stellaria nemorum l. and s. montana pierrat ( caryophyllaceae ) in the forest communities of...

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Folia Geobotanica 34: 115-125, 1999 STELLARIA NEMORUM L. AND S. MONTANA PIERRAT (CARYOPHYLLACEAE) IN THE FOREST COMMUNITIES OF SLOVENIA Igor Dakskobler, Andrej Seli~kar 1) & Branko Vre,~ 2) Institute of Biology, Centre for Scientific Research of the Slovenian Academy of Sciences and Arts, Novi trg 5, SI-1000 Ljubljana, Slovenia; tel. +386 61 1256 068, fax +386 61 1257 797, E-mail 1) [email protected]; 2) [email protected] Keywords: Distribution, Phytosociology, Stellaria nemorum group, Taxonomy,Forest vegetation Abstract: Morphological and coenological differences between closely related species Stellaria nemorum L. and S. montana PIERRAT are described. Stellaria montana is frequent in Slovenia and occurs mostly in the montane belt (altitude 600-1200 m). It has been recorded in thirty two forest communities (associations or lower syntaxa). Although widely distributed in Europe S. nemorum is more rare in Slovenia. It occurs mostly in upper-montane(altimontane) and subalpine belt (altitude 1200-1600 m) in ten forest or shrubby communities. Both, S. nemorum and S. montana are valuable diagnostic species for certain associations. They characterize sites with fresh soil, rich in nitrogen. As an example we present two syntaxa of beech forests of Slovenia, in which the species of the S. nemorum group have high constancy and cover value. INTRODUCTION In the present paper, two closely related species, Stellaria nemorum L. s. str. and S. montana PIERRAT,are discussed in connection with their relations to the forest communities of Slovenia. The aim of this work has been to search for new morphological characteristics of both species, their chorology and the relation of these plants in two beech forest communities, using the BRAUN-BLANQUET(1964) and MATUSZKmWICZ & MATUSZrdEWICZ(1981) approach. Stellaria nemorum was first described by LINNAEUS (1753). In 1880, the related species SteIlaria montana PIERRAT was described (PtERRAT 1880), and later (MURBECK 1891) the subspecies Stellaria nemorum subsp, glochidisperma MURB. (= S. glochidisperma (MuRB.) FREYN). In the past, they were subject to changes in their names as well as taxonomic rank (FREYN 1892, HEGI 1911, PETERSON 1936). Recent reports (GREEN 1954, ANDREAS 1956, KOOPMANS & SUM 1968, VRE~ 1991, BABIJ et al. 1997) show that both S. glochidisperma and S. montana represent the same taxon, which should be strictly distinguished from S. nemorum L. s. str. In the rank of species, the oldest valid name is Stellaria montana P1ERRAT (= S. nemorum subsp, montana (PIERRAT) BERHER). In Slovenia, recent chorological data show that Stellaria montana is more frequent than S. nemorum (Fig. I). In contrast, a different situation has been observed in relation to their whole distribution range in Europe. Throughout Europe, S. montana occurs rarely and is concentrated in Denmark and on the Balkan peninsula, being more or less dispersed elsewhere (JALAS & SUOMINEN1983). On the other hand, S. nemorum is a widely spread species. According to the herbarium (LJU, LJM), floristic and phytosociological data (e.g. MARIN~EK& DAKSKOBLER1988, MARINCEK et al. 1989, VRE~ 1991, ZUPANCI~ 1980), Stellaria

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Page 1: Stellaria nemorum  L. and  S. montana  Pierrat (  Caryophyllaceae  ) in the Forest Communities of Slovenia

Folia Geobotanica 34: 115-125, 1999

STELLARIA NEMORUM L. AND S. MONTANA PIERRAT (CARYOPHYLLACEAE) IN THE FOREST COMMUNITIES OF SLOVENIA

Igor Dakskobler, Andrej Seli~kar 1) & Branko Vre,~ 2)

Institute of Biology, Centre for Scientific Research of the Slovenian Academy of Sciences and Arts, Novi trg 5, SI-1000 Ljubljana, Slovenia; tel. +386 61 1256 068, fax +386 61 1257 797, E-mail 1) [email protected]; 2) [email protected]

Keywords: Distribution, Phytosociology, Stellaria nemorum group, Taxonomy, Forest vegetation

Abstract: Morphological and coenological differences between closely related species Stellaria nemorum L. and S. montana PIERRAT are described. Stellaria montana is frequent in Slovenia and occurs mostly in the montane belt (altitude 600-1200 m). It has been recorded in thirty two forest communities (associations or lower syntaxa). Although widely distributed in Europe S. nemorum is more rare in Slovenia. It occurs mostly in upper-montane (altimontane) and subalpine belt (altitude 1200-1600 m) in ten forest or shrubby communities. Both, S. nemorum and S. montana are valuable diagnostic species for certain associations. They characterize sites with fresh soil, rich in nitrogen. As an example we present two syntaxa of beech forests of Slovenia, in which the species of the S. nemorum group have high constancy and cover value.

