicate that B. racemosus is characteristic of MolinioArrhenatheretea meadows with good mineralisation and aftermath grazing. The optimum lies in grasslands of the alliance Alopecurion pratensis (Deschampsion cespitosae), but the species ranges from wetter Calthion palustris meadows to drier Arrhenatherion elatioris and Cynosurion cristati grasslands. It prefers intermediate nutrient levels and hydrological conditions (mesic sites), but within this range the highest cover is found in relatively nutrient rich and dry sites. Because of the absence of a seedbank and a low dispersal capability, B. racemosus is vulnerable to changes in grassland management. A management of mowing after 15 June and aftermath grazing is most suitable, since it enables fruit ripening and the maintenance of an open sward, needed for germination and development. The risk of extinction is likely to be higher in flat polders than in floodplain sites with natural relief, where the species may shift between belts in different years. Keywords: aftermath grazing, alluvial landscape, Alopecurion pratensis, grassland management, hydrology, meadows, Molinio-Arrhenatheretea, soil nutrients, winter annual Erweiterte deutsche Zusammenfassung am Ende des Artikels
Manuscript received 21 February 2016, accepted 07 March 2017 Co-ordinating Editor: Christian Berg
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1. Introduction Bromus racemosus L. is a grass species of moist meadows that has become rather rare in recent times in Northwest Europe. Its winter annual life-cycle is remarkable for a species of moist, permanent grasslands; it flowers in May and June, and the seed germinate in summer and autumn (WEEDA 1994). According to the Dutch Flora, the species can be divided into two subspecies: Bromus racemosus subsp. racemosus and Bromus racemosus subsp. commutatus (VAN DER MEIJDEN 2005). In the Netherlands, the latter is much rarer than the former (WEEDA 1994). In neighbouring countries these taxa are considered separate species: B. racemosus and B. commutatus (SPALTON 2002, SCHOLZ 2008). In this article we distinguish these latter species, and use the name Bromus racemosus s.l. in case data include both taxa, or there is uncertainty or possible confusion between the two taxa. Flora Europaea (TUTIN et al. 2001) is followed with regard to all other vascular plants, SIEBEL et al. (2006) for mosses and SCHAMINЙE et al. (1996) for plant communities. According to HULTЙN & FRIES (1986) both species are native in large parts of Europe, but the distribution range of B. commutatus reaches further eastward than that of B. racemosus. BЦHLING et al. (1998) suppose that B. commutatus was introduced into Central Europe in Roman times, due to transport of crops from Southern Europe. For B. racemosus no Central European records from pre-Roman times are known either. HULTЙN & FRIES (1986) state that both taxa spread towards Central and Northern Europe by man. In the course of the 20th century, B. racemosus is declining in several European countries because of agricultural intensification, including drainage, manuring, a high mowing frequency and ploughing of grasslands (WEEDA 1994, PRESTON et al. 2002). The abandonment of mowing of wet meadows in other regions further contributes to its decline (LUTZ 1996, BЦHLING et al. 1998, ROSENTHAL 2003). Within the Netherlands, B. racemosus was present before 1950 in most lowland areas and in some brook valleys in higher parts of the country. During the second half of the 20th century, it showed a strong decline (>50%) in distribution (WEEDA 1985, SPARRIUS et al. 2014). Nowadays it is largely restricted to small Nature Reserves and only in the Central riverine area still fairly common. In the Dutch Red List B. racemosus s.l. is assessed as Vulnerable (SPARRIUS et al. 2014). In the German Red List B. racemosus is assessed as Endangered, whereas B. commutatus is not considered Endangered (LUDWIG & SCHNITTLER 1996), even though the latter has a smaller distribution range (BFN 2013). Germany has a high international responsibility for the conservation of B. racemosus, since this country contains a high proportion of the world population (ca. 10­33%) and is situated in the centre of its distribution range (LUDWIG et al. 2007). Bromus racemosus is one of the few annuals in moist, permanent grasslands. Therefore different grassland management methods may be needed compared to other grassland species. The aim of this study is to gain insight into the habitat and management preferences of B. racemosus s.l. in the Netherlands and surrounding lowlands, by studying in which plant communities and under which site conditions the species occurs and has the highest abundance. Describing the phytosociological position of the species helps to demonstrate the intermediate environmental position of the species. This study should lead to a better understanding of the factors that have contributed to the species decline and of management measures that may contribute to its conservation. The research was conducted for the MSc thesis of the first author. The thesis report (SIMMELINK 2014) provides an extended overview of the research. 290
2. Study area The study deals with 28 nature reserves in the Netherlands where Bromus racemosus s.l. has been observed recently (Fig. 1, Supplement E2). All sites contain grasslands owned by nature conservation organisations. Most sites are situated in the Central Dutch riverine area, in floodplains and polders (former floodplains) along the rivers Nederrijn, Lek, Waal, Gelderse IJssel (Rhine branches), Zwarte Water and Meuse. A concentration of locations is situated in the polder Vijfheerenlanden. An additional location is the brook valley Het Merkske near the border with Belgium. Finally, three sites in the polders of the Wadden island of Texel have been studied. In these study sites, the average temperature ranges from about 3.0 °C in January to ca. 17.5 °C in July; the mean annual temperature is 10.0 °C. The precipitation is on average 800­850 mm per year, with a precipitation surplus of 200­300 mm (KNMI 2011). In all 28 reserves the soil is dominated by mineral materials deposited during the Holocene by rivers, brooks, tidal rivers or the sea. Some sites have slightly peaty soils. The floodplain sites are located in between the low summer dike and the higher winter dike, a landscape that may be flooded in winter. Most sites in the floodplains still have their origi- Fig. 1. The studied sites in the Netherlands. See also Supplement E2. Abb. 1. Die untersuchten Standorte in den Niederlanden. Siehe auch Anhang E2. 291
nal relief. Ditches were created in all of the polder grasslands. The polder sites have a loamy or clayey top layer, sometimes with a layer of peat underneath. In many polder reserves, a semi-natural water regime is maintained, with a higher water level in winter and a lower level in summer. Because of drainage, the peaty layer oxidised and subsidence occurred. The wetter edges of the lots along the ditches subsided less than the centre of the parcels, resulting in a bathtub shape of parcels. The sites on the Wadden sea island of Texel are flat polders from marine origin (KOOMEN & MAAS 2004) and still have slightly brackish groundwater (VAN GOETHEM & VAN ROOIJEN 2011). Data from surrounding countries relate to sites with B. racemosus s.l. from lowland areas, in the vicinity of a river or a stream (Fig. 2, Supplement S1): - the Bremen region in Germany, along rivers like the Weser and smaller streams (LUTZ 1996); - three areas in Flanders (Belgium): Snoekengracht, an area in a brook valley to the east of Leuven; the valley of the river Dijle near Leuven; Bourgoyen, an area in the valley of the river Leie near Gent (VLAVEDAT - Flemish Vegetation Databank: VANDENBUSSCHE & HOFFMANN (2001); CALLEBAUT et al. (2007)); - in southern Wallonia (Belgium): valleys along streams and rivers, including the Meuse, in the regions Famenne and Fagne (SOUGNEZ & LIMBOURG 1963); - four areas in northern France: the valley of the river Sambre, a tributary of the Meuse (GЙHU 1961); stream valleys in the Bocage Virois in Basse-Normandie (DE FOUCAULT 1980); stream valleys in the Boulonnais in the dйpartement du Pas-de-Calais (DE FOUCAULT 1986); areas along the river Seine-maritime downstream of Rouen (FRILEUX et al. 1988); - two areas in the valley of the English river Thames: the North Meadow near Cricklade and the surroundings of Oxford (unpublished data). Fig. 2. The studied sites in the Netherlands (see also Fig. 1) and surrounding countries. See also the footnotes of Supplement S1 for the names of the locations. Abb. 2. Die untersuchten Standorte in den Niederlanden (siehe auch Abb. 1) und den umliegenden Lдndern. Siehe auch die FuЯnoten der Beilage S1 fьr die Namen der Standorte. 292
3. Material and methods 3.1 Data gathering In the 28 Dutch nature reserves in total 144 vegetation relevйs were made in representative, homogenous plots in which Bromus racemosus s.l. occurred, in the period 19 May ­ 5 July 2013. Vegetation relevйs were made according to the Braun-Blanquet method (BRAUN-BLANQUET 1964, SCHAMINЙE et al. 1995) in plot sizes of 9 m2, using the cover-abundance scale of BARKMAN et al. (1964), and environmental and soil data were sampled and recorded. An additional 18 recent relevйs with B. racemosus were added from the Dutch National Vegetation Database (DNVD; SCHAMINЙE et al. 2012). The locations of these relevйs were visited to obtain additional environmental and soil data. Additionally, 958 relevйs containing B. racemosus s.l. from the DNVD were selected, in order to look for other plant communities than to those covered by the field sampling. Further, vegetation samples made by the Instituut voor Biologisch en Scheikundig Onderzoek van Landbouwgewassen, Wageningen (IBS) were analysed on relations between the abundance of B. racemosus s.l. and abiotic conditions and grassland management. These samples have been collected in the period 1934­1958, using a sampling method with around 100 subsamples of 25 cm2 per sampled parcel, and additional collection of environmental and management data (KRUIJNE et al. 1967). From these set 414 samples made in May and June were selected, of which 87 contained B. racemosus s.l. Outside that period B. racemosus s.l. may have been overlooked in the samples. Finally, 250 relevйs with B. racemosus s.l. were digitised from lowland areas near Bremen in northern Germany (LUTZ 1996), the mentioned nature areas in Belgium (SOUGNEZ & LIMBOURG 1963, CALLEBAUT et al. 2007), northern France (GЙHU 1961, DE FOUCAULT 1980, 1986, FRILEUX et al. 1988) and from southern England (unpublished relevйs by A. Corporaal and by M. Raman). Part of the Belgian relevйs were obtained in a digitised form from VLAVEDAT (Flemish Vegetation Databank; VANDENBUSSCHE & HOFFMANN 2001). For 81 of the sampled relevйs an earth auger was used to study humus and soil profile until a depth of around 95 cm. Additionally, for 152 relevйs a bread knife was used to inspect the upper 20 cm of the humus profile without destroying the structure. The humus characteristics were described according to VAN DELFT et al. (2006). For every relevй one composite soil sample was taken. With a gouge ten subsamples of 20 cm depth were taken in a systematic pattern within the relevй, and mixed in a bag. Soil chemistry of the composite samples was analysed according to the guidelines as described in HOUBA et al. (1995). The following measurements were carried out: Organic Matter percentage (OM%) with the loss of ignition method; pH-H2O and pH-KCl; Nitrogen total and Phosphorus total (N-total and P-total respectively; both in mmol/kg, by destruction); Phosphate status (Pw in mg/kg, by water extraction); Potassium and Calcium (K and Ca respectively; both in mg/kg, by water extraction, using the Pw extraction). K and Ca were measured in a spectrometer. N-total, P-total and Pw were measured in an auto analyser. Nitrate (NO3) and Ammonium (NH4) were not recorded, because the soil samples became too warm during fieldwork to measure these values in a reliable way. Measurements of the groundwater table of some sites were derived from www.dinoloket.nl (GDN 2013), and indications for the annual groundwater table fluctuations from www.bodemdata.nl (ALTERRA 2013). These data included measurements of the mean highest and mean lowest groundwater table, calculated over a period of eight years. Inundation frequencies of some floodplains were estimated using AGGENBACH et al. (2007), www.live.waterbase.nl (RIJKSWATERSTAAT 2013) and altitudinal data from www.ahn.nl (HET WATERSCHAPSHUIS 2013). Together with information from site managers and hydromorphic properties in the soil profiles, these data were used to describe the hydrology in a qualitative way. 293
3.2 data analysis All vegetation relevйs were entered in Turboveg (HENNEKENS & SCHAMINЙE 2001). Mean Ellenberg indicator values (ELLENBERG et al. 2001) were calculated for all relevйs, based on ordinal cover and on absence/presence. The vegetation relevйs were imported into Juice (TICHЭ 2002) and clustered with help of a modified version of the programme Twinspan (ROLECEK et al. 2009). The following settings were used: 2 cut levels: 0­25; number of clusters: 19; measure of cluster heterogeneity: total inertia; min. group size: 2. The resulting classification was improved by hand, by dividing, merging and translocating some clusters, and moving some relevйs. Species were assigned to clusters according to fidelity, measured by the phi coefficient (CHYTRЭ et al. 2002). The resulting clusters from the field data set were assigned to plant communities as described by ZUIDHOFF et al. (1996) and SCHAMINЙE et al. (2015) using expert knowledge. Indirect ordination in Canoco 5 (TER BRAAK & SMILAUER 2012) was used to analyse the relation between plant communities and environmental gradients for the field work data. Because of a short length of gradient (2.9), the linear method Principal Component Analysis (PCA) was applied. Some environmental variables were log-transformed before analysis. Besides, a representative subset of 237 of all available relevйs from the Netherlands and surrounding countries were selected for ordination to analyse the habitat diversity. The unimodal method Detrended Correspondence Analysis (DCA) was chosen, since the length of gradient was 3.24, with the option detrending by 26 segments. IBM SPSS Statistics 22.0 (IBM CORP. 2013) was used for univariate statistics, which have been applied to the dataset from field work. Because many variables could not be transformed to a normal distribution, non-parametric tests were used. The cover of B. racemosus s.l. was correlated to environmental variables with Spearman's rank correlation coefficient test. The Kruskal Wallis test with stepwise stepdown multiple comparisons was used to test the differences of environmental values between plant communities. The Mann-Withney U test was used in case two groups of relevйs (often groups with and without B. racemosus s.l.) were compared. This test has also been used to analyse the IBS dataset. 4. Results 4.1 Plant communities The classification of the Dutch relevйs resulted in eleven clusters of relevйs (Table 1). The relevй table and a synoptic table are included in Supplement S2 and E1, with the original Twinspan classification given in the header data. In Table 1 for each cluster the species composition and distribution are described, and it is mentioned whether, besides Bromus racemosus, also B. commutatus was observed in the community. In Table 2 environmental values and significant differences are presented for all clusters. All clusters are placed within the class Molinio-Arrhenatheretea Tьxen 1937, based on the large number of class character species in all relevйs, including Cerastium fontanum subsp. vulgare, Ranunculus acris, Rumex acetosa, Trifolium pratense, Cardamine pratensis and Holcus lanatus. Within this class, B. racemosus has been found in four of the five alliances that have been distinguished in the Netherlands: Calthion palustris Tьxen 1937, Alopecurion pratensis Passarge 1964, Arrhenatherion elatioris Koch 1926 and Cynosurion cristati Tьxen 1947. The same clusters are indicated in the PCA diagram, of which the first three axes are represented (Fig. 3a, b). The first axis can be interpreted as a moisture gradient, the second as a pH gradient and the third as a nutrient gradient. Bromus racemosus s.l. cover is mainly correlated to axis 1 (r = -0.3114) and 3 (r = -0.3262). 294
Table 1. Description of eleven vegetation clusters resulting from analysis of 162 Dutch relevйs with Bromus racemosus s.l., completed with three additional clusters from DNVD data. See Supplement E1 for the complete relevй table of the 11 clusters. Tabelle 1. Beschreibung von elf Gruppen von Vegetationsaufnahmen, die sich aus der Analyse von 162 niederlдndischen Vegetationsaufnahmen mit Bromus racemosus s.l. ergeben, ergдnzt mit drei zusдtzlichen Gruppen aus DNVD Daten. Siehe Anhang E1 fьr die komplette Vegetationstabelle der 11 Gruppen.
Cluster Cluster 1 Cluster 2 Cluster 3 Cluster 4 Cluster 5 Cluster 6 Cluster 7
Plant community (after ZUIDHOFF environmental factors et al. 1995, SCHAMINЙE et al. 2015)
Rhinantho-Orchietum morionis (Calthion palustris)
Like cluster 2, but fragmentary developed, without character species Anacamptis morio, and on slightly more productive soils. Bromus racemosus has a medium high cover in most relevйs.
Rhinantho-Orchietum morionis (Calthion palustris)
Restricted to polders on the island of Texel, on low productive soils, with a low cover of the herb layer and a high cover of the moss layer. Bromus racemosus has a low cover in most relevйs.
Lolio-Cynosuretum (Cynosurion cristati)
Restricted to the island of Texel. Sites with management of mowing with aftermath grazing, on slightly higher productive soils than cluster 1 and 2. Bromus racemosus has a relatively high cover in most relevйs.
Arrhenatheretum elatioris typicum (Arrhenatherion elatioris)
Community on relatively drier, but productive sites, often on higher ridges in floodplains. Bromus racemosus s.l. reaches a relatively high cover in most relevйs, but lacks in the higher zonation belts, especially when the loam content is lower than 40%. B. commutatus was present in six of 28 relevйs.
Basal Community Bromus racemosus[Alopecurion pratensis]
Most species-poor of all communities, found both in polders and floodplains. Stands belonging to this community have in general been fertilised heavily until 5 to 40 years ago, and have since then been mown twice a year without fertilisation, to decrease the nutrient availability. In this community B. racemosus reaches its highest cover.
Fritillario-Alopecuretum pratensis (Alopecurion pratensis)
On two sites in floodplains along the Zwarte Water, close to a former estuary, on relatively acid soils (compared to the Rhine branches). Sites are inundated by river water for a limited period, but water table fluctuations are reduced due to the vicinity to the estuary. Bromus racemosus has a low or medium high cover.
Sanguisorbo-Silaetum (Alopecurion pratensis)
Rare, species-rich community, found in two floodplains in the western river area, along the Meuse and Nieuwe Merwede (Hengstpolder). Bromus racemosus s.l. has a low cover on relatively low situated parts and a moderate cover on higher parts. Inundation occurs usually in winter, but the soil desiccates superficially in summer. From even higher parts of the Hengstpolder also B. commutatus is known (WEEDA 1991).
295
Cluster
Plant community (after ZUIDHOFF Environmental factors et al. 1995, SCHAMINЙE et al. 2015)
Cluster 8
Transition between SanguisorboSilaetum & Calthion palustris
Moderately species-rich community from floodplains along the Nederrijn and Lek. The groundwater table is relatively high during summer, because of respectively tidal influence and the presence of a weir. The soil is relatively nutrient rich. Bromus racemosus has a relatively high cover, on average.
Cluster 9
Basal Community Rhinanthus angustifolius-Lysimachia vulgaris[Calthion palustris]
This community was found in a clay pit along the Lek, on relatively nutrient rich soils with a high amount of tall herb species. All three relevйs contain B. commutatus, in low cover.
Cluster 10 Angelico-Cirsietum oleracei (Calthion palustris)
On a gentle slope of the stream valley of Het Merkske (Kempen region), where seepage of calcareous groundwater causes a neutral pH and a constant high groundwater table, but flooding does not occur. Bromus racemosus occurs with low cover and was more common in the past.
Cluster 11
Ranunculo-Senecionetum aquatici, subassociation juncetosum articulati* (Calthion palustris)
Stands on parcels that were bought by nature organisations around 25­50 years ago, and had never been heavily fertilised. Mainly situated in polders on overbank deposits in flood basins and on transitions to natural levees. In winter the groundwater table is high, and in some sites inundation by a mixture of rain and river water occurs for a short period. B. racemosus occurs on average with low cover and is usually rare or absent in lower situated, longer inundated parts of these meadows. B. commutatus was present in one relevй.
Additional cluster DNVD1
Trifolio fragiferi-Agrostietum stoloniferae Sэkora 1982 (LolioPotentillion anserinae Tьxen 1947)
Cluster with relevйs from around 1940, characterised by species indicating salty or desalinating conditions, like Alopecurus bulbosus, Juncus gerardii and Glaux maritima. This community developed on the former coast of the Zuiderzee in desalinated former salt marshes, after this sea had been dammed to create the IJsselmeer in 1932 (Boer 1955). B. racemosus was abundant in a succession stage between saline communities and fresh water, mesic grasslands.
Additional cluster DNVD2
Carici curtae-Agrostietum caninae Tьxen 1937 (Caricion nigrae Koch 1926 em. Nordhagen 1936)
A cluster indicating very wet, nutrient poor, slightly alkaline conditions in peat areas. Accompanying species are amongst others Pedicularis palustris, Carex diandra and Eriophorum angustifolium. The cover of B. racemosus is low. The relevйs are from the period 1936-1999 from several areas in the Netherlands, mainly in the west.
Additional cluster DNVD3
Ranunculo-Senecionetum aquatici, subassociation caricetosum paniceae* (Calthion palustris)
A cluster of wetter, more nutrient poor and more acid conditions than in community 11. All relevйs are from the period 1936­1978, mainly in the northeast of the Netherlands; they resemble former vegetation in the Bremen region in Germany.
*Synonym to Bromo-Senecionetum aquatici according to POTT (1992).
296
Table 2. Median values of several environmental variables per community, the names and descriptions of the communities can be found in Table 1. The communities 9 and 10 are omitted, since they were only observed in one site and few relevйs have been made. The similar communities 1 and 2 are combined to increase the Sample size. Total n = 153, d.f. = 7; Kruskal Wallis test with stepwise stepdown multiple comparisons. The letters indicate significant differences. The asymptotic significance (2 sided test) is < 0.001 for all shown variables. E: Ellenberg value, OM: organic matter, Pw: Phosphate status (water extraction). Tabelle 2. Medianwerte von mehreren Umweltvariablen pro Pflanzengesellschaft (Namen und Beschreibungen der Pflanzengesellschaften s. Tabelle 1). Die Gesellschaften 9 und 10 sind weggelassen, da sie nur auf einem Standort beobachtet wurden und nur einige Vegetationsaufnahmen vorliegen; die дhnlichen Gemeinschaften 1 und 2 sind kombiniert, um die Probenzahl zu erhцhen. Gesamtes n = 153, d.f. = 7; Kruskal Wallis-Test mit schrittweisen multiplen Rьckwдrtsvergleichen Die Buchstaben zeigen signifikante Unterschiede. Die asymptotische Signifikanz (2-seitiger Test) ist < 0,001 fьr alle gezeigten Variablen. E: Ellenberg-Wert., OM: Organische Substanz, Pw: Phosphat-Status (Wasser-Extraktion).
Plant communities from field work data
Predictors
1+2
3
(n = 8) (n = 15)
E_moisture
6.37c 5.93b
E_nitrogen
4.40a 5.04b
Pw (mg/kg)
0.8a
2.7b
P-tot (mmol/kg) 6.2a 10.0b
K (mg/kg)
15 a
46b
pH-KCl
5.83bc 5.81c
pH-H2O E_pH
6.35ab 5.48a
6.77b 5.57ab
Ca (mg/kg)
282ab 142a
OM (%)
9.0b
4.9a
Clay (%)
13.5ab
10a
Loam (%)
63ab
50a
E_light
7.14c 7.02c
Herb cover (%) 88.5a 95c
Moss cover (%) 50c
0a
Mean species
32bc
29b
number
4 (n = 28) 5.48a 5.78c 3.8b 20.1c 131c 7.05d 7.59c 6.44d 654c 10.1b 12ab 54a 6.86b 95c 2.5b 32bc
5 (n = 42) 6.33c 5.73c 7.4d 27.4e 253d 5.04b 6.02a 5.96c 330b 14.6cd 18c 73bc 6.76a 92.5bc 3b 25a
6 (n = 10) 6.83cd 5.54c 4.3bc 22.0cd 183cd 4.87ab 5.93a 6.21c 194a 11.8bc 17.5bc 67.5abc 6.78ab 90ab 10c 37.5d
7
8
11 Test
(n = 10) (n = 8) (n = 33) Stat.
