fulltext-Festuca eskia

Folia Geobotanica 36: 385-421, 2001

FESTUCA SECT. ESKIA (POACEAE) IN THE IBERIAN PENINSULA

Vicenta de la Fuente & Emma Orttifiez

Dpto. de Biologla, Facultad de Ciencias, Universidad Autrnoma de Madrid, Cantoblanco, E-28049 Madrid, Spain;fax +34 91 3978 344, e-mail [email protected], [email protected]

Keywords: Festuca, New combination, Nomenclature, Taxonomy

Abstract: A taxonomidal revision of the genus Festuca sect. Eskia in the Iberian Peninsula is presented. Seven species and two hybrid taxa are recognized in this section in the area studied. Karyological data show that six of the studied taxa (F. pseudeskia, F. scariosa, F. elegans subsp, elegans, F. burnatff, F. eskia, and F. xpicoeuropeana) are diploids, two of them are both diploid and tetraploid (F. gautieri and F. quadriflora), one is tetraploid (F. elegans subsp, merinoO and the ploidy level of one taxon (F. xsouliei) is unknown. A new combination F. elegans subsp, merinoi (PAU) FUENTE et ORI"0N~Z is published. A key based on data collected in this study allows for the identification of all taxa in this section. The complete description of each taxon is presented, as well as the ecology and chorology, including distribution maps. The illustrations summarize plant habits and the morphological and anatomical pattern observed on spikelets, ligules and leaf cross sections. Epidermal features were studied using scanning electron microscopy.

INTRODUCTION

WILLKOMM (1861: 95) described the section Eskia based on four species from the Iberian Peninsula (F. elegans Bolss., F. eskia RAMOND ex DC., F. pseudeskia Bolss. and 17. granatensis Bolss.), which were characterized as a group principally by the presence of oblong ligules, palea and lemma with wide scarious margins and apices.

Since then, further contributions concerned with the taxonomy of this section have been presented, such as HACKEL (1882) and MARKCRAF-DANNENBERG (1980), which are detailed later. The principal schemes proposed in the literature for section Eskia WILLK. in the Iberian Peninsula are summarized in Tab. 1.

Following the results o f our study, we also recognize the section Eskia in the Iberian Peninsula, which comprises a group of taxa with the following shared characters: presence of wide scarious margins and apices on the glumes and lemmas, and oblong ligules (0.4-8 mm long), characters previously given by WILLKOMM (1861). All taxa included in this section are generally confined to mountainous areas. The species possess distinctive morphological and anatomical characters that makes it easy to differentiate them.

In our scheme we recognize seven species (F. burnatii ST.-YVES, F. elegans BoIss., F. eskia RAMOND ex DC., F. gautieri (HACK.) K. RICHT., F. quadriflora HONCK., F. pseudeskia BoIss. and F. scariosa (LAG.) ASCH. et GRAEBN.) in the Iberian Peninsula, of which one (F. elegans) is divided into two subspecies (subsp. elegans and subsp, merinoi (PAU) FUENTE et ORT0~EZ). Furthermore, two hybrid taxa (F. ×picoeuropeana NAVA and F. ×souliei ST.-YVES) are presented; F. ×picoeuropeana is reported for the first time from the Pyrenees. Our scheme has been based on morphological, anatomical and cytological data that have been systematically

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Festuca sect. Eskia (Poaceae) in the Iberian Peninsula 387

collected from a large number of specimens. The complete descriptions, chorology and ecology are given for each taxon, as well as the micromorphological characters and the chromosome number. Distribution maps based on the studied material are given for each taxon, and a key for identification is presented.

MATERIAL AND METHODS

The results presented are based on the study of populations in the field and on the examination of herbarium specimens, including the analyses of type specimens.

During our field investigations, specimens of different populations have been collected, which have been deposited in our personal herbarium and cultivated in the greenhouse at the Universidad Aut6noma de Madrid.

In addition, material from the following herbaria was studied: ARAN, BC, BCF, BM, COI, FCO, FI, G, GDA, GDAC, JACA, K, LEB, MA, MAF, MGC, MPU, SALA, UNEX, VAL and W (herbarium abbreviations according to HOLMCREN et al. 1990). Altogether 764 herbarium specimens were studied (F. pseudeskia 38, F. scariosa 143, F. burnatii 56, F. elegans subsp. elegans 49, F. elegans subsp, merinoi 165, F. gautieri 176, F. quadriflora 6, F. eskia 104, F. xpicoeuropeana 23, F. xsouliei 4). The list of representative specimens is given in the Appendix.

The methods and terminology adopted for the morphological and anatomical analyses are the same as described by HACKEL (1882), SAINT-YVES (1913), AUQUIER (1974), ELLIS (1976) and WILKINSON & STACE (1991).

Epidermal characters have been studied using a scanning electron microscope, Philips XL30. The terminology and methods adopted are those of METCALFE (1960), ELLIS (1979), PALMER & TUCKER (1981) and FUENTE & ORTO~qEZ (1998).

Chromosome counts were made on root-tips of mature plants that were pretreated in 8-hydroxyquinoline and stained with acetic-orcein. Meiotic preparations were made on young florets fixed in a mixture of ethanol : acetic acid (3 : 1) and stained with acetic-orcein. The localities of analyzed plants are given in Tab. 2. The voucher specimens are deposited in our personal herbarium at the Universidad Aut6noma de Madrid (herb. F. & O.).

RESULTS

Epidermal micromorphology

Leaf blade abaxial surface (Fig. 1, p. 419). The studied taxa are characterized by the presence of a light cover of epicuticular wax, except in three taxa (F. pseudeskia, F. eskia and F. elegans) where the cover is dense; stomata absent except in F. pseudeskia; rectangular-elongated long cells with usually markedly sinuous cell walls, prominent and raised; square or narrow-rectangular short cells in the case ofF. burnatii, and, in some taxa, the presence of antrorse prickles. The study of the abaxial leaf surface shows some differential characters which allow one to separate easily some of the studied species (Tab. 3). F. pseudeskia is the only species with stomata on the abaxial leaf surface, F. burnatii is the only species with narrow-rectangular short cells and F. scariosa is the only species with narrow long cells and cell walls. F. scariosa and F. elegans are the only two taxa with abundant antrorse prickles along the entire surface. F. elegans subsp, elegans shows shorter long cells and denser antrorse prickles than F. elegans subsp, merinoi (Tab. 3). The rest of the

Tab

le 2

. L

ist

of k

aryo

logi

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y in

vest

igat

ed p

opul

atio

ns o

f F

. ps

eude

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, F

. sc

ario

sa,

F.

eleg

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F.

burn

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, F

. es

kia,

F.

gaut

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, F

. qu

adri

flor

a an

d F

. xp

icoe

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eana

(l

ocal

itie

s of n

ew c

ount

s ar

e m

arke

d by

ast

eris

k).

Spec

ies

2n

n L

ocal

itie

s B

ibli

ogra

phic

refe

renc

es

F. p

seud

eski

a 7

F. s

cari

osa

14

14

14

14

14

14

7 7

F. e

lega

ns s

ubsp

, el

egan

s 14

14

7

F. e

lega

ns s

ubsp

, m

erin

oi

28

28

28

28

14

F. b

urna

tii

14

14

7 F

. esk

ia

14

Spai

n, G

rana

da: P

icac

ho d

e V

elet

a, S

ierr

a N

evad

a, 3

0SV

G60

Sp

ain,

Gra

nada

: Sie

rra

Nev

ada,

30S

VG

60

Spai

n, G

rana

da: S

ierr

a Nev

ada,

137

0 m

Sp

ain,

Gra

nada

: Sie

rra

Nev

ada,

Pic

acho

de

Vel

eta,

30S

VG

60

wit

hout

loca

liti

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Spai

n, G

rana

da: S

ierr

a N

evad

a, B

arra

nco

de la

s V

ibor

as, 3

0SV

F69,

160

0 m

(S

EV

788

89)

wit

hout

loca

liti

es

Spai

n, G

rana

da: P

eflo

nes

de S

an F

ranc

isco

, Sie

rra

Nev

ada,

30S

VG

60,

2400

m

Spai

n, C

6rdo

ba: C

abra

, L

a N

ava,

30S

UG

75,

800-

1200

m (

UN

EX

133

9)

*Spa

in, G

rana

da: b

arra

nco

de S

an J

uan,

Sie

rra

Nev

ada,

30S

VG

60;

*Baz

a,

barr

anco

de

Bod

urri

a, s

ierr

a de

Baz

a, 3

0SW

G19

30,

1700

m;

*Baz

a, P

rado

s de

l R

ey, b

ase

del C

alar

de

Sant

a B

~bar

a, s

ierr

a de

Baz

a, 3

0SW

G13

37,

2000

m

*Spa

in, G

rana

da: L

oma

de la

s Y

egua

s, S

ierr

a N

evad

a, 3

0SV

G63

06,

1950

m

*Spa

in, Z

amor

a: S

esn~

ndez

de

T;ib

ara,

cam

ino

del C

asal

, sie

rra

de la

Cul

ebra

, 29

TQ

G45

34,

950

m

Port

ugal

, BA

: Man

teig

as-P

oqo

do In

fern

o, 2

9TPE

27;

MI:

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ra d

o G

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arri

s, 2

9TN

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Sp

ain,

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: Pue

rto

de T

omav

acas

, 30T

TK

76,

1000

m (

UN

EX

842

2)

Spai

n, A

vila

: Bec

edas

, pue

rto

del T

rem

edal

, 30T

TK

7674

, 12

20 m

; Zam

ora:

Se

snhn

dez

de T

~bar

a, c

amin

o de

l Cas

al, S

ierr

a de

la C

uleb

ra, 2

9TQ

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34,

950

m

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in, O

vied

o: T

ever

ga,

puer

to d

e V

enta

na,

29T

QH

4371

, 16

00 m

Sp

ain,

Le6

n: M

ampo

dre,

30T

UN

26

Spai

n, S

anta

nder

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pain

, San

tand

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Her

man

dad

de C

ampo

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o, s

ubid

a Pi

co T

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Mar

es,

KI3

PFER

(196

8: 8

8)

LEV

ITSK

Y &

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A (1

927:

22)

B

OR

RIL

L et

al.

(197

1: 6

) B

OR

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L et a

l. (1

977:

664

), M

OR

GA

N

et a

L (1

979:

117

) SE

AL

(198

3: 2

27)

RO

MER

O ZARCO (

1984

: 29

2)

BU

LIIq

SKA

-RA

DO

MSK

A et L

ESTE

R

(198

6: 1

55)

KO

PFER

(196

9: 3

3)

DEV

ESA

& L

UQ

UE

(198

8: 2

86)

FER

NA

ND

ES &

QU

EIR

0S (1

969:

33,

37)

D

EVES

A et

al.