INTRODUCTION

In the present paper, two closely related species, Stellaria n e m o r u m L. s. str. and S. montana

PIERRAT, are discussed in connection with their relations to the forest communities of Slovenia. The aim of this work has been to search for new morphological characteristics of both species, their chorology and the relation of these plants in two beech forest communities, using the BRAUN-BLANQUET (1964) and MATUSZKmWICZ & MATUSZrdEWICZ (1981) approach.

Stellaria n e m o r u m was first described by LINNAEUS (1753). In 1880, the related species SteIlaria montana PIERRAT was described (PtERRAT 1880), and later (MURBECK 1891) the subspecies Stellaria n e m o r u m subsp, glochidisperma MURB. (= S. g lochid isperma (MuRB.) FREYN). In the past, they were subject to changes in their names as well as taxonomic rank (FREYN 1892, HEGI 1911, PETERSON 1936). Recent reports (GREEN 1954, ANDREAS 1956, KOOPMANS & SUM 1968, VRE~ 1991, BABIJ et al. 1997) show that both S. glochidisperma

and S. montana represent the same taxon, which should be strictly distinguished from S. n emorum L. s. str. In the rank of species, the oldest valid name is Stellaria montana P1ERRAT (= S. nemorum subsp, montana (PIERRAT) BERHER).

In Slovenia, recent chorological data show that Stellaria montana is more frequent than S. n emorum (Fig. I). In contrast, a different situation has been observed in relation to their whole distribution range in Europe. Throughout Europe, S. montana occurs rarely and is concentrated in Denmark and on the Balkan peninsula, being more or less dispersed elsewhere (JALAS & SUOMINEN 1983). On the other hand, S. nemorum is a widely spread species.

According to the herbarium (LJU, LJM), floristic and phytosociological data (e.g. MARIN~EK & DAKSKOBLER 1988, MARINCEK et al. 1989, VRE~ 1991, ZUPANCI~ 1980), Stellaria

Page 2: Stellaria nemorum  L. and  S. montana  Pierrat (  Caryophyllaceae  ) in the Forest Communities of Slovenia

1 16 I. Dakskobler et al.

48 47

91

92

98 i /~

48 48 50 51 52 53 54 55 55 57 58 59 140 Ils*

6061 6 2 6 3 6 4 6 5

16" ; "?-,.~ ,V .

,,~, S L O V E N I A

~ k_ ~ Stettat~a montana

0"2 ~ e "~- ~lr~ _ _ () b"t~lar/, r~rnorum

04°3 - ' ' - - - 30km

Fig. 1. The distribution ofStel laria nemorum L. and S. montana PIERRAT in Slovenia, according to the mapping

of flora of Central Europe.

n e m o r u m is considerably rare in Slovenia. It has been recorded in forest communities of montane, upper-montane, and subalpine belt (600 to 1600 m), mostly at altitudes between 1200 and 1600 (-1800) m. The altitudinal range of the forest communities in which S. montana

occurs extends from the submontane to the upper-montane belt (200 to 1400 m), the median being in the montane belt (600 to 1200 m) (Fig. 2).

MORPHOLOGICAL DIFFERENCES BETWEEN STELLARIA NEMORUM S. STR. AND S. MONTANA

In Slovenia, the morphological features of the Stellaria n e m o r u m group were studied during the previous ten years. The specimens of S. nemorum and S. montana from herbarium LJU and LJM were examined and measured (VP, E~ 1991). The species usually differ clearly in their habitus (Fig. 3) and some morphological characters (Tab. 1). The most important differentiating characters are the ratio between the length and the width of the leaf at the node below the first branching of inflorescence, the length and pubescence of the sepals, the length of the papillae on ripe seeds and the presence or absence of hooks on the papillae of ripe seeds. Both species can be satisfactorily distinguished on the basis of the combination of these features. In Stellaria n e m o r u m , the leaf at the node under the first branching of inflorescence is sessile, ovate, truncate to angustate at base, (1.7-)2-2.6(-3.2) times longer than wide (2.4 + 0.3 times in average). In S. montana, leaves are stalked, cordate with a cordate basis, and only (1.3-) 1.6-2(-2.4) times longer than wide (1.8 -+ 0.2 times in average). In S. montana , the bracts decrease abruptly in size after the first dichotomy (the second pair is usually less than 1/3 as long as the first), but in S. n e m o r u m they decrease gradually (the second pair is usually almost 1/2 as long as the first). The sepals of flowers are slightly longer in S. n e m o r u m (5.6-7 mm), with a nude apex (pubescent mostly only at the base), compared to S. mon tana (4.6-5.5 mm), which has a pubescent apex. The ripe seeds are covered with cylindrical or hemispherical papillae. In addition, the two species differ from each other in the length, the width and the shape of papillae (Fig. 4). In S. nemorum, the seeds are furnished

