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Tropical Bryology 31:14-21, 2010

Using data of bryophyte mapping projects for nature

conservation purposes - a case study from Saxony

Frank Müller

Institut für Botanik, Technische Universität Dresden, Mommsenstr. 13, D-01062 Dresden, Germany

(frank.mueller@tu-dresden.de)

Abstract: The results of a bryophyte mapping project carried out in Saxony between 1980 and 2004 were

evaluated in cases of nature conservation. The following questions were analysed: the relation of frequency and

threat of species; the threat of species in different biotope types; differences in the ecological characteristics of

Red List species in comparison with unthreatened species using the Ellenberg’s indicator values and the

hemerobie status; main reasons for decline and threat of Red List species; changes in the classification in the

different editions of the Red Lists of Saxony and their reasons, mainly the decline and increase of species; the

allocation of distribution centres of threatened bryophytes of selected biotope types.

Keywords: Nature conservation, Red List, bryophytes, Saxony.

Introduction

In the years from 1980 to 2004 in Saxony a bryophyte

mapping project was carried out. The results,

distribution maps and comments to distribution,

ecology, threat, and protection for each species were

published in Müller (2004). The occurrence of the

bryophytes was recorded using grids with a cell size

of approximately 5*5 km. Altogether the bryophyte

inventory of 636 grids was investigated. In addition,

all available historic records were included. To get

this information herbarium specimens, literature

records and unpublished records of registers were

evaluated. In Saxony in the past years the threat of

bryophytes was regularly evaluated by Red Lists

(Borsdorf & Müller 1991, Müller 1998, Müller 2004,

Müller 2008). Unfortunately, the results of such

mapping projects are seldom thoroughly evaluated in

terms of conservation issues, although the analysis

options are very high.

In the following few examples the possibilities of the

evaluation of data of mapping projects and Red Lists

is demonstrated by the example of Saxony. This can

be addressed only on selected questions.

For a better understanding of the subject it is

particularly important to note that the evaluation of

the threat of species and the creation of Red Lists in

Germany are not made according to the guidelines of

the IUCN. In Germany for this purpose there is used

its own methodology developed by a team of experts

from the Federal Agency for Nature Conservation

(Ludwig et al. 2005). There are characteristics of the

new IUCN Criteria System (IUCN 2001, 2003)

hampering its application in Germany. Among this is

the IUCN criteria “rarity as an extinction risk". A rare

species does not necessarily be an endangered species.

In Germany there is used for the creation of Red Lists

a four-level system including the factors current

situation, long-term trend, short-term trend, and risk

factors (Ludwig et al. 2006). The evaluation of the

species of Saxony has been done according to these

regional guideline and not according the IUCN

criteria.

Statistical analysis was deliberately omitted, since

there are regardless of the good mapping stand many

uncertainties (mapping gaps in terms of the historic

distribution, taxonomic problems, low staff for

mapping). For these reasons in many cases only an

expert's classification of the species to Red List

categories was possible to done. But why it prohibits

supplement the results obtained by statistics.

Relation of frequency and threat of species

In Figure 1 the bryophytes recorded in Saxony are

arranged in groups in relation of their frequency. In

addition, in each group the share of Red List species

and unthreatened species is given. The general

information of this figure is: the rarer a species the

higher is their threat. Nearly all of the very rare

species recorded in only one to four grids are assigned

as endangered, whereas all species recorded in about

more as half of the grids of Saxony are all

unthreatened.

TROPICAL BRYOLOGY 31 (2010)


species number

350

300

250

200

150

100

50

0

MÜLLER: BRYOPHYTE MAPPING PROJECTS FOR NATURE CONSERVATION PURPOSES

1‐4 5‐20 21‐40 41‐300 301‐500 501‐636

frequency

0 1 2 3 R unthreatened

Figure 1. Correlation between frequency of the

bryophytes of Saxony and threat. The frequency is

ordered in groups according the number of grids with

records. Abbreviations: 0 – extinct, 1 – critically

endangered, 2 – endangered, 3 – vulnerable, R – rare.

The most frequent species in Saxony are Dicranum

scoparium, Brachythecium salebrosum, Aulacomnium

androgynum, Eurhynchium praelongum,

Rhytidiadelphus squarrosus, Brachythecium albicans,

Plagiomnium affine, Mnium hornum, Lophocolea

heterophylla, Polytrichum formosum, Tortula muralis,

Amblystegium serpens var. serpens, Bryum

argenteum, Brachythecium velutinum, Atrichum

undulatum, Dicranella heteromalla, Hypnum

cupressiforme var. cupressiforme, Pohlia nutans,

Brachythecium rutabulum, and Ceratodon purpureus.