(199

0: 1

56)

FER

RER

O &

FU

ENTE

(199

6: 3

06)

LITA

RD

It~R

E (19

54:1

80)

KO

PFER

(197

1)

GO

(30

o.

o "rl

t-

in

O

r"

tD

N

F. g

auti

eri

F. q

uadr

iflor

a

F.

xpic

oeur

opea

na

sier

ra d

e Pe

fla

Lab

ra, 3

0TU

N86

66,

2075

m

14

Fran

ce, A

rise

: B

rass

ac a

u Pi

cou,

31T

CH

85,

1400

m

14

Spai

n, G

eron

a: P

ico

de S

egre

, 31T

DG

29,

2300

m (

NE

U K

0152

5)

7 Sp

ain,

Ger

ona:

Pyr

dnde

s ori

enta

les v

ersa

nt e

spag

nol,

Puig

mal

, 31

TD

G29

, 23

00 m

; O

vied

o: C

ueto

de

Arb

as, 2

9TQ

H06

, 19

00 m

(N

EU

K01

524)

14

*S

pain

, L6r

ida:

Ari

stot

-Tol

oriu

, si

erra

de

las

Pina

s, 3

1TC

G84

96,

2150

m

7 Sp

ain,

Gra

nada

: Sie

rra

de la

Sag

ra, 3

0SW

H30

, 22

50 m

(N

EU

K01

586)

; H

uesc

a: S

oum

Bla

nc d

es E

sp6c

i&es

, 30

TY

'N33

, 230

0 m

(N

EU

K01

583)

; Ja

6n:

Sier

ra d

el P

ozo,

Cer

ro C

abaf

las,

30S

WG

08,

1950

m (

NE

U K

0157

3);

Nav

arra

: Pie

d'O

rhy,

30T

X'N

66, 1

900

m (

NE

U K

0157

5);

Ter

uel:

Sie

rra

de

Gfi

dar,

Pena

rroy

a, 3

0TX

K97

, 20

00 m

(N

EU

K01

576)

14

Sp

ain,

Hue

sca:

Tor

la, L

itro,

30T

YN

32,

1700

m (

NE

U K

0158

3);

Toz

al d

e G

uara

, 30

TY

M38

, 19

00 m

(N

EU

K01

582)

. T

erue

l: Si

erra

de

Java

lam

bre,

30

TX

K64

, 19

00 m

(N

EU

K01

578)

Fr

ance

, Ari

~ge:

Col

de

Pailh

&es

, Mija

n6s,

31T

DH

13,

1600

m

14

Spai

n, G

eron

a: N

uria

, 31

TD

G39

Sp

ain,

Ger

ona:

Nur

ia,

Pico

de

Segr

e, 3

1TD

G29

, 24

50 m

Fr

ance

, Pyr

6n6e

s-O

rien

tale

s: O

ss6j

a, 3

1TD

G19

, 19

00 m

14

28

28

14

14

28

14

14

Aus

tria

, Sty

ria:

Hoc

hsch

wab

; Fr

ance

, Alp

es-M

arit

imes

: St-

Dal

mas

-de-

Ten

de,

vall

on d

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aste

rino

; H

aute

s-A

lpes

: Val

l6e

de la

Rom

anch

e, M

alav

al, 1

370

m

wit

hout

loca

liti

es

wit

hout

loca

liti

es

wit

hout

loca

liti

es

Spai

n, L

e6n:

Veg

a H

uert

a, P

icos

de

Eur

opa,

mac

izo

Occ

iden

tal,

30T

UN

48,

2050

m

LITA

RD

II~R

E (19

50:

84)

KCT

PFER

(197

4: 9

0)

KO

PFER

(196

9: 3

3, 1

974:

90)

KO

PFER

(197

4: 8

7)

KO

PFER

(197

4: 8

7)

KER

GU

~LEN

& P

LON

KA

(198

9: 1

71)

KO

PFER

(196

8: 8

8, 1

972)

K

LvPF

ER (1

974:

87)

K

ERG

UI~

LEN

(197

5: 1

49),

KER

GU

I~LE

N

& P

LON

KA

(198

9: 1

69)

L/TA

RD

II~R

E (19

50:

84)

RO

UG

EOT

(197

4)

RO

UG

EOT

(197

4)

GU

TII~

RR

EZ V

ILLA

RIA

S & H

OM

ET

(198

4: 4

8)

NA

VA

(198

5: 2

4)

¢,o

t~

O

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--.i

e- ¢.O

390 V. de la Fuente & E. Ortufiez

taxa (F. eskia, F. gautieri, F. quadriflora and the two hybrid taxa, F. ×picoeuropeana and F. Xsoutiei) show more similarity; even so there are some differences concerning the long cell length, the silica body diameter and the presence of prickles (Tab. 3).

Lemma abaxial surface (Fig. 2, p. 420). All studied taxa are characterized by the presence of a slight cover of epicutelar wax; stomata are absent; long cells are rectangular with markedly sinuous cell walls all over the surface; they have hooks and usually antrorse prickles; silica bodies are rounded or kidney-shaped. The principal characters distinguishing the lemma micromorphology of these taxa are shown in Tab. 3. In this surface it is important to emphasize the length of the long cells, the shortest are present in F. scariosa. Different lengths are also observed in the two subspecies ofF. elegans, showing one taxon (subsp. etegans) with shorter long cells than the other (subsp. merinoi).

Palea abaxial surface (Fig. 3, p. 421). All studied taxa are characterized by the presence of rectangular long cells with markedly sinuous cell walls; papillae are present all over the surface; antrorse prickles are present over the surface and along the keels; silica bodies are sparse and rounded. Some diagnostic characters are observed on this surface (Tab. 3). Festuca scariosa shows the highest density of papillae. F. eIegans has abundant hooks all over the surface. Its two subspecies can be distinguished by the presence of shorter long cells and denser hooks in F. elegans subsp, elegans. The different number of rows of prickles along the keels can be also observed (Tab. 3): F. pseudeskia with only 1(-2) rows, but F. quadriflora with up to 3-6 rows. Further characters are given in Tab. 3.

Our micromorphological study has shown that the four studied taxa (F. pseudeskia, F. scariosa, F. elegans and F. burnatii), are easy to distinguish using the leaf surface characters. The rest of the taxa (F. eskia, F. gautieri, F. quadriflora and the two hybrid taxa), however, are more similar; even so there are some differential characters. Some of these differences allow one to observe the possible relationships between the two hybrid taxa (F. Xpicoeuropeana and F. xsouliei) and their respective parents (F. gautieri x F. eskia, and F. quadriflora x F. eskia), because some measurements for the hybrid taxa show intermediate values between those of the parents (see Tab. 3).

Karyology

Karyological studies of the taxa of this section has shown that six of them are diploid (F. pseudeskia, F. scariosa, F. elegans subsp, elegans, F. burnatii, F. eskia, F. xpicoeuropeana), two of them occur both as diploid and tetraploid populations (F. gautieri, F. quadriflora), one is tetraploid (F. elegans subsp, merinoi) and the ploidy level of one taxon (F. × souliei) is yet unknown. F. elegans is represented by two taxa, the diploid (subsp. elegans) and the tetraploid (subsp. merinoi), both with different distribution areas and ecology (Tab. 2). The studied karyotypes are symmetrical and the B-chromosomes are lacking.

DISCUSSION

WILLKOMM (1861: 95) described the section Eskia, which was characterized by the presence of "... palea inferior apice late scariosa, fructifera involuta. Folia (salten basilaria) convoluta angusta. Ligula oblonga integra vel lacera". He recognized four taxa from the Iberian Peninsula (F. etegans BOlSS., F. eskia RAMOND ex DC., F. pseudoeskia BOlSS. and F. granatensis BoIss.) in this section.

Festuca sect. Eskia (Poaceae)in the Iberian Peninsula 391

HACKEL (1881) compared the relationships among the European fescues and proposed a scheme of the different sections, but without descriptions. In this work and later in his monograph of the European species of Festuca (HACKEL 1882), he included the studied species by WILLKOMM (1861) into two different sections (sect. Variae HACK. and sect. Scariosae HACK.). HACKEL (1882) distinguished the section Variae from the section Scariosae principally by the following characters: obovate vs. pear-shaped ovary, oblong caryopsis free from the palea vs. obovate-oblong caryopsis highly adherent to the palea. Furthermore, the section Variae was divided into two series (ser. lntravaginales HACK. and ser. Extravaginales HACK.) based on the presence of intravaginal or extravaginal vegetative shoots respectively. The series lntravaginales was represented in the Iberian Peninsula by two species (F. varia HAENKE and F. elegans). The Spanish representatives ofF. varia came under three subspecies (subsp. eskia (RAMOND ex DC.) HACK., subsp, pumila (CHAIX) HACK. and subsp, scoparia A. KERN. et HACK.) and three varieties (subsp. pumila "vat. genuina", subsp, scoparia "var. genuina" and subsp, scoparia var. gautieri HACK.) The series Extravaginales was represented by one species (F. pseudeskia) in the same geographical area. Finally, in the section Scariosae HACI~L recognized only one species (F. granatensis) endemic to the south of Spain.

Later, MARKGRAF-DANNENBERG (1980) recognized seven species in the Iberian Peninsula (F. elegans, F. eskia, F. pseudeskia, F. scariosa, F. burnatii, F. gautieri and F. quadriflora). Although Markgraf-Darmenberg did not recognize sections, she distinguished two groups. One of these is characterized by the presence of non-flowering extravaginal shoots and stolons, and is represented in the Iberian Peninsula by two species (F. pseudeskia and F. scariosa), and the other group is characterized by non-flowering intravaginal shoots with absent stolons and represented by the rest of the species.

In the monograph of the French species ofFestuca, KERGU~LEN & PLONKA (1989) focused on three species which occur in the Spanish Pyrenees (F. eskia, F. gautieri and F. quadriflora). They were included in the section Eskia WmLK. (= Festuca sect. Variae HACK.). Furthermore they listed another section, Festuca sect. Amphigenes (JANKA) TZVELEV (= Amphigenes JANKA; = Festuca sect. l/'ariae ser. Extravaginales HACK.) based principally on the presence of extravaginal vegetative shoots.

Recently, CONERT (1994: 535) again recognized only the section Eskia WILLK. (= Festuca sect. Variae HACK.), which is, according to him, characterized principally by large ligules, lemma with wide scarious margins and densely pubescent ovary.

In our case all the studied specimens possess intravaginal shoots or mixed shoots (F. pseudeskia and F. scariosa), therefore these cannot be included in the section Amphigenes (JANKA) TZVELEV.