7.39d 5.31b
6.99d 5.63c
6.96d 100.274 5.03b 101.210
3.4b
6.2cd 5.8cd 69.434
18.7c 36.65e 24.9d 82.800
107bc 178cd 278d 59.305
7.11d 7.66c 6.47d 609cd 8.5b 10ab 55a
7.17d 7.70c 6.44d 856d 12.7cd 18.5c 76bc
4.64a 5.78a 5.85bc 291b 15.4d 20c 80c
72.309 76.867 83.232 66.289 79.119 59.700 54.155
6.88b 95bc 3b 41e
6.74a 94abc 2b 31.5bc
6.89b 90ab 10c 34cd
54.382 26.454 43.691 65.706
Clustering of the 958 older relevйs with B. racemosus s.l. from the DNVD revealed three additional plant communities in which B. racemosus s.l. was found in the past (Table 1). Of these clusters one relates to a Calthion palustris community, while two communities are not covered by the class Molinio-Arrhentheretea: one from the alliance Lolio-Potentillion anserinae Tьxen 1947 and one from the Caricion nigrae Koch 1926. The clustering of data from foreign countries revealed that most relevйs are quite similar to the Dutch relevйs and could be assigned to the same alliances. Supplement S1 shows which alliances where covered by the data of the different countries, and which species have a high frequency in the communities. The communities from northern Germany (D), Bel-
297
Fig. 3a. Principal Component Analysis (PCA) of 158 relevйs from fieldwork, that have been assigned to eleven communities with help of Twinspan, as shown in Table 1 and the relevй table (Supplement E1). The communities are depicted by symbols, as explained in the legend. Axis 1 and 2 are plotted, eigenvalues are 0.124 and 0.092 respectively. Only continuous environmental variables with a correlation of more than 0.30 with one of the plotted axes are shown. E: abbreviation for Ellenberg value. Br_cover: Bromus racemosus s.l. cover. Abb. 3a. PCA von 158 Vegetationsaufnahmen aus Feldarbeit, die mit Hilfe von Twinspan elf Pflanzengesellschaften zugeordnet wurden, wie in Tabelle 1 und der Vegetationstabelle (Anhang E1) gezeigt. Die Gesellschaften werden durch Symbole dargestellt (s. Legende). Achse 1 und 2 sind dargestellt, die Eigenwerte sind jeweils 0,124 und 0,092. Nur kontinuierliche Umgebungsvariablen mit einer Korrelation von mehr als 0,30 mit einer der dargestellten Achsen werden gezeigt. E: Ellenberg-Wert, Br_cover: Bromus racemosus s.l. Deckung. gium (B), northern France (F) and southern England (E) overlap largely with the Dutch relevйs and can be assigned to the same four alliances and additional Basal Community of the class Molinio-Arrhenatheretea. Bromus racemosus can reach a high cover in all of these alliances. In the DCA plot (Fig. 4) the same clusters from Supplement S1 have been indicated. The Dutch relevйs are distributed over a large part of the ordination space. Most relevйs from fieldwork are placed on the nutrient-richest half of the ordination space (the lower part), 298
Fig. 3b. Principal Component Analysis (PCA) of 158 relevйs from fieldwork, that have been assigned to eleven communities with help of Twinspan (see Fig. 3a for explanation). Here Axis 1 and 3 are plotted, eigenvalues are 0.124 and 0.077 respectively. Abb. 3b. PCA von 158 Vegetationsaufnahmen aus Feldarbeit, die mit Hilfe von Twinspan elf Pflanzengesellschaften zugeordnet wurden (siehe Abb. 3a fьr Erlдuterung). Hier sind Achse 1 und 3 dargestellt, die Eigenwerte sind jeweils 0,124 und 0,077. except for the relevйs from Texel (Dutch clusters 1 and 2), which do not resemble any foreign relevйs. The relevйs from Bremen (Germany) and Flanders (Belgium) indicate relatively wet and acid sites. They belong to the Calthion palustris and basal communities of nutrient-rich sites. The relevйs from Famenne & Fagne (southern Belgium) and the Bassin de la Sambre (northeastern France) are drier and indicate more basic conditions. This Alopecurion vegetation is characterised by the presence of Colchicum autumnale and a high cover of Alopecurus pratensis and B. racemosus. The French relevйs span a range of all four alliances, like the Dutch. The English relevйs overlap with the drier Alopecurion and Arrhenatherion relevйs. Although their average Ellenberg value for moisture is almost the same as for the Dutch relevйs, their position in the ordination indicates drier conditions. This may be explained from the more Atlantic (moist) climatic conditions, which enables growing in drier soils. 4.2 Hydrological conditions The Ellenberg value for moisture of the relevйs varied in the range 5­8. Bromus racemosus s.l. cover was negatively correlated to the Ellenberg value for moisture (r = -0.206, p = 0.009, n = 158), and reached an optimum around 6.2. 299
Fig. 4. Detrended Correspondence Analysis (DCA) of a selection of 237 relevйs with Bromus racemosus s.l. (74 from own fieldwork, 11 from the DNVD, 152 from nearby countries). The communities 8 and 9 are not included. Communities are depicted by the form of the symbols, countries are marked with a colour (Germany (D): green, Belgium (B): red, France (F): blue, border region of Belgium and France (BF): purple, Netherlands (N): transparent). The numbers between [ ] refer to the footnotes of Supplement S1. Axis 1 and 2 are plotted, eigenvalues are 0.327 and 0.221 respectively. Ellenberg values are plotted as environmental values. Encircled areas indicate relevйs from communities 1 and 2 (upper circle) and most relevйs from the Ranunculo-Senecionetum caricetosum paniceae (made in 1936­1974; lower circle); recent Dutch relevйs do not belong to the last group. Abb. 4. DCA von einer Auswahl von 237 Vegetationsaufnahmen mit Bromus racemosus s.l. (74 durch eigene Feldarbeit, 11 aus der DNVD, 152 aus umliegenden Lдndern). Die Pflanzengesellschaften 8 und 9 sind nicht enthalten. Gesellschaften werden durch die Form der Symbole dargestellt, Lдnder mit einer Farbe gekennzeichnet (Deutschland (D): grьn, Belgien (B): rot, Frankreich (F): blau, Grenzregion von Belgien und Frankreich (BF): lila, Niederlande (N): transparent). Die Zahlen zwischen [] beziehen sich auf die FuЯnoten von Beilage S1. Achse 1 und 2 sind dargestellt, die Eigenwerte sind jeweils 0,327 und 0,221. Ellenberg-Werte sind als Umgebungsvariablen angegeben. Umkreiste Bereiche zeigen Vegetationsaufnahmen der Gesellschaften 1 und 2 (oberer Kreis) und die Mehrheit der Vegetationsaufnahmen aus dem Ranunculo-Senecionetum caricetosum paniceae (1936­1974 erstellt; unterer Kreis) an; in der letzten Gruppe befinden sich keine neuen niederlдndischen Vegetationsaufnahmen. 300
The hydrological conditions differed between sites in polders and sites in floodplains. In polder grasslands (outside the direct influence of the river), the species was only found in parts with a mean highest groundwater table less than 40 cm below soil surface, in most sites less than 25 cm (ALTERRA 2013, GDN 2013). The species was most abundant on parts of parcels that are never or rarely inundated by rain or groundwater, and was lacking in parts that are inundated for a long period each winter. In areas where the soil consists of clay on top of peat, typically B. racemosus had a higher cover on the elevated borders of parcels with a bathtub shape. In the centre, where rain water stagnation occurs during winter, the species had a low cover or was absent. The mean lowest groundwater table was usually between 60 and 110 cm below soil surface. The species was absent or rare in areas with a higher lowest groundwater table. In floodplains, in sites that are flooded by river water at least once every ten years, the inundation frequency and duration determine the distribution of B. racemosus. Also here, the species occurred at relatively dry places with a mean highest groundwater table more than 80 cm below soil surface. In most sites, its upper distribution boundary was the belt that is inundated every few years. On its lower boundaries, B. racemosus appeared to be limited by long inundations, but it seemed to tolerate winter inundation with river water for a longer period (probably almost two months) than inundation with rain and groundwater in the polders. Inundation for a couple of days of flowering B. racemosus was observed in Cortenoever (along the Gelderse IJssel) in early June 2013. Bromus racemosus died and was not able to produce fruits in zones were the water table had been more than about 80 cm above the surface. The lower the water depth had been, the more individuals survived and produced fruits. Inundation in the flowering period is rare in the Netherlands, but it became more frequent during the 20th century (AGGENBACH et al. 2007). 4.3 Soil conditions Data from fieldwork (n = 158) indicated that B. racemosus s.l. is relatively abundant on relatively nutrient-rich, productive sites. Its cover was correlated negatively to the Ellenberg value for light (r = -0.359, p < 0.001) and positively to the Ellenberg value for nitrogen (r = 0.252, p = 0.001), N-total (r = 0.211, p = 0.008), P-total (r = 0.289, p < 0.001), Pw (r = 0.250, p = 0.002) and K (r = 0.162, p = 0.041). Bromus racemosus s.l. cover was positively correlated (n = 162) to herb cover (r = 0.358, p < 0.001) and negatively to moss cover (r = -0.183, p = 0.021). It was also negatively correlated to species richness (r = -0.288, p = 0.001). From these correlations it may be concluded that B. racemosus s.l. cover is higher under more nutrient-rich conditions. However, these conclusions hold only for the mesotrophic range of the study sites. Bromus racemosus s.l. was not observed in more nutrientrich (eutrophic) sites. The Ellenberg value for nitrogen of relevйs with B. racemosus s.l. varied between 4.36­6.42. Hardly any of the visited sites had been fertilised within the last five years. In the IBS dataset B. racemosus s.l. is negatively correlated to measures for the availability of K (r = -0.258, p < 0.001, n = 395) and P (r = -0.121, p = 0.015, n = 406), and the species was absent from sites with very high values. From the remarks about fertilisation it appears that its cover is somewhat higher in fertilised than in unfertilised sites. This indicates that a moderate gift of fertiliser has a positive effect on the species, whereas heavy fertilisation has a negative effect. 301
The acidity of the plots varied in the following ranges: pH-H2O 5.1­7.9, pH-KCl 4.0­7.5, Ellenberg value for reaction lie in the range 4.95­7.13. The pH and the Calcium concentration were not significantly correlated to the cover of B. racemosus s.l. Bromus racemosus occurs normally on soils with a fine texture. In the data from fieldwork, loam percentage (clay + silt) was 20­90% and clay 4­26%, but B. racemosus s.l. cover was not significantly correlated to those parameters. If only the communities of drier habitat (cluster 3­5) are selected, the species cover shows a positive correlation to loam percentage (r = 0.223, p = 0.040, n = 85). In the IBS dataset the species is more often present on sites with a finer texture (MannWithney U: 16402.5, p = 0.011, n = 407), and it was also found on sites with a clay percentage much higher than 26%. On the other hand, it was found a few times on sites with a coarse texture as well. In almost all relevйs the humus form was a type of mull, a humus form with a thin or absent ectorganic horizon and a good mineralisation. No soils with podzol were found. 4.4 Grassland management In the IBS dataset B. racemosus s.l. was more frequent in meadows with aftermath grazing than in grasslands that are only mown or grazed (Test stat. = 77.741, p < 0.001, d.f. = 5, n = 380). In the data set, the species was found in 12.5% of the 48 meadows that were only mown, in 48.3% of the 116 meadows with aftermath grazing and in 4.3% of 92 pastures. This makes B. racemosus s.l. the species with the strongest preference for meadows with aftermath grazing of around 200 common grassland species studied by the IBS (KRUIJNE et al. 1967). Fig. 5. Cover-abundance of Bromus racemosus s.l. on the ordinal scale in meadows with and without aftermath grazing (Mann-Withney U: 1292.0, p = 0.007, n = 151). Abb. 5. Abundanz/Dominanz von Bromus racemosus s.l. auf der Ordinalskala in Wiesen mit und ohne Nachbeweidung (Mann-Withney U: 1292,0; p = 0,007; n = 151). 302
With the data from fieldwork, it was tested whether B. racemosus s.l. reaches a higher cover on meadows with aftermath grazing, compared to meadows that are only mown. This was indeed the case (Mann-Withney U: 1292.0, p = 0.007, n = 151, Fig. 5). Within the 120 relevйs with aftermath grazing, it was tested whether the yearly number of mowings was related to the cover of B. racemosus s.l. (92 relevйs were mown once, 28 twice). The cover was higher in sites that were mown twice (Mann-Withney U: 1689.5, p = 0.010, n = 120). Most sites visited during fieldwork are mown in the second half of June or later. Only two productive, species-poor sites were mown earlier, around the first of June; here B. racemosus was locally abundant. The species was found quite abundant in two extensively grazed pastures, with mosaics of long and short vegetation. It grew in intermediate grazed sites, but was not found in shortly grazed vegetation and very rare in tall vegetation. 5. Discussion 5.1 Plant communities In the Netherlands Bromus racemosus is considered a weak character species of the Alopecurion pratensis Passarge 1964 (ZUIDHOFF et al. 1996). In terms of hydrology, this alliance is intermediate between the drier Arrhenatherion and the wetter Calthion palustris. The phytosociological analysis of the relevйs from fieldwork revealed that B. racemosus reaches its optimum (highest cover) in the Netherlands in a basal community of the alliance Alopecurion pratensis, occurring on relatively nutrient rich, moderately moist soil. Furthermore it occurs in Alopecurion associations, relatively dry parts of Calthion sites, relatively wet Arrhenatherion sites and moist, mown Cynosurion sites. Our analyses confirm that the optimum of the species is in the Alopecurion pratensis, but also indicates that the range of alliances is wider than suggested in literature. The same range of alliances of the class Molinio-Arrhenatheretea was found in our analysis of relevйs from England, Germany, Belgium and France. In Germany, B. racemosus is considered a character species of the Calthion palustris, an alliance of wet, mesotrophic meadows (LUTZ 1996, BЦHLING 1998, BURKART et al. 2004, PДZOLT & JANSEN 2004). OBERDORFER (1957) describes four types of B. racemosus meadows typical for base-rich soils with a low lime content along streams and rivers in the lowlands, which besides typical Calthion species, also contain species that are characteristic of the Alopecurion pratensis. In the sites near Bremen, at the end of the 20th century, B. racemosus had disappeared from many Calthion palustris grasslands and was mainly found in species poor basal communities (LUTZ 1996), that are similar to the Dutch basal community from cluster 5. It is remarkable that relevйs from the Senecioni-Brometum racemosi (Calthion palustris) near Bremen are quite similar to the relevйs of the Ranunculo-Senecionetum aquatici caricetosum paniceae from the DNVD, made in the period 1936­1978 in the northeast of the Netherlands, as currently B. racemosus seems to be absent from sites in this latter region. In Austria WILLNER et al. (2013) found B. racemosus with a low cover in an association belonging to the Calthion. BOTTA-DUKБT et al. (2005) studied lowland wet meadows in a large part of Central Europe. They conclude that the Alopecurion pratensis cannot be floristically separated from the alliances Agrostion albae Soу 1941, Cnidion venosi Balбtovб-Tulбckovб 1966, Deschamps- 303
ion cespitosae Horvati 1930 and Veronico longifoliae-Lysimachion vulgaris BalбtovбTulбckovб 1981. Therefore they propose to use the name Deschampsion cespitosae for all these alliances. They mention B. commutatus as a diagnostic species for mesic, continental Deschampsion cespitosae meadows in Hungary and southern Slovakia. In England, GOWING et al. (2002) state that B. racemosus occurs in the community MG4 Alopecurus pratensis - Sanguisorba officinalis grassland (RODWELL 1992), a floodplain meadow comparable to the Dutch Alopecurion pratensis. This is confirmed by the analysed British relevйs. Also French authors describing floodplain communities (SOUGNEZ & LIMBOURG 1963, DE FOUCAULT 1986, FRILEUX et al. 1988), indicate that B. racemosus is absent or rare in the wettest flood meadows (like the Ranunculo-Alopecuretum geniculati) and the driest sites (Arrhenatherion sites), having its optimum in between. In France and Wallonia communities with B. racemosus are often placed in the alliance Bromion racemosi (GEHU 1961, SOUGNEZ & LIMBOURG 1963, DE FOUCAULT 1986), which corresponds to the Calthion palustris. However, in some case the species composition is closer to the Alopecurion pratensis. Besides, the species has been observed in the Arrhenatherion elatioris (DE FOUCAULT 1988, FRILEUX et al. 1988), and also in our data the French data covered the relatively driest alliances Arrhenatherion and Cynosurion. Overall, although B. racemosus occurs regularly in communities of the Calthion palustris, our data demonstrate that its optimum in NorthWestern Europe lies in communities of somewhat drier habitats, where the species reaches the highest covers. Bromus racemosus can be considered as a character species of the Molinio-Arrhenatheretea, with a preference for moderately moist, mesotrophic Alopecurion pratensis meadows. This is in accordance with the situation described by BOTTA-DUKБT et al. (2005) for Central Europe. Outside these alliances of semi-natural, mesic grasslands, B. racemosus occurs sometimes in desalinating sites, as described for the Netherlands (BOER 1955), and reported from Belgium (ZWAENEPOEL 2006) and Germany (LUTZ 1996). This is however a rare situation, and the only example of more `natural' plant communities. It is noteworthy that the genus Bromus (s.str., comprising only annual species) is a group of species confined to anthropogenic habitats (anecophytes), and many Bromus taxa having probably evolved in Holocene times in the context of agriculture (SCHOLZ 2008). Some of them are weeds in arable land and mimic crops in their floral characters, like B. secalinus (Secale cereale, Hordeum vulgare), B. bromoideus (Triticum spelta), and B. interruptus (Onobrychis viciifolia, Trifolium spp.). The latter two had a very restricted area (East Belgium and England respectively) and are extinct in the wild nowadays (SMITH 1980). Other Bromus species are adapted to grassland management, like B. racemosus, whose ecology suggests an origin as a hay-field specialist. 5.2 Hydrology Since moisture could not be measured in the field during our study, mean Ellenberg values were used. This method has been criticised (ZELENЭ & SCHAFFERS 2012), but in practice it works well for comparing vegetation plot data (DIEKMANN 2003). Bromus racemosus has a quite narrow hydrological amplitude. It grows only on sites with moist conditions in winter or limited periods of inundation. The species cannot survive long inundation (GALL 1995, cited in LUTZ 1996), probably because the wintering plants need oxygen. In floodplains, spring flooding may be disastrous for B. racemosus, as was observed along the river IJssel in Cortenoever and confirmed by GREVILLIOT (1996). Also flooding in summer may cause a decline (LUTZ 1996). Many floodplain plants are more 304
tolerant to (regular) winter flooding than occasional summer floods (VAN ECK et al. 2006). The species seems to tolerate inundation with river water better than inundation with rain and ground water, which may be explained by the higher nutrient content and alkalinity of river water. In floodplains, the species was found under drier circumstances than in polders, provided the soil was sufficiently loamy (> 40%) and inundation occurred at least every 5­10 years. The cover of B. racemosus was higher in the relatively drier sites, which is confirmed by LUTZ (1996) and KALUSOVБ et al. (2009). Probably B. racemosus benefits from summer desiccation, in competition with other species, since it can survive as a fruit until moist conditions occur. GOWING et al. (2002) suggest that the winter annual life cycle of B. racemosus could be an adaptation to avoid summer drought and/or to fruit before the first hay-cut. In even more dry sites the species may lack because of sensibility to desiccation in spring, or because of a weaker competitive ability. The hydrologic range of the species is smaller in polders than in floodplains. But in polders, the species reached more often high covers (> 12,5%). Here a more stable water table enables the species to build up a large population with a high cover. However, the risk of extinction with changing hydrology is higher in flat polders than in floodplains, as in the latter the population may shift between higher and lower places within the natural gradient. 5.3 Soil conditions The cover of B. racemosus s.l. was higher in relatively nutrient-rich plots, but the species is absent from even more nutrient-rich (eutrophic) sites. It's likely that other grass species outcompete B. racemosus under eutrophic soil conditions. Besides, in heavily manured parcels the first hay cut is usually already in April or May, before ripening of fruits of B. racemosus (GREVILLIOT et al. 1998). When nutrient richness of former agricultural grasslands is decreased by mowing without fertilisation, B. racemosus can be among the first species that recolonise the grasslands, provided fruits are present in the vicinity, as was demonstrated in the Drentsche Aa region by VAN DUUREN et al. (1981). Under moderate nutrient-rich conditions it can compete well with perennial, nitrophilous grasses, and here the species has its optimum (KALUSOVБ et al. 2009). A further reduction of soil fertility may lead to a decline of B. racemosus. Nowadays in the Drentsche Aa region the species has completely vanished, possibly due to continued lowering of nutrient availability and/or rising of the water table. Bromus racemosus is probably not dependent on soil texture, but on correlated factors like moisture and nutrient availability. Heavy clay for example seems unsuitable for B. racemosus, because its low permeability may lead to long stagnation of water. A relatively open substrate is important for germination of B. racemosus. During fieldwork the species was only observed on sites with mull humus, characterised by a good mineralisation, few litter and without a well-developed root math. Moss cover was negatively correlated to B. racemosus s.l. cover. 5.4 Grassland management Bromus racemosus flowers in May and early June and fruits in June, before the traditional mowing dates. It is monocarpic, dying soon after ripening (or failure) of the seeds. In this way it is not sensitive to superficial drying of the soil in summer (WEEDA 1994). After shedding, the seeds germinate as soon as they become moist. Light does not influence germina- 305