The last one is the commonest one recorded in nearly

all of the investigated grids.

The threat of species in different biotope types

The information of the ecology of each species were

evaluated and to each species the mean biotope types

of their occurrences were assigned (Fig. 2). Due to

multiple selections the total sum exceeds the number

of the 712 species recorded in Saxony. Most species,

nearly 70%, are able to inhabit rock and immature soil

biotopes. In Saxony these formations play the most

important role for bryophyte biodiversity. A

comparative high proportion of bryophytes are

characteristic for forest biotopes. All the other biotope

types are characterized by relative low species

diversity. The species inventory of the biotope types

“bogs and fens”, “bodies of standing water”, and

“bodies of flowing waters” is limited, but contain a

high proportion of specialized species, whereas the

inventory of the biotope types “settlements,

infrastructure and industrial areas’, “arable land,

gardens, areas with specialised cultivations”,

“heathlands and neglected grassland”, “populations of

perennial herbs”, “grassland”, “shrubberies,

hedgerows and bosks” are also limited, but contains

mostly species with a wide ecological range.

It should be noted here that a part of the bryophytes

recorded in Saxony show a very limited ecological

TROPICAL BRYOLOGY 31 (2010)

range and some are only known from man-made

substrates. Only on open soil on paths were recorded

the species Bryum oblongum, Bryum knowltonii (the

recent record), Campylopus subulatus, Jungermannia

paroica, and Meesia uliginosa (the recent record).

Only from walls there are known Didymodon

cordatus (the recent records), Didymodon luridus,

Didymodon umbrosus, Ditrichum flexicaule (the

recent record), Grimmia crinita, Isopterygiopsis

pulchella, Jungermannia confertissima, Scapania

cuspiduligera, and Tortella densa. Restricted in their

occurrence to mining areas, e. g. quarries, opencast

pits, there are Aloina aloides, Bryum funckii (the

recent records), Bryum radiculosum, Cephaloziella

massalongi, Cephaloziella phyllacantha, Dicranella

crispa, Lophozia guttulata, Nardia insecta,

Palustriella commutata var. falcata (the recent

record), Pohlia tundrae, and Riccia ciliata (the recent

record).

Figure 3 shows the share of unthreatened and Red List

species for each mean biotope type. The share of Red

List species is remarkably high in bog and fen

biotopes. More then two-thirds of the species

occurring in this biotope type are listed in the Red

List. In the biotope types “forests”, “rock and

immature soil biotopes”, “bodies of standing water”,

“bodies of flowing waters”, “heathlands and neglected

grassland” the share of Red List species and

unthreatened species is nearly identical. The share of

Red List species is significantly low in the biotope

types “settlements, infrastructure and industrial

areas”, “arable land, gardens, areas with specialised

cultivations”, “populations of perennial herbs”,

“grassland”, and “shrubberies, hedgerows and

bosques”.

Differences in the ecological characteristics of Red

List species in comparison with unthreatened

species

As one example to estimate the different ecological

amplitude of Red List species in comparison with

unthreatened species the Ellenberg’s indicator values

of both groups were compared (Ellenberg et al. 2002).

The Ellenberg's indicator values are simple ordinal

classes of plants with a similar realized ecological

niche along a gradient. The latest edition of

Ellenberg's indicator values contains values on a 9

point scale for soil acidity, soil humidity,

continentality, temperature, and light.

The evaluation of the diagrams (Figs. 4–8) shows that

in all five examined indicator values differences in in

the spectrums between Red List species and

unthreatened species exist. Under extreme habitat

conditions the proportion of Red List species in

comparison with unthreatened species is usually

higher. Under average habitat conditions, the

15


16

arable land, gardens, areas with specialised cultivati ons

biotope type

settlements, infrastructure and industrial areas

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rock and immature soil biotopes

heathland and neglected grassland

populations of perennial herbs

grassland

bogs and fens

bodies of standing water

bodies of flowing water

shrubberies, hedgerows and bosks

forests

42

76

79

100

98

112

126

127

159

325

species number

Figure 2. Species number of bryophytes in the eleven

main biotope types of Saxony.

species number in %

45

40

35

30

25

20

15

10

5

0

T1 T2 T3 T4 T5 T6 T7 T8 T9

indicator value for temperature

unthreatened species red list species

Figure 4. Indicator values for temperature of Red List

bryophytes in comparison with unthreatened

bryophytes in Saxony.

species number in %

25

20

15

10

5

0

R1 R2 R3 R4 R5 R6 R7 R8 R9

indicator value for soil acidity

unthreatened species red list species

Figure 6. Indicator values for soil acidity of Red List

bryophytes in comparison with unthreatened

bryophytes in Saxony.