Page 3: Stellaria nemorum  L. and  S. montana  Pierrat (  Caryophyllaceae  ) in the Forest Communities of Slovenia

Stellaria nemorum and S. montana in the forest communities of Slovenia 117

2ooi-21oo 19Ol-2OOO

18o1-19oo 17o1-18oo

16o1-170o

15o1-16oo

14o1-15oo

13o1-14oo

12o1-13oo

11o1-12oo

~- 1oo1-110o

~ 901-1000 701--800 601-71~

401--500

101--200 I!-11111

-~.o

m m

m

m

m

l i b

m

ibm

I I S. nemorum

D S mom~a

]

]

]

-1

-~0.0 -15.0 -10.0 -5.0 0.0 5.0 10,0 15.0

Frequency (%)

20.0

Fig. 2. The distribution frequency of Stellaria montana and S. nemorum s. str. in Slovenia in correlation to altitude (data sources: literature, herbarium data), n=35 (S. montana), n=67 (S. nemorum).

with hemispherical unarmed papillae, being (0.3-)0.6-1.5(-3) times as long as wide (1.0 + 0.4 times in average), whereas in S. montana the seeds have long cylindrical papillae with barbate caps, being (1.2-)2-4(-12) times longer than wide (3.8 + 1.4 times in average).

THE COMPARISON OF BOTH SPECIES AFFINITY TO VEGETATION TYPES

Characteristic habitats of S. montana are in communities with Fraxinus excelsior, Acer pseudoplatanus, and Ulmus glabra. It is an important diagnostic species for the association Hacquetio-Fraxinetum MARINCEK in WALLNOFERet al. 1993 and for the suballiance Polysticho setiferi-Acerenion pseudoplatani BORHIDI et KEVEY 1996. This species also grows in humid forms of the montane and upper-montane beech, fir-beech, and maple-beech forests. It can be found in fir forests on colluvial soil, in humid moderately acidophilic fir-beech forests, or in the moderate freezing dolines and medium acidophilic spruce forests of the Dinaric phytogeographic region. It is very scarce in hornbeam forests and submontane beech forests.

Stellaria nemorum is recorded in the forest communities of the montane, upper-montane, and subalpine belt but mostly thrives in the subalpine maple-beech forests or in the more humid upper-montane and subalpine beech communities. In the Alpine and pre-Alpine phytogeographical region of Slovenia, it grows in the subalpine spruce forests or in shrubby communities of Alnus viridis on moderately fresh and acid soil.

THE PRESENCE OF STELLARIA N E M O R U M AND STELLARIA MONTANA IN THE FOREST COMMUNITIES OF SLOVENIA

Generally, the Stellaria nemorum group most frequently thrives in the communities of the order Fagetalia sylvaticae PAWLOWSKI et al. 1928. Stellaria nemorum s. str. is classified as a characteristic species of the alliance of the hygrophilic communities of central Europe (Alnion incanae PAWLOWSKt et al. 1928 (=Alno-Ulmion BR.-BL. et R. Tx. 1943), e.g.

Page 4: Stellaria nemorum  L. and  S. montana  Pierrat (  Caryophyllaceae  ) in the Forest Communities of Slovenia

1 18 I. Dakskobler et al.

Table 1. Mean, standard deviation, range (in brackets), in mm, number of observations of morphological characters of Stellaria nemorum L. s.str, and S. montana PIERRAT in Slovenia; all differences among samples are highly significant (P < 0.001).