tion. In moist meadows all seeds usually will germinate, and therefore no seedbank is formed (LUTZ 1996, JENSEN 2004). In this respect, B. racemosus behaves like a cereal, like the closely related B. secalinus. The seeds have a good germination ability: 86% respectively 93% germinated in experiments of LUTZ (1996) and JENSEN (2004). Bromus racemosus responds rapidly to changes in environmental conditions. The species disappeared from some sites where it had been abundant one or a few years before, and on the other hand it was able to establish and increase remarkably on a parcel within a few years (VAN DUUREN et al. 1981, LUTZ 1996). An appropriate and stable management is necessary to assure that the population survives the two most important bottlenecks of the species lifecycle each year. The first bottleneck is that it should not be mown or grazed before the seeds have ripened. Mowing after 15 June is advisable (see also LUTZ 1996). Since the fruits are relatively heavy (LUTZ 1996, JENSEN 2004), they may be unable to travel large distances by wind transport, and dispersal by mowing machines may be important. The second bottleneck is the establishment of the seedlings in summer and autumn. The vegetation should not be too dense and tall, which can be guaranteed by aftermath grazing and/or mowing twice a year. Aftermath grazing generates open spots in the sward through trampling, which are a good substrate for germination and development of seedlings. MEISEL (1969) also states that B. racemosus has the highest frequency in meadows with aftermath grazing. OBERDORFER (1994) mentions that B. racemosus is more frequent in meadows that are only mown than in meadows with aftermath grazing. The advantage of mowing twice a year is questionable. Since this management practice is mostly applied on nutrient rich parcels, B. racemosus may reach a higher cover partly because of the high nutrient availability. Continuation of mowing with the same frequency will decrease nutrient availability faster and may lead to a lower B. racemosus cover on the longer term, without any change in management. 6. Conclusion Bromus racemosus has its optimum in moderately moist, moderately nutrient-rich Molinio-Arrhenatheretea meadows with good soil mineralisation. In the Netherlands its highest cover is in a basal community of the Alopecurion pratensis (=Deschampsion cespitosae). Further it occurs in other Alopecurion communities, relatively dry parts of wet meadows (Calthion palustris), and relatively moist parts of drier meadows and pastures (Arrhenatherion and Cynosurion). In the lowlands of surrounding countries the species occurs in a similar range of vegetation types, with the same optimum. Since it is a winter annual species without a seedbank, fruit ripening, germination and seedling development should be successful every year to maintain a population. Mowing (once or twice) after 15 June is essential, while a relatively open sward in summer and autumn is critical for establishment, which can be guaranteed by aftermath grazing. Bromus racemosus requires a high groundwater table and/or river flooding during winter for a limited period at least every few years. Under these conditions the species competes successfully with nitrophilous grasses. Within its optimum, its cover is highest under relatively dry, nutrient-rich conditions. The species can establish in restored grasslands within a few years, but dispersal over larger distances is probably very limited. 306
Erweiterte deutsche Zusammenfassung Einleitung ­ Bromus racemosus L. ist eine ziemlich seltene Grasart der Feuchtwiesen. In den Niederlanden wird sie in zwei Unterarten unterteilt, subsp. racemosus und subsp. commutatus, die in anderen Lдndern meist als separate Arten aufgefasst werden (SPALTON 2002). Die vorliegende Arbeit befasst sich hauptsдchlich mit B. racemosus s.str. (= B. racemosus subsp. racemosus). Bromus racemosus gehцrt innerhalb der Gattung zu Bromus s.str., mit einjдhrigen, auf anthropogene Standorte beschrдnkte Arten (Anцkophyten). Das Areal von B. racemosus umfasst weite Teile Europas. In vielen Lдndern steht sie auf der Roten Liste, weil sie aufgrund der Intensivierung der Landwirtschaft in den letzten Jahrzehnten in starkem Rьckgang begriffen ist. Ihr winterannueller Lebenszyklus ist bemerkenswert fьr eine Sippe des Dauergrьnlands. Die Samen keimen sofort nach der Reifung, sobald sie feucht werden. Dies verhindert die Bildung einer Samenbank (LUTZ 1996, JENSEN 2004). Die vorliegende Studie soll das Wissen ьber die Habitat- und Managementprдferenz von B. racemosus in den Niederlanden und den benachbarten Gebieten vergrцЯern. Dazu wurden der Einfluss von abiotischen Bedingungen und der Grьnlandbewirtschaftung auf ihre Abundanz und ihre syntaxonomische Position untersucht. Untersuchungsgebiet ­ Die Studie berьcksichtigt 28 Naturschutzgebiete mit Bromus racemosus in den Niederlanden (Abb. 1 und Anhang E2). Die meisten Fundorte befinden sich in den Auen und Poldern der niederlдndischen Flusslandschaft. Der Boden wird dominiert von Mineralschichten, die wдhrend des Holozдns von Flьssen (einschlieЯlich Tide-Flьssen) oder vom Meer abgelagert wurden. An einigen Stellen wдchst B. racemosus auf torfigen Bцden. Daten aus den umliegenden Lдndern wurden analysiert und mit den Daten aus den Niederlanden verglichen. Es handelt sich um Aufnahmestellen im Bremer Raum, vier Naturschutzgebieten in Belgien, vier in Nordfrankreich und zwei in Sьdengland (Abb. 2). Sie liegen ausnahmslos im Tiefland, in der Nдhe eines Flusses oder Baches. Material und Methoden ­ Es wurde die Braun-Blanquet-Methode angewendet. An Wuchsstellen von Bromus racemosus wurden Vegetationsaufnahmen von 9 m2 gemacht. Der Aufbau der Boden- und Humusprofile wurde mit einem Erdbohrer und einem Brotmesser sichtbar gemacht. Fьr eine bodenchemische Analyse wurde fьr jede Vegetationsaufnahme eine integrierte Bodenprobe aus zehn Teilproben bis 20 cm Tiefe entnommen. Die Daten wurden in Juice mit Twinspan klassifiziert, und mit verschiednen Ordinationsmethoden in Canoco sowie mit univariaten Statistiken analysiert. Vegetationsaufnahmen aus der niederlдndischen Vegetationsdatenbank (SCHAMINЙE et al. 2012) und aus den umliegenden Lдndern wurden mit Vegetationsaufnahmen aus der eigenen Feldarbeit verglichen. Die Hydrologie wurde anhand hydromorpher Bodeneigenschaften und Online-Daten ьber Grundwasserschwankungen und Ьberschwemmungen beschrieben. Ergebnisse ­ Die Klassifizierung ergab elf Gruppen von Vegetationsaufnahmen (Tabelle 1, 2), die auch in der PCA sichtbar werden (Abb. 3a, b). Die erste Achse kann als Feuchtigkeitsgradient interpretiert werden, die zweite als pH-Gradient und die dritte als Nдhrstoffgradient. Alle Vegetationsaufnahmen sind in die Klasse Molinio-Arrhenatheretea einzuordnen. Sie verteilen sich ьber Gesellschaften des Alopecurion pratensis (synonym: Deschampsion cespitosae), Calthion palustris, Arrhenatherion elatioris und Cynosurion cristati, wobei die Art innerhalb des Calthion relativ trockene, im Arrhenatherion und Cynosurion die feuchteren Ausbildungen bevorzugt. Die meisten Vegetationsaufnahmen aus benachbarten Lдndern дhneln den niederlдndischen Aufnahmen und kцnnen den gleichen Verbдnden zugeordnet werden (Beilage S1, Abb. 4). In den meisten Gesellschaften kann Bromus racemosus eine hohe Deckung erreichen, aber gerade im Calthion erscheint die Deckung in allen untersuchten Gebieten relativ gering. Das Optimum der Art liegt eher im Alopecurion, insbesondere in mдЯig nдhrstoffreichen Basalgesellschaften. In Wiesen ohne Flussьberflutungen wurde B. racemosus an Standorten mit einem mittleren hцchsten Grundwasserspiegel von weniger als 40 cm unter der Bodenoberflдche gefunden. Die Art fehlt an Standorten, die jeden Winter ьber einen lдngeren Zeitraum ьberflutet werden und erreicht die hцchste 307
Deckung an relativ trockenen Standorten. An Standorten, die mindestens einmal in zehn Jahren von Flusswasser ьberflutet werden, wдchst die Art auch auf trockeneren Bцden mit einem mittleren hцchsten Grundwasserspiegel von mehr als 80 cm unter der Bodenoberflдche. Wдhrend B. racemosus die hцchste Deckung an relativ nдhrstoffreichen Standorten erreicht, kommt dieses Gras jedoch nicht an sehr nдhrstoffreichen Bцden vor. Es wдchst vor allem auf Bцden mit Mull als Humusform, gekennzeichnet durch eine gute Mineralisierung. Bromus racemosus hat auf Wiesen mit Nachweide hцhere Deckungen als auf ausschlieЯlich gemдhten Wiesen (Abb. 5). Die meisten Wiesen mit B. racemosus werden ab der zweiten Junihдlfte gemдht. Nur selten tritt die Art in Weiden auf. Diskussion ­ Bromus racemosus wдchst nur an Standorten mit feuchten Bedingungen im Winter oder kurzzeitiger Ьberflutung, die Lьcken in der Vegetation schafft. Wahrscheinlich wird B. racemosus von leichter Sommeraustrocknung in der Konkurrenz mit anderen Arten begьnstigt, weil die Samen im Ruhezustand ьberleben, bis feuchte Bedingungen auftreten. Die aktuelle hydrologische Amplitude ist geringer innerhalb als auЯerhalb der Deiche. Die Deckung innerhalb der Deiche ist aber durchschnittlich hцher als auЯerhalb. Ein konstanter Wasserhaushalt ermцglicht der Art, eine groЯe Population mit hohem Deckungsgrad aufzubauen. Wenn die Hydrologie sich дndert, ist das Risiko des Verschwindens im flachen Polder hцher als in Auen mit reichem Relief, wo die Population im Hцhengradient pendeln kann. In den Auen besteht aber die Gefahr der Ьberflutung in der Blьtezeit, wodurch die Samenproduktion verhindert wird. Das Fehlen von B. racemosus in sehr nдhrstoffreichen Wiesen ist wahrscheinlich der starken Konkurrenz zuzuschreiben; darьber hinaus werden diese durchweg zu frьh gemдht fьr ausreichenden Fruchtansatz der Art. Wird der Nдhrstoffreichtum von ehemaligem Wirtschaftsgrьnland durch Mahd ohne Dьngung verringert, so kann B. racemosus die Wiesen innerhalb weniger Jahre wiederbesiedeln. Auf lдngere Sicht erscheint eine niedrige Dьngergabe alle paar Jahre wohl vorteilhaft fьr B. racemosus. Das Fehlen einer Samenbank und eine geringe Ausbreitungsfдhigkeit gefдhrden die Art. Voraussetzung fьr die Erhaltung der Populationen ist erfolgreiche Samenreifung und Etablierung der Keimlinge in jedem Jahr. Eine Bewirtschaftung mit Mahd nach dem 15. Juni und Nachweide eignet sich am besten, da diese Nutzungsweise die Samenreifung zulдsst und anschlieЯend eine ausreichend lьckige Krautschicht erhдlt. Acknowledgements We are grateful for the permissions to visit the grassland reserves of Staatsbosbeheer, Zuid-Hollands Landschap, Utrechts Landschap, Geldersch Landschap and Vereniging Natuurmonumenten. Rein de Waal helped with the humus and soil description. Dick Kerkhof, Albert Corporaal and Piet Schipper gave excellent suggestions about study sites, the species ecology and existing datasets. Valuable data were received from Maud Raman (INBO, Belgium), Rob Geerts (PRI-WUR), and Gert Rosenthal. Supplements Supplement S1. Shortened synoptic table of all studied vegetation, grouped by country and syntaxa. Beilage S1. Gekьrzte Stetigkeitstabelle aller untersuchten Vegetation, gruppiert pro Land und Syntaxa. Supplement S2. Complete synoptic table of relevйs from own field work. Beilage S2. Komplette Stetigkeitstabelle der Vegetationsaufnahmen von eigener Feldarbeit. 308
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Simmelink et al.: Habitat and management preference of Bromus racemosus L. Supplement S1. Shortened synoptic table of all studied vegetation, grouped by country as columns (D = Germany, N = Netherlands, B = Belgium, F = France, E = England) and syntaxa as rows (sensu Schaminйe et al. 1996). Per combination of a country and a syntaxon, a unique subset of relevйs was used, shown in a framed section of the column. The number of relevйs per group is given above in italics (in total 412 relevйs are included). In the footnotes, given between [ ], the locations, the original names of the syntaxa and references are given; only the Dutch relevйs are mainly from own field work. The table contains percentage frequency values and average non-zero cover in % (as superscript). Beilage S1. Gekьrzte Stetigkeitstabelle aller untersuchten Vegetation, gruppiert pro Land als Spalten (D = Deutschland, N = Niederlande, B = Belgien, F = Frankreich, E = England) und Syntaxa als Reihen. Pro Kombination aus einem Land und einem Syntaxon wurde eine Selektion von Vegetationsaufnahmen verwendet; diese wird in einem gerahmten Abschnitt der Spalte gezeigt. Die Zahl der Vegetationsaufnahmen pro Kombination ist oben in kursiv angezeigt (insgesamt sind 412 Aufnahmen enthalten). In den FuЯnoten, angegeben zwischen [], werden die Standorte, die ursprьnglichen Namen der Syntaxa und die Referenzen beschrieben; nur die niederlдndischen Vegetationsaufnahmen sind hauptsдchlich von eigener Feldarbeit. Die Tabelle enthдlt prozentuale Frequenzwerte und mittlere nicht-null Deckungswerte in % (hochgestellt).
Country Calthion palustris Bromus racemosus Caltha palustris Lychnis flos-cuculi Lotus pedunculatus Dactylorhiza majalis Cirsium palustre Alopecurion pratensis Bromus racemosus Sanguisorba officinalis Fritillaria meleagris Lathyrus pratensis Silaum silaus Alopecurus pratensis Arrhenatherion elatioris Bromus racemosus Arrhenatherum elatius Dactylis glomerata Trisetum flavescens Tragopogon pratensis Peucedanum carvifolia Cynosurion cristatus Bromus racemosus Cynosurus cristatus Lolium perenne Bellis perennis Trifolium dubium Agrostis capillaris Basal communities MolinioArrhenatheretea Bromus racemosus Alopecurus pratensis Ranunculus repens Alopecurus geniculatus Lolium perenne Bromus hordeaceus
D
N
B
F
E
22 [1]
53 [3]
34 [8]
16 [11]
100 12 100 4 100 3 100 5
82 10 26 4 62 7 44 3
91 4 75 3 91 3 63 4
41 8 58 6 47 9 50 3
23 5 8 2 12 1 0
5 1 64 3 15 3 25 2
19 [4] 100 63 37 84 5 100
21 [9] 4 100 10 0 30 4 95 3 57 5 67
13 [12]
23 [16]
12 100 8 100 2
0
70 10
0
65 3
5 69 5 65 2
50
74 3
13 100 17 65 4
25 [5] 100 5 80 13 92 8 68 7 36 3 20 5
16 [13] 100 6 94 23 94 6 63 3 81 2 13 3
14 [6] 100 9 93 8 100 11 57 2 79 6 64 20
16 [14] 100 25 75 13 88 16 56 3 88 14 38 16
8 [2]
42 [7]
41 [10]
40 [15]
100 17 100 8 100 4 100 8
63 4 95 7 66 14 18 5
75 15 98 10 98 30 98 13
50 7 33 13 37 6 23 3
50 11 93 9 80 16 98 12
88 8 62 4 49 5 15 2
Origin of relevйs: 1 Senecioni-Brometum racemosi (Calthion palustris ), near Bremen, Lutz 1996 (relevйs from several authors, 1951-1996) 2 BC / Basalgesellschaft Molinio - Arrhenatheretea , near Bremen, Lutz 1996 (relevйs from several authors, 1951-1996, but most from Lutz 1996) 3 Communities 1, 2, 10, 11 (own field work) and eleven additional relevйs from the DNVD (Schaminйe et al. 2006) from 1936-1974, assigned to the Ranunculo-Senecionetum caricetosum paniceae 4 Communities 6, 7 (own field work) 5 Community 4 (own field work) 6 Community 3 (own field work) 7 Community 5 (own field work) 8 Snoekengracht near Leuven, Dijlevallei near Leuven & Bourgoyen near Gent (Flanders), Callebaut et al. (2007) & VLAVEDAT (Flemish Vegetation Databank; Vandenbussche & Hoffmann 2001); assigned to the Calthion palustris based on an analysis of the species composition. 9 Colchico-Brometum racemosi (Bromion racemosi , Molinietalia ), Femenne & Fagne (southern Wallonia), Sougnez & Limbourg 1963; assigned to the Alopecurion pratensis based on an analysis of the species composition. 10 Dijlevallei near Leuven & Bourgoyen near Gent (Flanders), Callebaut et al. (2007) & VLAVEDAT (Flemish Vegetation Databank; Vandenbussche & Hoffmann 2001); classified as basal communities based on the absence of character species on the alliance level. 11 Brometo-Senecietum (Bromion racemosi , Molinietalia ), Bassin de la Sambre, Gehu 1961 & Cirsio dissecti - Scorzoneretum , Bocage Virois, Basse-Normandie, De Foucault 1980; assigned to the Calthion palustris based on an analysis of the species composition; in the relevйs of De Foucault 1980 additionally some Junco-Molinion species are present. 12 Arrhenatheretum Colchicetosum (Cynosurion cristati ), Bassin de la Sambre, Gehu 1961; assigned to the Alopecurion pratensis based on an analysis of the species composition; species from the Cynosurion were rare and Arrhenatherion species not very frequent; several moisture indicating species were present. 13 Hordeo secalini-Arrhenatheretum elatioris & Galio veri-Trifolietum repentis (both Arrhenatherion ), Seine-maritime downstream of Rouen, Frileux et al. 1988 14 Oenantho peucedanifoliae - Brometum racemosi (Cynosurion cristati ), Bocage Virois, BasseNormandie, De Foucault 1980 15 Senecio aquatici-Oenanthetum mediae , Seine-maritime, Frileux et al. 1988 & Senecio-Brometum racemosi (Bromion racemosi , Agrostietalia-stoloniferae , Agrostio-Arrhenatheretea elatioris ), Boulonnais, De Foucault 1986; classified as basal communities based on the absence of character species on the alliance level. 16 Unpublished relevйs from A. Corporaal, near Cricklade, 1991, all with Fritillaria meleagris & from M. Raman, near Oxford, 2010; assigned to the Alopecurion pratensis based on an analysis of the species composition.
Simmelink et al.: Habitat and management preference of Bromus racemosus L. Supplement S2. Complete synoptic table of relevйs from own field work, with percentage frequency values. All relevйs are from the Netherlands and belong to the Molinio-Arrhenatheretea . The communities are described in Table 1 in the article. The assignment of species to the communities is according to fidelity, this is explained in supplement E1. Species that are diagnostic for one community are in dark grey, species that are diagnostic for a group of communities are in light grey. Layer: h: herb, m: moss. Beilage S2. Komplette Stetigkeitstabelle der Vegetationsaufnahmen von eigener Feldarbeit, mit prozentualen Stetigkeitswerten. Alle Vegetationsaufnahmen sind aus den Niederlanden und gehцren zu den Molinio-Arrhenatheretea . Die Gesellschaften sind in Tabelle 1 im Artikel beschrieben. Zuordnung von Arten zu den Gesellschaften entsprechend der Treue (s. Anhang E1). Fьr eine Gesellschaft diagnostische Arten sind dunkelgrau, fьr eine Gruppe von Gesellschaften diagnostische Arten hellgrau hervorgehoben. Schichten: h: Kraut, m: Moos. Communities / Pflanzengesellschaften: 1: Fragment of / Fragment des Rhinantho-Orchietum morionis 2: Rhinantho-Orchietum morionis 3: Lolio-Cynosuretum 4: Arrhenatheretum elatioris typicum 5: BC / Basalgesellschaft Bromus racemosus -[Alopecurion pratensis ] 6: Fritillario-Alopecuretum pratensis 7: Sanguisorbo-Silaetum 8: Transition between /Ьbergang zwischen Sanguisorbo-Silaetum & Calthion palustris 9: BC / Basalgesellschaft Rhinanthus angustifolius-Lysimachia vulgaris -[Calthion palustris ] 10: Angelico-Cirsietum oleracei 11: Ranunculo-Senecionetum aquatici juncetosum articulati
Community Number of relevйs
Bromus racemosus Bromus commutatus
1
Hydrocotyle vulgaris
Leontodon taraxacoides subsp. taraxacoides
Juncus articulatus
Juncus conglomeratus
Rhytidiadelphus squarrosus
2
Danthonia decumbens
Luzula campestris
Carex ovalis
Ophioglossum vulgatum
Hypochoeris radicata
Orchis morio
Dactylorhiza majalis subsp. praetermissa
Carex flacca
1-3 Odontites verna subsp. serotina Juncus gerardi Lotus tenuis Sagina procumbens Euphrasia stricta Cynosurus cristatus Agrostis capillaris Rhinanthus minor Carex nigra Triglochin maritima Ranunculus bulbosus Vulpia bromoides Ranunculus sardous Potentilla anserina Plantago major
3-5 Bromus hordeaceus Lolium perenne Cirsium arvense Hordeum secalinum Elymus repens
4
Dactylis glomerata
Achillea millefolium
Trisetum flavescens
Tragopogon pratensis subsp. pratensis
Equisetum arvense
Arrhenatherum elatius
Anthriscus sylvestris
Senecio jacobaea
Peucedanum carvifolia
Galium mollugo
Heracleum sphondylium
Potentilla reptans
Crepis biennis
Pimpinella major
Convolvulus arvensis
Eryngium campestre
Carex spicata
Medicago lupulina
6
Fritillaria meleagris
Poa palustris
Leontodon autumnalis
Ranunculus ficaria
Eleocharis palustris
Poa pratensis
Senecio aquaticus
Leontodon hispidus
6-7 Festuca pratensis Sanguisorba officinalis Stellaria palustris Centaurea jacea
7
Symphytum officinale
Achillea ptarmica
Mentha aquatica
Silaum silaus
6-10 Rumex crispus Thalictrum flavum Filipendula ulmaria Galium palustre Myosotis scorpioides Carex acutiformis Valeriana officinalis
6-11 Carex acuta Carex disticha Calliergonella cuspidata Equisetum palustre Lychnis flos-cuculi Lysimachia nummularia Cardamine pratensis Polygonum amphibium Lathyrus pratensis Phalaris arundinacea Agrostis canina Vicia cracca Caltha palustris Phragmites australis Carex riparia
9
Lysimachia vulgaris
10 Juncus acutiflorus Cirsium oleraceum Crepis paludosa Primula elatior Stellaria uliginosa Plagiomnium affine
11 Equisetum fluviatile Pedicularis palustris Lotus pedunculatus Cirsium palustre Glyceria fluitans Rhinanthus angustifolius Myosotis laxa subsp. caespitosa Ranunculus flammula Carex hirta Juncus effusus Prunella vulgaris
Companions Poa trivialis Cerastium fontanum subsp. vulgare Rumex acetosa Ranunculus acris Taraxacum officinale s.l. Ranunculus repens Brachythecium rutabulum Trifolium pratense Alopecurus pratensis Trifolium repens Plantago lanceolata Holcus lanatus Agrostis stolonifera Anthoxanthum odoratum Trifolium dubium Bellis perennis Phleum pratense subsp. pratense Festuca rubra Glechoma hederacea Veronica arvensis Kindbergia praelonga Alopecurus geniculatus Glyceria maxima Geranium dissectum Allium vineale Lotus corniculatus Cerastium glomeratum Veronica serpyllifolia Poa annua Fraxinus excelsior Leucanthemum vulgare x Festulolium loliaceum Urtica dioica Lythrum salicaria Stellaria graminea Galium aparine Juncus bufonius Galium verum Crataegus monogyna Festuca arundinacea Rumex obtusifolius Myosotis discolor Stellaria media Veronica chamaedrys Cardamine hirsuta Epilobium tetragonum Oenanthe fistulosa Drepanocladus aduncus Dactylorhiza incarnata Rorippa palustris Cirsium vulgare Carum carvi Lathyrus tuberosus Amblystegium serpens Vicia sepium Rorippa sylvestris Juncus tenuis Climacium dendroides
1
2
3
4
5
6
7
8
9
10 11
4
4
15
28
42
10
10
8
3
2
33
h 100 50
93
68 100 90 100 88
100 97
h
21
100
3
h 100
.
.
.
.
.
.
.
.
.
.
h 100 25
.
.
.
.
.
.
.
.
.
h
75
25
7
.
.
.
20
.
.
.
3
h 100 50
20
.
5
.
.
.
.
.
6
m 100 50
13
4
.
.
.
.
33
.
9
h
50
50
7
.
.
.
.
.
.
.
.
h
50 100 20
7
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Rare species in only one column (column/frequency): Herb layer (h): Acer campestre:5/2; Aegopodium podagraria: 5/3; Campanula rotundifolia: 4/4; Capsella bursa-pastoris: 3/7; Cardamine flexuosa: 10/50; Carex distans: 2/25; Carex oederi subsp. oedocarpa: 5/2; Carex otrubae 2/25; Chamomilla recutita: 3/7; Chamomilla suaveolens: 3/7; Dactylorhiza majalis: 3/7; Dactylorhiza majalis subsp. majalis; Daucus carota: 4/4; Deschampsia cespitosa: 5/5; Epilobium parviflorum: 10/50; Eupatorium cannabinum: 10/50; Filaginella uliginosa: 3/7; Galium boreale: 7/10; Iris pseudacorus: 7/20; Juncus compressus: 5/2; Juncus filiformis: 6/10; Juncus inflexus: 11/6; Knautia arvensis; Lotus species: 3/13; Matricaria maritima:5/2; Medicago sativa subsp. falcata: 4/4; Mentha arvensis: 5/2; Polygonum aviculare: 3/7; Polygonum persicaria: 3/7; Prunus spinosa: 4/4; Quercus robur:11/3; Rumex cripsus x obtusifolius: 5/2; Saxifraga granulata: 10/50; Scirpus sylvaticus: 7/20; Sparganium erectum: 10/50; Triglochin palustris: 7/20; Vicia hirsuta: 4/7; Vicia sativa: 4/7. Moss layer (m): Amblystegium varium: 11/3; Fissidens taxifolius: 9/33; Oxyrrhynchium hians: 5/2; Physcomitrium pyriforme: 11/3; Plagiomnium species: 7/10; Plagiomnium undulatum: 9/33.
Simmelink et al.: Habitat and management preference of Bromus racemosus L.