All the characters given by WILLKOMM (1861) are present in all of the taxa studied. Although each taxon shows typical characters, some of these are shared with other taxa. So, 17. pseudeskia and F. scariosa possess short rhizomes and sclerenchyma corresponding to the veins, characters absent in the rest of the species. F. pseudeskia is the only species with stomata present in the leaf blade abaxial surface and F. scariosa is the only taxon with the caryopsis completely adherent to the palea. The presence of caryopsis adherent to the palea is one of the characters used by HACKEL (1882) to separate the section Variae from the section Scariosae, but there is one species (F. elegans) with palea slightly adherent, which is a character also cited for this species by HACKEL (1882:182). Furthermore, when HACKEL (1882) included F. pseudeskia into the section Variae (series Extravaginales) he considered it

¢D

Tab

le 3

. P

rinc

ipal

ch

arac

ters

of

leaf

bla

de

abax

ial

surf

ace,

lem

ma

abax

ial

surf

ace,

and

pal

ea a

baxi

al s

urfa

ce d

isti

ng

uis

hin

g th

e ta

xa b

elo

ng

ing

to

Fes

tuca

sec

tion

r~

E

skia

WlL

LK

.

Cha

ract

ers

F. p

seud

eski

a F

. sca

rios

a F

, ele

gans

F

. el

egan

s F

. bur

natii

F

. esk

ia

F g

auti

eri

F. q

uadr

iflor

a F

. ×

pico

- F

, X

soul

iei

subs

p, e

lega

ns

subs

p, m

erin

oi

-eur

opea

na

Lea

f b

lad

e ab

axia

l su

rfac

e

Lon

g ce

lls (

Inn)

: -

long

(3

1.2-

)68.

7-92

.5

42.5

-73.

5 31

-43

68.5

-142

.5

110-

150

(50-

)60-

165

88-2

00

82-2

20

60-1

60

68-2

00

- w

ide

19-2

6 9.

2-18

,7

26-3

0 24

-30

25-3

5 25

-27

21-2

6 24

-25

21-2

6 20

-30

- w

alls

6.

2-8

3-4

7-8(

-10)

7-

9 6-

8 (7

-)9-

12

6-9,

5 7-

9.5

6-9.

5 6-

12

Sho

rt c

ells

: -

shap

e sq

uare

sq

uare

sq

uare

sq

uare

na

rrow

- sq

uare

sq

uare

sq

uare

sq

uare

sq

uare

-r

ecta

ngul

ar

- de

nsity

sp

arse

ab

unda

nt

spar

se

spar

se

abun

dant

sp

arse

sp

arse

ab

unda

nt

spar

se

spar

se

Silic

a bo

dies

: -

shap

e ro

unde

d ro

unde

d ro

unde

d ro

unde

d no

t see

n ro

unde

d ro

unde

d ro

unde

d ro

unde

d ro

unde

d -

dens

ity

spar

se

spar

se

abun

dant

ab

unda

nt

abun

dant

ab

unda

nt

abun

dant

ab

unda

nt

abun

dant

Pric

kles

: -

dens

ity

abse

nt

very

very

ab

unda

nt

abse

nt

spar

se

abse

nt

spar

se

abse

nt o

r so

met

imes

ab

unda

nt

abun

dant

an

tror

se

anlr

orse

an

tror

se

som

etim

es

spar

se

antr

orse

an

tror

se

pric

kles

pr

ickl

es

pric

kles

sp

arse

an

tror

se

pric

kles

pr

ickl

es

antr

orse

pr

ickl

es

pric

kles

Stom

ata

pres

ent

abse

nt

abse

nt

abse

nt

abse

nt

abse

nt

abse

nt

abse

nt

abse

nt

abse

nt

Lem

ma

abax

ial

surf

ace

Long

cel

ls (Inn):

-lo

ng

30

-50

12-3

1 25

-50

(28-

)32-

75

(44-

)50-

75(-

125)

30

-87.

5 30

-70

(50-

)60-

125

38-1

00

(30-

)44-

-88(

-100

) -

wid

e 16

-19

16-1

9 19

-25

19-2

5 12

-19

15-2

2 15

-20

16-1

8 18

-20

14-2

0 -

wal

ls

5.2-

6.2

4.5-

5 6-

9 6-

9 3-

6 6-

10

7-9

(3-)

4-5

6-10

3-

6

.<

-n

t-

in

o c N

Hoo

ks b

ase

in

vert

ical

dia

met

er (

~an

) 7-

8

Pal

ea a

baxi

al s

urfa

ce

Lon

g ce

lls (

0m):

-

long

50

-90

- w

ide

18-2

0 -

wal

ls

5-7.

5

Pric

kles

or

spar

se

hook

s on

pr

ickl

es

the

surf

ace

all

over

sur

face

Silic

a bo

dies

: -

shap

e ro

unde

d

- de

nsity

sp

arse

Pric

kles

on

the

keel

s:

shor

t -

row

s I (

-2)

- po

siti

on

over

the

basa

l 1/

4

11-1

3 6

-7

6-7

6-7

7-12

9-

12

6-7

6-11

5-

6

25-5

0 18

-35

(18-

)33-

70

(21-

)36-

112

34

-75

(-9

0)

32

-90

5

0--

18

0

37-1

20

35-1

00

20-3

0 15

-20

18-2

0 13

-18

12-1

5 2

5-2

8

10-1

5 12

-15

9-12

5-

7.5

6-7

6-

7 6-

7 5-

6 1

0-1

2

3--4

3-

4 3-

5

spar

se

very

ab

unda

nt

spar

se

pric

kles

lo

ng-

abun

dant

lo

ng-

abun

dant

ho

oks

and

pric

kles

ar

id s

pars

e -p

rick

les

pric

kles

al

l -p

rick

les

hook

s an

d sp

arse

at

the

tip

hook

s at

at

the

tip

over

sur

face

al

l ov

er s

urfa

ce

spar

se p

rick

les

pric

kles

th

e tip

roun

ded

roun

ded

or

roun

ded

or

roun

ded

roun

ded

roun

ded

roun

ded

kidn

ey s

hape

d ki

dney

sha

ped

spar

se

abun

dant

ab

unda

nt

spar

se

spar

se

spar

se

spar

se

shor

t sh

ort

shor

t sh

ort

shor

t lo

ng

long

1-

2 2

-3(-

4)

2-3

(-4

) 1-

2 1-

3 1 -

3

3-6

over

the

alon

g w

hole

al

ong

who

le

alon

g w

hole

al

ong

who

le

at th

e tip

al

ong

who

le

basa

l 1/

4 ke

els

keel

s ke

els

keel

s ke

els

spar

se

pric

kles

at

the

tip

roun

ded

spar

se

long

2-

3 al

ong

who

le

keel

s

pric

kles

at

the

tip

roun

ded

spar

se

long

2

-4

over

the

basa

l 1/

4

O~

o .oq

5"

¢0

o-

2.

to

o~

r'- ¢D

60


\

Fig. 4. Festuca pseudeskia BOISS., Spain, Granada, Sierra Nevada, Veleta (herb. F. & O.): a - habit, b - leaf blade section, c - ligule, d - spikelet.

Ecology. Festuca pseudeskia occurs on sunny slopes and sandy substrates, over 2500 m. It is a characteristic species o f the grassland ofFes tuce tum moleroi-pseudoeskiae QUEZEL 1953 (Ptilotrichion purpurei QU~ZEL 1953) (RWAS MARTtNEZ et al. 1991: 33).

2. Festuca scariosa (LAG.) ASCH. et GRAEBN.

Fesmca scariosa (LAG.) ASCH. et GRAEBN., Syn. Mitteleur. F1.2(1): 502, 1900. - Poa scariosa LAG., Gen. Sp. Pl.: 3, 1816. Ind. loc.: "Hab. circa Gades". Typus ignotus.

= F. granatensis Bolss., Elench. P1. Nov.: 93, 1838. Ind. loc.: "Species habitu distinctissima, omnium mont ium Granatensium incola ab alt. 4000' usque ad 7500' ". Lectotypus (BURDET et al. 1981: 566): In omnibus montibus Regn. Granat., alt. 4000 ' -9500 ' (VI I . 1837 BOISSIER G-BOISS !).


1;'. pseudeskia l~lr iblaJon area

• S~udled material

Fig. 5. Distribution of Festuca eskia RAMOND ex DC. and F. pseudeskia BOISS.

Iconography: (Fig. 6).

Description. Tufted grass. Vegetative shoots intravaginal and extravaginal, with short rhizomes. Culms up to 60-120 cm long, glabrous-scaberulous. Leaf-sheaths fused for 1/4 of their length, glabrous or scabrid (retrorse, appressed and very short prickles). Ligules 3-6(-7) mm long, oblong, acute, 2-veined. Leaf-blades 26-72 cm long, erect, rigid, scabrid (antrorse and very short prickles); apex of the leaves acute and sharp; in cross section orbicular to suborbicular in outline, about 1-1.4 mm in diameter, with 13-17 veins, with sclerenchyma forming a complete ring and extending to the adaxial surface, with 13-17 adaxial grooves and 13-17 adaxial ridges. Panicle (15-)20-30(-50) cm long, erect, with about 119-215 spikelets, with 9-11 branches compressed to the rachis (of the which the first two usually shorter than the respective internode, the first 7-8 branches branched). Spikelets 5-7 mm, with 3-6 fertile florets. Glumes unequal, shortly acuminate, with wide scarious margins; lower glume 3-4.5 mm, 1-veined, lanceolate; upper glume 3.5-5.8 mm, 3-veined, ovate-lanceolate. Lemma 3.5-5 mm long, 5-veined, ovate-oblong, with scarious margins, mucronate. Palea 3-5 mm long, scabrid on the keels. Anthers about 2 mm long. Ovary pubescent. Caryopsis adherent to the palea.

Chromosome number. 2n=2x = 14. Chorology. Mountains of south-east of Iberian Peninsula (Fig. 7). The presence of this

taxon in the North Morocco cited by some authors (MArnE 1955: 164) is based on one sheet collected by Gandoger (N Tetouan, Mont Yakmil), which was determined by SAl~qT-YvEs (1922: 47). BORRILL et al. (1971: 3) and BORRILL (1972: 530) did not find this taxon in the North Morocco. We have not found another sheet or reference from this area.

Ecology. Festuca scariosa grows in xerophilous grazing lands on calcareous and dolomitic substrates. It is a characteristic species of Festucion scariosae MARTINEz-PARRAS et al. in

400 V. de la Fuente & E. OrtuSez

I

Fig. 8. Festuca burnatii ST.-YVES, Spain, Le6n, Puerto Ventana (herb. F. & O.): a - habit, b - leaf blade section, c - ligule, d - sheath, e - spikelet.

4. Festuca elegans BOlSS.

Festuca elegans BOISS., Elench. PI. Nov. : 92, 1838. Ind. loc.: "Hab. in declivitatibus in Sierra Nevada loco Pra de la Yegua dicto. Alt. 6500' ". Lectotypus (BURDET et al. 1981: 565): In Sierra Nevada loco dicto Pra de la Yegua, a!t. 6500' (VII.1837 BOISSIER G-BOISS, no. 196!). Isolectotypus: (G!).