523

species number

300

250

200

150

100

50

0

180

forests

143

50

shrubberies, hedgerows and bosks

28

54 57

bodies of flowing water

38 37

bodies of standing water

50

bogs and fens

108

73

biotope type

grassland

52

TROPICAL BRYOLOGY 31 (2010)

39

populations of perennial herbs

2

48 47

heathland and neglected grassland

unthreatened species red list species

263

253

Figure 3. Comparison of the share of unthreatened

and Red List species for each of the eleven main

biotope types of Saxony.

species number in %

30

25

20

15

10

5

0

L1 L2 L3 L4 L5 L6 L7 L8 L9

indicator value for light

unthreatened species red list species

Figure 5. Indicator values for light of Red List

bryophytes in comparison with unthreatened

bryophytes in Saxony.

species number in %

35

30

25

20

15

10

5

0

F1 F2 F3 F4 F5 F6 F7 F8 F9

indicator value for soil humidity

unthreatened species red list species

Figure 7. Indicator values for soil humidity of Red

List bryophytes in comparison with unthreatened

bryophytes in Saxony

rock and immature soil biotopes

67

arable land, gardens, areas with specialised

cultivati ons

32

103

settlements, infrastructure and industrial areas

21


species number in %

45

40

35

30

25

20

15

10

5

0

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K1 K2 K3 K4 K5 K6 K7 K8 K9

indicator value for continentality

unthreatened species red list species

Figure 8. Indicator values for continentality of Red

List bryophytes in comparison with unthreatened

bryophytes in Saxony.

succession

drainage

intensive forestry

eutrophication of waterbodies and bogs

intensive utilisatio n of grassland

measures of rock and slope protection

decline of humidity

air pollution

unexpected events

hydro‐engineering measures

specifics of species and areal/biological risk factors

clean‐up of walls

diffuse nutrient contaminatio n and eutrophication

elemination of trees

removal by science and teaching

mining

intensive fish farming

soil sealing and house building

deletion of stone walls and drywalls

intensive agriculture

tourism

flood

changes of macroclimate

reason of decline unknown

TROPICAL BRYOLOGY 31 (2010)

8

6

6

6

23

19

41

40

38

34

34

33

75

69

66

65

61

59

percentage of species

40

35

30

25

20

15

10

5

0

Hemerobie

red list species unthreatened species

Figure 9. Hemerobic status of Red List bryophytes

in comparison with unthreatened bryophytes in

Saxony.

87

98

species number

0 50 100 150 200 250

Figure 10. Reasons for the decline and the threat of the Red List bryophytes of Saxony.

proportion of unthreatened species in comparison

with endangered species is higher. The Red List

species are often specialists restricted in their

distribution to extreme habitats, such as very dry or

very wet sites, very base rich soils, poor alkaline soils,

very shaded sites, very light-rich sites, very cold sites,

very warm sites, sites with atlantic climate, and sites

with continental climate. The conservation of such

extreme habitats is therefore of particular importance

to protect endangered mosses.

In addition, differences between both groups exist in

the hemerobie (Fig. 9). The ratio of a-hemerob and

oligo-hemerob species, that means species

characterized for habitats with none or low influence

by man, is higher among the Red List species,

whereas the share of meso-hemerob, eu-hemerob and

113

109

129

poly-hemerob is higher among the unthreatened

bryophytes.

Reasons for decline and threat of species

For each species listed in the Red List of Saxony the

main reasons for decline and threat were analysed

(Fig. 10). In a few cases these reasons are unknown.