Supple me nt E1. Table of relevйs from own field work. All relevйs are from the Netherlands and belong to the Molinio-Arrhenatheretea . T he relevйs that were made before 2013 were also visited in 2013 to obtain soil data. T he relevйs were ordered by T winspan, this classification was improved by hand. T he communities are described in T able 1 in the article. All two relevйs of cluster 16 and one relevй of cluster 14 were removed from the table. Species were assigned to clusters according to fidelity, measured by the phi coefficient (using presence/absence data, within the whole dataset, and the size of all groups was standardised to equal size). Only species with a phi coefficient value of 25.0 or higher for one community or a group of communities were assigned to clusters. Species that had a slightly higher fidelity to small clusters of less than five relevйs than to a big cluster were assigned to the big cluster. Diagnostic species are arranged in an order that creates a diagonal structure (first criterion) and further they were ordered by strength of fidelity (second criterion). Companions were arranged by decreasing frequency. Species that are diagnostic for one community are in dark grey, species that are diagnostic for a group of communities are in light grey. Layer: h: herb, m: moss. Altitude (in meters): between -0.8 and 8.0, MK1&2: 14.6. Slope in degrees if >3 + exposition: BB2: 5, NW. WA6: 7, N. ABa: 10, N. WL7: 15, NNE. BB4: 28, SSE. Land use: G0: only grazing. G1: mown once and grazed afterwards. G2: mown twice and grazed after the second mowing. N1: mown once, not grazed. N2: mown twice, not grazed. Cover-values according to the Braun-Blanquet scale for cover-abundance adjusted by Barkman et al. (1964). Anhang E1. T abelle der Vegetationsaufnahmen von eigener Feldarbeit. Alle Vegetationsaufnahmen sind aus den Niederlanden und gehцren zu den Molinio-Arrhenatheretea . Die Vegetationsaufnahmen, die vor 2013 hergestellt wurden, wurden auch im Jahr 2013 besucht, um Bodendaten zu erhalten. Die Vegetationsaufnahmen wurden von T WINSPAN geordnet, diese Klassifizierung wurde von Hand verbessert. Die Pflanzengesellschaften werden beschrieben in T abelle 1 des Artikels. Alle zwei Vegetationsaufnahmen aus Gruppe 16 und eine der Gruppe 14 sind aus der T abelle entfernt. Arten wurden den Pflanzengesellschaften zugeordnet nach T reue, gemessen nach dem Phi-Koeffizienten (mit den folgenden Einstellungen: presence/absence data, within the whole dataset, size of all groups standardised to equal size). Nur Arten mit einem PhiKoeffizientenwert von 25,0 oder hцher fьr eine Pflanzengesellschaft oder eine Gruppe von Pflanzengesellschaften wurden Gruppen zugeordnet. Arten, die eine etwas hцhere T reue zu kleinen Gruppen von weniger als fьnf Vegetationsaufnahmen haben als zu einer groЯen Gruppe, wurden der groЯen Gruppe zugeordnet. Kennarten sind in einer Reihenfolge angeordnet, die eine diagonale Struktur erstellt (erstes Kriterium) und weiter sind sie nach Stдrke der T reue (zweites Kriterium) geordnet wurden. Begleitarten sind in absteigender Hдufigkeit angeordnet. Arten, die charakteristisch sind fьr eine Pflanzengesellschaft, sind in dunkelgrau abgebildet, Arten, die fьr eine Gruppe von Pflanzengesellschaft charakteristisch sind in hellgrau. Schicht: h: Kraut, m: Moos. Hцhe (in m): zwischen -0,8 und 8,0, MK1 & 2: 14.6. Hangneigung in Grad, wenn > 3 + Hangexposition: BB2: 5, NW. WA6: 7, N. ABa: 10, N. WL7: 15, NNO. BB4: 28, SSO. Bewirtschaftung: G0: nur Beweidung. G1: einmal gemдht mit Nachbeweidung. G2: zweimal gemдht mit Nachbeweidung nach der zweiten Mahd. N1: einmal gemдht, nicht beweidet. N2: zweimal gemдht, nicht beweidet. Deckungswerte entsprechend der angepassten Braun-Blanquet Skala fьr Abundanz/Dominanz nach Barkman et al. (1964). Namen der nummerierten Pflanzengesellschaften: siehe die Englische Ьberschrift. Communities / Pflanzengesellschaften: 1: Fragment of / Fragment des Rhinantho-Orchietum morionis 2: Rhinantho-Orchietum morionis 3: Lolio-Cynosuretum 4: Arrhenatheretum elatioris typicum 5: BC / Basalgesellschaft Bromus racemosus -[Alopecurion pratensis ] 6: Fritillario-Alopecuretum pratensis 7: Sanguisorbo-Silaetum 8: T ransition between /Ьbergang zwischen Sanguisorbo-Silaetum & Calthion palustris 9: BC / Basalgesellschaft Rhinanthus angustifolius-Lysimachia vulgaris -[Calthion palustris ] 10: Angelico-Cirsietum oleracei 11: Ranunculo-Senecionetum aquatici juncetosum articulati
Co mmun it y Relevй number
1
2
3
4
5
6
7
8
9
10
11
144 143 142 145 157 150 149 156 160 21 158 162 161 146 148 159 155 22 154 153 147 151 152 127 128 102 110 112 24 25 111 99 113 16 104 103 121 11 119 120 117 114 118 13 2 65 66 44 12 90 3 23 130 1 74 20 17 60 6 73 26 59 34 76 91 94 67 70 10 55 95 71 69 18 15 68 51 31 30 129 7 80 14 45 72 54 19 62 77 5 75 131 8 98 108 97 109 96 106 105 107 100 101 122 123 124 125 126 136 134 138 135 137 4 43 50 47 49 32 46 52 139 141 140 116 115 83 86 9 39 87 92 84 41 93 82 88 132 40 36 37 42 38 133 35 89 85 28 27 63 64 61 78 29 79 53 58 48 33
WA4 WA3 WA9 WA1 OH4 SB5 OH3 BG3 BG6 BG5 SB3 SB4 RG2 RG6 SC2 KB7 RG9 SC1 RG8 RG7 RG11 KB6 RG5 LA3 SC6 RG12 RG1 SC5 RG4 RG10 KBc RG3 LA4 MK1 MK2 KD2 KD3 KD1 WL2 WL6 WL7 WL5 WL9 WL8 DW1 AB1 GU4 GU2 GU5 GU1 GU3 HP5 HP4 HP3 HP2 HP1 GB5 GB4 BR4 BR2 BR3 BR1 BR6 GB1 BR5 GB2 COc KB5 PA8 SW1 OH2 BG4 AHa WA2 PA5 DW3 ABi LA1 COa KB3 DW2 WL4 WL1 PA1 COb AHf PA2 PA4 ZK2 WA5 KBb PA3 LL1 ZK1 SC4 OH1 WA6 BG1 SB2 PA6 SB1 BG2 AHe AHd PA7 AH1 KB4 AH2 ABc SC3 ABb WL3 MW2 MW1 ABd ABa BB2 CO8 BB1 BB4 BB3 ABf BB5 CO2 CO3 COd CO7 GB3 CO5 AB3 AB2 CO6 CO4 CO1 KB2 KB1 WB1 DB6 DB2 WB2 WB3 DBa WB4 WB8 DB3 DB1 WB10 WB11 WB7 WBa WB9 WB5 DB4 DB5 WB6 DH4 DH1 DH2 DH3
Relevй name T WINSPAN, modified, 19 clusters Year (20..) Plot size (m2) T otal cover (%) Cover herb layer (%) Cover moss layer (%) Land use Number of plant species
Bromus racemosus Bromus commutatus
1
Hydrocotyle vulgaris
Leontodon taraxacoides subsp. taraxacoides
Juncus articulatus
Juncus conglomeratus
Rhytidiadelphus squarrosus
2
Danthonia decumbens
Luzula campestris
Carex ovalis
Ophioglossum vulgatum
Hypochoeris radicata
Orchis morio
Dactylorhiza majalis subsp. praetermissa
Carex flacca
1-3 Odontites verna subsp. serotina Juncus gerardi Lotus tenuis Sagina procumbens Euphrasia stricta Cynosurus cristatus Agrostis capillaris Rhinanthus minor Carex nigra T riglochin maritima Ranunculus bulbosus Vulpia bromoides Ranunculus sardous Potentilla anserina Plantago major
3-5 Bromus hordeaceus Lolium perenne Cirsium arvense Hordeum secalinum Elymus repens
4
Dactylis glomerata
Achillea millefolium
T risetum flavescens
T ragopogon pratensis subsp. pratensis
Equisetum arvense
Arrhenatherum elatius
Anthriscus sylvestris
Senecio jacobaea
Peucedanum carvifolia
Galium mollugo
Heracleum sphondylium
Potentilla reptans
Crepis biennis
Pimpinella major
Convolvulus arvensis
Eryngium campestre
Carex spicata
Medicago lupulina
6
Fritillaria meleagris
Poa palustris
Leontodon autumnalis
Ranunculus ficaria
Eleocharis palustris
Poa pratensis
Senecio aquaticus
Leontodon hispidus
6-7 Festuca pratensis Sanguisorba officinalis Stellaria palustris Centaurea jacea
7
Symphytum officinale
Achillea ptarmica
Mentha aquatica
Silaum silaus
6-10 Rumex crispus T halictrum flavum Filipendula ulmaria Galium palustre Myosotis scorpioides Carex acutiformis Valeriana officinalis
6-11 Carex acuta Carex disticha Calliergonella cuspidata Equisetum palustre Lychnis flos-cuculi Lysimachia nummularia Cardamine pratensis Polygonum amphibium Lathyrus pratensis Phalaris arundinacea Agrostis canina Vicia cracca Caltha palustris Phragmites australis Carex riparia
9
Lysimachia vulgaris
10 Juncus acutiflorus Cirsium oleraceum Crepis paludosa Primula elatior Stellaria uliginosa Plagiomnium affine
11 Equisetum fluviatile Pedicularis palustris Lotus pedunculatus Cirsium palustre Glyceria fluitans Rhinanthus angustifolius Myosotis laxa subsp. caespitosa Ranunculus flammula Carex hirta Juncus effusus Prunella vulgaris
Companions Poa trivialis Cerastium fontanum subsp. vulgare Rumex acetosa Ranunculus acris T araxacum officinale s.l. Ranunculus repens Brachythecium rutabulum T rifolium pratense Alopecurus pratensis T rifolium repens Plantago lanceolata Holcus lanatus Agrostis stolonifera Anthoxanthum odoratum T rifolium dubium Bellis perennis Phleum pratense subsp. pratense Festuca rubra Glechoma hederacea Veronica arvensis Kindbergia praelonga Alopecurus geniculatus Glyceria maxima Geranium dissectum Allium vineale Lotus corniculatus Cerastium glomeratum Veronica serpyllifolia Poa annua Fraxinus excelsior Leucanthemum vulgare x Festulolium loliaceum Urtica dioica Lythrum salicaria Stellaria graminea Galium aparine Juncus bufonius Galium verum Crataegus monogyna Festuca arundinacea Rumex obtusifolius Myosotis discolor Stellaria media Veronica chamaedrys Cardamine hirsuta Epilobium tetragonum Oenanthe fistulosa Drepanocladus aduncus Dactylorhiza incarnata Lotus species Rorippa palustris Vicia sativa Vicia hirsuta Cirsium vulgare Carum carvi Lathyrus tuberosus Amblystegium serpens Vicia sepium Deschampsia cespitosa Rorippa sylvestris Juncus tenuis Scirpus sylvaticus T riglochin palustris Iris pseudacorus Climacium dendroides Juncus inflexus Carex distans Carex otrubae Dactylorhiza majalis Dactylorhiza majalis subsp. majalis Chamomilla recutita Polygonum persicaria Filaginella uliginosa Capsella bursa-pastoris Polygonum aviculare Chamomilla suaveolens Campanula rotundifolia Medicago sativa subsp. falcata Prunus spinosa Daucus carota Knautia arvensis Acer campestre Aegopodium podagraria Oxyrrhynchium hians Rumex cripsus x obtusifolius Matricaria maritima Carex oederi subsp. oedocarpa Mentha arvensis Juncus compressus Juncus filiformis Galium boreale Plagiomnium species Plagiomnium undulatum Fissidens taxifolius Sparganium erectum Eupatorium cannabinum Epilobium parviflorum Saxifraga granulata Cardamine flexuosa Quercus robur Physcomitrium pyriforme Amblystegium varium
15 15 15 15 15 1 1 1 2 2 2 2 2 2 2 2 3 3 3 3 3 3 3 4 4 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 6 6 6 6 6 6 6 6 7 7 7 7 7 7 7 7 7 7 7 8 8 8 8 8 8 8 8 8 8 8 8 9 10 10 10 10 10 10 10 10 10 10 10 18 18 18 18 18 18 12 12 12 12 12 12 13 13 13 13 6 14 14 14 14 14 14 14 14 14 14 6 14 14 14 14 6 14 14 13 13 13 11 11 13 17 17 17 17 17 17 17 17 17 17 17 17 17 17 17 17 17 17 17 17 19 19 19 19 19 19 19 19 19 19 19 19 13 13 13 13 13 13 13 13 13 7 13 13 13 13 13 13 13 9 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 9 13 13 13 8 13 13 13 13 13 8 8 13 13 13 8 13 8 13 13 13 13 10 10 13 13 13 13 13 13 13 13 13 13 13 7 13 13 13 13 10 9 13 13 13 13 13 7 13 8 13 13 13 10 13 13 13 13 13 7 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 7 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 9 9 9 9 9 9 9 9 9 25 9 9 9 9 9 9 9 25 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 4 9 9 9 16 9 9 9 9 9 16 16 9 9 9 16 9 16 9 9 9 9 25 25 9 9 9 9 9 9 9 9 9 9 9 4 9 9 9 9 25 4 9 9 9 9 9 25 9 16 9 9 9 24 9 9 9 9 9 25 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 4 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 10 95 95 92 95 93 90 95 90 97 100 95 95 95 95 95 98 95 100 98 92 85 98 85 90 90 97 95 95 95 95 95 90 97 95 95 93 90 100 95 90 95 93 90 100 100 90 88 90 100 97 100 98 87 92 93 100 100 95 92 92 80 95 95 90 90 93 92 90 0 95 90 90 93 100 100 95 85 90 90 93 99 85 100 99 92 100 97 93 80 88 95 93 99 93 95 95 90 90 90 80 90 95 93 97 95 90 90 90 95 93 95 95 95 93 80 95 95 95 95 95 95 95 92 93 95 95 90 90 0 95 92 95 92 97 95 93 85 95 95 98 95 90 95 90 98 80 90 93 85 85 95 90 85 90 85 82 97 80 90 85 90 75 90 87 90 95 80 97 100 95 95 95 95 95 98 95 100 98 92 85 98 85 90 90 95 95 95 95 95 95 90 97 95 95 93 90 100 95 90 95 93 90 100 100 88 87 90 100 97 100 98 87 90 93 100 100 95 90 92 80 92 95 80 88 93 90 90 0 95 90 88 93 100 100 95 85 90 90 93 99 85 100 99 92 100 97 92 70 85 95 93 99 90 95 95 90 90 85 70 85 92 92 97 95 90 90 90 95 93 95 95 95 93 80 95 95 95 90 90 95 85 90 93 70 80 80 90 0 95 90 95 92 97 93 85 85 95 95 98 95 90 95 88 95 75 90 87 80 80 95 87 80 85 80 82 97 80 80 50 50 70 45 60 0 1 60 0 1 15 0 7 0 0 0 1 0 2 0 0 2 0 3 5 20 3 1 7 1 2 3 1 0 3 7 4 0 1 3 1 10 3 0 1 10 10 2 1 2 0 3 0 4 10 1 5 10 3 3 0 30 2 45 20 0 2 1 0 7 3 7 7 0 0 3 1 2 1 1 1 3 5 10 2 1 5 3 45 3 25 3 1 7 3 7 10 10 50 60 35 20 7 3 5 10 10 1 3 3 2 3 1 3 1 1 0 5 10 10 1 70 10 1 80 70 50 10 0 1 5 10 10 1 30 50 3 7 10 0 3 10 0 7 15 15 10 20 35 10 7 20 50 40 40 2 10 0 10 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 N2 N2 G1 G1 G1 G1 G1 G1 N2 G1 G1 G1 G1 G0 G1 G0 G0 G0 G1 G0 G1 G1 G1 G1 N1 G1 G1 G1 G2 N2 G2 G0 G2 G2 G2 G1 G2 G1 G2 G1 G2 G1 N2 G1 G2 N2 G1 N2 G2 G2 G2 G1 G2 G1 G1 G1 N2 G1 N1 G1 G1 G2 G1 G2 G1 G2 G1 G0 N2 G1 N2 N2 N2 N2 N2 N2 N2 N2 N2 N2 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 N1 N1 N1 N1 N1 N1 G2 N2 G2 G2 G1 G2 G2 G1 N1 G2 N2 G2 G2 G2 G1 G2 N2 G1 G2 G2 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 34 31 31 34 46 29 33 20 27 29 34 26 32 23 26 24 37 30 25 32 31 30 26 36 40 32 30 27 23 24 30 31 33 17 35 28 28 28 34 34 32 40 32 31 30 45 44 35 33 37 31 21 20 15 31 12 20 20 25 24 17 22 34 28 30 19 27 24 35 24 21 25 27 15 24 33 27 28 28 23 17 36 31 23 26 23 26 31 41 25 32 26 22 32 37 36 41 40 40 38 44 32 35 43 44 35 43 29 41 41 39 43 40 34 25 29 35 34 35 29 29 44 41 36 43 53 42 34 24 26 33 45 34 37 37 40 35 40 37 30 30 33 30 42 25 27 33 36 40 34 36 37 27 36 33 30 35 32 28 h 1 1 1 1 + . 2a . 2b 2a 2a . 2b 2m 2m 2b 2m 1 2b 2a + 2a 1 . . 1 2m . 2m + 2a + 2a 1 1 . . 2m . . . 2m . 2m 2m 1 1 + 2m 2m 2a 3 + 2a 2m 2a 2b 2a 3 2a 1 2a 1 2a 1 2m + 2m 2m 2m + 2a 2a 2b 2a 3 2m 2m 2a 2a 1 1 1 2m 2b 2m 1 2a 2m 2m 2m 2a 2m 1 + 2a . + + 2m + 2b 1 1 2m + 1 + + + 1 + 1 3 2a + 2m 1 1 . 2m . . . + + + 1 1 1 1 2m 2m 2m 1 1 1 . 1 1 2m 2m + + 2m + 2a 1 1 2a 2m 2m 2m + 1 1 2a + 1 h . . . . . . . . . . . . . . . . . . . . . . . 2b + . . . . . . . . . . . . . 2a 2a 2m . 2a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + 1 2m . . . . . . . . . . . . . 2a . . . . . . . . . . . . . . . . . . . . . h1111 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h 2a 2m 1 + 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h 1 1 1 . 2a . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . + . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . h 2a 1 2a 2b r . . + . . . . . . . . + . r 2a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . r . . . . . . . . . . . . . . . m 2b + 2a 1 2b . . 4 . . 1 + . . . . . . . . . . . . 2m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . 2m . 2a . . . . h 1 . 1 . + . . 2b . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h 1 . . + 1 2a 1 2a . . . . + . 1 . + . . . . . . . + . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . h+ . . . + . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . 1 2a 1 r . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . r 1 2a + 1 1 . + 2m . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . .11r+ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . +1 r . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . 2a 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . 2m 2a 1 . . . . + . + . . . . 2m . 1 . r + . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h+1 . . . . . . . . . . . . . . r . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . 2b 1 . 2m . . . 2a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . +++ . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . + + . 2a . . . . . 1 . . . . . . + . 1 . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h 2a 1 1 2a 2a 1 2a . . 2a 2a 2a 2a 2a 2a 1 2b 2a 1 2a + 2a 2a 1 2a . . . . . . . . . . . . . . . . . . . . . . . . 1 . . + . . . . . . . . . 2b . + . . 1 1 2a . 2a 1 . . 1 . . . 1 . + . . . . . 1 . . . 2a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2a 2m . . 2a 1 2a 1 2m 1 2a 1 + 2a 2m 1 2a 2a + 1 2a 1 . 1 . 2a . + . . 2a 1 1 h 2a 2b . 2b 1 3 2b 2b . 2m 2b 2b 2b 2b 3 . 2a . 3 2b . . 2b . 2a . . . . . . + . . . . . . . . . 2a . . . . . . . . . . . . . . . . . . . . 1 1 . . . . . 1 . . . 2b . . . . . . . . . . . 1 . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . 2a . . . . 2a 2m . + . . . . . . . . . . . 2a . . . . h . + + 2a + 2a 2b 1 + 1 2b 2b 2b 2a 2b 2a 2b . 2a . + 1 . . . 2m 2m 2m . . . . 1 . 2a 2m . 2a . . . 1 . + + . . . 2m . 2m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . r . . . . . . 2a . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . 1 . . + . . 2b . . . . . . . . 2a . . 2a . . 2b . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . + . . . . . . . . . . . . . . . . 2b . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . r . . + . . 2m + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . 1 + . . 2m + 2m 1 . 1 . . 2m . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . + . . . . . . . + . . . . . . + . ++ . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . + . . 1 2a . 2m . . . . . . . . . . . . . . . . . . . . . . 1 1 . . . . . . . + . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . 2m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . . . . . .1 . . . . . .1 .+. .+. . . . . . . . . . . . . . . . . .+. . . . . . . . . . . . . .1 . . . . .1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1 . . . . . . . . . . . . . .r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . r + . 1 1 r 1 1 + . 1 + 2m 1 . . 1 . 1 1 1 + 2m 2b 2b 2m 2m 1 1 + 1 + 2m . + . 2m . . . + . 1 . + . 1 . . . 2a . . 2a 1 . . 2b . + . + 2a . + + + + . + 1 1 + 1 . 2a + 1 1 2m + 2m 1 + . . . . . . . . . . . + + 1 . . . . . . . . . . . + . . . + . . . . r . . . . . . . + . + . . . . . . . . . . . . . . 1 . + . . . . . . . . h 1 . + + 1 . 2a . 3 2m 1 2a 2b 2a 2a 3 2a 2m 2a 1 2a 1 + . 1 2a 2a 2a 2b 2a 2a 2a 2a + 2a 2a 2a 2a 1 2a 1 2m 2a 2a 2b 2m 2m 2m 2a 2a 2a 2a + 1 2b 2b 2a 2b 2a 2a 1 2a 1 2a 2a . 2m 1 1 2a . 2m 2a 2b 2a 2a 2a 2b 2a 1 2b 2b 2m 2b 2a 2a 2m 2b 2a 2a 2a 2b . . . . . . . . + 2a 2a + + . . . . . . . . 1 2a . 1 2m 2m . 2a . + 1 . . . 1 + . 1 2m 1 1 1 . + 2b 1 1 + + 1 1 . . 2m 1 2m 2m 1 2a 1 1 2a 2m 1 . . h . . . . . . . . 1 2a . . + 1 + . + + . . r . . r . + . . + r . . + . 1 + + 1 1 1 2a . + . + 2a 2m . + . . . . . 2a . . . . . . . + . . . 1 . r . . . . r . . 1 + 2a . . + r + 1 . . . . . + . . . . . . . . . . . . . . . r . . + r + . . . . . . r . + . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . + . + . . . . . . . . 2a 2a 1 . . . . . . + . 1 . . . 2b . . . 1 . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . + . . . . 3 . . . . . . . . . . . 2a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . 1 + + 1 . . + . . + + + . + . . + + 1 1 1 1 . + . + . 2a . 1 1 2b + 1 . . 1 2m 2a . 1 . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . + . + + . + . . . . . . . + 2a + . . . . . . . + . . . . . + . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . 1 1 2a 2a 2a + 2a 2a + 1 1 2a 2a 1 2m 2b 2a 2b 2a 2b 2a 2m 1 2a . 1 2b . . . . . . . . . . . . r . 2a r . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . . r . . . . . . . . . . . . . . . . . + . . 1 2a 2a . . . . . . . 2a . . 1 . . + . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . + 1 . 1 1 1 2a r . + . . . + 1 . 1 . . . 2m 2m . . + 2m . 1 . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . 2b 2b 2b . 2b . + + . + . 2a 2a 3 2m . 2b 2m . + 2m 2m + + . 2m . 2m . . . . . . . . . . . 2b . . . . . 1 . . . . . . . . . . . . . 2m . . . . . . . . . . . . . . . . . . . . + + . . . . . . . . + . . . . . . . . . . . . r . . . . . . . . . . 2m . . . . . 1 . . . . . + + + . . . . . . . h . . . . . . . . . . . + . . . . . . . . . . . . . + 2a + . . r . 2a . + 1 . r . . . 1 . + . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + 1 2b 2a 2a + + + 2a 2m 2a . . . 2a . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . 2b 2b 2a 1 2b 1 1 1 2b . 3 2b 3 . 2b 2a . 1 + . 3 2a 2a 2b . 2a 2b . . . . . . . . . . . . + . 1 . 1 . + . . . . . . . . 2a . . . 1 . . . . . . . . . . . . . . . . 2a . 2b . 1 . . . . . . . . . . . . . . . . . . 2b 3 + . + . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . + . . . . . . . . . . . . . 1 + . . . . + . r 1 + . + r . . . . . . . . . . + . . . . . . . . . . . . . . r 2b . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . 1 2a . . . . . . . . . . . 1 r . + r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . 2a . . . . 1 . . 2a 1 2a . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . . + 2a 2m . 1 . . 2m 1 . . . 2a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . 2a . . . . . . . . . . . . . 2a + . . . . 2b 2a 2a 2a . 2a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . 2a . . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . 1 1 . + . . . + . 1 . 1 . . . 2m . 2a . 2a . . . 2m 2a . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . 2a . . . . . . . . . . . . . + . . . . . 1 + . . . + r . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . + r . . . . . . . 