Descr ipt ion. Densely tufted grass. Vegetative shoots intravaginal. Culms 50-112 cm long, scabrid (antrorse prickles). Leaf-sheaths fused for 1/4 of their length, scabrid (antrorse


Fig. 9. Distribution ofFestuca l~rnatii ST.-YV~ and F. gautieri (HACK.) K. RlCtrr.

~eea

~ I Ploidy level

? Uncertain distrib.

F. burnatii DLctribldio~l are~

• .~Jdled material

NX Plotdy level

prickles). Ligules 2-7(-8) mm long, oblong, acute. Leaf-blades (11-)14--43 cm long, densely scabrid (antrorse prickles), becoming spiral to the tip; apex of the leaves obtuse; in cross section oval in outline, (0.4-)0.5-0.7 mm in diameter, with 5(-7) veins, with sclerenchyma forming a complete ring, with 2 adaxial grooves and 1 adaxial ridge without sclerenchyma. Panicle (5.4--)6-17 cm, lax, with about 40-77 spikelets. Spikelets (4.7-)5-8 mm long, with (3-)4--6(-8) fertile florets, green or slightly variegated-purple. Glumes unequal, with wide scarious margins; lower glume 2-3.4(-3.7) ram, 1-veined; upper glume 2.5-3.6(-3.9) mm, 3-veined, oval-lanceolate. Lemma 3.4-4.6(-5.2) rnm long, 5-veined, oval-oblong, with wide scarious margins, mucronate. Palea 3.4--4.4(-5) mm, scabrid on the keels to the tip. Anthers about 2-3 mm long. Ovary glabrous or sparsely pubescent. Caryopsis slightly adherent to the palea.

4.a. Festuca elegans BOISS. subsp, elegans

Iconography: (Fig. lOad, f). Description. Culms (54-)59-112 cm long. Leaf-blades densely scabrid (antrorse prickles),

(15-)20-43 cm long. Panicles (8.3-)9.3-17 cm. Spikelets 6-8 mm long, with (4-)5-6(-8) fertile florets.

Chromosome number. 2n=2x=14. Chorology. Endemic to he Sierras B6ticas (Spain) and Rif Mountains (North Morocco)

(Fig. 11). Ecology. Festuca elegans subsp, elegans grows forming grassland mostly on feldspar

micaschist and on serpentine soils, sometimes on base rich soils, at 1200-2000 m altitude. It occurs in Festucion scariosae MARTINEZ-PARRAS et al. in ALCARAZ 1984 communities.


f g

C

d e

Fig. 10. Festuca elegans BOISS. subsp, elegans, Spain, Granada, Sierra Nevada, Domajo (herb. F. & 0.): a - habit, b - leaf blade section, c - ligule, d - spikelet, f - leaf blade abaxial surface. F. elegans subsp, merinoi (PAU) FUENTE et ORT01~Z, Spain, Avila, Tomavacas, puerto de Tomavacas (herb. F. & 0.): e - spikelets, g - leaf blade abaxial surface.

4.b. Festuca elegans subsp, merinoi (PAU) FUENTE et ORTUI~EZ, comb. nov., hoc Ioco

= F e s t u c a m e r i n o i PAU, Bol. Soc. Aragonesa Ci. Nat. 1(3): 51, 1902. Ind. loc. : "Bosque de Roqueim (Galicia): P. MERINO S. J.". Typus ignotus.

Iconography: (Fig. 10e, g).


c~

) Dialrtbution area

Studled material

~ L E quadriflora I ~ Dlst~buti~n area

I • Studied material

~ _ N x P I o ~ level

Fig. 1 |. Distribution of Festuca elegans BOISS. subsp, elegans, F. elegans subsp, merinot (PAU) FUENTE ¢t O R ~ and F. quadriflora HONCK.

Description. Differs from subsp, elegans principally by: culms 50-84(-90) cm long; leaf-blades scabrid (with fewer antrorse prickles), (11-)14-26(-30) cm long; panicles (5.4-)6-15 cm long; spikelets shorter, (4.7-)5-7(-7.5) mm long; with fewer fertile florets, (3-)4-5(-6).

The principal characters distinguishing the epidermal micromorphology of these two subspecies (subsp. elegans and subsp, merinoi) are given in Tab. 3.

Chromosome number. 2n=4x=28. Chorology. Endemic to Iberian Peninsula (central and northwest mountains) (Fig. 11). Ecology. Festuca elegans subsp, merinoi forms silicicolous grasslands with preponderantly

tufted grasses in Festucion elegantis RrVAS MARTtNEZ et SANCHEZ MATA in RIVAS MART~NEZ et al. 1986 communities (Stipo giganteae-Agrostietea castellanae RIVAS MARTiNEZ et al. 1999).

5. Festuca gautieri (HACK.) K. RICHT.

Festuca gautieri (HACK.) K. RICHT., PI. Eur. 1: 105, 1890. = F. varia subsp, scoparia vat. gautier i HACK., Monogr. Festuc. Eur.: 181, 1882. Ind. loc.: "In Pyrenaeis orientalibus: Col de Nourry l[g]. GODRON; Canigou, Montfort (Vall6e de la Boulsane) Col de la Porteille de Mantet, 2400 m lg. GAUTIER". Lcctotypus (FUENTE et al. 1988: 520): Col de Nourry (Pyr6n6es orient.) (s.a. GODRON W!).

= F. varia subsp, scoparia A. KERN. et HACK. in HACK., Monogr. Festuc. Eur.: 181, 1882. Ind. loc.: "in Pyrenaeis centralibus: Gavamie, Port de Paill~res etc. lgg. BORDII~RE [sic!], TIMBAL-LAGR.". Lectotypus (FUENTE et al. 1988:517): Gavamie, Htes. Pyr6n6es (VII. 1876 BORDI~RE W !).

404 V. de la Fuente & E. Ortuffez

d

c

v

Fig. 12. Festuca gautieri (HACK.) K. RICHT., Spain, Huesca, Chia, Sierra de Chia (herb. F. & O.): a - habit, b - leaf blade section, c - ligule, d - spikelet.


Description. Densely tufted grass. Vegetative shoots intravaginal, with erect or prostrate innovations. Culms (18.4-)20-48(-50) cm long, usually glabrous. Leaf-sheaths fused for 1/2-2/3 of their length, usually glabrous. Ligules 0.4-0.6 mm long, oblong, obtuse-truncate to acute-obtuse. Leaf-blades (4-)5-19(-22) cm long, glabrous, curved or curved-erect, frequently curved-prostrate; apex of leaves acute, very sharp, glabrous; in cross section oval-polygonal in outline, (0.3-)0.4--0.7 mm in diameter, with 5-7 veins, with sclerenchyma forming a complete ring or interrupted forming 7 strands (occasionally decurrent), with 2 adaxial grooves and 1 adaxial ridge without sclerenchyma. Panicles 4-7.4 cm long, erect, with


(6-)9-18 spikelets, with (1-)2-3(--4) branches; the first branch (1.8-)2-3.7(-4.3) cm long, with 3-5(-6) spikelets; the first internode (0.7-) 1.4-2 cm long. Spikelets (6-)7-11 (-11.6) mm long, with (3-)4-6 fertile florets. Glumes unequal, with wide scarious margins, sometimes variegated-purple-spotted; lower glume 3-4.5 mm, 1-veined; upper glume (3.8-)4-5.2(-5.5) mm, 3-veined, oval-lanceolate. Lemma (4.7-)5-7.2 mm long, 5-veined, lanceolate, acuminate, with scarious margins and tip, sometimes variegated-purple, mucronate or with short awns 0.2-0.7 mm. Palea 4.5-6.8 mm long, scabrid on the keels. Anthers 2.5-3.5 mm. Ovary pubescent. Caryopsis free from the palea.

Chromosome number. 2n=2x=14; 2n=4x=28. Chorology. Endemic to Iberian Peninsula (Cordillera Can~brica, Pyrenees, Sistema

Ibrrico and Cordillera Penibrtica) (Fig. 9). The presence of this taxon in North Morocco as cited by some authors (SAINT-YVES 1922: 40, JAHANDIEZ 8~ MAIRE 1931: 74, MAIRE 1955: 161) is based on only one sheet collected by Gandoger (Tetouan: "Djebel Zerka"). We have not yet found another sheet or a more recent reference from North Morocco.

Ecology. Festuca gautieri occurs in perennial xerophytic grassland and vegetation of cryophilous geliturbated soils, above 1400 m altitude. It is a characteristic species of Festuco hystricis-Ononidetea striatae PdVAS MARTINEZ et a1.1991 communities.

Observations. KERGUELEN & PLONKA (1989) recognized two subspecies ofF. gautieri in the Pyrenees: Festuca gautieri subsp, gautieri, tetraploid, growing on acid soils and distributed in the Eastern Pyrenees; and F. gautieri subsp, scoparia (A. KERN. et HACK.) KERGU~LEN, diploid, growing on basic soils and distributed in the rest of the area of the species. These authors distinguished F. gautieri subsp, gautieri from F. gautieri subsp. scoparia principally by the following characters: spikelets (9.5-)10-12 mm vs. 9(-10) mm long, with 3~5 vs. 3-8 fertile florets and lemma 6-7.7 mm vs. 5.2~.2 mm long. We have studied numerous material and we have not found differences allowing one to separate these subspecies, in fact we have observed variability in these characters both for specimens from the same locality and from different localities, and both on basic soils and on acid soils. Moreover, we have found the plants of diploid level in the Eastern Pyrenees on acid soils (Tab. 2).

6. Festuca quadriflora HONCK.

Festuca quadriflora HONCK., Verz. Gew. Teutschl.: 271, 1782. Ind. loc.: "BliJhet auf den Alpen im Brach und Heum". Neotypus (MAP~GRAF-DANNENBERG 1979: 374): Mt. Taveyannaz, Vaud, Suisse, Weld. Ausl~iufer der Diablerets, 1900 m (23.VII.1977 I. & F. MARKGRAF & M. WELTEN Z).