The most important factor in this relation is the

succession of habitats. A high proportion of

bryophytes is characteristic for open habitats not

colonized by vascular plants. These habitats are often

of short life, and will, if management measures are

stopped, become colonised by vascular plants which

will displace the bryophytes. Regularly measures of

biotope conservation are e.g. essential for the biotope

types arid and semi-arid grasslands, grassland, fen,

193

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MÜLLER: BRYOPHYTE MAPPING PROJECTS FOR NATURE CONSERVATION PURPOSES

and heathland to stop the succession and to conserve

the characteristic species inventory.

To rank second there is drainage as a further reason

for threat. By this action, especially the many species

of moist sites are negatively affected. Another

important reason of threat is intensive forestry. This

has especially negative effects on species of rotten

wood, species of matured forest, epiphytic species and

species characterized for humid forests.

Changes in the classification in the different

editions of the Red Lists of Saxony

Since 1991, the threat of the bryophytes of Saxony is

documented by different editions of Red Lists

(Borsdorf & Müller 1991, Müller 1998, Müller 2004,

Müller 2008). Figure 11 gives an overview of the

development of the shares of the different categories

of the Red Lists over the years. By the evaluation of

these data it is important to note that the values from

2008 are not directly comparable with the values of

the editions of the previous years since they were

created using a new methodology (Ludwig et al.

2006).

Over the years there can be seen an increase of rare

species, but this increase does not reflect the real

situation of threat. It is more caused by an

intensification of the research of the bryophyte flora

of Saxony during the last years whereby many species

originally classified as extinct were rediscovered.

Among the species of the category critically

endangered an increase in the number of species is

noticeable. Many species originally classified in the

category rare had to be regrouped in category

critically endangered, because their habitats are more

species number

160

140

120

100

80

60

40

20

0

0 1 2

red list category

3 R G

1991 1998 2004 2008

Figure 11. Changes in the numbers of bryophytes of

the different Red List categories in the different

editions of the Red Lists of Saxony (Borsdorf &

Müller 1991, Müller 1998, Müller 2004, Müller

2008). Abbreviations: 0 – extinct, 1 – critically

endangered, 2 – endangered, 3 – vulnerable, R – rare,

G – indeterminate.

threatened then previously thought. This applies

particularly for many rare rock bryophytes which are

more threatened in recent years by route protection

measurements. Bryophytes on siliceous rocks are

further more threatened in recent years by

measurements of liming of forests.

Positive developments have mainly arisen due to an

improvement of air quality through measures of

prevention of air pollution (Tab. 1). Saxony belonged

until 1990 to the areas of Germany most affected by

acidic air pollution, so that especially the epiphytic

bryophytes have been strongly limited in their

distribution. In the last years a lot of epiphytic species

were able to recolonise the area in relation to

improvement of air quality. A sample of species of

this ecological group, e. g., Orthotrichum pulchellum,

O. rogeri, and Zygodon dentatus, were first

recognized in Saxony during the past years. A spread

of species and a downgrading of their threat in the

Red Lists in connection with a better air quality were

also noted for selected bryophytes on forest floor and

for selected rotten wood inhabitants.

An increase of threat often associated with a decrease

of localities or a decrease in population sizes, is

especially observed for calciphile bryophytes,

bryophytes of sources, species of bogs and fens, on

silicate rocks and of arid and semi-arid grasslands

(Tab. 2).

Distribution centres of threatened bryophytes of

selected biotope types

Coniferous forests. Pure coniferous forests in Saxony

are by nature rare. The natural range of the principal

coniferous trees, Pinus sylvestris and Picea abies, has

been substantially increased by the work of forestry,

so that nowadays in the Saxon forests conifers are the

dominant forest trees. Fig. 12 shows the weighted

distribution of species of the Red List of Saxony,

which are typical for coniferous forests. An

accumulation can be seen in the high altitude areas of

the western Erzgebirge. Weaker accumulations are

situated in the central Erzgebirge, in the Saxon

Switzerland, and in the upper Vogtland. In this

context, it is interesting to point out, that the centres

of occurrences of rare and endangered species of

coniferous forests are located mainly in areas with

natural coniferous forests. These are in the areas

mentioned above high altitude forests of Norway

spruce. Threatened species with their centres of

distribution in this habitat are e. g., Anastrepta

orcadensis, Barbilophozia kunzeana, Scapania

umbrosa, and in wet forests Sphagnum affine, S.

riparium, Splachnum sphaericum, and S.

ampullaceum.