1 . + 1 . + 2a 2a 2m 2m 1 1 . 1 . . . . . . . . . . . . . . . + . . . . . . . . . + + 1 . . + + + . . . . . . . . . . . . . . . . . . . 1 1 . . + . . . . . + . . r + r . + 2m . . . . . . . . . . . . . + . 1 . . . . . . . . . . . . . . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1 . .+. . . . . . .1 .+. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .+. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . 2a . . 2a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . .r . . . . . . . .+. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 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. r 1 + 1 2m 1 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b 1 + 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . 1 + + 1 + . + . + . + + . 1 + + . + + . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . 1 1 2a 1 1 + 1 1 2m + . . + + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . 1 . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . 2a . 1 2m 1 2m 1 + 2m . . . . . . . . . . . . . . . + . 1 . . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . 2m r + r 2m 1 . . . . . . . . + . . . . 2m + . + . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . r . . . . . . . . h . 1 . . + . . . 2a . 1 . 1 . . + . . . . . . . + . . + . . . 1 + 1 . . . . . . + . . . . . 1 1 + 2a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . r . r 1 2a 1 1 2a . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1 .1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 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. . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 + 2m . + . . . . . . . 1 + . 2m . + 2m + . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . 2a 1 2a 2a . 1 + 1 . . . . . . . . . . . . . . . . . 2b . . . . . . . . . . . . . 2a . . . . . . . . . . . . . . . + + 2a + 1 1 2a . 1 + . + 3 + 1 . . . . . . . . 1 . . . . . + + . 2a . 1 . 2b . 2m 2b . . . + . 2a . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + 1 + . . r . . . 1 . . . . . . . . . . + . . r . . . . . . . . . . . . . . . . . . . . . . . . . r . . r . . + . . + . + + + 2a 1 . 2a 2a . + + . . . . . + + + . . . . . . . . . . . . . . . r . . . r . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ++ . 1+++ . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . + . + . . . . . . . . 1 . + 2a r + 2a . . . . + . . . . . . . . 1 . 1 + . . . . . . . . . . . . . . . . . . . . . 2m . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . r . . . . . . . r . . . . . . + . + . + . . . . + . . . . r r + . . . . . . + . . . . . . . . . . r . r . . . . . . . . . . . . . . . + . . . . . 1 r + + 2a r 1 r r r . + . . . . . . 2b . 1 . + . . . . . . . r + + + + + . r r + . + + + r + 1 . . + . . r . . . . r . . . r . + . . + r r . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . 1 + . 1 1 . + . . . + . . . + . 2a . + . . . . . . . . . . . . . . . . . 2b . . 4 . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . + + . . . . + + + 2a 2a 3 . 1 + 1 . . r 1 + + . . 1 . . 2b 2b . . . . . + . . . . . . . . . + . + . . . . 1 + . . . + . . . . . h + 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . + . + . . . . 2m 2m 1 2m 1 . + . . . . . 2m 2m 2m 1 + + 2m 1 . . + + . . 1 . . . . 1 2m . 1 . . + . . . . . . . 1 . . . . . . + + . . . + + 1 + . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . 2m 2m 2m 2m . . . r . . . . + . . 1 1 + 2m 1 . . . . . . . . . . . 2m 2a r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2a 2a 3 2a . 2a 2b 2a 1 2b . . . . . . . . . . . 2b 2b . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . r . . . . . . . . ++ . + . . . . . . . . . . 111+1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b 2a 2a 2a 1 2a 2b 2a . . + + . 2a 3 . . . . . 3 2a 3 1 2m 1 2b 1 + . . 1 1 2a . 3 2a 2a + 2b 3 2a 2b 2a + . . 1 2b 1 2b 2b . 2b 2b 1 2m 4 2b 3 2a 1 2a 3 2m 1 h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2a 1 + . 2b 2a . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . 1 1 2b 2a 2a 1 1 1 . 2b 1 2b 2b 2a 2a 2a 2b 2b 2b 3 2b 1 + 2a 2b . 2b . 2m 1 2a + + 1 1 2m 2a 3 . . 2m 2a . 2a 1 . 3 . 3 + . 2b . . . . . . . . . . . 1 2a + + m 3 1 2b 3 2a . 1 . . . 1 . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . 1 . . . . . 2m . . . . . . . . . . . . . . . . . + . . . . . . . . . . . + . . 2m . . . . . . 2m . . 2b 1 2m 2m 2m . 2a 2m 3 3 3 1 . 2m 2m 2a 2a 1 2m 2m 1 2m 1 . + 1 2m 1 2a 2m + 3 2m 1 4 4 3 2a . 2m . 2m 2a 2m 2a 3 1 1 2a . 2m + + 1 2a 2a 2a 2b 2b 2a 2a 2a 2b 3 2b . . 2m 1 h . . . . . . . . . . . . . . . . . . . . . . . 2a + . . . . . . . . . . . . . . . . . + . . . . . . 2a . . 2b . + . . . . . . . + . 2b . . . + + . . . . . . 1 1 + 1 . 1 2m . . + . + 1 . + + + . 1 . + . + + 1 . . + 1 2m 2m 1 + + + 1 + . + + + 1 + + + + 1 + 2m 2m 1 1 r 2m 1 2a . 1 2a 1 1 + + 2a + . 2m + + 1 . 2b 2a 2m 1 1 + 2b . 1 1 1 1 h + 1 + . 1 1 1 . . . . . . . . . 1 + . 1 . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b . . . . . . . 1 . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + 1 + 2a . + . + . + . + r . 1 + 1 + + 1 . + 1 + . 1 . + + . 1 + + 2m 1 . 1 + 2m . . 1 2m . + . . + . 2m 1 . 1 2a 2m + 1 1 . 2m + . . 1 1 h . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . + . 1 . r . . . . . r 1 r + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . 1 2m + 1 1 + + . . . + 1 1 2a + + + + + . . . . r . . + . + 1 . . r + 1 . 1 1 1 + + . . 1 . 2a . 1 . . . . + 2m . + . 1 . . . 1 . . . . h 1 . . + 1 1 1 . . + 1 1 1 . 1 . 1 . 2m 1 . . 2m 2m + 1 1 . . . 1 1 1 . 1 1 . . 1 . 1 1 1 . 1 r + + . 2m 2m + 1 + + . 2m . . 1 + 1 1 2m 1 1 + 1 r . 1 1 1 2m 1 1 1 2m + 1 . + . r 1 . 2m + 1 2m 1 1 1 1 1 2m 2m 1 1 2m 1 1 1 1 1 + 1 + 1 2m 1 2m 1 1 1 1 + + 1 2m 2m + 1 1 + 1 1 1 + + 1 1 + 1 + 1 1 1 1 1 1 1 1 1 1 2m 2m + 1 1 1 2m 1 1 1 1 1 1 1 h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . r . . 1 + + . . 2m . . . + . . . . 1 . 1 . 1 1 . r + r . . + . . . + 1 1 1 r . . 1 + 1 . . + + + r + . . 2a 1 1 1 + 1 . + . + + + 1 1 1 1 + 1 1 1 . . . + + 1 1 r 1 . . . + + . 1 . 1 . 1 + 1 + + 1 + + + . + . 1 r . . . + 1 . . + . 1 . . h . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . 2a . 3 . . . . . 2m 1 . . . + + 2m 2m 2a 1 1 1 . . . . . . . . . . . 2a . . . . . + . . . . . 2a 1 + + 1 . . . + + . . . . . . . . . . + . . . 1 2m 1 2m 1 1 1 1 1 2a 1 2a 1 2a 1 2m . + 1 1 + 1 . + 1 1 + + 1 . . + . 1 . + . 1 . + + + . . . . . + . + 1 . . . . . . . 1 . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . 1 . + . . 2b . 1 . 2b . 2a . + 1 2b 2b . 2a 1 . . . 1 . . . . . + + . . . . 1 1 1 1 . + + . r 1 . . . + . + . . . . . 1 + . + 1 . 2a + . + . . . . 2a + . . 2a 1 2a 2a + + . 2b . 1 . + 2b . 2a . + + . + + . + + 1 . 1 . . 1 2a . . h 2a 1 2b 1 . . . 2a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . 2m 2a + 2a 1 + . + . . . . . . . . . . . . . . . . . . . . . . . . . . 2m . . 2m . . 1 . . 2a . 2a . . . . 2b . 2b . + 2a . . . 2a 1 2a . . . 1 h + . . 2a . + + . . + . + 1 . 1 . + . . + . 1 2m + + . . . . . . . 2b . . . . . . . . + . 1 + + 1 + . + 1 . . . . . . . . . . . . . 1 . . . r . . . . . . . + + . . . . + . . . . . . . . . . . . . r 1 . . . . 2m 2a 1 . 1 2a 1 . 1 + + + . . + . + + 2a 1 + 1 + + . + r 2a 1 1 r 1 1 + 1 + 2a 2a 3 + + 2m . 2m 2a . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 + . 2a . . . . . . . . . . . . . . . . . . 1 r . . 2a . . . . 1 + . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . r . . . h 2b 2a 2a 2a 1 . 2a r . . . . . . . . 1 . 1 1 . 1 . . . . . . . . . . . . . . . . . . . . . . . 1 1 . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . 1 . . . . . . . . . . . . 2a . . 1 1 1 . . . 1 1 1 1 1 . . 2a . . . + 2a 2m 1 2a 2a 1 1 1 + 1 2a 2a . . . . . . . . 2a . . . . . . . . . . . r + . . . . . . . 2b . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .r1 . . . . .+. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .+. . . .r .+. . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2a + 2b . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . 2a + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 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. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .11 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b 2a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . + . . . . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2m 1 . . . . . . . . . . . . . . . . . . . . . . 1 1 1 + . . . + . . + h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . 2a . . r . . 2b 2a 1 1 . . 2a . + + 2a 1 h . . . . . . . . . . . . . . . . 1 . . . . . . + 1 . . . . . . . . . . . . . . . . . . . + 1 . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . 1 . 1 1 2a . . . . . . . . . . . . . . . . . 1 + . + 2b . . . . . . . . + 1 . 1 1 1 . r 2m 1 . 1 2a + 2a 1 + 2a 2b 2b + 3 1 1 2a 1 . . . . 1 . . . + . . . h 2b + + 2a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . r . . 1 . . . . . . r . . . . . . r . . . . . . . . . . . . 1 + 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . + . . + . 1 + . . + . + . . + + . . . + + 1 + + r 1 2a + . 1 r 1 h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b 1 + . . . 2m 2a 3 . . . . . . . . . . + + . . 1 . . . . . . . . + . . 1 1 1 r . . + . . 2a . . . . . . . . . . . . . . . . . . . . . . + . . . . + . . . . . . + . + . . . . 1 . r . . + . . . + 1 2m . 1 + 1 + 2a 1 . . h . . . . 2b + + . 2a 1 1 . . 1 + 2a . 3 . + 2b 1 1 . . . . . . . . . . . . . . . . . . . . . . r . . . 2a . . . . . . . . . . . . + 1 2b . . . . . . . 2b . . + . . r . . 1 . + . 2b . 1 . 2b . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . 2b 2a 3 . + 2b . . . . 2b 2b 2a 3 2a . . + 2b + 1 + . . + 2a 3 2a 2a 1 1 2a 2a 2b 2a + 2b 2b h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . + r . . . . . . . . . . r + . . + . . . . . 1 . r + . . r . 1 + 2m . . . . 2m . 1 . 1 1 h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .+ . . . . . . .1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1+ . . . .+ . .1 .1 . r . .1 .1 . . . . . . .+ .1 .+ . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b . 2a 3 . . 2a 1 . . . . . . . 3 . . . . . 1 . + . + . + 1 . . . 1 1 . . 1 . . . . . 1 . . + + + 2a + . 2b . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . 1 . . 1 . 1 . + 1 1 . 2b + + . . + 1 r r . 1 . 2a 1 2m . . 2a . . . . h 1 1 + 2a . . . + . . . . . . . . + . r r . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . r . . . + . . . + . . . . . . . + . 2a . 1 . 2b . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . 1 r + 1 + . 2a r r 2m + 2a 2b 1 . 2a 1 . 2b . + 1 1 . 1 1 2a . + + 1 2a h . . . . 2a . . . . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 1 1 . . 1 + . 1 . . . . . . . . . . . . . . 2a . . . . + 2b + . 1 . 2a . . r 2m + . . . 1 r + 2m . 2a 1 + . . . . 2m 1 . + . . h + 1 + 2a 1 + 2a . 3 2m 2a 2a 1 2b 2a 3 2b 2a 3 2a 2a 2a 2m 1 1 2b 2b 2b 2b 3 2b 2b 2a 2b 2b 3 2a 2a 2a 2m 2a 2b 2a 2a 2a 2a 2a 2a . 2a 2a 3 1 3 2a 2a 2b 3 3 2a 2b 3 1 2a 2b 2b 2a 2a 1 2m 2a 2a 2a 2m 2a 3 2a 3 3 2a 2b 2b 2m 2m 2b 1 2m 3 2b 3 2b 2a 2a 2a 2b 2b 2a + . . . 2a 2a 2m 2m 2b 2a 2a 2b 2b 2a 2b 2b 2b 3 3 3 2b 3 2b 2b . 1 2a 2m 2a 2a . 1 2m 1 2b 2a 2b 1 2a 1 2a 2b 3 2a 2a 3 1 2b 2a 2a 2a 2b 3 2b 3 2b 2a 1 3 2b 2a 2a h 1 . + + 1 1 1 . 1 + . 1 1 1 1 2m 1 2m 1 + 1 1 . . + + + 1 1 1 + 1 + . 1 1 + 2m + + 1 1 1 2m 2m 2m 2m + 2m 1 . 1 + 1 2m 1 1 1 2m 1 1 2m + 1 2m 1 2m 1 . 1 2m 1 1 1 + 1 + 1 1 1 . + 1 + 1 + 1 2m 1 1 1 1 . . . + + + 1 1 1 1 1 + 1 + + . + 1 + 1 + + + 1 2m 2m 1 1 1 + + + + 1 + . . + 1 1 1 + 1 + . 1 . + . + + + 1 . 1 1 2a 2m 1 2m 1 2m 1 + 1 + 1 h 1 . . 1 1 1 2m + . 2m 1 2a 1 2m 2m . 2m 2m 2a + 2m 1 1 1 1 1 1 1 . . 2a 1 2a 1 2m 2a 1 . 1 . + 1 1 r . + 1 . 1 2m + 1 . 1 . 2b 1 . 2m 1 . . 1 2m 2m 2a . 1 + 1 1 1 1 2m . 2a . + 1 + + 1 1 1 2a 1 1 1 2m 2m . 1 . + 1 1 1 1 1 1 1 1 1 1 1 1 1 + 1 1 1 1 1 1 . + 1 1 + 1 2m + 1 + . + 1 + . 2a 2a 2m 1 1 1 1 2m 1 1 1 1 1 + 1 2m 1 1 1 2a 2m 2m 2m 2a 2m 2m + 1 1 2a h 1 + 1 1 2m 1 1 . . + 1 . + 2m 1 . 2a + 1 1 1 1 + + + 2a 2a 2a . . 2b 1 2a 2a 2a 2a + 2a + + 1 2a 1 1 1 2m 2m 1 2a + 2m + . . . . 1 . 2a 2a . . 2a 1 1 1 1 1 2a 2b 1 2a 2a 2b . + 2a 2a + + 2a 1 + 2b 1 2b . 2a 1 2a . 2a . + 1 1 + 2a 2m 2m 2m 2a 2a 1 1 . 1 + + 1 + + + 2a . + 1 + 2a . + + + + + + 1 1 r 1 1 + 2a + 2a 1 2a + + 2m . 2a 1 1 . . + 1 2a 2a 2a 2a 1 2a 2a . 2a 1 1 h + . . . 2m 1 1 . 1 2a + 2m + 1 . 1 1 . 2a . . . . + . 2a 2b 2a 1 + 2a 2a 2a 2a 2a 2a 2a . 1 1 2m 2a 1 1 . 1 1 1 2m 2a 2m 2m . 1 + 2a 1 + + 1 1 1 1 1 2m 2a 1 1 r 1 1 1 1 1 1 2m 1 + 1 + 1 1 + 1 1 1 1 1 1 + . + + . + 1 . + 1 1 1 1 1 1 1 . + . + + + + + 1 + + 1 1 1 + 1 1 + 1 . + + 1 . . + 1 1 + 1 + + + + . . + + r . . 1 . 1 . + 1 1 + 2m . 2m + . h . + + + . . . . . 2a 2a . . . 1 2b 2a . 1 . . . . . . 1 1 . 1 1 . . 1 . + + . . 2m . 1 1 2a 1 1 1 1 + 1 + 2a 2b 1 2b 2a 2b 2b + 2a 2b 2b 2b + 2a + . 2a 2a r 2a 2a 2a 2a 2a 3 2a 1 2a 2a 2a 2b 1 2b 2a 2a 2a 1 2a 2a 2b 2a 2a 2b 2a 2a + 1 1 . + . 2a 1 + + + + 1 + + 1 + 1 1 1 1 2b 3 2a 2b 2b . . . + + 1 2a 2a 1 2a 2b 1 1 . 1 2a 2a 1 2a 2a . + 1 + 1 2a 1 2a 2a 2a 2a 1 2a 2a 2a 1 2b 2a m . . . 2b 2b 1 1 . + 1 2a 1 2a + . . 1 . 2m . . 1 . 2m 2m 2b 2m 1 2a 1 2m 2m 1 . 2m 2a 2m . + 2m 1 1 2m . 1 2a 2a + 1 2m . 2m . 1 2a 2m 2a 2a 2m 2m . 3 2m 3 2b . 1 1 + 2a 2m 2a 2a . . 2m 1 2m 2m 1 . 2m 2a 2a 2m + 2a 2m 3 2m 2a 2m . 2m 1 2a . 2a 2a 2b 1 2b 2a . 2m . . . + . . . 1 2m 1 . + 2m 2m + . + 2m + . . 1 . . 2m 2a 2m + + 2a 2m 2m 2a . + 1 2a + 2m 2a 1 1 + 2b 2a 2m 2a 3 2a 2m 2m 2a 1 2a h . . . . 2a 1 1 + . r . 2a 1 + 1 2a 2m 1 . + . 2a + 1 2a 2a 2m 2b 2a 2a 1 2m 1 3 2a 2a 1 2a + + + 2a + 2m 2a 1 2a 2b 2a 1 2a . . . 2a . . + . 2a . + 1 . 2m . 1 . + 1 + 2a 2a + 1 1 2a 2a 1 . 2b 1 2a 3 1 3 . 2b + 1 2a . 1 1 . 2m . + 1 + + 1 2a 1 1 . . . + + . 1 + 2a 1 1 1 + 1 1 + 1 + . . . + + . . + 2m 2a . 2m 1 1 2a 1 + . + . + . . + 2m 1 1 1 1 . + 1 1 + + . h . . . . . . . . . . . . . . . . . . . . . . . 1 . 1 2a + . . 2a 1 1 1 1 1 . . 1 + + + 1 . . 1 + 2a . 2a 1 2a + 1 1 1 2a 1 1 2a + . 1 2a 1 2b 2a 2a 2m 3 2b 1 1 1 2m 2a 1 1 1 + 1 2a 2m 1 2b 1 . 2m 2a 2a 1 2b 1 2a 2a 2a 1 2a 2a 1 2a 2a 2a + + + + 1 + + 1 + 1 + + + + 1 + 1 . r 2a 2a . + . + . 2a 1 2b + 2b + + + 2a 2a 1 2a 1 2a . . 1 1 1 1 2a 1 2m 1 2a 1 2a 1 + . h + 2a 2a 1 1 . 1 . 2a 1 3 2a 2a 1 + 2a 1 2a . . 1 1 + . . 1 2m 1 2m 2a 1 1 1 1 1 + 1 . 2m 1 1 1 2m 1 . 2m 2m . 1 + . 1 1 1 2a . 1 2m 1 2a 2m 2a . 2m . . 1 1 . . . 1 1 . 3 1 1 2m + 2a 1 2m 2a . 1 + 2b . 2a 2m 2a 2a 2m + . 1 + + . 2m 1 1 1 2m 1 + 1 + 1 1 + + + 2m . . + + 2m + 1 . . . . . 1 + . . 1 . 1 + + 1 1 2m + . . . + + . 2m + 1 2m 2m . 2m 1 2m 2m . . + . h + . . + 2a 1 2a + + 1 + 2a + 1 1 1 2a 2b 1 1 1 2a . 1 1 2a 2a 2a . 1 2a 1 2a 2a 2a 2a 2a 2a 1 1 1 1 2a 2m 2a 1 1 2a 2a 2a 2m . . . . . r . . . . 1 2a 1 2a . 1 . r + 2a . + . 2a . 2a 2a 2a . 2m 2a 1 2a . 1 . . + . . . r . 1 1 + 2a 2a 2a 2a 2a 2a 1 1 1 1 . 1 2a 1 2a 2a 1 r . + 1 1 1 1 + . + + + 1 + . . 1 1 1 . + + 2a 1 . . . 1 . 1 + . + . . 1 1 1 + . 1 . + + + h 2a 2a 2a 2a 1 2a 2a 1 2a 2b 1 2a 2a 2a 2a + 2b 2m 2b 2b 1 2b 2a 2a 1 . . . . . . 2a . . . . . 2m . . . . . . 2b 1 1 + 2a 2a 2m 2b 1 2b 2a 2b 1 1 2b 2a 2a 2a + 1 1 2a 2a 2a 2b 2m 2a 1 1 2a . 1 . . . 2b . 1 2b . 1 . 2b 1 2a 2b 2a 2b . 1 . 2a . . 2a 1 2a 2b 3 . r . . . . 2m + . . 1 . . . . . . . 1 1 + 1 2m 1 2m + 2b 2m 2a 2m 2a 1 1 1 2a 1 2a 2a 1 2a 1 1 1 1 1 2a 1 1 2m 1 2b 1 1 1 1 + h 1 2a 2a . . . . . + 2b 1 . . 2a . + . 2b 1 1 2a 2a 2b 2a 1 2b 2b 2b 2b 2a 2b + 2b . 1 1 2b . 2m 2b 2a 2a 1 . . 1 + 2a . 2a . 2b 2a . 2a . 2m 1 1 1 1 2a . 2a . . 2b 1 1 1 1 2a 1 . 3 1 2a . 2m 2a . 1 2a + 1 2m 2m 2a 1 2a 2a 2a 2b . . 2a . 2a 2a 2b 2a 2a 2a . 2m 2a 2a 2a 2a 2b 2a 2a 2b 1 2a + 2m 2m . 2a 2b + 1 2a . . . + . . . 2b + 1 . 1 + 1 2a 1 1 . 2a 1 . 1 1 1 . . 1 . + . 1 2m . 2a . h 2a + 1 2m 2m 2a 1 2b + 2a 2m 1 1 2a 2a . 2m 2m 2m 2a + 2a 2a + 2m . . 1 . . . 2a . . . . . 2m . . . . . 2m . 2m 1 . 2m 2m 2a 1 . . . . 2a . 1 . . . 2a 2a 2b . + 1 . 2a 2b 2a 2a . . 1 . . . . . 2a 2a . + 1 2b 2a . 2b . 1 . 1 1 2m 1 2a 2m 2m 2m 2a 2a 2m + . . . . 2m 1 1 . 2b . . . . . . . 2a . . 1 + 2m 2a + 2m 2b 2a 2a 2m 2a 1 2a 1 2m 2m 2m 2m 2m 2m 1 2a 2m 1 2m 2a 1 2b 2a 2m 2a 2m 2m 2a 2b h + . + 1 2a . 2m . 2a 2a 2a + 2b + + 2a 1 1 1 . . 1 . . . 1 1 1 1 1 + 1 . 1 1 + + . . . . + . 1 2a 1 1 1 2m 1 2a 1 1 2a . . 2b 2a . 1 . 2m 2a . 2a . . . . + . . 2a . + 2a 2a 1 1 2b . + 1 2b 1 + . 2a + 2a 1 1 . . . . . . . . . + 1 . + . . . . + . . . 2m . + . 2b 1 r 2b . . . + . + 2m . . 1 1 2a . + 1 2a + . . . . . + . . 1 2a 2m 2a . 1 . 2m 2m 1 1 . . h . . . . 2m . . . . r 1 . 1 1 . 1 r 1 . . . + . . + 1 1 . 2a 2m . 1 1 + 2m + 1 . . 1 . 1 1 2m 2m . . 1 2m . 1 . . . 1 . + . . 2a . . 1 r . . . + . 1 . 1 2a . . 2m 2a 2m 2m . . . + 1 2m . . 1 . 1 2m 1 . + . 1 r 1 2m 2m 1 . 1 + + . . . . . . . . 1 . . 2m 2m 1 r 1 . . . . . . . . . . + 1 . . . . . . . . . . . . . . + 2m 1 + r . 1 1 . . . . h . . . . . . . . 3 . . . + . . + . . . . . . . 1 . . . . . . . . . . . . 2a . 1 1 . + 2a . . . . . . + . + + . 2b . . . . . . . . 1 . . . . + . . . . . . . . . . 1 . r . . . + . . . . 2a 1 . . 2a + + + + . + . 1 . . . . . + . + 1 + . 1 . . . . . + 1 2a 2a . . . . . 1 . 1 . 1 + . . 2a + + . . 1 1 . . . + . . . . . + . . . . . h 2a 2b 2b 2m 1 2a 2a . . . 1 . 1 . 2b . . . . 1 . 2b 2m 2m 2b . 2m 1 . . . 2a 1 . 1 2m 2a . . 2m . 2a . . . 2b 3 . . 2a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2m . . 2a . 2a 1 2m 2m 2a . 2m . . . 1 . 1 . . . . . . . . 2b 2a . 1 2m 1 . . . . + . . 1 1 . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . 1 1 1 1 1 r . 1 1 + . 1 . . . . 1 . . . . . . . r + . 2a . 1 . . 1 1 . . . 2m . . 2m . . . . . . . . . + . + . . . . + . . . 1 + . + . . . . . . + + . + 1 . r . . . . + . . . . . . . . . . + + . . . . . . . . + . . . . 1 . 1 . . + . + 1 . + . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . + 1 2m . . . . + . . . . . . . . . . . . + . . . 1 . . . 1 + + 2m . . + + r 1 . . . . . . . 1 . . 1 2m 1 . . . . . . + . + 2m + . . . . . . . . . . . . . . . . . . . . . . . . . . . . + 1 . . . . . + + . . . . . . + . . . . 2m . . . . . . . . . + . . . r . . . . . + . m . . . + . . . . . . + . . . . . . . . . . . . . . . . . . . . + . . . . . . + . 1 . . . . . . . . + . + . . + . . . . . . + + . + + . + . + . . . . . . . + . 1 . . . . . . . . . . + + . . . . . 1 . . . . . . . . . . . . . . . . . . + + . . . . 2m 1 . . . . . . . 1 . . 1 . . + . . . . . + . . . . . . . . . . . . + . . h . . . . . . . . 1 . + . . 1 . 1 . 2m . . 2b . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2a 3 2b 2m . 2m 2a 2m + 4 2a . . . . . . . . . . . . . . . . . . . . . . . . 2m . 2a . 1 1 . r . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . 2a . . . . . 2a . . + h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . r 1 . . 1 . . . . . . . . . . 1 . + . + . . 2a . . 2b . . . . . . . . . . . . . . . . . + . . . . . + . . . . . . . . 2a . . . + . . . . + . + . . . . . . . . . . . . . . . + . . h . . . . . . . . . + . . . . . . . . . . . . . + + . . . 2a 2m . . . . . . . . r + + . . . . . . . . . . . . . . . . . . . . . + . . . + . r + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . h . . . . . . . . . . . . . . . . . . . . . . . . + 1 . . . . . + . . + 1 + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . + . + . + + . . . . . . . . . . . . . . r . . . 2m 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . + . 2m . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . r 1 1 + + . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2m . 1 . . . . . . . . . . . . . . . . . . . . . . . . 1 . . 2m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . 1 . + + + . . . + . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .r . . . . . . . . . . . . . . . . . . .+. . . .+. . . . . . . . . . . . . .+. . . . . . . . . .+.+. . .+. . .r+. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . 2m . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 1 . r 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . .r . . . . . . . . . . . . . . . . . . . . . . . . . . .r . . . . . . . . . . . . . . . . . . . . . . . . . .r .+. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1 . . . . . . .+. . . . .+. . . . . . . . . . .r . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . . . + 2a . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . + . . 2a . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . + 2m . . . 2m . . . . . . . . . . . . . . . . . . . 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. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .+. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .+. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .+. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .+. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .+. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .+. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .+. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .r . . . . . . . . . . . . . . . . . . . . . . . . . . . m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . +
Simmelink et al.: Habitat and management preference of Bromus racemosus L.