Variable

Length of the first stem leaf under branching of inflorescence

Length of the second stem leaf under branching of inflorescence

Length of peduncle of the first stem leaf under branching of inflorescence

Length of peduncle of the second stem leaf under branching of inflorescence

Length of bracts in the first branching of inflorescence

Length of bracts in the second branching of inflorescence

Length of bracts in the third branching of inflorescence

Petal limb length

Calyx length

Fruit length

Peduncle length

Papillae length

Papillae width

Stellaria nemorum

66.7-+17.5 (27-115); n=93

66.8'19.4 (32-117); n=89

0.6-+1.8 (0-10); n=93

6.9+-8.1 (0-31); n=89

53.8~16.2 (15-92); n=90

30.5-+12.2 (4-64); n=185

13.4-+7.5 (2.5-42); n=303

11.0-+0.7 (9-12); n=59

6.1-+0.6 (4.5-7.8); n=535

8.5-+1.0 (6-11); n=260

24.2~6.3 (10-50); n=403

0.06-+0.02 (0.03-0.13 ); n=65 l

0.06--+0.01 (0.03-0.10); n=595

S. montana

47.5-+15.6 (11-90); n=95

47.5-+13.4 (17-82); n=95

18.7-+9.6 (2-50); n=94

29.9-+9.8 (10---60); n=95

43.3-+16.1 (6-85); n=95

13.0!--8.0 (1.7-36); n=156

3.4+2.0 (1-10); n=86

11.8-+1.5 (7.5-15); n=101

4.9-+0.5 (3.5-7); n=562

7.4+1.2 (4.5-10); n=284

19.5+7.7 (5-55); n--469

0.10~.02 (0.04-0.15); n=752

0.03-+0.01 (0.01-0.07); n=685

OBERDORb'ER (1983: 370-371), ELLENBERG (1991: 145), WALLNOFER et al. (1993: 91)). Stellaria mon tana has, due to its considerable distribution in south-eastern Europe, certain diagnostic value in mesophilic beech communities (alliance Aremon io -Fag ion BORHIDI in TOROK et al. 1989), and especially in the communities of sycamore maple (suballiance Polys t icho set i fer i-Acerenion BORHIDI et KEVEY 1996) -- BORHIDI & KEVEY (1996: 114). Stellaria n e m o r u m s. str. mostly thrives in similar communit ies both in Central and western

Europe (e.g. ACCETrO 1991, Tab. 2). The presence of both species in the forest communities of Slovenia is represented in Tab. 2.

Sources for the table were published phytosociological data. The forest associations were grouped on the basis of floristic and ecological similarity (columns 1 and 2). Authors of the sources are stated in the column 3. The next two columns specify the frequency (Fr) of each species (0-100%), except when there exists only one relev6 or synoptic table published, and the presence degree class (C1) ( I -V) . The importance of each species (Im) for alliances and suballiances is indicated by values 1-5 (1 means rare and diagnostically unimportant, 5 means frequent and characteristic for group of communities).

It was not possible to verify completely the reliability of the determination of both taxa of the Stellaria n e m o r u m group in the phytosociological material applied. The herbarium material, gathered in different regions of Slovenia and in the stands of numerous syntaxa, indicates that the determination was generally correct. We established false determination with certainty

Page 5: Stellaria nemorum  L. and  S. montana  Pierrat (  Caryophyllaceae  ) in the Forest Communities of Slovenia

Stellaria nemorum and S. montana in the forest communities of Slovenia 119

Fig. 3. Habitus of Stellaria montana (a) and Stellaria nemorum (b) (del. B. VRE~).

only in two cases, considering the corrigenda in the Tab. 2. It is possible that both taxa of the S. nemorum group occur together in the stands of some communities, thriving in the upper-montane belt, while the authors identified only one. In the synoptic table (Tab. 3) there is an example of two beech communities in Slovenia, in which the species of the SteUaria

nemorum group grow optimally. The first one (Tab. 3, column 1) is classified in the association Lamio orvalae-Fagetum

(HORVAT 1938) BORmDI 1963. The stands of this community are located in the area of Kalski gozd (9948/2, UTM VM00), a rather extensive forest complex in the northern part of the Banj~ice plateau in western Slovenia. They thrive on diversified rocky High Karst surface, mostly on sinkhole edges at the altitude of 800 to 1050 m. The parent material is Jurassic and partly Cretaceous limestone. The prevailing soil type is brown calcareous soil (Chromic Cambisols). Stellaria montana has high constancy and cover value in herb layer of these stands (see also DAKSKOBLER 1996a: 31"34).

As an example of beech forest, where both species of the S. nemorum group treated thrive, we present upper-montane beech community, which is classified in the association Ranunculo

platanifolii-Fagetum MAR~EK et al. 1993 (Tab. 3, column 2). The stands of this community have been found in the south-eastern part of the Julian Alps, mainly on the north-western slopes of Porezen (9849/2, UTM VM21) in the Ba6a Valley. They thrive in the upper-montane belt, up to the border of the subalpine belt (altitude 1200-1450 m). Parent material is Cretaceous

Page 6: Stellaria nemorum  L. and  S. montana  Pierrat (  Caryophyllaceae  ) in the Forest Communities of Slovenia

Tab

le 2

. T

he p

rese

nce

of S

tella

ria

nem

orum

and

S. m

onta

na i

n th

e fo

rest

com

mun

ities

of

Slov

enia

, Fr

. -

freq

uenc

y; C

I. -

cl

ass,

Im

. -

Impo

rtan

ce o

f sp

ecie

s.

