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
Go
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he p
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(198
0)
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seud
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seud
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AC
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. var
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T.-Y
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. R
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ON
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~LEN
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. gau
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AC
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ICH
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ON
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seud
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ASC
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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
call
y in
vest
igat
ed p
opul
atio
ns o
f F
. ps
eude
skia
, F
. sc
ario
sa,
F.
eleg
ans,
F.
burn
atii
, F
. es
kia,
F.
gaut
ieri
, F
. qu
adri
flor
a an
d F
. xp
icoe
urop
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
es
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:
Ser
ra d
o G
eres
, C
arri
s, 2
9TN
G72
Sp
ain,
Oie
eres
: 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
G45
34,
950
m
*Spa
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
*S
pain
, San
tand
er:
Her
man
dad
de C
ampo
o de
Sus
o, s
ubid
a Pi
co T
res
Mar
es,
KI3
PFER
(196
8: 8
8)
LEV
ITSK
Y &
KU
ZMIN
A (1
927:
22)
B
OR
RIL
L et
al.
(197
1: 6
) B
OR
RIL
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
e C
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
--%
"lJ
--.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
396 V. de la Fuente & E. Ortufiez
\
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 !).
Festuca sect. Eskia (Poaceae) in the Iberian Peninsula 397
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
Festuca sect. Eskia (Poaceae) in the Iberian Peninsula 401
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.
402 V. de la Fuente & E. Ortufiez
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).
Festuca sect. Eskia (Poaceae) in the Iberian Peninsula 403
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.
Iconography: (Fig. 12).
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
Festuca sect. Eskia (Poaceae) in the Iberian Peninsula 405
(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.
Iconography: (Fig. 13).
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.
Festuca sect. Eskia (Poaceae) in the Iberian Peninsula 407
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).
Iconography: (Fig. 14).
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.
408 V. de la Fuente & E. Ortufiez
) 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
Festuca sect. Eskia (Poaceae) in the Iberian Peninsula 409
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;
Festuca sect. Eskia (Poaceae) in the Iberian Peninsula 411
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.!).
Iconography: (Fig. 16).
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.
Festuca sect. Eskia (Poaceae) in the Iberian Peninsula 413
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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
Festuca sect. Eskia (Poaceae) in the Iberian Peninsula 415
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,
Festuca sect. Eskia (Poaceae) in the Iberian Peninsula 417
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.).
418 V. de la Fuente & E. Ortufiez
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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"o
¢D
t.- .=~
¢,O
Fig.
2. L
emm
a aba
xial
sur
face
oft
axa b
elon
ging
to th
e se
ctio
n E
skia
WIL
LK~ i
n th
e Ib
eria
n Pe
nins
ula
(Vid
eo g
raph
ic p
rint
er U
P-85
0, S
ON
Y):
a-
F. p
seud
eski
a (S
pain
, L
oma
del r
io S
eco,
Mul
haee
n, h
erb.
F.
& 0
.),
b -F
. sc
ario
sa (
Spai
n, C
6rdo
ba, S
ierr
a T
ifio
sa, P
rieg
o, M
A 1
8738
7), e
-F
. bu
rnat
ii (
Spai
n, C
anta
bria
, Hor
cadi
na d
e C
ovar
obre
s, M
aciz
o C
entr
al P
icos
de
Eur
opa,
her
b. F
. & 0
.),
d-
F.
eleg
ans
subs
p, e
lega
ns (
Spai
n, G
rana
da, S
ierr
a N
evad
a, b
arra
neo
de S
an Ju
an, h
erb.
F. &
O.),
e -
F.
eleg
ans
subs
p, m
erin
oi (
Spai
n, Z
amor
a, S
esnk
adez
de
T~tb
ara,
herb
. F. &
0.)
, f-
F. g
auae
ri (
Spai
n, T
erue
l, Si
erra
de
Java
lam
bre,
her
b. F
. & 0
.),
g -
F.
quad
rifl
ora
(Spa
in,
Hue
sca,
Col
d'A
fiis
elo,
Fan
l-B
iels
a, J
AC
A 3
7278
4),
h -
F.
eski
a (S
pain
, Lle
ida,
Lie
s, S
tany
s de
la
Pera
, her
b. F
. & O
.), i
- F
. xp
icoe
urop
eana
(S
pain
, C
anta
bria
, E
spin
ama,
JA
CA
290
488)
, j -
F.
xsou
liei
(Sp
ain,
Hue
sea,
Pue
rto
de B
arro
sa, B
iels
a, J
AC
A 1
2749
8).
O .<
tO
-11
t--
tD
.m
O
tO
N
Fig.
3. P
alea
aba
xial
sur
face
of t
axa
belo
ngin
g to
the
sect
ion
Esk
ia W
ILLK
. in th
e Ib
eria
n Pe
nins
ula
(Vid
eo g
raph
ic p
rint
er U
P-85
0, S
ON
Y):
a-
F. p
seud
eski
a (S
pain
, L
oma
de1
rio
Seco
, Mul
hace
n, h
erb.
F. &
0.)
, b
-F.
scar
iosa
(Spa
in, C
6rdo
ba, S
ierr
a T
ifios
a, P
rieg
o, M
A 1
8738
7),
c -F
. bu
rnat
ii (S
pain
, Can
tabr
ia, H
orca
dina
de
Cov
arob
res,
Mac
izo
Cen
tral
Pic
os d
e E
urop
a, h
erb.
F. &
O.),
d-
F. e
lega
ns su
bsp,
ele
gans
(Spa
in, G
rana
da, S
ierr
a Nev
ada,
bar
ranc
o de
San
Juan
, her
b. F
. & O
.), e
- F
. ele
gans
subs
p, m
erin
oi (S
pain
, Zam
ora,
Ses
nfin
dez
de T
~iba
ra, h
erb.
F. &
0.)
, f-
F. g
auti
eri (
Spai
n, T
erue
l, Si
erra
de J
aval
ambr
e, h
erb.
F. &
O.),
g -
F. q
uadr
iflo
ra
(Spa
in,
Hue
sca,
Col
d'A
fiis
clo,
Fan
lo-B
iels
a,
JAC
A 3
7278
4),
h -
F. e
sk/a
(Spa
in,
Lle
ida,
Lie
s, St
anys
de
la P
era,
her
b. F
. & 0
.),
i - F
. xp
icoe
uro
pean
a (S
pain
, C
anta
bria
, Esp
inam
a, J
AC
A 2
9048
8), j
-F
. xs
ouli
ei (S
pain
, Hue
sca,
Pue
rto
de B
arro
sa, B
iels
a, J
AC
A 1
2749
8).
E"
?0 O9
O
O
-'1
::7
O
"a
"O
O