The pine forests on sand dunes in the lowland of the

Upper Lusatia are not characterised by a particular

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MÜLLER: BRYOPHYTE MAPPING PROJECTS FOR NATURE CONSERVATION PURPOSES

Table 1. Decline of bryophytes in Saxony as

expressed by the different evaluation of their threat in

the different editions of the Red Lists of Saxony

(Borsdorf & Müller 1991, Müller 1998, Müller 2004,

Müller 2008). Abbreviations: 0 – extinct, 1 – critically

endangered, 2 – endangered, 3 – vulnerable, R – rare,

V – near threatened, - – not recorded, G –

indeterminate, * – least concern, n.e. – not evaluated.

TROPICAL BRYOLOGY 31 (2010)

Red List of Saxony 1991

Red List of Saxony 1998

Distribution atlas 2004

Species of calcareous habitats

Amblystegium confervoides R 1 1 1

Anomodon longifolius 3 R R 1

Bryum funckii R R 1 1

Conardia compacta - R R 3

Eucladium verticillatum R R R 2

Palustriella commutata var. commutata 3 2 2 2

Philonotis calcarea 3 2 2 1

Rhynchostegiella teneriffae

Species of spring habitats

R R R 1

Hookeria lucens 2 1 1 1

Philonotis seriata

Species of bogs

3 2 2 2

Cephalozia pleniceps 3 G 0 0

Sphagnum fuscum

Species on rocks

2 1 1 1

Cynodontium tenellum 0 R 1 1

Grimmia laevigata * 3 2 2

Grimmia longirostris n.e. R 3 2

Pohlia elongata * 2 1 1

Grimmia ovalis * * 3 3

Reboulia hemisphaerica 1 R 1 1

Tortula atrovirens

Species of arid and semi-arid grasslands

R 1 1 1

Entodon concinnus 2 1 1 1

Rhytidium rugosum 2 1 1 1

Thuidium abietinum R 2 2 2

Red List of Saxony 2008

Table 2. Increase of bryophytes in Saxony as

expressed by the different evaluation of their threat in

the different editions of the Red Lists of Saxony

(Borsdorf & Müller 1991, Müller 1998, Müller 2004,

Müller 2008). Abbreviations: 0 – extinct, 1 – critically

endangered, 2 – endangered, 3 – vulnerable, R – rare,

V – near threatened, - – not recorded, G –

indeterminate, * – least concern, n.e. – not evaluated.

Red List of Saxony 1991

Red List of Saxony 1998

Distribution atlas 2004

Epiphytes

Dicranoweisia cirrata 3 * * *

Dicranum flagellare 2 3 3 V

Dicranum montanum 3 * * *

Frullania dilatata 1 1 3 3

Hypnum pallescens 0 0 2 2

Orthotrichum affine 1 2 * *

Orthotrichum lyellii 0 0 3 *

Orthotrichum obtusifolium 1 1 3 *

Orthotrichum pallens 0 0 3 *

Orthotrichum patens 0 - 3 *

Orthotrichum pulchellum - - 3 *

Orthotrichum pumilum 1 2 * *

Orthotrichum rogeri - - 1 *

Orthotrichum scanicum 0 - 1 3

Orthotrichum speciosum 0 0 3 *

Orthotrichum stramineum 1 1 3 *

Orthotrichum striatum 1 1 3 *

Orthotrichum tenellum 0 0 2 *

Pterigynandrum filiforme 2 3 3 V

Ptilidium pulcherrimum 2 3 3 *

Pylaisia polyantha 1 2 3 V

Radula complanata 2 2 3 V

Tortula papillosa 0 0 3 V

Ulota bruchii 0 2 3 *

Ulota coarctata 0 0 1 1

Ulota crispa s.str.

Species on rotten wood

0 1 3 *

Riccardia latifrons

Species on the forest floor and on slopes

0 1 2 2

Buxbaumia aphylla 2 3 3 3

Hylocomium splendens 2 3 3 V

Hylocomium umbratum 0 0 1 2

Plagiothecium undulatum 2 3 3 *

Ptilium crista-castrensis 1 2 3 V

Rhytidiadelphus loreus 1 2 3 *

Rhytidiadelphus triquetrus

Species on rocks

2 2 3 V

Hedwigia ciliata var. ciliata 2 3 * *

19

Red List of Saxony 2008


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Figure 12. Weighted frequence of Red List species

of the biotope type coniferous forest in Saxony.