Alli

ance

, su

balli

ance

, as

soci

atio

n;

geog

raph

ical

va

rian

t, su

bass

ocia

tion

Ref

eren

ces

]~lio

-Ace

rion

KLI

KA

195

5 H

acqu

etio

-Fra

xine

tum

MA

RIN

~EK

in W

ALL

NO

FER

et a

l. 19

93;

vat.

geog

r. D

enta

ria

pent

aphy

llos

MA

RIN

(~EK

(199

0)

Hac

quet

io-F

raxi

netu

m M

AR

IN~E

K in

WA

LLN

OFE

R et

al.

1993

; vat

. ge

ogr.

Den

tari

a pe

ntap

hyllo

s, v

at.

Car

pinu

s be

tulu

s M

AR

IN(~

EK (1

995b

) A

cer

pseu

dopl

atan

us-U

lmus

gla

bra

ass.

(no

m.

prov

.)

MA

P,IN

C~-E

K (1

995a

) C

oryd

alo

cava

e-A

cere

tum

pse

udop

lata

ni M

OO

R 1

938;

var

. ge

ogr.

Den

tari

a en

neap

hyllo

s ZU

PAN

(~I(~

(199

6)

Ery

thro

nio.

Car

pini

on

(Hop

,VA

T 19

58)

MA

RIN

(~E

K in

WA

LL

NO

FER

et

al.

1993

P

rim

o pa

di-C

arpi

netu

m b

etul

i MA

RIN

~EK

199

4

Lam

io o

rval

ae-F

agen

ion

BO

RH

1DI e

x M

AR

IN(~

EK

et a

l. 1

993

Lam

io o

rval

ae-F

aget

um (

Hop

,VA

T 19

38)

BO

RH

/DI 1

963;

vat

. ge

ogr.

Dem

aria

pen

taph

yllo

s La

mio

orv

alae

-Fag

etum

(H

op,V

AT

1938

) B

OP,

HID

I 196

3; v

ar. g

eogr

. D

enta

ria

pent

aphy

llos,

ste

llari

etos

um m

onta

nae

Lam

io o

rval

ae-F

aget

um (

Hop

,VA

T 19

38)

BO

RH

IDI 1

963;

vat

. ge

ogr.

Den

tari

a po

lyph

yllo

s O

mph

alod

o-F

aget

um (T

RE

GU

BO

V 1

957)

MA

P,IN

(~EK

et a

l. 19

93;

acer

etos

um

Om

phal

odo-

Fag

etum

(T

P, EG

UB

OV

195

7) M

AR

IN~E

K et

al.

1993

; ac

eret

osum

fac

ies

Pet

asite

s al

bus

Om

phal

odo.

Fag

etum

(T

RE

GU

BO

V 1

957)

MA

RIN

(~EK

et a

l. 19

93;

typi

cum

O

mph

alod

o-F

aget

um (T

REG

OB

OV

195

7) M

AR

IN~E

K et

al.

1993

; ga

lieto

sum

odo

roti,

vat

. H

acqu

etia

epi

pact

is

Om

phal

odo-

Fag

etum

(TR

EG

UB

OV

195

7) M

AP,

INdE

K et

al.

1993

; th

elyp

tere

tosu

m li

mbo

sper

mae

C

arda

min

i sav

ensi

-Fag

etum

Kog

m 1

962

Car

dam

ini s

aven

si-F

aget

um K

o~IR

196

2; v

at. g

eogr

. A

bies

alb

a ls

opyr

o-F

aget

um K

o~m

196

2; v

at.

geog

r. A

rum

mac

ulat

um

lsop

yro-

Fag

etum

KO

glR

196

2; v

at. g

eogr

. Ade

nost

yles

alli

aria

e

Saxi

frag

o ro

tund

ifolia

e-F

agen

ion

MA

RIN

t~E

K et

al.

199

3 H

omog

yno

sylv

estr

is-F

aget

um M

AR

IN(~

EK et

al.

1993

; va

t. St

ella

ria

mon

tana

R

anun

culo

plat

anifo

lii-F

aget

um

MA

RIN

CEK

et a

l. 19

93; v

at.

geog

r. H

epat

ica

nobi

lis

Ran

uncu

lo p

lata

nifo

lii-F

oget

um M

AR

IN~E

K et

al.

1993

; va

r. g

eogr

. H

epat

ica

nobi

lis, c

ortu

seto

sum

R

anun

culo

plat

anifo

lii-F

aget

um

MA

RIN

CEK

et a

l. 19

93

Stel

lari

o m

onta

nae-

Fag

etum

(Zt

JPA

N~:

I(~) M

AR

IN~E

K et

al.

1993

K

naut

io d

rym

eiae

-Fag

etum

ZU

PAN

~Id (

1969

) 19

94

Aco

nito

pan

icul

ati-

Fag

etum

(ZU

PAN

~I(~

1969

) M

AP,

~I(~

EK et

al.