Supplement E2. Overview table with header data of relevйs from own field work. Anhang E1. Ьbersichtstabelle mit Kopfdaten der Vegetationsaufnahmen von eigener Feldarbeit.
Relevй name AB1 AB2 AB3 ABa ABb ABc ABd ABf ABi AH1 AH2 AHa AHd AHe AHf BB1 BB2 BB3 BB4 BB5 BG1 BG2 BG3 BG4 BG5 BG6 BR1 BR2 BR3 BR4 BR5 BR6 CO1 CO2 CO3 CO4 CO5 CO6 CO7 CO8 COa COb COc COd DB1 DB2 DB3 DB4 DB5 DB6 DBa DH1 DH2 DH3 DH4 DW1 DW2 DW3 GB1 GB2 GB3 GB4 GB5 GU1 GU2 GU3 GU4 GU5 HP1 HP2 HP3 HP4 HP5 KB1 KB2 KB3 KB4 KB5 KB6 KB7 KBb KBc KD1 KD2 KD3 LA1 LA2 LA3 LA4 LL1 MK1 MK2 MW1 MW2 OH1 OH2 OH3 OH4 PA1 PA2 PA3 PA4 PA5 PA6 PA7 PA8 RG1 RG10 RG11 RG12 RG2 RG3 RG4 RG5 RG6 RG7 RG8 RG9 SB1 SB2 SB3 SB4 SB5 SC1 SC2 SC3 SC4 SC5 SC6 SW1 WA1 WA2 WA3 WA4 WA5 WA6 WA7 WA8 WA9 WB1 WB10 WB11 WB2 WB3 WB4 WB5 WB6 WB7 WB8 WB9 WBa WL1 WL2 WL3 WL4 WL5 WL6 WL7 WL8 WL9 ZK1 ZK2
Date X-coordinate Y-coordinate Length
(year/month/ (x 1000) (x 1000) relevй
day)
(m)
20130521 158.600
444.677
3
20130603 157.413
443.823
3
20130603 157.402
443.820
3
20080527 157.816
444.233
4
20080526 158.996
444.412
4
20080526 158.573
444.663
4
20080526 158.713
444.453
4
20080526 159.169
444.390
4
20080605 159.461
445.169
4
20130519 169.656
444.522
3
20130613 170.256
444.515
3
20100427 169.827
444.459
6
20100427 170.082
444.818
5
20100426 170.298
444.507
5
20100426 170.345
444.757
5
20130617 132.110
425.131
3
20130617 132.082
425.139
3
20130617 132.018
425.077
3
20130618 20130618 20130525
132.170 132.165 136.674
425.444 4,5
425.453
3
440.903
3
20130525 136.693
440.915
3
20130525 136.157
441.183
3
20130525 136.153
441.197
3
20130526 135.711
441.462
3
20130526 135.699
441.433
3
20130606 204.001
510.288
3
20130611 203.945
510.257
3
20130611 203.952
510.271
3
20130611 203.939
510.266
3
20130611 204.017
510.299
3
20130611 204.001
510.300
3
20130610 211.578
457.593
3
20130610 211.589
457.576
3
20130610 211.592
457.545
3
20130612 211.560
457.344
3
20130612 211.545
457.346
3
20130612 211.583
457.340
3
20130612 211.555
457.368
3
20130629 211.485
457.269
3
20070516 211.300
457.560
5
20090527 211.432
457.270
2
20070516 211.399
457.577
5
20090527 211.606
456.975
2
20130702 121.357
568.508
3
20130702 121.318
568.485
3
20130702 121.287
568.274
3
20130702 121.195
568.346
3
20130702 121.202
568.334
3
20130702 20090618 20130704
121.176 121.319 119.456
568.350 4,5
568.452
5
564.051
3
20130704 119.448
564.040
3
20130704 119.405
564.072
3
20130704 119.439
564.055
3
20130531 130.365
442.890
3
20130531 130.390
442.870
3
20130531 130.414
442.895
3
20130606 203.742
508.428
3
20130606 203.774
508.418
3
20130607 203.654
508.433
3
20130607 203.712
508.445
3
20130607 203.691
508.427
3
20130620 20130620 20130620
132.744 132.742 132.720
413.855 4,5
413.850
3
413.838
3
20130620 132.715
413.841
3
20130620 132.627
413.789
3
20130618 115.301
424.255
3
20130618 115.250
424.260
3
20130618 115.283
424.008
3
20130618 115.256
423.923
3
20130618 115.250
424.084
3
20130619 140.967
421.906
3
20130619 20130619 20130619
140.967 140.973 140.958
421.913
3
421.885
3
421.878 4,5
20130621 20130621 20130621
140.978 140.561 140.559
421.810
3
422.486
3
422.432 4,5
20070620 140.679
422.006
2
20070620 20130626 20130626
140.668 129.659 129.524
422.181
2
443.328 4,5
433.376 4,5
20130626 20130529 20130529
129.529 135.802 135.811
443.386 4,5
443.044 4,5
433.048
3
20130529 136.021
433.105
3
20130529 136.040
433.142
3
20130527 123.701
441.576
3
20130614 116.255
381.689
3
20130614 20130526 20130526
116.266 132.977 132.939
381.700
3
444.326 4,5
444.335 4,5
20130530 135.433
438.428
3
20130530 135.423
438.363
3
20130530 135.296
438.424
3
20130530 135.342
438.417
3
20130528 128.111
441.023
3
20130528 128.136
440.995
3
20130528 128.178
441.075
3
20130528 128.205
441.042
3
20130528 128.405
441.288
3
20130528 128.421
441.262
3
20130626 128.009
440.459
3
20130626 127.951
440.325
3
20130605 145.443
437.605
3
20130624 146.790
437.842
3
20130624 146.665
437.909
3
20130624 146.610
437.924
3
20130605 145.482
437.605
3
20130605 145.396
437.536
3
20130605 145.683
437.387
3
20130605 145.618
437.380
3
20130605 145.472
437.839
3
20130624 146.154
438.316
3
20130624 20130624 20130524
146.578 146.756 137.925
437.938
3
437.841 4,5
437.262 4,5
20130524 20130524 20130625
137.937 137.796 137.760
437.273
3
437.128 4,5
437.129
3
20130625 137.762
437.149 4,5
20130527 136.284
434.249
3
20130531 136.320
434.270
3
20130627 136.211
435.119
3
20130627 20130627 20130627
136.219 136.044 136.004
435.105
3
434.842 4,5
434.728
3
20130523 131.114
440.012
3
20130527 138.710
438.591
3
20130531 138.794
438.486
3
20130531 138.690
438.553
3
20130531 138.680
438.600
4
20130531 138.508
438.688
3
20130531 138.420
438.600
3
20130625 138.691
438.576 4,5
20130625 20130625 20130701
138.668 138.546 115.845
438.620
3
438.642
3
565.762 4,5
20130705 20130705 20130701
115.638 115.641 115.798
566.446
3
566.479
3
565.756 4,5
20130703 116.796
566.811
3
20130703 20130703 20130703
116.790 116.801 117.314
566.870 4,5
566.879
3
567.226
3
20130703 117.298
567.281
3
20130705 116.081
566.454
3
20130705 116.109
566.623
3
20070619 116.233
566.806
5
20130527 120.185
439.262
3
20130527 120.227
439.268
3
20130531 120.145
439.237
3
20130531 120.150
439.230
3
20130531 120.162
439.206
3
20130531 120.161
439.200
3
20130531 120.160
439.160
3
20130531 20130531 20130606
120.163 120.134 204.990
439.163
3
439.179
3
507.692 4,5
20130606 204.910
507.684
3
Width relevй (m) 3 3 3 4 4 4 4 4 4 3 3 4 5 5 5 3 3 3 2 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 5 2 5 2 3 3 3 3 3 2 5 3 3 3 3 3 3 3 3 3 3 3 3 2 3 3 3 3 3 3 3 3 3 3 3 3 2 3 3 2 2 2 2 2 2 2 3 3 3 3 3 3 2 2 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 2 2 3 2 3 2 3 3 3 3 2 3 3 3 3 3 2,5 3 3 2 3 3 2 3 3 2 3 2 3 3 3 3 3 5 3 3 3 3 3 3 3 3 3 2 3
Surface Exposition
relevй
(mІ)
9
N
9
9
16
N
16 16 16
16 16 9
9
24 25 25
25
9
9
NW
9
9
SSE
9
9
NW
9
SW
9
SE
9
SE
9
9 9 9
9 9 9
9
9
9
NE
9
9
N
9
W
9
9
NW
9
E
25
4
25 4 9
9
NW
9
9
N
9
9 25 9
9
W
9
9
W
9
9 9 9
9
9
SW
9
E
9
SE
9 9 9
9
9 9 9
9 9 9
9
S
9
S
9
9
9
9
9
E
4
4
9
9
W
9
W
9
NE
9
S
9
E
9
9
S
9
9
9
SE
9
SE
9
9
E
9
9
W
9 9 9
9
9 9 9
9 9 9
9
9 9 9
9 9 9
9
9
9
E
9
9 9 9
9
9
9
9
W
9
S
9
9
W
9
9
SW
9
9
10
9
9
N
9
S
9
S
9
9
9
N
9
9
9
9
NW
9
NW
9
9
N
9
SW
9
25
9
9
E
9
SE
9
SE
9
SE
9
9
NNE
9
N
9
9
9
NE
Inclination (degrees) 1 0 0 10 0 0 0 0 0 0 0 0 0 0 0 0 5 0 28 0 1 0,5 2 0,5 0 0 0 0 0 0 0 0 0 0,5 0 0,5 2 0 1 0,5 0 0 0 0 0 0,5 0 0,5 0 0 0 0 0,5 0 0,5 0 0 0 0 0 1 0,5 1 0 0 0 0 0 0 0 0 0 0 1 2 0 0 0 0 0,5 0 0 0 0,5 2 0,5 0,5 0,5 0 1 0 0 1 1 0 0,5 0 0,5 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 1 0 2 0 0,5 0 0 0 0 7 0,5 0,5 0 0 0,5 0 0 0 1 0,5 0 1 0,5 0 0 0 1 1 1 1 0 15 0,5 0 0 0,5
Heigt above NAP (AHN2) 5,3 6,14 6,26 7,38 7,03 5,3 6,99 6,36 6,01 5,42 5,19 5,29 5,08 5,14 5,12 3,19 2,65 3,05 2,58 3,01 -0,11 0,09 0,18 0,31 0,23 0,08 -0,07 -0,02 0,18 0,14 -0,12 -0,13 7,62 7,94 7,52 7,53 7,25 7,99 7,31 7,26 6,47 6,36 6,15 7,82 -0,6 -0,68 -0,67 -0,79 -0,72 -0,81 -0,72 -0,49 -0,5 -0,47 -0,46 0,99 1,32 1,46 0,16 0,1 0,86 0,32 0,4 0,96 0,9 0,85 0,92 0,95 0,8 0,79 0,73 0,75 0,71 1,85 1,97 1,56 1,47 1,61 1,4 1,27 1,3 1,25 1,25 -0,42 1,1 0,6 0,68 0,92 0,88 2,42 14,51 14,69 1,67 2,11 0,06 0,07 0,12 0,15 0,09 0,12 0,04 0,02 0,16 0,09 0,05 0,09 1,25 1,3 1,32 1,3 1,3 1,24 1,37 1,29 1,28 1,48 1,31 1,34 0,21 0,07 0,07 0,12 0,07 0,02 0,03 0,28 0,23 0,17 0,19 0 0,06 0,38 0,1 0,11 1,03 0,4 0,05 0,01 0,1 -0,43 -0,45 -0,45 -0,39 -0,68 -0,64 -0,72 -0,71 -0,58 -0,58 -0,45 -0,44 1,15 1,06 1,29 1,04 1,11 1,06 1,09 1,03 1,09 0,29 0,41
Total cover (%) 93 95 95 100 100 100 100 100 100 92 98 97 100 100 100 95 90 95 90 90 95 95 90 93 85 95 90 80 90 90 95 90 97 93 95 95 95 95 97 93 99 100 99 95 95 85 95 95 90 98 100 92 95 95 95 80 90 99 95 93 90 95 93 93 95 95 95 95 97 95 90 90 90 90 90 93 87 93 95 90 0 0 95 93 92 85 75 93 90 92 95 95 90 88 90 80 85 85 95 93 90 90 92 92 93 95 92 95 95 97 90 90 92 85 80 98 95 95 92 80 85 93 90 90 98 97 90 95 95 88 82 100 80 90 95 95 85 85 97 85 95 95 92 98 95 90 93 95 98 97 100 85 95 90 90 95 95 95 95 95 93 90
Cover Cover Cover litter
herb layer moss layer layer (%)
(%)
(%)
93
3
0
95
7
40
95
1
35
100
0
15
100
1
10
100
0
0
100
1
0
100
0
10
100
5
40
90
4
0
98
3
40
97
5
35
100
1
40
100
5
35
100
0
40
95
1
15
90
3
30
95
1
25
90
3
5
90
4
10
92
30
25
95
10
40
87
20
5
92
3
10
80
10
20
95
7
50
90
10
40
70
60
5
85
50
10
85
35
10
95
3
30
90
10
50
95
20
5
93
7
5
95
3
5
95
3
8
95
2
7
95
1
20
97
1
30
93
10
7
99
1
5
100
0
5
99
1
5
95
0
5
95
0
20
85
0
7
95
0
30
95
1
10
90
0
35
98
2
30
100
0
15
75
70
20
90
50
30
85
50
30
90
45
25
80
1
10
90
1
10
99
10
10
95
7
30
90
7
60
90
3
35
92
20
30
92
7
20
93
3
20
95
3
10
95
3
15
95
1
10
95
2
15
97
3
10
95
5
10
90
10
5
90
10
10
90
1
10
90
3
15
90
5
20
93
1
10
87
0
30
93
3
10
95
7
10
88
7
10
0
0
27
0
0
30
85
70
10
93
1
7
90
10
10
85
3
5
75
7
10
85
50
25
80
50
25
90
2
20
80
70
5
70
80
5
88
10
10
87
10
10
80
45
10
70
45
30
80
50
60
80
40
30
95
3
40
93
7
20
90
1
50
88
7
25
92
2
20
92
3
25
93
10
20
95
25
30
92
10
10
95
1
10
95
10
15
97
1
15
90
10
10
90
10
10
90
5
20
85
3
40
75
15
50
98
0
10
95
3
10
95
0
20
90
3
30
80
0
5
80
35
20
87
20
20
85
40
8
90
10
20
95
15
20
97
2
7
88
20
5
95
10
8
93
30
5
85
3
10
82
2
15
100
1
10
80
0
10
80
10
15
95
7
25
95
2
20
75
40
10
80
35
10
97
10
20
85
0
75
95
7
5
95
0
15
92
0
75
98
2
15
95
1
5
80
60
40
87
60
5
95
15
7
98
0
5
97
0
7
100
1
7
85
1
5
95
1
10
90
2
40
90
2
15
95
5
20
90
10
20
90
10
10
95
1
5
95
0
20
93
0
70
90
3
60
Cover bare soil (%) 7 2 2 0 0 0 0 0 0 5 15 0 0 0 0 20 20 10 30 20 5 5 10 8 12 5 5 30 15 20 10 15 20 30 10 20 10 20 8 15 0 0 0 0 8 25 3 10 10 5 15 1 1 3 5 20 10 2 5 5 20 5 15 7 7 6 10 10 10 10 20 20 10 20 20 15 20 20 10 25 0 0 7 35 20 15 20 5 5 8 5 5 10 10 5 10 5 10 3 5 5 10 7 8 15 10 8 20 10 10 10 10 5 10 8 15 10 8 10 20 20 10 20 10 5 15 20 20 15 10 15 10 20 20 5 5 30 15 10 1 20 20 1 7 10 3 7 8 20 20 15 15 5 10 10 5 7 10 5 5 10 15
Average height high herb layer (cm) 50 40 40 80 70 40 70 80 60 50 70 70 70 40 55 70 70 70 75 60 40 50 60 50 50 65 60 50 70 70 90 65 70 80 70 65 75 60 70 60 50 30 60 30 55 30 50 60 45 60 40 55 55 60 65 55 50 45 60 50 80 50 55 65 50 80 80 80 60 70 60 50 80 80 80 75 50 65 75 70 0 0 70 70 100 50 50 60 50 70 80 55 50 50 50 50 70 45 60 45 60 45 70 55 50 100 40 90 70 90 60 35 45 45 50 90 75 80 65 35 40 65 75 70 65 70 60 80 55 50 50 55 50 50 50 50 50 40 80 50 65 55 60 60 55 40 40 45 65 70 40 35 60 50 45 60 65 45 80 40 90 70
Average height low herb layer(cm) 30 15 15 0 0 0 0 0 0 30 30 15 25 0 20 30 10 25 25 10 25 40 30 25 30 45 25 10 20 20 35 20 25 35 30 20 20 25 25 20 0 0 0 0 10 3 12 20 7 20 7 5 10 10 15 30 30 20 40 25 35 25 25 20 20 25 30 30 30 30 10 15 30 25 15 25 10 12 25 15 0 0 10 40 40 30 15 25 20 50 30 15 30 30 20 20 40 20 30 25 40 20 45 20 10 10 15 40 30 30 15 15 15 15 10 25 30 30 40 15 20 20 10 40 40 33 15 35 12 25 30 30 20 30 25 30 5 5 35 8 20 10 15 18 15 5 5 8 20 20 0 15 20 25 15 20 40 30 50 30 45 20
Maximum height herb layer (cm) 65 65 80 130 0 0 100 110 0 70 100 90 110 90 90 100 100 100 100 80 60 95 90 80 100 95 110 90 100 110 120 110 95 120 110 90 120 90 110 85 0 0 0 0 73 60 68 115 68 102 60 95 100 87 110 80 90 75 90 90 105 95 90 110 110 120 120 120 105 100 100 100 100 130 110 105 110 90 100 100 0 0 120 130 135 95 90 100 85 100 135 110 65 60 100 100 100 85 95 70 95 80 85 90 95 140 80 125 125 120 100 90 90 70 90 115 120 110 90 45 90 100 130 100 85 120 100 135 90 95 80 95 80 80 90 80 90 75 170 72 88 87 100 110 95 70 82 62 90 95 0 60 100 95 80 100 90 80 90 80 120 120
Location (with parcel number) Amerongse Bovenpolder p1 Amerongse Bovenpolder p6 Amerongse Bovenpolder p6 Amerongse Bovenpolder p4 Amerongse Bovenpolder p3 Amerongse Bovenpolder p2 Amerongse Bovenpolder p1 Amerongse Bovenpolder p3 Amerongse Bovenpolder p5 Achterbergse Hooilanden p1 Achterbergse Hooilanden p2 Achterbergse Hooilanden p4 Achterbergse Hooilanden p5 Achterbergse Hooilanden p2 Achterbergse Hooilanden p3 Brakelse Benedenwaard zuid Brakelse Benedenwaard zuid Brakelse Benedenwaard zuid Brakelse Benedenwaard noord Brakelse Benedenwaard noord Bolgerijen oost Bolgerijen oost Bolgerijen perceel 1/6c Bolgerijen perceel 1/6c Bolgerijen perceel 1/34A Bolgerijen perceel 1/34A De Brommert, Grote Aanwas De Brommert, Hoge Brommert De Brommert, Hoge Brommert De Brommert, Hoge Brommert De Brommert, Grote Aanwas De Brommert, Grote Aanwas Cortenoever perceel 404 G2 Cortenoever perceel 404 G2 Cortenoever perceel 404 G2 Cortenoever perceel 404 G Cortenoever perceel 404 G Cortenoever perceel 404 G Cortenoever perceel 404 G Cortenoever perceel 404 G Cortenoever perceel x Cortenoever perceel 404F Cortenoever perceel 404F Cortenoever perceel 401A4 De Bol noordelijk van plas De Bol noordelijk van plas De Bol ver zuidelijk van plas De Bol net zuidelijk van plas De Bol net zuidelijk van plas De Bol net zuidelijk van plas De Bol noordelijk van plas Dijksmanshuizen Dijksmanshuizen Dijksmanshuizen Dijksmanshuizen Dertienmorgenwaard Dertienmorgenwaard Dertienmorgenwaard Genninger Buitenlanden Genninger Buitenlanden Genninger Buitenlanden Genninger Buitenlanden Genninger Buitenlanden Gansooiense uiterwaard p1 Gansooiense uiterwaard p1 Gansooiense uiterwaard p1 Gansooiense uiterwaard p1 Gansooiense uiterwaard p2 Hengstpolder Hengstpolder Hengstpolder Hengstpolder Hengstpolder Komgrondenres Bommelerwaard p1 Komgrondenres Bommelerwaard p1 Komgrondenres Bommelerwaard p2 Komgrondenres Bommelerwaard p2 Komgrondenres Bommelerwaard p2 Komgrondenres Bommelerwaard p3 Komgrondenres Bommelerwaard p3 Komgrondenres Bommelerwaard p4 Komgrondenres Bommelerwaard p4 Kleiput Dertienmorgenwaard Kleiput Dertienmorgenwaard Kleiput Dertienmorgenwaard Lingeuiterwaard Asperen ZW Lingeuiterwaard Asperen ZW Lingeuiterwaard Asperen NO Lingeuiterwaard Asperen NO Lekuiterwaard bij Lopik Het Merkske, Waterbeemd Het Merkske, Waterbeemd Middelwaard Middelwaard Overheicop perceel 1/16A Overheicop perceel 1/16A Overheicop perceel 1/11A Overheicop perceel 1/11A Polder Achthoven perceel 26 Polder Achthoven perceel 26 Polder Achthoven perceel 28 Polder Achthoven perceel 28 Polder Achthoven perceel 36 Polder Achthoven perceel 36 Polder Achthoven De Kikker Polder Achthoven De Kikker De Regulieren noordwest perc.1 De Regulieren zuidoost perc. 2 De Regulieren zuidoost perc. 1 De Regulieren zuidoost perc. 1 De Regulieren noordwest perc.1 De Regulieren noordwest perc.2 De Regulieren zuidwest De Regulieren zuidwest De Regulieren noord De Regulieren noordoost De Regulieren zuidoost perc. 1 De Regulieren zuidoost perc. 2 Sonsbrug nieuw hooiland Sonsbrug nieuw hooiland Sonsbrug oud hooiland Sonsbrug oud hooiland Sonsbrug oud hooiland Schaayk perceel 2/31A Schaayk perceel 2/31A Schaayk perceel 3/14A Schaayk perceel 3/14A Schaayk perceel 2/3A Schaayk perceel 2/3A Scharperswijk perceel 1B De Waai midden De Waai oost De Waai midden De Waai midden De Waai noordwest De Waai zuidwest De Waai midden De Waai midden De Waai westkant van midden Waal en Burg Blok Waal en Burg centraal p4 Waal en Burg centraal p4 Waal en Burg Blok Waal en Burg ten N v Molenkil Waal en Burg ten N v Molenkil Waal en Burg ten N v Molenkil Waal en Burg Westerkolk Waal en Burg Westerkolk Waal en Burg centraal p3 Waal en Burg centraal p2 Waal en Burg centraal p1 Willige Langerakse Waard Willige Langerakse Waard Willige Langerakse Waard Willige Langerakse Waard Willige Langerakse Waard Willige Langerakse Waard Willige Langerakse Waard Willige Langerakse Waard Willige Langerakse Waard Zijlkolk noordelijk perceel Zijlkolk zuidelijk perceel
Natura Cover B.