= F. pumila CHA]X, P1. Vapinc.: 12, 1785; CHAIX in VmL., Prosp. Hist. P1. Dauphin6 1: 316, 1786. Ind. loc.: "in alpinis, h Chaudun". Typus ignotus. = F. varia HAENKE subsp.pumila (CHA]X) HACK., Bot. Centralbl. 8: 408, 1881.


l)eseription. Tufted grass. Vegetative shoots intravaginal. Culms (6-)10-20(-30) cm long, scabrid or scabrid-pubescent (antrorse prickles). Leaf-sheaths fused for 0 -1 /4 of their length, glabrous to scabrid (retrorse prickles). Ligules 0.5-1 mm long, oblong, acute-truncate. Leaf-blades 8.5-12 cm long, scabrid (antrorse prickles), erect, thread-like; in cross section polygonal in outline, 0.4-0.6(-0.8) mm in diameter, with 5-7 veins, with

406 V. de la Fuente & E. Ortu~ez

d

Fig. 13. Festuca quadriflora HONCK., Spain, Huesca, Col d'Afiisclo, Fanlo-Bielsa (JACA 372784): a - habit, b - leaf blade section, c - ligule, d - spikelet.

sclerenchyma usually interrupted and forming 7 strands, with 2-4 adaxial grooves and 1 (rarely 3) adaxial ridge. Panicles 2-4 cm long, erect, dense, with 6-10 spikelets, with 1-2 branches. Spikelets 6.5-9(-10) mm, with 3(-4) fertile florets. Glumes unequal; lower glume 3-3.5 mm long, 1-veined; upper glume 3.6-5 mm long, 3-veined, ovate-lanceolate, widely variegated-purple below scarious margins. Lemma 4-6 mm long, 5-veined, lanceolate, variegated-purple-spotted below scarious margins, with an awn 0.2-1.3 mm long. Palea 4-5.5 mm, oblong, scabrid on the keels. Anthers 2-2.5 mm. Ovary pubescent. Caryopsis free from the palea.


C h r o m o s o m e number. 2n=2x=14; 2n=4x=28. Chorology. Jura, Alps and more sparsely in the Central Pyrenees (Fig. 11). Ecology. Festuca quadriflora grows forming grassland on cryophilous geliturbated and

basic soils, above 1800-2000 m altitude. It occurs in Elyno-Seslerietea BR.-BL.1948 communities.

Observations. HACKEL (1882: 177) first cited F. quadriflora (as F. varia subsp, pumila (CHAIX) HACK.) from the Central Pyrenees. Later, other authors (MARKGRAF-DAr, n'~ENBERG (1980), KEI~GU~LI~N & PLONKA (1989) and PORTAL (1999)) confirmed the presence of this taxon in the Central Pyrenees. However, concrete citations of localities from the Iberian Peninsula are very scarce in the herbaria and literature. During our research in this section we have found specimens in herbaria W and JACA from two new localities in the Pyrenees, which correspond to F. quadriflora. Three sheets are from France "Hautes Pyrrnres, Col d' Aure, alt. 2500 m" (25.VIII.1888 BORD~RE W), originally identified as F. varia subsp, pumila and revised by Markgraf-Dannenberg, and another, without determination, is from Spain "Huesca, Col d'Afiisclo, Fanlo-Bielsa, 2480 m, 31TBH6027" (6.VIII. 1984 M. SAULE JACA 372784).

7. Festuca eskia RAMOND ex DC.

F. eskia Ramond ex DC. in Lam. et DC., FI. Franq. 3 : 52, 1805. Ind. loc.: "Cette plante est originaire des Pyrrnres; elle occupe les pentes s~ches des hautes montagnes, oh cUe forme des tapis ~'pais et glissans. Les habitans du pays la nomment eskia. Cette esp~ce a &6 drcouverte par M. Ramond". Lectotypus (N~GRE 1975: 317): Festuca eskia LAM. ft. fr. (= F. varia SCH~D. non HOST), Pyrrn6es (s.a. RAMOND G).

= F. eskia var. orientalis NI~GRE, Candollea 30: 318, 1975. Ind. loc.: "Pyrrnres orientales, Tosse du Puigmar'. Holotypus: Pyrrnres orientales, Tosse du Puigmal (s.a. N~GPa~ RN 5966 herb. N~GRE).


Description. Densely tufted grass. Vegetative shoots intravaginal. Culms 27-50 cm long, erect, glabrous or slightly scabridulous-scabrid below panicle. Leaf-sheaths fused for 0--1/3 of their length, glabrous, rarely scabrid (retrorse prickles). Ligules (2-)2.5-7(:7.2) mm, oblong, acute, scarious. Leaf-blades (14-)17-29(-36) cm long, erect-curved, usually glabrous, sometimes scabrid below (antrorse prickles); apex of leaves acute, very sharp and glabrous; in cross section oval in outline, (0.8-)0.9-1.2 mm in diameter, with 11-15(-17) veins, with sclerenchyma forming a complete and wide ring, with (8-)9-11(-13) adaxial grooves and (7-)9-11(-12) adaxial ridges with sclerenchyma and with abundant trichomes. Panicles (4.8-)6-8.5(-9.6) cm long, erect, lax, with (11-)16-26 spikelets; the first branch (2-)2.7--4.2(-4.5) cm long, with 3-8 spikelets; the first internode 1.1-2 cm long. Spikelets (7.5-)8-I 1 mm long, with 4-6(-7) fertile florets, purple-variegated. Glumes unequal, with wide scarious margins, glabrous to shortly scabridulous; lower glume 3-4.5(-5.2) mm long, 1-veined, sometimes slightly mucronate; upper glume 4-5.3(-5.9) mm long, 3-veined, obovate. Lemma (5.2-)5.5-7.3 mm long, 5-veined, oval-lanceolate, with wide scarious margins, glabrous or shortly scabridulous, mucronate to shortly aristate, with an awn 0.1-1.1 mm long. Palea 5-7.3 mm long, scabrid on the keels to the tip. Anthers 3-3.5 mm. Ovary pubescent. Caryopsis free from the palea.

C h r o m o s o m e number. 2n=2x= 14.


) i t c5

d /L_~.

Fig. 14. Festuca eskia RAMOND ex DC. , Spain, Lleida, Al to Ar~n, Montgar r i (herb. F. & O.): a - habit , b - l ea f blade sect ion, c - l igule, d - spikelet .

Chorology. Pyrenees, Cordillera Cantabrica mountains and Montes de Le6n (NW Spain) (Fig. 5).

KgAJINA (1933: 10) first reported F. eskia from the Carpathians (Romania) and cited only one sheet collected by Ivan Kl~i~tersk~, ("in montibus Retezatu apud Malajesti prope Hateg, altitudine?"). He compared this specimen with others from the Pyrenees and did not find essential differences in respect to the characters given by HACKEL (1882) for F. eskia from the Pyrenees. The presence of this taxon in Romania cited by KOPFER (1974: 90) is based on the work by K.gAJINA (1933). Later, MARKGgAF-DANNENBERG (1980:133) recognized F. eskia for the Pyrenees, but doubted the presence of this taxon in Romania. The description given by


KRAJINA (1933) shows two important differences from the specimens studied by us from the Pyrenees and from NW Spain namely leaf-sheaths fused for 2/3 (vs. base-I/3) and the presence of lax tuft (vs. dense). KRAnNA (1933) did not specify the presence ofintravaginal or extravaginal vegetative shoots for the specimen from Romania which is an important character. It would be important to study more populations from the Carpathians in order to study their variation.

Ecology. Festuca eskia occurs in psychroxerophilous grazing land on acid soils, above 1400 m altitude. It also grows in stone quarries and on stony slopes. It is a characteristic species of Festucion eskiae BR.-BL.1948, Juncetea trifidi HADA~ 1946 (RIVAS GODAY 8~ RIVAS MARTtNEZ 1963:212)

Observations. NI~GRE (1975:318) described F. eskia var. orientalis from the Pyrenees based principally on the presence of scabrid and cylindrical-subcylindrical culms, mucronate-aristate glumes and lemmas always with an awn. We have studied numerous material from these mountains and observed the existence of a high degree of variability in the characters cited by N~.GRE (1975), both for specimens from the same locality and from different localities, and both in the Western Pyrenees and in the Eastern Pyrenees. So we do not consider vat. orientalis worth recognizing.

N~GRE (1975: 318) described another variety, F. eskia var. tenuifolia, from "Montes Cantabriae, l'Espiguete, Collado de los Arran, 1900 m .... "(Spain). He separated this taxon from the other varieties by the thin leaf-blades, and said that it was taxonomically close to F. scoparia (A. KERN. et HACK.) NYMAN (F. gautieri (HACK.) K. RICHT.). We have analyzed the type material of vat. tenuifolia and found that it corresponds with F. ×picoeuropeana NAVA (NAvA 1985: 23) (F. gautieri x F. eskia).

8. Festuca × picoeuropeana NAVA (F. eskia RAMOND ex DC. × Fo gautieri (HACK.) K. RICHT.)

Festuca ×picoeuropeana NAVA, Fontqueria 7: 23, Jan 1985, pro sp. Ind. loc.: "Asturias, Picos de Europa, Macizo del Corni6n, Tone de los Traviesos, 30TUN48, 2300 m". Holotypus: Asturias, Picos de Europa, Macizo del Corni6n, Torre de los Traviesos, 30TUN48, 2300 m (8.IX.1983 NAVA FCO 10880!)

=F. xpicoeuropeana GUTII~RREZ VILLARIAS et HOMET, Bol. Ci. Naturaleza I.D.E.A. 34: 146, Mar 1985, nora. iUeg. (Art. 53). Ind. loc.: "Cantabria: Collada de las Nieves: Macizo Central de los Picos de Europa, 2100--2200 m, 30TUN5180". Holotypus: Cantabria: Collada de las Nieves: Macizo Central de los Picos de Europa, 2100-2200 m, 30TUN5180 (26.VIII. 1983 H.S. NAVA FERNANDEZ FCO 10875!)

= F. eskia var. tenuifolia N~GRE, Candollea 30:318, 1975. Ind. loc.: "Montes Cantabriae, l'Espigiiette, Collado de los Arran, 1900 malt.". Holotypus: Montes Cantabriae, l'Espigtiette, Collado de los Arran, 1900 malt . (12.VIII.1953 P. MONTSERRAT JACA 2053!)

-F . Xsouliei auct. non. ST.-YvEs (1924): CATALAN, Taxon 39: 661, 1990. Iconography: (Fig. 15).

Description. Densely tufted grass. Vegetative shoots intravaginal. Culms (13-) 15-36(-47) cm long, erect, glabrous or slightly scabridulous below panicle. Leaf-sheaths fused for (1/4-)1/3-1/2(-2/3) of their length, glabrous, sometimes scabridulous (retrorse prickles).

410 V. de la Fuente & E. OrtuSez

Fig. 15. Festuca xpicoeuropeana NAVA, Spain, Cantabria, La Vueltona, Pieos de Europa (herb. F. & O.): a - habit, b - leaf blade section, c - ligule, d - spikelet.