1993

P

olys

ticho

lon

chiti

s-F

aget

um (H

OR

VA

T 193

8) M

AR

1N(~

EK in

PO

LD

INI e

t N

AR

DIN

I 199

3; v

at.

geog

r. S

alix

wal

dste

nian

a P

olys

ticho

lon

chiti

s-F

aget

um (H

oP, V

AT

1938

) M

AR

INgZ

EK in

PO

LD

INI e

t N

AR

OIN

I 199

3; v

at.

geog

r. A

llium

vic

tori

alis

LmTu

lo-F

agen

ion

LO

ItM

. et

R.

'Ix

. 19

54

Luzu

lo-F

aget

um M

EU

SEL

193

7; v

at. g

eogr

. C

arda

min

e tr

ifolia

, abi

etet

osum

Lu

zulo

-Fag

etum

ME

USE

L 1

937;

vat

. geo

gr.

Car

dam

ine

trifo

lia, a

biet

etos

um, v

at.

Gal

ium

rot

undi

foliu

m

Vac

cini

o-P

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on B

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Page 7: Stellaria nemorum  L. and  S. montana  Pierrat (  Caryophyllaceae  ) in the Forest Communities of Slovenia

Stellaria nemorum and S. montana in the forest communities of Stovenia 121

0, b

Fig. 4. The form and papillae length: Stellaria montana (a), S. nemorum (b) (del. B. VREg).

platy l imestone with admixture of red marl and chert. The soils are fresh, humose, loamy (brown calcareous soil and eutric brown soil). These beech stands are characterized by high cover of the species o f the Stellaria nemorum group. Stellaria montana predominates at the altitude of 1200 to 1350 m, whereas above 1350 m, in the transition area to the subalpine beech stands of the associat ion Polyst icho lonchi t is -Fagetum (HORVAT 1983) MARmffEK in POLDINI et NARDINI 1993, Stellaria nemorum is dominant. In one relev6 an intermediate form between both related species was reported. However, due to a lack of readi ly available plant material it can only be analyzed in the future.

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ANDREAS C.H. (1956): Notes on the Stellaria nemorum L. subsp, glochidisperma MURB. in the Netherlands. Acta Bot. Neerl. 5: 145-156.

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9-166. FREYN J. (1892): Flora von Oesterreich-Ungarn. Osterr. Bot. Z. 42: 356-359. GREEN P.S. (1954): Stellaria nemorum L. subspecies glochidisperma MURBECK in Britain. Watsonia 3: 122-126. HEGI G. (1911): Zwei gute Arten von Stellaria nemorum. Mitt. Bayer Bot. Ges. 2: 340-341. JALAS J. & SUOMINEN J. (eds.) (1983): Atlas florae europaeae. Distribution of vascular plants in Europe 6.

The Comittee for Mapping the Flora of Europe & Societas Bioiogica Fennica Vanamo, Helsinki. KOOPMANS A. & SLIM G.K. (1968): Some data on the karyotypes ofStellaria nemorum L. ssp. nemorum and

Stellaria nemorum L. ssp. glochidisperma MURB. in the Netherlands. Acta Bot. Neerl. 17: 90-92. KO~m P,. (1979): Ekolo~ke, fitocenolo~ke in gozdnogospodarske lastnosti Gorjancev v Sloveniji (Ecological,

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KO~IR Z. (1994): Ekolo~ke in fitocenolo~ke razmere v gorskem in hribovitem jugozahodnem obrobju Panonije (Ecological and phytosociological conditions in hilly outlying regions of Pannonia). Zveza gozdarskih dru~tev Slovenije, Ljubljana.

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Stellaria nemorum and S. montana in the forest communities of Slovenia 123

MATUSZKIEWICZ W. & MATUSZKIEWICZ A. (1981): Das Prinzip der mehrdimensionalen Gliederung der Vegetationseinheiten, erl~iutet am Beispiel der Eichen-Hainbuchenw~lder in Polen. In: DIERSCHKE H. (ed.), Syntaxonomie, Bet. Int. Syrup. Int. Vereinig. Vegetationsk. Rinteln, 1980, J. Cramer, Vaduz, pp. 123-148.

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VRES I . (1991): Stellaria nemorum agregat najugovzhodnem obrobju Alp. (Stellaria nemomm aggregate on the south-eastern margins of Alps). Ms.Thesis, Prirodoslovno-matemati~ki fakultet, Zagreb.

WALLNOFER S., MUCINA L. & GRASS V. (1993): Querco-Fagetea. In: MUCINA L., GRABHERR G. & WALLNOFER S. (eds.), Die Pflanzengesellschaften Osterreichs 3, WiiMer und Gebiische, Gustav Fischer Verlag, Jena, pp. 85-236.