Grid records of critically endangered species are

included in the calculation with the factor three,

records of endangered and rare species with the

factor two, records of vulnerable species with the

factor one, and records of near threatened species

with the factor 0.5.

Figure 14. Weighted frequence of Red List species

of the biotope type calcareous fen in Saxony. Grid

records of critically endangered species are

included in the calculation with the factor three,

records of endangered and rare species with the

factor two, records of vulnerable species with the

factor one, and records of near threatened species

with the factor 0.5.

richness of rare or Red List species. The higher values

of individual grids in this area are based in particular

by a few occurrences of Dicranum spurium.

Bogs. Focus for the protection of bog bryophytes in

Saxony are the high altitude areas of the western and

central Erzgebirge (Fig. 13). In the high altitude areas

of the eastern Erzgebirge, characterised by lower

precipitation, bogs are less common and the condition

of the few remaining ones is not optimal due to

different negative influences in the past (drainage,

peat cutting, high sulphur dioxide emissions).

Correspondingly lesser is in them the proportion of

Figure 13. Weighted frequence of Red List species of

the biotope type bog in Saxony. Grid records of

critically endangered species are included in the

calculation with the factor three, records of

endangered and rare species with the factor two,

records of vulnerable species with the factor one, and

records of near threatened species with the factor 0.5.

Figure 15. Weighted frequence of Red List species of

the biotope type arid and semi-arid grasslands in

Saxony. Grid records of critically endangered species

are included in the calculation with the factor three,

records of endangered and rare species with the

factor two, records of vulnerable species with the

factor one, and records of near threatened species

with the factor 0.5.

endangered species. Outside the high altitude areas of

the Erzgebirge there exists three areas in the Saxon

lowland with a higher proportion of threatened bog

species: the Dübener Heide with the bogs

Zadlitzbruch and Wildenhainer Bruch, the

Königsbrück-Ruhlander Heide with a high quantity of

small bogs, and the Muskauer Heide with a couple of

bogs situated in dune valleys.

Calcareous fens. Calcareous fens were in the Saxony,

an area mainly characteristic by soils with insufficient

base nutrients, never very common. By melioration,

intensification of land use and afforestation, they

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MÜLLER: BRYOPHYTE MAPPING PROJECTS FOR NATURE CONSERVATION PURPOSES

belong nowadays to the rarest fen and bog biotopes of

Saxony. With the decline of the biotope type

underwent a decline of bryophytes typical for it.

Formerly more widespread species such as Paludella

squarrosa, Tomentypnum nitens, Hamatocaulis

vernicosus etc. are therefore now of absolute rarity

and their survival is in extreme danger. The few

remaining remnants of this biotope type are usually

characterized by the occurrence of a whole set of

highly endangered species on a very limited space.

Most of the reported remaining calcareous fens are

located in the mountainous areas of the Erzgebirge

and Vogtland (Fig. 14). In the Saxon lowland one fen

in the Upper Lusatian Heath and Pond area is

conspicuous by a particular richness of species of this

ecological group.

Arid and semi-arid grasslands. Arid and semi-arid

grasslands are in Saxony in comparison with

neighbouring regions (Bohemia, Thuringia,

Brandenburg) relatively rare and are mostly

developed in only small quantities. This is due mainly

to the fact that in Saxony extremely low rainfall areas

(at or below 500 mm rainfall per year) will reach

nowhere. The region of Saxony characterized by the

least precipitation, with rainfall about 550 mm per

year, is the Elbe Valley and its side valleys, especially

the Ketzerbachtal, below Meissen. The occurrence of

loess is another reason for the enrichment of the

existing arid and semi-arid grasslands of this area

with rare short-living bryophytes (Fig. 15). In this

area there is one of two occurrences of Hilpertia

velenovskyi in Germany (Müller 2000), along with

sites of other characteristic species, e. g.,

Pterygoneurum subsessile, P. lamellatum, Acaulon

triquetrum, and Aloina ambigua.

The other small areas characterized on the map by the

occurrence of threatened bryophytes of this biotope

type are small limestone areas with occurrences of

bryophytes of limestone arid and semi-arid

grasslands, for example with Entodon concinnus,

Rhytidium rugosum, and Thuidium abietinum. The

occurrences of these species are less connected with

low precipitation, rather than with the limestone

substrate as subsoil.

TROPICAL BRYOLOGY 31 (2010)

References

Ellenberg, H., H. E. Weber, R. Düll, V. Wirth & W.

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