Ligules 0.5-4(-6.8) mm, oblong, with variable shape (acute-obtuse to obtuse or obtuse-tnmcate). Leaf-blades (8-)9-17(-22.5) cm long, in general glabrous; apex of leaves acute, very sharp and glabrous; in cross section oval-polygonal in outline, sometimes oval, (0.5-)0.6-0.8(-0.96) mm in diameter, with 7-9(-11) veins, with sclerenchyma forming a complete ring, with (2-)4-6(-8) adaxial grooves and (3-)4-7 adaxial ridges with or without sclerenchyma and with trichomes. Panicles (3-)4.5-6.3(-7.2) cm long, erect, lax, with 10-16(-25) spikelets, with 2-4 branches; the first branch 2.2-3.5(-3.7) cm long, with 3-5(-7) ;pikelets; the first intemode 1-2.1(-2.3) cm long. Spikelets 7-11 mm long, with (3-)4-5(-6) fertile florets, sometimes purple-variegated. Glumes unequal, with wide scarious margins;


lower glume (3-)3.6-4.7 mm, 1-veined; upper glume (3.8-)4-5.6 mm, 3-veined. Lemma (5.5-)6-7 mm long, 5-veined, with scarious margins and tip, mucronate or with short awns (0.1-)0.24).7(-1) mm long. Palea of similar size to the lemma. Anthers (1.5-)2.1-3.8 mm. Ovary pubescent. Caryopsis free from the palea.

Chromosome number. 2n=2x=l 4. Chorology. Endemic to Iberian Peninsula (Picos de Europa and Pyrenees) (Fig. 7). Ecology. Festuca ×picoeuropeana grows on basic soils at 1900-2500 m. In the Pyrenees it

occurs together with F. eskia and F. gautieri on basic or acid soils. Observations. In the Picos de Europa this fescue grows without the parents (F. esl~'a and

F. gautieri). In the rest of the distribution area it grows in populations with the two parents and shows a high degree of variability.

9. Festuca × souliei ST.-YVES (Festuca eskia RAMOND ex DC. × F. quadriflora HONCK.)

Festuca ×souliei ST.-YVES, Bull. Soc. Bot. France 71: 126, 1924. Ind. loc.: "France. Pyrrnres: Hras au pic de Gabiedou, leg. SOULII~ (cum ssp. pumila) (herb. Coste); vallre de Campbieil (cure ssp. pumila) leg. SENNEN (herb. COSTE) et leg. BORDI~RE (herb. J. CHEVALIER); Aragnouet pros du port de Baroude, leg. SOULII~ (herb. COSTE et herb.St.-YvEs); Pie Blanc, leg. BORDI~RE in herb. GAUTIER et in SCHULTZ, herb. norm., no. 981 sub "F. eskia" (herb. Deless.)". Lectotypus (GUTIl~RREZ VILLARIAS et al. 1992b: 77): Htes.Pyrrnres: Aragnouet; PrOs du port de Barroude, 2500 m, (10.VII.1913 SOULIE G-Herb. ST.-YVES s.n.!). Isolectotypus: (MPU-CoSTE s.n.!).


Description. Tufted grass. Vegetative shoots intravaginal. Culms 15-26.5 cm long, scabrid below the panicle. Leaf-sheaths fused for 1/4 of their length, glabrous or scabrid (antrorse prickles). Ligules 1.2-2 mm long, oblong, acute or acute-truncate. Leaf-blades about 12-17 cm long, glabrous to scabrid to the base (antrorse prickles); apex of leaves acute, not or slightly sharp; in cross section oval-polygonal in outline, 0.6-0.7 mm in diameter, with 7-10 veins, with sclerenchyma forming a ring, with 4-6 adaxial grooves and 3-5(-7) adaxial ridges unequal with or without sclerenchyma. Panicle 4-5 cm long, lax, with about 10 spikelets, with 2 branches; the first branch about 2.7 mm long, with 3-4 spikelets. Spikelets 7.8-10 mm long, with 4-5 fertile florets. Glumes unequal, variegated-purple-spotted below scarious margins, margins glabrous; lower glume 3-4 mm long, 1-veined; upper glume 4-5 mm, 3-veined, acuminate. Lemma 5-6 mm, variegated-purple-spotted below scarious margins, glabrous or scabridulous to the tip, mucronate or with short awns about 0.5 mm. Palea 5-6 mm, scabrid on the keels. Anthers about 3 mm. Ovary pubescent. Caryopsis free from the palea.

Chromosome number. Unknown. Chorology. Central Pyrenees. The distribution of this taxon is insufficiently known; the

same applies to its ecology. Observations. This taxon shows much similarity with F. ×picoeuropeana. Both are hybrid

taxa and apparently with one common parent, F. eskia. We have studied the type of F. ×souliei (G!) together with some specimens from Spain (Pto. de Barrosa, Bielsa, Huesca, 2570 m, 31TBH6935 (26.VIII.1998 P. CATAL,~N JACA 127498)) and these present similar characters to those of F. ×picoeuropeana, both anatomically and morphologically (see also Tab. 3). The presence ofscabrid culms on F. xsouliei (v. glabrous or slightly scabridulous on

412 V. de la Fuente & E. Ortu5ez

I

b

d

16. Festuca xsouliei ST.-YVES, France,.Pyr6n6es, port de Barroude (G s.n.): a - habit, b - leaf blade section, d - ligule, e - spikelet; Spain, Huesca, puerto de Barrosa (JACA 127498): c - leaf blade section.

F. xpicoeuropeana), leaf-sheaths fused for 1/4 (vs. fused ( I /4-)1/3-1/2(-2/3)) , glabrous or

scabrid leaf-sheaths (v. glabrous, sometimes scabridulous), leaf-blades glabrous and slightly

scabridulous to scabrid at the base (vs. in general glabrous) and lemmas 5-6 mm long (vs. (5 .5-)6-7 nun) are the principal characters observed by us that allow one to separate

F. xsouliei from F. xpicoeuropeana.

Acknowledgements: We thank Prof. C. Navarro (Madrid) for her advice and criticism concerning nomenclatural questions, Mrs. M. C. de la Fuente for the English revision, Prof. R. Gamarra (Madrid) for the revision of the text and chorological advice and Dr. L. M. Ferrero for the chromosome counts, as well as the directors and curators of the consulted herbaria for providing the plant material.


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diploid species of Festuca. Chromosoma (Berlin) 75:117-127. NAVA H.S. (1985): El grupo Festuca varia en Picos de Europa. Fontqueria 7: 21-24. NI~GRE R. (1975): Observations morphologiques sur les gentianes du groupe alpina-acaulis, star Festuca

paniculata et F. eskia en Pyr6n6es. Candoltea 30: 301-321. PALMER P.G. & TUCKER A.E. (1981): A scanning electron microscope survey of the epidermis of East African

grasses. 1. Smithsonian Contr. Bot. 49: 1-84. PORTAL R. (I 999): Festuca de France. Imprimerie Jeanne-D'Arc, Le Puy-en-Velay. RIVAS GODAY S. & RIVAS MARTINEZ S. (1963): Estudio y clasificacidn de los pastizales espa~oles. Pub.

Ministerio Agricultura, Madrid. RIVAS MARTtNEZ S., ASENSI A., MOLERO MESA J. & VALLE F. (1991): Endemismos vasculares de Andalucia.

Rivasgodaya 6: 5-76. ROMERO ZARCO C. (1984): NOmeros cromos6micos para la flora espafiola. Nfirneros 337-341. Lagascalia 12:

292-294. ROUGEOT M.C. (1974): Contribution ~ l'6tude taxonomique du Festuca varia HAENKE ssp. eu-varia HACK.

des Alpes franqaises. Ann. Sci. Univ. Besan¢on, Bot., 15: 59-108. SAINT-YVES A. (1913): Les Festuca de la section Eu-Festuca et leurs variations dans les Alpes Maritimes.

Annuaire Conserv. ,lard. Bot. Gen~ve 17: 1-218. SAINT-YVES A. (1922): Les Festuca (subg. Eu-Festuca) de rAfrique du Nord et des lies Atlantiques.

Candollea i: 1-63 SEAL A.G. (1983): DNA variation in Fesmca. Heredity 50: 225-236. WILKINSON M. J. & STACE C. (1991): A new taxonomic treatment of the Festuca ovina L. aggregate (Poaceae)

in the British Isles. Bot. J. Linn. Soc. 106: 347-397. WILLKOMM M. (1861): Faro 19. Gramineae JUSS. In: WILLKOMM M. & LANGE J. (eds.), Prodromusflorae

hispanicae L Stuttgart, pp. 33-118.

Received 11 May 2000, revision received 10 November 2000, accepted 21 June 2001 Encl. Appendix - Material studied pp. 415-418; Figs. 1-3 pp. 419-421


APPENDIX

Material studied

F. pseudeskla BOlSS. Spain. Almeria: 30SW019 0dTM coordinate), Sierra Nevada, Laujar, 2000 m (2.VIII.51 s. coll. MA 176234). Granada: 30SVG60, in sumnfis glareosis Sierra Nevada, 1000' (VIII. 1837 BOISSIER G s.n., K s.n.). Loma de Rio SeeD, lagtma de Rio Seed (3.V]].1990 V. DE LA FUENTE herb. F. & O.). Sierra Nevada, Veleta (14.VIII.1992 V. DE LA FUENTE herb. F. & O.). Sierra Nevada, Cueva de Panderone, 2300 m (30.VII.1876 HACKLE W s.n.). 30SVG70, Sierra Nevada, in glareosis ad Pefi6n de San Francisco, 2400 m (VII. 1891 PORTA & RICO MA 12491 ). Sierra Nevada, Loma del rio SeeD, Mulhac6n (3.VII. 1990 V. DE LA FUENTE herb. F. & O.). Sierra Nevada, pefias de Vacates (31 .VIII. 1923 GROS BC 81769).