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ZUPANI~It~ M. (1967): Der dinarische Bergahom-Buchenwald (Aceri-Fagetum dinaricum) im slowenischen Hochkarstgebiet. Mitt. Ostalpin-Dinarischen Pflanzensoziol. Arbeitsgem. 7: 89-96.

ZUPAN(~I(~ M. (1969): Vergleich der Bergahom-Buchengesellschaften (Aceri-Fagetum) im alpinen und dinarischen Raume. Mitt. Ostalpin-Dinarischen Pflanzensoziol. Arbeitsgem. 9:119-131.

ZUPAN~I~ M. (1980): Smrekovi gozdovi v mrazig~ih dinarskega gorstva Slovenije (Spruce forests in the freezing ravines of the Dinaric mountains of Slovenia). Dela IV. Razr. Slov~ Akad. Znan. Umetn. 24: 1-262.

ZUPAN~I~ M. (1994): Popravki imen nekaterih rastlinskih zdru~.b v lu~i novega kodeksa (Revision to the names of some plant communities in the light of the new code). Hladnikia 2: 33-40.

ZUPAN(~[(~ M. (1996): European maple association in Slovenia Corydalo cavae-Aceretum pseudoplatani MOOR 1938). Razpr. IV. Razr. Slov. Akad. Znan. Umem. 37: 189-205.

Encl. Appendix pp. 124-125

Page 10: Stellaria nemorum  L. and  S. montana  Pierrat (  Caryophyllaceae  ) in the Forest Communities of Slovenia

124 I. Dakskobler et al.

A P P E N D I X

Table 3. Synoptic table of two beech communities from western Slovenia in which there is a growing optimum of the species of the Stellaria nemorum group. 1 - Lamio orvalae-Fagetum (HoRVAT 1938) BORrlIDI 1963; 2 - Ranunculo platanifolii-Fagetum MAPaN~EK et al. 1993. Nomenclature of vascular plants in this paper follows TRnN & VRF~ (1995) and that of mosses DOLL (1991).

Association number 1 2 Association number 1 2 Number of relev4s 18 13 Number of rolev6s 18 13

Stellaria montana 100 54 Stellaria nemorum 100 Stellaria nemorum × S. montana (?) 8

Aremonio-Fagion Lamium orvala 100 38 Dentaria enneaphyllos 100 69 Helleborus odorus 89 Cardamine trifolia 61 85 Cyclamen purpurascens 39 Aremonia agrimonoides 28 lsopyrum thalictroides 28 Rhamnusfallax E2 17 Anemone trifolia 6 Primula vulgarls 6 15 Polystichum setiferum 8

Saxifrago rotundifoliae.Fagenion Adenostyles glabra 11 85 Saxifraga rotundifolia 11 85 Polygonatum verticillatum 100 Adenostyles alliariae 77 l-atzula sylvatica subsp, sylvatica 38 Cicerbita alpina 15 Ranunculus platanifolius 15

Fagetalia sylvaticae Fagus sylvatica E3 100 100 Fagus sylvatica E2 72 100 Fagus sylvatica El 78 8 Dentaria bulbifera 100 62 Corydalis cava 100 69 Dryopterls filix-mas 1 O0 92 Anon maculatum 100 Galium odoratum 100 15 Polystichum aculeamm 100 77 Adoxa moschatellina 100 77 Geranium robertianum 100 38 Paris quadrifolia 100 100 Circaea lutetiana 94 8 Myosotis sylvatica agg. 94 46 Scrophularia nodosa 94 69 Galeobdolanflavidum 94 85 Milium effusum 94 62 Epilobium montanum 94 62 Actaea spicata 94 85 Epipactls helleborine 94 Mycelis muralls 94 31 Sambucus nigra E2 89 Viola re&henbachiana 89 Daphne mezerum E2 89 8 Chrysosplenium alternifolium 78 31 Polygonatum multiflorum 78 Acer pseudoplatanus E3 6 62 Acerpseudoplatanus E2 39 46 Acer pseudoplatanus El 78 62 Fraxinus excelsior E3 6

Fraxinus excelsior E2 17 Fraxinus excelsior E1 72 Mercurialis perennls 72 15 Phyllitis acolopendrium 72 Cardamine impatiens 72 31 Dentaria pentaphyllos 67 Pulmonaria officinalis 61 31 Polystichum xluerssenii

(= P braanii x P. aculeatum) 56 23 Veronica montana 56 Corydalis solida 50 8 Sanicula europaea 50 8 Carex sylvatica 50 8 Symphytum tuberosum 44 92 Lathyrus vernus 44 8 Festuca altissima 39 8 Neottia nidus-avis 28 31 Prunus avium E3 6 Prunus avium El 22 Melica nutans 17 Brachypodium sylvaticum 17 Asarum europaeum subsp, caucasicum 17 Salvia glutinosa 17 Circaea intermedia 17 Stachys sylvatica 11 Euporbia dulcls I 1 Dryopterls affinls subsp, borreri 11 Galium laevigatum 11 38 Campanula trachelium 6 31 Prenanthes purpurea 6 31 Ranunculus lanuginosus 6 62 Poa nernoralis 6 31 Lilium martagon 92 Leucojum vernum 31