2000- racemosus
area
66
7
66
4
66
2
66
4
66
4
66
5
66
4
66
4
66
3
no
5
no
7
no
3
no
5
no
6
no
6
71
4
71
5
71
5
71
5
71
0
no
5
no
5
no
4
no
5
no
5
no
4
36
2
36
4
36
2
36
2
36
2
36
0
38
3
38
0
38
3
38
4
38
5
38
0
38
5
38
4
38
3
38
5
38
4
38
3
2
4
2
2
2
4
2
5
2
0
2
5
2
3
2
3
2
3
2
3
2
3
no
5
no
5
no
4
36
5
36
3
36
2
36
6
36
3
no
2
no
2
no
2
no
3
no
3
112
3
112
4
112
2
112
3
112
2
no
6
no
2
no
5
no
2
no
5
no
5
no
2
no
4
no
3
no
2
no
4
no
3
70
3
70
2
70
3
70
2
no
2
no
2
no
2
no
3
no
3
no
5
no
4
no
4
no
3
105
7
105
5
105
4
105
5
105
6
105
5
105
4
105
4
no
4
no
3
no
3
no
4
no
5
no
3
no
3
no
3
no
2
no
3
no
4
no
2
70
7
70
3
70
3
70
3
70
2
70
4
70
4
70
4
70
3
70
4
70
3
70
4
no
3
no
4
no
2
no
3
no
4
no
3
no
0
no
0
no
5
2
3
2
6
2
0
2
5
2
6
2
4
2
0
2
2
2
5
2
6
2
6
2
5
no
4
no
4
no
2
no
4
no
3
no
0
no
3
no
2
no
4
36
4
36
2
Phenology B. racemosus begin flowering mid flowering mid flowering mid flowering mid flowering begin flowering begin flowering mid flowering mid flowering begin flowering end flowering end flowering end flowering end flowering end flowering end flowering end flowering end flowering end flowering begin flowering begin flowering begin flowering begin flowering begin flowering begin flowering mid flowering end flowering end flowering end flowering end flowering mid flowering mid flowering end flowering end flowering end flowering end seed dead dead dead - begin seed begin seed mid seed mid seed begin seed begin seed begin seed begin seed begin seed begin seed begin flowering begin flowering begin flowering mid flowering mid flowering mid flowering mid flowering mid flowering begin seed end flowering mid seed mid seed mid seed end flowering end flowering end flowering end flowering end flowering begin seed begin seed begin seed begin seed begin seed begin seed begin seed begin flowering begin flowering begin flowering begin flowering mid flowering begin flowering begin flowering begin flowering end flowering end flowering begin flowering begin flowering mid flowering begin flowering begin flowering begin flowering begin flowering begin flowering begin flowering begin flowering begin flowering begin flowering mid seed mid seed mid flowering mid seed mid seed mid seed mid flowering mid flowering mid flowering mid flowering mid flowering mid seed mid seed mid seed begin flowering begin flowering begin flowering end seed end seed begin flowering begin flowering end seed end seed end seed end seed begin flowering begin flowering begin flowering begin flowering begin flowering begin flowering begin flowering mid seed end flowering begin seed end flowering begin seed begin seed begin seed begin seed begin seed begin seed begin flowering begin flowering begin flowering begin flowering begin flowering begin flowering begin flowering begin flowering mid flowering mid flowering
Community 8 4 4 4 4 4 4 4 5 5 5 5 5 5 5 4 4 4 4 4 5 5 11 5 11 11 6 6 6 6 6 6 4 4 4 4 4 4 4 4 5 5 5 4 3 3 3 2 2 3 3 1 1 1 1 8 5 5 6 6 4 6 6 8 7 7 7 7 7 7 7 7 7 4 4 5 5 5 11 11 5 11 9 9 9 5 11 11 5 10 10 4 4 5 5 11 11 5 5 5 5 5 5 5 5 11 11 11 11 11 11 11 11 11 11 11 11 5 5 11 11 11 11 11 4 5 11 11 5 11 5 11 11 5 5 11 11 11 3 3 3 3 3 3 2 2 3 3 3 3 5 8 4 5 8 8 8 8 8 5 5
pH-H2O 7,61 7,55 7,38 7,67 7,58 7,6 7,77 7,63 5,84 6,57 5,76 5,56 5,37 5,37 5,72 7,87 7,73 7,7 7,86 7,61 5,61 5,81 5,7 7,16 6,12 6,12 6,23 5,92 6,01 6,51 6,15 5,6 7,48 7,6 7,54 7,62 7,08 7,03 7,37 7,91 7,17 6,69 7,09 7,44 7,04 6,93 6,34 7,4 6,42 7,23 6,77 6,22 6,36 6,06 6,33 7,63 7,53 7,41 5,6 5,63 5,75 5,46 5,94 7,59 7,36 7,65 7,76 7,81 7,66 7,67 7,65 7,74 0 7,23 7,39 6,06 5,75 6 6,12 5,5 6,52 6,62 7,75 7,64 7,72 7,61 7,56 7,56 7,62 7,82 6,39 6,62 7,66 7,55 5,1 5,38 5,22 5,26 6,08 5,57 5,75 5,86 6,04 6,03 6,31 6,27 5,78 5,77 5,68 5,59 5,64 5,8 5,69 5,42 5,45 5,67 5,7 5,8 5,45 6,46 6,28 6,61 5,92 6,32 7,27 5,6 5,68 5,53 5,33 5,84 5,66 6,16 5,87 5,87 5,3 6,61 5,49 5,67 5,83 5,88 6,4 5,87 6,36 6,04 7,39 5,08 7,84 7,69 7,61 7,15 6,67 7,61 7,6 7,66 7,83 7,86 7,58 7,83 7,77 7,88 5,9 5,84
pH-KCl OM (%) K (mg/kg) K (mg/L) Ca (mg/kg) Ca (mg/L) Pw (mg/kg) P2O5 (mg/L) P-total N-total CN-ratio (mmol/kg) (mmol/kg)
7
13,5
179
2,36
724
9,55
4,5
7,11
10
146
2,07
656
9,3
7,1
6,6
8,2
134
2,43
247
4,49
6,6
7,2
10,4
190
2,61
872
11,99
3,6
7,21 11,6
77
1,07
844
11,72
1,4
7,04 14,7
335
5,26
939
14,73
3,8
7,02 10,6
116
1,48
822
10,5
3,1
7,02 12,4
126
1,42
652
7,34
4,4
4,83 12,5
272
4,04
214
3,18
6,1
6,11 23,3
151
2,05
341
4,64
41,9
5,1
30,9
95
1,08
380
4,33
35,1
5,1
33,9
102
1,13
269
2,99
6,3
4,95 36,7
401
4,62
289
3,33
10,4
4,64
37
321
3,74
284
3,31
7,7
4,64 27,7
141
1,98
114
1,6
4,3
7,09 10,1
117
1,83
708
11,11
3,2
7,11
11
169
2,61
835
12,87
4,5
7,24
10
135
2,06
669
10,22
4,6
7,34 6,3
112
1,49
608
8,1
9,8
7,26 6,8
229
2,96
519
6,71
17,8
4,58 14,6
587
8,48
325
4,7
7,6
4,84 19,2
415
5,07
278
3,4
14,7
4,81 17,4
473
7,55
387
6,18
6,9
6,55 13,4
96
1,16
509
6,15
10,8
5,35 12,1
272
3,17
235
2,74
19,7
5,29 20,2
276
4,33
332
5,21
11,5
5,32
12
391
4,72
257
3,1
4,1
5,06 7,9
214
3,09
141
2,04
2,1
4,96 6,6
75
1,16
127
1,97
1,9
5,41 10,8
121
1,54
197
2,52
4,7
5,36 14,5
286
3,78
318
4,19
4,5
4,78
14
169
2,37
187
2,63
2,1
6,9
11,9
100
1,41
448
6,34
3,5
7,11 7,9
62
0,81
587
7,66
3,8
7,02 11,4
129
1,51
668
7,79
3,4
7,09 8,2
380
4,59
521
6,3
3,3
6,55
8
83
1
305
3,69
4,1
6,53 6,8
39
0,65
205
3,39
3,6
6,9
7,9
79
0,98
364
4,5
3,2
7,2
7
47
0,62
569
7,51
0,8
6,59 11,1
113
1,47
434
5,66
5,4
5,92 10,8
185
2,78
379
5,68
5,3
6,5
11,3
325
4,56
468
6,57
4,3
6,91 7,5
61
0,81
283
3,74
6,1
6,51 4,9
46
0,88
169
3,21
2,1
6,63 5,8
19
0,29
230
3,48
4
5,27 4,3
12
0,16
0
0
3
7,01
8
10
0,14
294
4,13
0,7
5,2
2,8
16
0,28
0
0
1,1
7,04 8,8
37
0,41
2893
32,14
0,9
5,81 5,8
22
0,3
138
1,88
6,6
5,74 9,6
41
0,48
265
3,13
0,8
6,03 11,8
4
0,04
377
4,29
0
5,87 12,1
18
0,21
295
3,48
0,8
5,79 13,6
14
0,24
270
4,77
0,6
7,1
25
91
1,27
1044
14,64
6,4
7,07 13,8
130
1,55
962
11,49
9,2
6,82 14,5
134
1,77
538
7,1
3
4,6
11,6
163
2,06
112
1,41
5,5
4,35 12,7
143
1,85
214
2,76
7
4,88 7,3
57
0,83
118
1,7
2,8
4,51 11,2
235
3,07
191
2,5
3,8
4,76 12,6
196
2,58
348
4,59
6,1
7,18 7,4
56
0,85
533
8,15
3,9
7,01 8,1
28
0,32
428
4,82
3,6
6,9
8,5
107
1,26
591
6,98
3,4
7,21 8,4
63
0,85
474
6,38
3
7,07 8,2
45
0,61
609
8,29
0,7
7
11
129
1,56
810
9,79
2,5
7,14 9,2
142
1,85
804
10,5
2,3
7,23 13,8
173
2,12
979
11,97
4,1
7,11
12
434
5,18
927
11,07
5
0
0
0
0
0
0
0
6,62 11,3
465
8,65
761
14,17
4,3
6,79 10,3
440
5,93
842
11,35
3,7
4,97 11,5
308
3,94
337
4,3
5,5
4,72 11,7
315
3,67
335
3,91
6,9
5,02 11,3
319
4,47
217
3,05
10
5,33 14,8
410
5,29
368
4,75
7,7
4,6
15,9
289
3,25
252
2,83
4,4
5,19 11,9
642
8,92
465
6,46
5,8
5,43 13,2
473
6,57
436
6,05
5,8
7,26 8,8
69
0,79
601
6,85
1,8
7,07 11,4
192
2,48
966
12,48
3,1
6,81
10
418
6,27
655
9,82
6,7
7,17 9,9
72
0,81
596
6,7
7,1
7,15 10,3
0
0
0
0
3,7
7,33 13,9
101
1,55
1309
20,18
7,1
7,32 12,8
118
1,48
1241
15,51
7,2
7,33 5,5
122
2,03
771
12,85
5,4
5,71 42,6
97
1,05
258
2,8
10,2
6,06 29,7
41
0,47
195
2,25
0,9
7,08 11,5
145
2
794
10,92
2,2
7,09 11,9
165
1,99
832
10,05
2,5
4,28 20,8
194
2,42
230
2,88
7,2
4,52
16
569
7,98
301
4,22
6,4
4,02 17,8
436
5,81
215
2,86
3,8
4,24 20,6
354
5,06
329
4,7
5,6
4,97 16,5
312
3,77
345
4,17
11,6
4,6
17,6
350
4,86
449
6,23
9,4
4,71 19,5
256
2,84
134
1,49
6,3
4,8
15,7
262
3,46
353
4,66
8,3
4,88 17,6
415
5,07
322
3,94
26,2
4,93 19,2
298
4,6
270
4,17
14,9
5,24 18,3
281
3,44
191
2,34
5,7
5,52 17,2
328
4,88
394
5,85
9,4
4,57 15,1
521
5,93
349
3,98
4,4
4,64 14,6
305
4,37
291
4,17
11,2
4,47 16,6
203
2,74
181
2,44
6,7
4,48
16
248
2,82
284
3,24
7
4,59 16,1
193
2,36
182
2,23
6,5
4,24
15
367
4,84
210
2,77
4,5
4,35 18,7
351
4,39
140
1,75
4,8
4,07 13,9
283
3,34
280
3,3
4,2
4,27 15,4
501
7,38
248
3,65
4,8
4,58 13,3
417
6,89
315
5,2
6,7
4,54 14,5
255
2,94
285
3,29
7,8
4,58 13,2
279
4,19
322
4,83
12
4,87
24
167
2,11
325
4,11
9,5
5,66 24,8
189
2,63
594
8,25
25,2
5,21
17
335
4,47
399
5,32
5,3
5,69 17,7
278
4,13
431
6,4
8,1
4,87 17,6
229
2,83
307
3,8
4
5,98
12
28
0,34
184
2,22
1,7
6,75 8,6
59
0,7
521
6,22
1,7
4,5
15,4
259
3,71
254
3,64
4,2
4,38 15,1
251
2,89
288
3,32
6,1
4,17 15,1
210
2,95
217
3,04
4,3
3,97 12,1
148
1,77
171
2,04
2,5
4,65 12,5
365
6,65
342
6,22
5,5
4,96 25,8
74
0,99
359
4,78
5,3
5,35 7,6
127
2,26
135
2,4
5,6
4,84 19,6
104
1,33
142
1,81
54
4,96 15,8
107
1,6
216
3,24
3,3
4,27 8,4
256
3,63
155
2,19
16,2
5,7
10,4
226
3,04
272
3,66
4,5
4,75 38,1
168
2,19
353
4,61
2,3
5,14
45
31
0,37
569
6,87
1,7
4,91 16,4
312
4,16
437
5,83
10,5
4,73 4,3
142
1,62
14
0,16
2,6
5,61 5,5
83
1,09
159
2,07
1,5
4,49 3,5
28
0,39
0
0
0,7
5,3
6,9
324
5,8
79
1,41
5
5,56 9,9
1
0,01
142
1,81
2,3
7,02
7
75
0,93
463
5,72
2,4
4,16 6,3
11
0,16
0
0
0,7
7,48 8,3
28
0,46
308
5
1,2
6,99 4,8
30
0,55
80
1,47
2,7
7,21 4,1
114
1,77
248
3,86
7,1
6,38 4,2
49
0,69
171
2,42
7,1
5,55 4,5
63
0,86
137
1,86
5,1
7,06 10,6
137
1,54
685
7,71
8
7,19 11,5
183
2,44
868
11,57
6
7,05
9
252
4,3
742
12,67
4,7
7,15 11,8
191
2,33
756
9,24
13,9
7,2
13,6
254
3,32
939
12,26
9,2
7,17 17,5
262
3,49
1317
17,56
1,5
7,25 10,4
140
2,17
782
12,17
5,1
7,14 11,9
176
2,54
843
12,17
17,3
7,17 11,6
143
2,44
710
12,13
8,2
5,05 7,7
107
1,31
167
2,04
4,1
5,13 5,3
195
3,2
91
1,49
10,4
8,3
26,7
439,3 10,99
13,8
22,2
309
11,57
16,5
20,7
284,8 10,34
6,9
20,3
260,5 14,32
2,8
20,7
339,2 12,26
8,3
22,5
392,3 13,35
5,5
23,6
302,8 12,51
6,9
23,5
319,5
13,8
12,4
18,6
326,7 13,63
78,4
39,7
505
16,47
55
64,7
846,7 13,04
9,6
40,6
967,9 12,52
16,5
52,9
1190,3
11
12,4
44,2
1156,1 11,43
8,3
29,8
655,3 15,09
6,9
20,8
253
14,24
9,6
24,2
326,6 11,99
9,6
22,6
261,5 13,61
17,9
18,9
96,2
23,2
31,6
17,3
86,7
27,91
15,1
29,2
401,9 12,93
24,8
44,8
554,2 12,34
15,1
23,1
419,9 14,75
17,9
34,1
371,3 12,85
31,6
67,1
409,2 10,53
24,8
36,3
439,6 16,36
6,9
23,4
387,7 11,09
4,1
14,1
223,8 12,64
4,1
13,2
179,5 13,08
8,3
20,6
312
12,4
8,3
19,5
353,8 14,62
4,1
20,5
468,4 10,71
6,9
22,6
344
12,37
6,9
19,4
295,1
9,56
5,5
21,4
323
12,64
5,5
18
277,7
10,6
6,9
18,2
248,1 11,58
8,3
14,9
231,9 10,45
5,5
16,8
241,3 11,74
1,4
15,7
245,3 10,12
9,6
23,7
296,1 13,42
11
23,7
357,3
10,8
8,3
23,6
315,4 12,83
11
18,6
226,7 11,77
5,5
5,9
93,6
18,6
8,3
11,8
189,6 10,94
5,5
4,4
115,2 13,31
1,4
10
190,9
15
2,8
2,1
54,5
18,1
1,4
8,1
165
19,07
12,4
12,7
184,8 11,19
1,4
7,4
272
12,63
0
5,2
157,8 26,67
1,4
7,5
296,6 14,51
1,4
7,1
234,3 20,68
12,4
45,8
735,8 12,13
15,1
36,4
444,6 11,05
5,5
26,1
401,2 12,88
9,6
38,5
355,1 11,68
12,4
39,5
396
11,46
5,5
19,7
236,9 11,02
6,9
36,6
399,3 10,04
11
39
406,2 11,11
8,3
19,5
236,3 11,22
5,5
18,5
214
13,55
5,5
17,8
234,5 12,91
5,5
15
188,6 15,85
1,4
13,6
202
14,56
4,1
20,1
317,4 12,42
4,1
18,7
286,3 11,43
6,9
23,7
331,3 14,88
8,3
25,3
323,5 13,27
0
0
0
0
11
18,8
348,8 11,53
6,9
14,5
243
15,13
9,6
21,4
392,2 10,47
11
24
368,9
11,3
19,3
20,8
453,8
8,92
13,8
25,9
425,7 12,43
6,9
20,8
481
11,81
11
14,5
318,3 13,31
11
14,4
349,4 13,49
2,8
13,5
211,9 14,78
5,5
16,8
248,2 16,42
13,8
18,3
260,7 13,66
11
20,7
364,8
9,68
6,9
18,5
349,4 10,56
15,1
20,3
499,7
9,96
12,4
20
459,7
9,94
12,4
16,6
127,5
15,3
15,1
29,1
888,4
17,1
1,4
13,5
588
18,02
4,1
18,7
364
11,3
4,1
19,9
379,9 11,22
12,4
39,7
757,1
9,79
12,4
24
473,3 12,03
6,9
23,9
522,1 12,15
11
26,7
596,6 12,32
19,3
27,4
510,6 11,51
17,9
28,9
608,7 10,34
9,6
27,2
637,6 10,91
15,1
23,2
525
10,7
44
47,3
588,3 10,69
31,6
39
672,5 10,17
9,6
24,8
470,3 13,86
19,3
27,3
478,9 12,85
6,9
24,3
508,9 10,58
22
26,1
349,4
14,9
12,4
32,9
471,5 12,59
11
32,1
469,6 12,13
11
29,7
466,6 12,29
8,3
17,6
346,7 15,49
8,3
23,6
552,8 12,08
6,9
27
527,4
9,44
9,6
22,1
498,2 11,04
15,1
27,9
404,4 11,77
12,4
31,3
429,2 12,03
24,8
32,7
389,8 12,12
16,5
37,2
671,4 12,75
48,1
56,7
801,3 11,07
9,6
20,9
467,2
13
16,5
24,9
467,1
13,5
6,9
17,2
427,9 14,71
2,8
13,3
279,2 15,36
2,8
11
239,2 12,87
8,3
28,7
448,6 12,26
9,6
28,2
428
12,57
8,3
25,2
446,8 12,03
4,1
19,4
337,6 12,79
13,8
27,1
507,9
8,82
9,6
34
685,4 13,45
13,8
53,7
255,5 10,64
94,9
34
691,1 10,11
6,9
21,9
449,1 12,52
31,6
25,7
249,8 12,02
8,3
23,3
347,6 10,63
4,1
28,7
1000,4 13,59
2,8
31,9
1423,3 11,28
19,3
26,8
540,5 10,85
4,1
8,8
222,6
6,83
2,8
10
148,4
13,3
1,4
8,6
140,9
8,96
12,4
8,7
175,7 13,97
4,1
34,5
306
11,51
4,1
11,4
197,7 12,69
1,4
2,3
67,9
32,91
2,8
3,5
102,1 29,06
6,9
5,6
68,9
24,93
15,1
12,2
117,7 12,52
13,8
13,2
128,9 11,62
9,6
13,2
137
11,62
12,4
30,6
367,8
10,3
11
33,5
321,5 12,82
11
22,1
234,8 13,62
23,4
38
379,7
11,1
16,5
36,9
369,4 13,18
2,8
50,3
420
14,92
11
27,7
210,5 17,57
34,4
49,8
290,7 14,66
19,3
36,4
307,3 13,44
6,9
27,2
283,9
9,72
23,4
18,9
127,6 14,82
Light (Ellenberg value) 6,71 6,85 6,87 7 6,98 6,83 6,8 7,07 6,82 6,67 6,71 7,06 6,7 6,55 6,63 6,74 6,75 6,83 7,09 7,13 6,59 6,87 6,74 6,86 6,88 6,76 6,72 6,79 6,94 7,01 6,72 7,08 6,86 6,86 6,83 6,77 6,79 7,05 6,82 6,73 6,95 6,84 6,85 7,05 7,17 7,26 7,04 6,94 7,04 7,16 7,21 7,41 7,3 7,28 7,03 6,76 6,66 6,67 6,75 6,85 6,78 6,76 6,75 6,94 6,92 6,88 6,88 6,91 6,87 6,91 6,81 6,86 6,93 6,88 7,04 6,69 7,07 6,95 6,8 7,11 7,04 6,97 6,99 6,73 6,81 6,77 6,95 7,08 7,01 6,83 6,78 6,91 6,93 6,88 6,57 6,81 6,75 6,78 6,71 6,82 6,66 6,78 6,74 6,8 6,94 6,99 6,89 6,91 7,01 6,94 6,66 6,89 6,75 6,91 6,98 6,87 6,89 6,89 6,8 7,21 6,86 7,04 6,96 6,73 6,67 6,93 6,6 6,72 6,85 6,68 6,81 6,84 6,96 6,96 6,67 6,69 7,13 7,3 6,77 7,22 6,92 7,02 7,09 6,71 6,99 7,23 7,05 6,96 6,93 6,98 6,98 6,74 6,79 6,81 6,68 6,65 6,69 6,69 6,76 6,81 6,44 6,23