F. scariosa (LAG.) ASCH. et GRAEBN. Spain. Almerfa: 30SWG 1 I, barranco de Fifiana (5.VII. 1908 ST.-LAGER K s.n.). 30SWG41, Sierra de G~dor, Sierra Nevada, Filab(es (V. 1992 A. PALLARI~S MA 546486). 30SWG6617, Sierrade los Filabres, Benizalbn, 800 m (6.V.1988 A.B. ROBLES & C. MORALES GDAC 28358). 30SWG66, Chirivel, Francos, 1650 in (1 .VII. 1980 A. SEGURA ZUBIZARRETA MA 232742). 30SWG76, V61ez-Rubio, Sierra de Maim6n, 1300-1600 m (VII.1899 E. REVERCHON MA 265485). 30SWG77, Mafia, Sierra Maria, 1500 m (26.VI.1984 A. SEGURA ZUBIZARRETA MA 591347). Cidiz: 30STF86, Benaocaz (24.VI.1925 GROS BC 111113). 30STF87, Grazalema (24.V.1884 Pl~REZ LARA MA 12591). 30STF88, Algodonales, Sierra de Lijar, 1000-800 m (31 .V. 1980 A. APARICIO & B. CABEZUDO MGC 11341). C6rdoba: 30SU093, Cabra, ermita de la Virgen de la Sierra, 800-1200 m (13.VI. 1984 J. ARROYO, MEJIA & S. TALAVERA UNEX 1339). Sierra Tifiosa, Priego (20.VI.1961 J. BORJA MA 187387). Granada: 30SVF0787, Sierra Tejeda, entre Barracones, 1700 m (26.V.1982 J.M. NIETO MCrC 20453). 30SVG41, Silla del Moro (5.VII.1876 M. WINKLER K s.n.). Solaria Alfacar (18.VI1.1975 F. VALLE GDAC 9892). 30SVGS0, Diechar, carretera desvio a Diechar km. 1 (2.VII.1990 V. DE LA FUENTE & E. ORTI3"~EZ herb. F. & O.). 30SVG60, in omnibus montibus regn. Granat., 4000-9500' (V11.1837 E. BOISSIER VI].1837 G s.n., MA 12593). Base de Sierra Nevada (26.VI.1851 E. BOURGEAU K s.n., MA 12601). Antequera et Sierra Nevada, 1000 m (16.VI. 1879 HUTER, PORTA & RIGO K s.n.). 30SVG61, Encinar de Giiejar-Sierra (s.d.P. PRIETO & P. ESPINOSA GDAC 2950). 30SWG02, Sierra de Baza, Rambla del Agua (28.VI. 1984 J. TORRES, G. BLANCA & C. MORALES GDAC 26303). Ja6n: 30SVG35, Carboneras, barr. Atanor, 1050 m (6.VI.1926 CUATRECASAS BC 69945). 30SV047, Sierra de Ja6n (20.VI. 1941 E. GUINEA MA 447775). 30SV057, Sierra M~gina, Cerro la Vieja (15.VI. 1926 J. CUATRECASAS MAF 27532). 30SV068, La Serrezuela de Bedmar, in declivibus Jodar, 900 m (7.VI. 1926 CUATRECASAS BC 69943). Milaga: 30SVF08, Sierra Tejeda (9.VI.1935 M. LAZA MA 12596, MAF 27531). 30SVF17, Los Hornajos, Canillas de AIbaida (8.VI.1919 GROS BC 69946, MA 12594). 30SVF18, Sierra Almijara (23.VI.1935 M. LAZA PALACIOS MA 12597).

F. burnatii ST.-YVES Spain. Astarias: 29TQH4371, Puerto Ventana (Teverga), 1600 m (17.VII.1995 V. DE LA FUENTE & L. M. FERRERO herb. F. & O.). Cantabria: 30TUN57, Espinama, Picos de Europa, Fuente D6 - Aliva (3.VII.1987 B. CABEZUDO MGC 42439). 30TUN58, Pefia Vieja, pr6s Espinama, 2500 m (31.VII. 1914 J. SOULI~ MPU-Coste s.n.). Puerto de Aliva, Casaseca (15.VII. 1976 LADERO & G. LOPEZ GDA 6344). Montes Picos de Europa, circa Aliva (VII. 1879 L. LERESCHE & E. LEVER FI s.n.). Le6n: 29TPH90, Ferradillo, 900 m (7.VII. 1984 J. ANDRI~S LEB 28663). 29TQG09, Santiago de Pefialba (9NII. 1983 F. LLAMAS & J. ANDRI~S LEB 22129, LEB 25456, LEB 37354). 29TQH26, Puerto de Somiedo (28.VII. 1984 E. PUENTE LEB 24278). 29TQH36, La Cueta, Picos Blancos (31.VIII.1984 E. PUENTE LEB 26228). 29TQH46, La Majua (7.V11.1973 C. ROMERO LEB 28469, LEB 4962). 29TQH47, Puerto Ventana, 1600 m (17.VII.1995 V. DE LA FUENTE & L. M. FERRERO herb. F. & O.). 30TTN75, Collada de Aralla (7.VII. 1973 C. ROMERO LEB 18473). Puerto Aralla (10.VI. 1983 C. ROMERO LEB 37329). 30TTN95, Valporquero (22.VII. 1973 J. ANDRI~S LEB 46808). 30TUN05, Hoces de Valdeteja (14.VII.1978 M.J. LOPEZ PACHECO LEB 16297). 30TUN25, Cr6menes (11.VI.I972 J. ANDRl~S & R. CARB0 LEB 21774). 30TUN26, Marafia (6.VII. 1980 FERNANDEZ ARECES & Pl~REZ CARRO LEB 1125 l, LEB 11252). Palencia: 30TUN86, Puerto de Piedrasluengas, ad Aquilonem, 1400 m (26.V1.1954 LAiNZ BC 126623, K s.n.).

416 V. de la Fuente & E. Ortuffez

F. elegans BOISS. subsp, elegans Spain. Granada: 30SVF08, Sierra Tejeda, Fuente del Robledal (21.VI.1980 J. M. NIETO MGC 20471). 30SWG13, Sierra de Baza, Calar de Santa Barbara, 2000 m (28.VIII.1997 R. GAMARRA & E. ORTI31~F.Z herb. F. & O.). Milaga: 30SUF16, Ronda, Sierra de las Nieves, Puerto de los Pilones, 1750 m (23.VI.1995 B. CABEZUDO, A. PI~REZ LATORRE, P. NAVAS, D. NAVAS & Y. GIL MGC 43474). 30SUF26, Sierra de las Nieves, Puerto del Oso, 1700 m (9.VI.1998 A. ASENSI, B. DtEZ GARRETAS & V. E. MART1N OSORIO MGC 24652, MGC24672, MGC 24671). 30SUF2163, Tolox, P. N. Sierra de las Nieves, Vereda al TorreciUa, cercania al Puerto de los Pilones, 1720 m (12.VI.1997 A. PI~RF.Z LATORRE, P. NAVAS, Y. GIL & D. NAVAS MGC 45974). 30SVF27, Yunquera, Sierra de las Nieves, E1 Pinsapar - La Pefia de Enamorados, 1500 m (4.VII. 1991 B. CABEZUDO, J.M. NIETO, A. PI~REZ LATORRE & J. J. VERA MGC 32942). Morocco. 30SUD27, in cedretis montis Tiziren, 1800-2000 (18.VI.1928 MAIRE MPU-Afrique Nord s.n.). 30SUD42, Aflante rhiphaeo, Bu-Mesiat, 1600 m (13.VI. 1929 FONT QUER BM s.n., MPU-Afrique Nord s.n.). 30SUD82, in Atlante rifano: Bab Amegas, 1500-1600 m (17.VI. 1928 MAIRE MPU-Afrique Nord s.n.).

subsp, merinoi (PAU) FUENTE ell OITT~I~IEZ Spain. Avlla: 30TTK7361, Tomavaeas, Puerto de Tornavacas, 1270 m (10.VI.1989 S. SARDINERO herb. F. & O.). 30TUKI 7, Navarredonda (26.VI. 1966 RIVAS GODAY, MAYOR & IZCO FCO 6780). 30TUK1681, Puerto de Chia, Sierra de Villafranca, 1663 m (7.VIII.1990 E. ORT~EZ herb. F. & O.). 30TUK3563, Puerto de los Serranillos, 1580 m (VII. 1991 E. ORTOREZ herb. F. & O.). Ciceres: 30TTK76, Puerto de Tornavacas, 1000 m (21.IV.1988 J.A. DEVESA & A. ORTEGA UNEX 8422). Le6n: 30TTN74, Puerto de la Magdalena, 1457 m (27.VII. 1982 AGUILELLA VAL 09994). Zamora: 29TPG95, a t Km SW de Lobeznos y a 4 Km SW de Puebla de Sanabria, 1000 m (18.V.1972 R. K. BRUMMITT & A. O. CHATER K s.n.). Portugal. Beira ARa: 29TPE26, sanatorio de Covilha, Covilha - Penhas da Saude, Serra da Estrela, 1250 m (26.V. 1995 V. DE LA FUENTE, C. DEL PALACIO & L. M. FERRERO herb. F. & O.). Douro Litorai: 29TNF9465, subida a la Fraga da Ermida, Serra do Marao (22.VII.1996 L. M. FERRERO herb. F. & O.).

F. gautieri (HACK.) K. RICHT. Spain. ,~lava: 30TWN2916, Pipa6n, Sierra de Cantabria, W Cervera, 1300 m (5.VII.1985 J. A. ALEJANDRE MA 339921). Alicante: 30SYH38, Sierra de Aitana (4.VII.1949 RIVAS MARTINEZ & BOP, JA K s.n., MA 170010, MAF 25071). Almeria: 30SWG77, V61ez Blaneo, Sierra de Maria, above Maria, 1800 m (26.VIII.1981 M.F. & S.G. GARDNER K s.n.). Barcelona: 31TDG06, Bag/L, 1000 m (15.VI.1908 ST.-YVES MPU-Coste s.n.). Cantabria: 30TUN58, Pico Cort6s, 2130 m (30.VIIL 1982 S. RIVAS MARTINEZ et al. herb. F. & O., LEB 28622). Ibidem, (14.VIII.1982 E. PUENTE LEB 14310). Castell6n de la Pinna: 30TYK0331, Alt Palancia, Pina de Montalgrao, Cerdafia, 1050-1200 (27.V.1989 J. RIERA VAL 21632). 30TYK37, Alt Maestrat, Vilafranca Penya Bermeja (2.VIII.1981 AGUILELLA VAL 31216). 31TBF50, Els Ports, Coratx~ (7NII. 1988 AGUILELLA VAL 31230). 31TBF61, Montes de Fredes, 1200 m (VI. 1979 MATEO & MANSANET MA 463001, MA 463002). Gerona: 31TCG8383, Sierra del Cadi, pr. Arseguel, 1890 m (23.VII. 1993 C. AEDO et al. MA 529426). 31TDG09, Pyr6n6es-Orientales, Montagne de la Font de Comps (8.VII.1897 SENNEN BC-Sennen s.n.). 31TDG39, Pyr6n6es/t Nuria, 2000 m (20.VII.1914 SENNEN MA 12531, MPU-Coste s.n., W s.n.). Granada: 30SWGI8, Sierra de Castril, 1800 m (VII.1903 REVERCHON MA 12535). 30SWH30, in Flechtenwiilderm bei der Stadt Pobla de Don Fadrique in der Sierra de la Sagra, 1900 m (VII. 1900 REVERCHON MA 263335, W s.n.). 30SWG3597, Sierra de la Sagra, 1700 m (1900 REVERCHON MPU-Coste s.n.). Guipuzcoa: 30TWN5159, Ofiate, Sierra de Aizkorri, Arizamburu, 1350 m (6.VII. 1985 J. A. ALEJANDRE MA 339953). 30TWN5357, Segura, Sierra de Aizkorri, 1350 m (10.VI1.1985 J. A. ALEJANDRE MA 339932). Huesca: 30TXN8152, Ans6, Linza, Loma del Sobrante, 1500-1600 m (1.VII.1991 D. GOMEZ & P. MONTSERRAT JACA 121291). 30TYN24, Bafios de Panticosa, Zubia (12.VII.1882 s. coll. MA 12532). 30TYM28, Nocito,/l la Sierra de Guara, 1800-2000 (3.VIII.1909 J. SOULII~ MPU-Coste s.n.). 31TBH52, Monte Perdido, 2700 m (20.VII. 1965 S. RIVAS MARTINEZ et al. BCF 00213). 31TCH0111, Valle de Llauset, subida al Collado de Vallibiema, 2100-2600 m (17 .VIII. 1984 AIZPURU & P. CATALAN ARAN 2748 l, ARAN 27485). Ja~n: Sierra de Segura, 1700 m (VII.1906 REVERCHON MA 12534). La Rloja: 30TWMI 5, Sierra de Urbi6n, La Traici6n (2.VIII. 1981 G. NAVARRO herb. F. & O.). Lleida: 3 ITCG25, Montsec de R6bies, obaga del Mira Pallars, 1500 m (2.Vll. 1978 A. ROMO BC 687368). 31TCG3049, La Noguera, Serra de Sant Mamet, Barranc de Sant Mamet, 1100 m (11.VI.1980 A. ROMO BC 687347). 3 ITCG8282, Sierra del Cadi, pista desde Adra6n, 2200-2300 m (7.VIII. 1987 L PEDROL & C. PEDROL MA 439238). 31TCG8496, Aristot-Toloriu,