Querco-Fagetea Anemone nemorosa 1 O0 1 O0 Camptothecium lutescens 89 31 Isothecium alopecuroides 89 54 Ctenidium molluscum 83 54 Gagea lutea 67 31 Anemone ranunculoides 56 69 Lonicera xylosteum E2 33 Corylus avellana 33 Irmca minor 28 Clematis vitalba E2 28 Hypericura montanum 17 Crocus napolitanus (= C. vittatus) 17 Sesleria autumnalis 17 Moehringia trinervia 11 8 Arabls turrita 11 Platanthera bifolia 11 Hepatica nobilis 11 Carex digitata 11 Calamintha sylvatica 11 Moehringia muscosa 11 Galanthus nivalis 6 8

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Stellaria nemorum and S. montana in the forest communities of Slovenia 125

Association number 1 2 Association number 1 2

Cirsium erisithales 31 Polypodium vulgare 50 8 Laburnum alpinum E2 15 Cymbalaria muralis 28

Sedum hispanicum 17 Asplenium viride 46 Vaccinio-Piceetea

Oxalis acetosella Luzula luzuloides Gyranocarpium dryopteris Dryopteris eapansa Hypnum cupressiforme Dryopteris carthusiana Polytrichum formosum Maianthemum bifolium Gentiana asclepiadea Luzula luzulina Saxifraga cuneifolia Picea abies E3 Picea abies E2 Abies alba E3 Abies alba E2 Atrichum undulatum Thelypteris phegopteris Dryopteris dilatata Aposeris foetida Calamagrostis arundinacea Veronica urticifolia Valeriana tripteris Thelypteris lirabosperma Vaccinium myrtillus

A denostyletalia Senecio ovatus Urtica dioica Athyrium filix-femina Impatiens noli-tangere Scrophularia vernalis Angelica sylvestris Ribes alpinum E2 Veratrum album subsp, labelianum Thalictrum aquilegiifolium Phyteuma ovatum Chaerophyllum hirsutum subsp, hirsutum Rumex arifolius Aconitum vulparia Doronicum austriacum Carduus personata Alnus viridis Myrrhis odorata Geum rivale Crepis paludosa Melandrium rubrum Salix appendiculata E2 Chaerophyllum hirsulum subsp, viltarsii

Asplenietea trichomanis Cystopteris fragilis Asplenium trichomanes

89 83 56 83 56 61 39 56 28 50 28 44 22 39 17 33 11 33 11 17

28 11 28

17 11 17 11 11 6 11

6

100 100 83 50 39 28 17 I I

78 72

77 85 Other species 38 Galeopsis pubescens 15 Rubus hirtus E2 31 Cardamine flexuosa

8 Galeopsis speciosa 8 Rubus ideus E2

Solarium dulcamara - -

62 Sambucus racemosa E2 8 Atropa bella-donna

Sorbus aucuparia E2 38 Sorbus aucuparia E1

8 Fragaria vesca 8 Verbascum lanatum

Deschampsia cespitosa 8 Galeopsis tetrahit 8 Bromus ramosus

85 Dactylis glomerata 69 Hypericum hirsutum 54 Polystichum xillyricum 54 Dactylorhiza maculata s.l. 23 1.amium maculatum 15 Crocus albiflorus 15 Poa alp ina

Betonica alopecuros

100 Mosses and lichens 46 Grimmia pulvinata 94

100 Brachythecium velutinum 89 31 Plagiochila asplenioides 67

8 Collema sp. 50 46 Neckera crispa 44

Plagiothecium denticulatum 39 92 Plagiomnium affine 39 62 Thamnobryum alopecurum 39 62 Brachytheciura rutabulum 39 54 Mnium marginatum 39 46 Mnium sp. 33 38 Madotheca platyphylla 33 38 Anomodon viticulosus 28 38 Plagiomnium undularum 28 31 Bryum capillare 22 23 Conocephalum conicum 22 23 Plagiomnium cuspidatum 22 15 Peltigera canina 17 15 Cladonia sp. 11 15 Metzgeria sp. 1 I 15 Anomodon attenuatus 11

Fissidens cristatus 6 Plagiothecium sylvaticum 6

77 Paraleucobryum sauteri 23 Tortetla tortuosa

54 54

8

23

46

8 38 23 15 15 15 15 15

54 31 46

8

23

31 31

8

8 23 38 23

8 15

8 46 31 46 38