Moisture (Ellenberg value) 6,82 5,45 5,41 5,36 5,17 6,53 5,5 5,28 6,26 6,51 6,29 6,68 6,04 6,08 6,35 5,46 5,74 5,63 5,1 4,88 6,6 6,3 6,74 6,33 6,73 7,41 7,1 6,69 6,33 6 7,57 7,92 5,53 5,31 5,48 5,52 5,6 5 5,68 5,47 6,28 6,5 7,12 5,62 5,93 6,11 5,47 5,98 5,33 6,04 5,73 6,74 6,9 6,83 6,47 7,16 6,38 6 6,96 7,63 5,45 6,38 5,89 7,42 7,65 7,6 7,38 7,33 6,67 6,43 7,71 7,39 7,48 5,33 5,07 6,11 7 6,1 6,09 7,07 6,76 7,81 6,54 6,22 6,38 6,33 6,4 7,04 6,92 6,14 7,32 7,51 6,28 6 6,39 6,84 6,96 6,44 6,5 6,04 6,47 6,42 6,3 6,55 6,27 6,53 6,42 7,38 7,48 7,25 6,6 7,06 6,51 6,87 7,13 6,54 7,14 6,62 6,47 6,93 6,95 6,68 6,89 7,09 7,02 6,12 6,02 6,32 6,04 6,48 7,35 6,17 7,11 7,1 5,88 5,68 7,88 8,25 7,3 6,09 5,3 5,18 6,38 6,43 6,38 6,26 5,94 5,71 5,93 5,78 5,71 6,08 6,72 5,93 6,14 6,66 7,52 6,65 7,68 7,2 6,2 6,45
Reaction (Ellenberg value) 5,94 6,45 6,37 6,27 6,33 6,23 6,29 6,42 5,82 6,06 5,81 5,44 5,92 5,83 5,38 6,52 6,4 6,5 7,13 6,78 6 6,15 5,78 6,03 5,85 6,07 5,72 6,32 6,22 6,36 6,11 5,85 6,39 6,7 6,36 6,38 6,44 6,46 6,48 6,35 6,38 6,29 6,9 6,44 5,62 6,17 5,2 5,81 5,46 6,11 5,87 5,49 5,49 5,06 5,22 6,47 6,05 6,18 5,8 6,19 5,85 6,26 6,32 6,53 6,47 6,45 6,44 6,47 6,73 6,61 6,26 6,35 6,22 6,6 6,36 5,84 6,42 5,97 5,94 5,5 6,61 6,38 6,55 6,79 6,8 6,22 6,36 6,51 6,43 6,24 6,33 6,26 6,46 6,53 5,93 5,51 5,54 5,64 6 6 5,95 5,89 5,63 5,83 6,05 5,72 6,08 6,44 5,79 5,91 5,97 5,7 5,37 5,32 5 5,39 6,06 5,59 5,68 6,35 5,6 5,95 5,53 6,27 6,39 6,52 6,04 6,03 5,72 5,93 5,88 5,89 5,65 5,31 5,67 5,96 4,95 5,31 6,2 4,95 5,41 5,42 5,1 5,13 5,25 4,24 5,88 5,68 6,23 6,22 5,57 5,9 6,38 6,62 6,16 6,13 6,41 6,5 6,56 6,46 6 5,94
Nitrogen (Ellenberg value) 5,28 5,84 5,82 6,05 5,8 5,29 5,57 5,57 5,37 6,12 5,79 5,33 5,67 6,06 5,44 5,77 5,64 5,75 5,34 5,57 6,13 6,36 5,28 5,6 5,31 4,94 5,58 5,5 5,48 5,59 5,42 5,26 5,95 6,06 5,93 5,96 5,94 5,76 5,78 5,46 5,76 5,71 5,41 6,07 5,17 5,67 4,58 4,76 3,79 5,08 4,96 4,46 4,42 4,36 4,67 5,95 5,78 5,77 5,43 5,64 5,82 5,71 5,83 5,37 5,31 5,19 5,42 5,37 5,36 5,19 4,98 4,86 5,11 5,19 5,08 5,38 5,7 5,65 5,6 4,85 5,7 5,07 4,9 5,43 5,69 5,79 5,6 5,03 4,99 5,92 5,09 4,91 5,86 6,08 6 5,52 5,17 5,02 5,86 5,53 5,68 5,9 5,74 5,77 6,09 5,63 4,96 5,55 5,18 5 5,2 4,54 5,1 4,57 4,57 4,85 5,26 5,09 5,91 6,24 5,03 4,63 4,8 4,88 5,1 5,62 5,38 5,29 4,68 5,5 5,37 5,11 4,62 4,77 5,55 5,32 3,59 4,14 5,47 4,59 5,04 4,75 4,62 5,17 4,61 3,48 4,38 5,05 5,35 5,42 4,96 5,62 5,7 5,57 5,95 5,64 5,47 5,81 5,42 5,61 6,42 6,21
GLG (Wamelink value) 72,15 88,48 88,96 86,4 89,5 72,72 86,05 91,43 78,05 75,6 79,93 71,41 86,23 80,22 81,6 89,77 79,65 85,81 88,44 93,51 81,01 79,93 68,84 73,74 66,37 57,9 65,89 69,8 75 76,59 58,06 45,62 86,06 89,67 86,96 83,49 83,15 90,26 80,25 86,61 76,06 66,75 59,08 84,53 81,79 70,03 88,75 83,64 93,08 75,76 73,9 67,07 68,32 71,26 78,77 65,05 78,89 80,82 65,42 51,83 89,11 77,15 82,02 62,53 56,05 56,24 62,3 61,6 68,45 73,45 53,43 58,53 53,53 88,53 96,04 79,24 58,88 74,57 77,81 62,89 71,24 47,74 69,44 74,3 76,65 77,33 73,61 60,51 62,79 79,17 61,68 57,91 79,86 84,4 80,98 75,05 67,66 72,02 73,74 75,36 77,49 74,22 76,49 71,49 76,63 70,23 68,19 59,57 53,65 58,89 66,14 58,3 69,47 64,26 58,7 69,75 62,02 66,55 77,16 61,23 64,55 60,37 60,84 64,22 63,66 78,49 81,93 74,76 79,62 71,11 59,41 76,71 60,48 58,92 84,31 86,46 44,6 35,38 64,3 73,59 89,51 91,77 73,73 83,66 75,93 91,12 79,83 85,98 76,5 81,48 82,55 77,71 68,59 80,63 80,54 73,21 56,29 73,69 55,76 63,17 85,4 82,19
GVG Humus profile (names Texture (above 20
(Wamelink exist only in Dutch)
cm, in Dutch)
value)
37,01 51,92 52,36 51,74 49,12 37,67 48,12 52,58 39,58 42,17 43,08 35,13 47,84 42,51 43,36 51,1 45,76 51,68 49,66 50,84 41,94 45,77 36,71 40,32 36,27 32,34 39,05 38,16 41,55 45,42 36,07 26,37 50,33 53,39 51,32 47,11 47,85 51,19 43,19 48,73 45,77 37,78 32,17 52,27 40,66 36,3 41,56 39,48 38,4 36,55 36,86 30,65 30,45 32,98 35,87 36,93 44,45 43,2 35,33 29,13 48,85 41,03 45,53 32,98 30,77 29,71 32,63 31,95 34,56 35,8 27,72 29,3 27,49 49,57 49,95 41,71 32,76 40,54 41,01 31,82 38,52 25,72 33,96 41,99 45,14 43,79 40,35 30,19 29,93 44,33 36,36 29,74 44,97 48,81 41,81 39,07 35,05 37,46 40,82 38,68 42,13 39,8 41,46 38,82 42,66 37,18 34,54 29,68 28,63 30,57 35,05 31,02 34,63 31,32 30,54 34,82 30 33,15 42,31 35,49 37,4 32,56 33,28 32,43 30,48 44,57 43,24 39,65 39,42 36,48 32,14 39,78 31,5 30,01 44,75 46,84 25,18 23,27 33,83 30,4 44,2 44,97 31,05 42,1 36,36 33 34,58 40,78 41,44 43,67 41,53 41,89 38,88 44,95 45,05 40,77 31,17 43,58 31,8 35,77 56,87 46,7
Kleihydromull Kleihydromull Zure wormmull Kalkwormmull Kalkwormmull Kalkwormmull Kalkwormmull Kalkwormmull Schrale zure wormmull Moereerdmull Moereerdmull Kleieerdmoder Schrale moereerdmull Moereerdmull Moereerdmull Nesvaagmull Kalkwormmull Kalkwormmull zandige Nesvaagmull zandige Nesvaagmull Beekvaagmull Zure wormmull Beekhydromull Verstoorde Kalkwormmull Nesvaagmull Kleihydromull Beekhydromull Zure wormmull Zure wormmull Zure wormmull schrale Zure wormmull Zure wormmull Kalkwormmull zandige Kalkwormmull zandige Kalkwormmull zandige Kalkwormmull Kalkwormmull Vlakvaagmull Kalkwormmull Nesvaagmull Kalkwormmull Kleihydromull Kleihydromull Kleihydromull Vlakhydromull Vlakhydromull Kleihydromull Kleihydromull Beekhydromull Kleihydromull Vlakhydromull Kleihydromull Kleihydromull Kleihydromull Kleihydromull Kleihydromull Kleihydromull Schrale Beekhydromull Schraalwormmull Zure wormmull Beekhydromull Zure wormmull Kalkwormmull Kalkwormmull Kalkwormmull Kalkwormmull Kleihydromull Kalkwormmull Kalkwormmull Kalkwormmull Kalkwormmull
matig lichte zavel zeer lichte zavel zeer lichte zavel matig lichte zavel zeer lichte zavel zware zavel zeer lichte zavel matig lichte zavel lichte klei zeer lichte zavel zeer lichte zavel zeer lichte zavel matig lichte zavel matig lichte zavel zware zavel matig lichte zavel matig lichte zavel matig lichte zavel kleiig zand kleiig zand zware zavel zware zavel lichte klei zware zavel zware zavel zware zavel zware zavel zeer lichte zavel matig lichte zavel matig lichte zavel matig lichte zavel zware zavel zeer lichte zavel kleiig zand zeer lichte zavel kleiig zand zeer lichte zavel kleiarm zand zeer lichte zavel matig lichte zavel matig lichte zavel zware zavel matig lichte zavel zeer lichte zavel kleiarm zand kleiarm zand kleiarm zand matig lichte zavel kleiig zand matig lichte zavel kleiarm zand matig lichte zavel matig lichte zavel zeer lichte zavel zware zavel matig lichte zavel matig lichte zavel matig lichte zavel zware zavel zware zavel zeer lichte zavel matig lichte zavel zware zavel zeer lichte zavel zeer lichte zavel zeer lichte zavel zeer lichte zavel zeer lichte zavel matig lichte zavel zeer lichte zavel matig lichte zavel matig lichte zavel
Kleihydromull Kalkwormmull Zure wormmull Zure wormmull Zure wormmull Zure wormmull Zure wormmull Kleihydromull Kleihydromull Kleihydromull Kleihydromull Nesvaagmull Kalkwormmull Kalkwormmull Kalkwormmull Kalkwormmull Verstoorde Nesvaagmull Moereerdmoder Moereerdmoder Kalkwormmull Kalkwormmull Beekhydromull Beekhydromull Beekvaagmull Beekvaagmull Zure wormmull Beekhydromull Beekhydromull Beekhydromull Beekhydromull Zure wormmull Beekhydromull Zure wormmull Beekhydromull Beekhydromull Zure wormmull Zure wormmull Beekhydromull Beekvaagmull Beekhydromull Beekvaagmull Beekhydromull Beekvaagmull Zure wormmull Zure wormmull Zure wormmull Zure wormmull Beekhydromull Zure wormmull Zure wormmull Kleihydromull Zure wormmull Zure wormmull Beekhydromull Beekhydromull Beekhydromull Beekeerdmull Zure wormmull Kleieerdmull Zure wormmull Zure wormmull Zure wormmull Wormeerdmull Wormeerdmull Zure wormmull schrale Zure wormmull zandige Kleihydromull zandige Beekhydromull schrale Zure wormmull Beekhydromull Kleihydromull Beekhydromull Vlakhydromull Vlakhydromull Kleihydromull Kalkwormmull Zandhydromull Kleihydromull Kleihydromull schrale Zure wormmull zandige Zure wormmull
lichte klei zware zavel zware zavel zware zavel zware zavel zware zavel zware zavel lichte klei lichte klei matig lichte zavel matig lichte zavel zware zavel zware zavel zware zavel matig lichte zavel matig lichte zavel matig lichte zavel kleiig zand? kleiig zand? zware zavel zware zavel matig lichte zavel zware zavel zware zavel zware zavel zware zavel zware zavel lichte klei lichte klei zware zavel lichte klei zware zavel matig lichte zavel zware zavel zware zavel zware zavel zware zavel zware zavel zware zavel zware zavel lichte klei lichte klei zware zavel lichte klei zware zavel zware zavel matig lichte zavel zware zavel lichte klei zware zavel zeer lichte zavel kleiig zand zware zavel zware zavel zware zavel zware zavel zware zavel matig lichte zavel matig lichte zavel zware zavel matig lichte zavel matig lichte zavel zware zavel zware zavel matig lichte zavel zware zavel matig lichte zavel zeer lichte zavel zeer lichte zavel zware zavel zeer lichte zavel matig lichte zavel matig lichte zavel kleiarm zand kleiarm zand zeer lichte zavel zeer lichte zavel zeer lichte zavel zware zavel zware zavel zware zavel zware zavel zware zavel zware zavel matig lichte zavel zware zavel zware zavel zeer lichte zavel Kleiig zand
Clay % 15 12 12 14 10 21 12 15 12 12 12 14 14 18 16 14 14 8 7 24 18 26 18 19 18 18 8 14 17 15 18 10 8 10 8 10 3 9 14 17 18 17 10 4 4 13 6 15 4 15 16 10 18 17 15 16 20 20 9 16 20 8 10 8 9 10 12 10 17 16 26 20 23 22 17 20 25 16 12 19 19 20 13 17 10 8 8 18 18 17 20 21 22 23 24 25 25 18 25 18 16 22 20 18 20 22 20 18 25 26 23 25 20 22 17 20 25 24 12 7 20 21 19 18 24 15 15 20 17 12 19 19 14 20 17 10 12 18 10 16 14 4 4 10 10 10 18 20 18 18 21 19 17 18 19 12 7
Silt % 50 50 40 45 40 60 40 55 50 45 40 50 55 60 40 50 50 35 35 65 50 65 50 65 60 50 40 40 50 45 60 40 35 40 40 35 16 35 45 50 55 50 30 20 30 55 25 45 20 45 50 40 50 60 50 50 60 60 40 45 50 40 45 50 40 40 45 45 50 50 65 55 60 60 50 60 60 50 40 55 60 60 45 60 40 30 40 55 55 50 65 60 65 50 60 60 65 55 65 55 60 60 55 50 60 60 55 55 65 60 65 60 60 65 60 60 65 65 40 40 55 60 45 40 60 60 45 60 50 50 55 60 55 60 40 45 40 65 40 60 70 20 15 30 45 30 55 55 50 50 60 55 50 60 60 40 30
Loam % 65 62 52 59 50 81 52 70 62 57 52 64 69 78 56 64 64 43 42 89 68 91 68 84 78 68 48 54 67 60 78 50 43 50 48 45 19 44 59 67 73 67 40 24 34 68 31 60 24 60 66 50 68 77 65 66 80 80 49 61 70 48 55 58 49 50 57 55 67 66 91 75 83 82 67 80 85 66 52 74 79 80 58 77 50 38 48 73 73 67 85 81 87 73 84 85 90 73 90 73 76 82 75 68 80 82 75 73 90 86 88 85 80 87 77 80 90 89 52 47 75 81 64 58 84 75 60 80 67 62 74 79 69 80 57 55 52 83 50 76 84 24 19 40 55 40 73 75 68 68 81 74 67 78 79 52 37
Aftermath grazing 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 1 1 1 1 1 1 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 0 0
Number of mowings 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 0 0 0 0 0 2 2 1 1 1 1 2 2 2 2 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 1 1 1 1 1 1 1 1 1 1 1 1 2 2 1 1 2 2 2 2 2 2 0 0 2 2 2 2 2 2 2 2 2 2 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 2
Management G1 G1 G1 G1 G1 G1 G1 G1 G1 G2 G2 G2 G2 G2 G2 G0 G0 G0 G0 G0 G2 G2 G1 G1 G1 G1 N2 N2 N2 N2 N2 N2 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 N2 N2 N2 N2 N2 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 N2 N2 N2 N2 N2 N2 N2 N2 N2 N1 N1 N1 N1 N1 N1 N1 G1 N1 N1 G1 G1 G2 G2 G1 G1 G2 G2 G2 G2 G2 G2 G0 G0 G2 G2 G2 G2 G2 G2 G2 G2 G2 G2 G2 G2 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 G1 N1 G1 G1 G1 G1 G1 G1 N2 N2
Mowing date Number of
(first allowed plant
date)
species
25.6
34
15.6
23
15.6
24
25.6
31
25.6
33
25.6
31
25.6
30
25.6
28
15.6
31
15.6
15
15.6
21
15.6
26
15.6
12
15.6
20
15.6
15
32
32
34
34
28
20.5
22
20.5
20
25.6
37
25.6
31
1.7
34
1.7
36
15.6
40
15.6
38
15.6
40
15.6
44
15.6
37
15.6
41
15.6
32
15.6
28
15.6
35
15.6
30
15.6
30
15.6
27
15.6
33
15.6
40
15.6
17
15.6
24
15.6
22
15.6
17
1.7
23
1.7
31
1.7
26
1.7
33
1.7
29
1.7
30
1.7
30
1.7
31
1.7
31
1.7
34
1.7
34
15.6
25
15.6
28
15.6
23
15.6
36
15.6
32
15.6
31
15.6
32
15.6
35
25.6
41
25.6
43
25.6
41
25.6
40
25.6
39
1.7
43
1.7
44
1.7
35
1.7
43
1.7
29
15.6
36
15.6
40
15.6
23
15.6
20
15.6
26
15.6
40
15.6
42
15.6
35
15.6
24
1.8
44
1.8
36
1.8
41
15.6
36
15.6
32
15.6
40
15.6
42
15.6
27
20.7
53
20.7
43
1.8
45
1.8
44
1.6
28
1.6
41
1.7
27
1.7
33
20.6
33
20.6
27
15.6
24
15.6
25
20.6
26
20.6
24
31
32
15.6
34
15.6
26
15.6
37
15.6
37
15.6
33
15.6
34
15.6
33
15.6
35
15.6
27
15.6
30
15.6
30
15.6
30
15.6
25
15.6
17
1.7
40
1.7
36
1.7
36
15.7
33
15.7
25
1.7
37
1.7
30
1.7
45
1.7
37
1.7
25
15.7
30
15.7
23
15.7
32
15.7
28
1.7
24
1.7
34
15.7
29
15.7
34
15.7
35
1.7
26
1.7
32
1.7
26
1.7
32
1.7
25
1.7
37
1.7
20
1.7
46
1.7
34
1.7
24
1.7
27
1.7
29
15.6
27
15.6
29
15.6
35
15.6
28
15.6
34
15.6
29
15.6
35
15.6
29
15.6
35
15.6
19
15.6
21

MR Simmelink, JAM Janssen, JHJ Schaminée

File: habitat-and-management-preference-of-bromus-racemosus-l-a-rare.pdf
Author: MR Simmelink, JAM Janssen, JHJ Schaminée
Published: Tue Sep 19 19:31:04 2017
Pages: 28
File size: 1.21 Mb


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