Sierra de las Pinas, 2150 m (13.VII. 1995 R. GAMARRA herb. F. & O.). 31TCH23, Salardfl, massif de Ruda, 2200 m (8.VIII. 1912 SOULII~ MPU-Coste s.n.). Navarra: 30TXN6261, Ochagabia, cima del Orhy, 1800 m (22.VII.1987 AIZPURU & P. CATALAN ARAN 27490). 30TXN85, Puerto de La Piedra de San Martin, 1700 m (2.VII. 1991 V. DE LA FUENTE herb. F. & O.). Saria: 30TWMI5, Santa ln6s, Zorraquin-Majadarrubia, 1500 m (24.VI. 1964 SEGURA MA 363084, MA 363084). Teruel: 30TXK64, Sierra de Javalambre, 1700 rn (V. 1988 V. DE LA FUENTE herb. F. & O.). 30TXK83, Manzanera, Fuente Tejeda (23.VI.1984 AGUILELLA VAL 9222). 30TXL9305, Aliaga, rio Laval, 1100 rn (11.IV.1990 P. MONTSERRAT JACA 66090). 30TYK07, Sierra de Gfidar, Monegro de Linares (30.VI. 1960 BORJA MAF 68497). Valencia: 30SXK81, Sierra de Sacane [Saeafiet]

la Salada, 1800 m (VII.1891 REVERCHON W s.n,), Zaragoza: 30TXM02, Sierra de Moncayo, E1 Morr6n (20.VI. 1987 G. NAVARRO herb. F. & O.). Andorea, 3 ITCH7602, St Juli/~ de L6ria, 940 m (12.V. 1997 V. DE LA FUENTE & L. M. FERRERO herb. F. & O.). 3ITCH7311, Pal, Col de la Botella, 2100 m (16.VIII.1998 R. GAMARRA & E. ORTf~EZ herb. F. & O. ). France. 30TYN43, aufKalkger611 des Cirque de Gavarnie, ca. 1600 m (5.VIII.1902 BORD~RE MA 263336, K s.n.). 31TBH53, Gavarnie, hautes Pyr6n6es, Environs de la Hourquette d'Alians [d'Alans], 2200-2400 m (28.VII.1947 B. DE RETZ W s.n.). 31TDG39, Pyr6n6es-Orientales, Col de Nourry (Pyr6n6es orientales) (s.d. GODRON W s.n.). 31TDH10, Pyr6n6es-Orientales, Cerdagne, Val d'Angoustrine, 1600 m (4.VII SENNEN MA 12528).

F. quadriflora HONCK. Spain. Huesea: 31TBH52, Col d'Afiisclo, Fanlo - Bielsa, 2480 m (6.VIII.1984 M. SAULE JACA 372784). France. 31TBH63, Gabiedou (Hantes-Pyr6n6es) (VIII. 1869 BORDgRE K s.n.). Sommet du Gabiedou a He, as, Hantes-Pyr6n6es (VIII. 1863 BORDE"P,E K s.n.). 31TBH64, Hantes Pyr6n6es, Col d'Aure, 2500 m (25.VIII. 1888 BORDI~RE W s.n.). Italy. Bolzano, Selva di Val Gardena, Tyrol m6ridional, hauteur desservie par le t61eph6rique Wolkenstein-Dantercepies, 2200 m (17.VIII.1986 G. VAN BUGGENHOUT VAL 016687). Switzerlnd. Zermatt, Rottenboden, 2750 m (13.VIII.1990 M LUCEI~O & P. VARGAS MA 526332).

F. eskia RAMOND ex DC. Spain. Asturia$: 29TQH06, Pico de Canellas (VII. 1835 DURIEU K s.n.). Cantabrin: 30TUN8566, Pico Tres Mares, sierra de Pefia Labra, 2175 m (19.V11.1995 V. DE LA FUENTE & L. M. FERRERO herb. F. & O.). 30TUN96, Pico (]el Cordel, supra Hotel La Corza Blanca (14.VII.1969 BORJA & VALDI~S GDA 7110, LEB 4979). Gerona: 31TDG39, Pyrenees ~ Nuria, 2100 (3.VIII.1914 SENNEN MPU-Coste s.n.). Huesea: 30TXN93, Sayerri, 2000 m (3.VI. 1982 V. DE LA FUENTE herb. F. & O.). 30TYN06, Jaca, Ib6n de las Truchas, 2150 m (15.VIII.1984 M.I. GUTIERREZ VILLARIAS FCO 10891, FCO 10890, FCO 10897). 30TYN33, Bujaruelo, 2100 m (25.VILI982 F. BOISSET VAL 01200). 31TBH6022, Puertolas-Forquela, hacia Cuello Viceta, 2000 m (4.VIII.1991 J.V. FERRANDEZ JACA 330091). 31TBH9927, Benasque, Valle de Lliterola, subida a Fites de Perdiguero, 2260 m ( 19.VII. 1987 J.A. SESI~ et al. JACA 773087). 31TCH92, Sommet du port de Venasque dans les Pyr6nees (VIII.1863 TIMBAL-LAGRAVE K s.n.). Le6n: 29TPH74, Pico Cuifia (15.VIII.1982 L. HERRERO LEB 14966). 29TQH06, Laguna de Arras, Leitadegos (19.VII.1984 M.I. GUTIERREZ VILLARIAS FCO 10892). 30TTN96, Pico Huevo ( 19.VII. 1979 M.J. LOPEZ PACHECO LEB 17079). 30TUNI6, Puebla de LiUo (18.VII.1977 E. HERNANDEZ LEB 50232, LEB 10272). 30TUN46, Portilla de la Reina (27.VII.1972 J. ANDR1LS LEB 10164, LEB 10165). 30TTN96, Redilluera (31.VII.1978 M. J. L6PEZ PACHECO LEB 16212, LEB 17236). Llelda: 31TCH02, Pyrenaei eentrales, Port de la Picadeq 2000 m (27.VII.1924 CUATRECASAS GDA s.n.). 31TCH12, Lago Red6 (23.VII.1980 J. ANDR]~S LEB 13303). 3 ITCH1922, Bol Estany Rius, 2360 m (28.VII.1988 J.V. FERRM~EZ JACA 454088). 31TCH31, Estany de Munyidera, Espot-Boi, 2250 m (27.VII.1976 A. CARRILLO & J.M. NINOT JACA 643291). Orense: 29TPG79, Casayo (28.VIII. 1983 CASTROVIEJO et al. SALA 43400). 29TQH37, Pico Ferreirfia (22.VII. 1972 L. HERRERO LEB 14851). Paleaein: 30TUN86, Pefia Labra, 1800 m (26.VII.1952 P. LMNZ GDA s.n., K s.n.). Andorra. 3ITCH7621, El Serrat, Arcalis, 2400 m (17.VIII.1998 R. GAMARRA & E. ORT01qEZ herb. F. & O.). 3ITCH9410, Port d'Envalira, Soldeu, 2500-2400 m (9.VIII.1978 F. FILLAT & L. VILLAR JACA 320778). France. 31TBH53, Htes. Pyr6n6es, G6dre, Pic Blanc, 2000 m (VIII.1874 E. BORDI~RE K s.n.). 31TDG99, Recasens au dessous de l'hermitage (29.V.1847 E. BORDI~RE K s.n.). 31TDHI3, Port de PaiU6res (Ari6ge), 1800 m (3.VIII. 1879 A. GUILLON K s.n.).


F. x picoeuropeana NAVA Spain. Asturias: 30TUN4186, Torte de los Traviesos, Macizo Occidental de los Picos de Europa, 2300 m (8.IX.1983 H. NAVA FCO 10880). 30TUN68, Pico Castrillo Grajal (6.VI.1983 H. NAVA LEB 24537). Cantabria: 30TUN5180, Collada de las Nieves, Macizo Central de los Picos de Europa, 2100-2200 m (26.VIII.1983 H. NAVA FCO 10875). 30TUN68, Cargadero de las minas de Andara - Vegas de Anadara, Pefiarrubia (4.IX. 1988 P. MONTSERRAT, M. LA[NZ & L. VILLAR JACA 298088). Huesca: 30TXN9041, Pico de Ag/ierri, Siresa, Hecho, 2200 m ( 17.VII. 1974 L. VILLAR JACA 10054574). 30TYN 13, Pico Tres Hombres, Formigal, Pirineo Central, 2200 m (29.VI.1990 V. DE LA FUENTE herb. F. & O.). 31TCHI017, barranco de Llauset, Llauset, 2390 m (13.VII.1989 M. B. GARCIA JACA 585692). Le6n: 30TUN48, Cuetalbo, 2140 m (7.VIL1983 H. NAVA LEB 24538). Torte de Pefialba (26.VI11.1981 M.T. PONGA LEB 21241). 30TUN57, Torre de1 Regalliz (29.VII.1981 M.T. PONGA LEB 21245). Lleida: 3ITCH3426, Alto Aneu, puerto de la Bonaigua, camino de la Collada de Muntany6, 2100 m ( 11.V. 1997 V. DE LA FUENTE & L. M. FERRERO herb. F. & O.). Paiencia: 30TUN5257, Valverde de la Sierra, Pico del Espigiiete, Collado de Arras, 1900 m (12.VIII.1953 P. MONTSERRAT JACA 2053).

F. xsoullei ST.-YVES Spain. Huesca: 31TBH6935, Puerto de Barrosa, Bielsa, 2570 m (26.VIII. 1998 P. CATALAN JACA 127498). France. 31TBH63, Hautes Pyr6n6es, Aragnouet, pr6s du port de Barroude, 2500 m (10.VII.1913 SOULI'//G s.n., MPU-Coste s.n.). Hautes Pyr6n6cs, G/tdros, vallon de Campliel (7.VIII. 1903 SENNEN MPU-Coste s.n.).

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