Bocconea 7 - 1997 427
Abies pinsapo Boiss.: a p o ec ed species in a p o ec ed a ea
Mon se a A is a, Ja ie He e a & Sal ado Tala e a
In oduc ion
Abs ac
A is a; M., He e a,
J.
& Tala e a, S.: Abies pinsapo Boiss.: a p o ee ed speeies
in
a p o ee ed
a ea. -Boeconea 7: 427-436. -lSSN 1120-4060.
This pape p esen s he
e
sul s
o
a s udy on he ep oduc i e eyele
o
Abies pinsapo. The main
ep odue i e aspec s which a ee he quan i y and quali y
o
h
e seed
s,
such as he
lo
w pollen
dispe sal capaei y, he subdioecious condi ion and
h
e combina ion g ow
h
o n- ee
a chi ec u e a e
di
seussed. The equi emen s
(l
seed ge mina ion, seedling
e
e ui men and
some aspec s
o
h
e species a e analysed using
ali
he a a
il
able da a.
Abies pinsapo Boiss. is one
o
he 46 species
o
i s which a e discon inuously
dis ibu ed
in
he No he n hemisphe e (Fa jon 1990). Abies pinsapo was desc ibed o he
i s ime by Clemen e
in
he i s decade
o
he 19 h cen u y. Howe e , his desc ip ion
was
lo
s and he species
wa
s la e desc ibed by Edmund Boi
ss
ie
in
1938 (Abies pinsapo
Boiss., Biblio h. Uhi . Génè e, sé .
2,13:
402,406. 1839).
Abies pinsapo
is
a coni e endemie o he sou h
o
Spain a
nd
belongs o a ci cum-
Medi e anean g oup
o
i s which
is
conside ed
o
be a elic species. The wes e n
Medi e anean species a e sepa a ed in o wo sec ions: sec ion Abies
is
ep esen ed by A.
alba Mille (S. Eu ope) a
nd
A. neb odensis (Lojac.) Ma ei (Sicily), and sec ion Piceas e
(Fig. I) by A. pinsapo Boiss. (S. Spain), A. ma ocana T ab. (N.W. Mo occo), A.
azao ana C6za ex Hug. Villa (N.W. Mo occo), and A. numidica Ca . (N.W. Alge ia).
The spec
ie
s
in
sec ion Piceas e a e mo phologically simila , ha ing igid Iea es, and
b ac s smalle han he o uli e ous scale
s.
Due o such simila
i i
es, many au ho s ha e
g ouped hese axa in o h ee spec
ie
s:
A. pinsapo, A.
a
zao ana and A. numidica (including
A. ma ocana as subspecies
o
A.
pinsapo; Mai e 1952; F anco 1950)
o
in o wo specie
s:
A. pinsapo and A. numidica (Ceballos & Bolanos 192
8;
Fa jon 1990), including A.
ma ocana and
A.
a
zao ana as a ie ies
o
A.
pinsapo (Fa jon 1990).
428 A is a & al.: Abies pinsapo Boiss.: a p o ee ed speeies ...
x
A.
pinsapo
Ào.
A.
numidica
•
A.
ma ocana •
A.
neb odensis
o
A.
azao ana
~
A.
alba
Fig. I. Dis ibu ion a ea
o
he sec ion Abies acco ding o Mai e
(1952)
and
Jalas &
Suominen
(1973).
E en hough he speei ie limi s
o
he genus
Abi
es a e no clea , he pinsapo popula ions
o
sou he n Spain ha e some di e en eha ae e s om hose
o
N.
A ica. Fo example,
A.
pinsapo has ema gina e, mue ona e b ae s (subula e
in
A.
ma o
ca
na and
A.
azao ana).
The b ae s
o
A.
pinsapo a e 1
/3
-1
/4 sho e han he o uli e ous seales (e. 1/2 sho e
in
A.
ma ocana a
nd
A.
azao ana;
pe so
obse . Fig. 2). O he eha ae e s such as numbe
o
le
a
s oma a o numbe a
nd
di
sposi ion
o
esin canals a y be ween ees and hey eanno be
used o dis ing
ui
sh hese axa (Ma inez & Aisa 1995). Due o sueh mo phologieal
di e enees,
we
eonside A. pinsapo Boiss.
is
an
endemie species
o
he dolomi ic and
se pen ine moun ains
o
sou he n Spain. The in o ma ion he e p esen ed come om
h
e
Doc o al hesis 'Biologia del pinsapo (Abies pinsapo Boiss.)', Uni e si y
o
Se ille, Spain,
1993.
Dis ibu ion
o
Abies
pinsapo
Cu en ly, Abies pinsapo occu s a 900-1600 m
in
he Sub-Be ie moun ains, occupying
a o al
o
only 1200 ha. The mos impo an o es s a e
in
Sie a de G azalema
(Cidiz
P o ince), Sie a de las Nie es and Reales de Sie a Be meja (Malaga p o ince).
Howe e , isola ed indi iduals o sma
ll
popula ions
o
se e al indi iduals can be ound a
Ronda, Pa au a, Is {m, Monda, Ojén, Tolox, Yunque a, Es epona, Casa abonela and Co es
Bocconea 7 - 1997
429
de la F on e a (Malaga), and a Algodonales, Zaha a de la Sie a, G azalema and
Benacoaz (Cadiz).
A TAZAOTANA A MAROCANA A PINSAPO
I
l
l
2
l
3
Fig.
2.
Scales anù B ac s
o
Abi
es pinsapo (Spain: Càùiz, Sie a ùe G azalcma. 1200
m.
12 Oc
1992,
M.
A is a,
S.
Tala e a &
J.
He e a 139392, SEV), Abies ma ocana (Mo occo: Jbel
Talamsen ane, 1800 m. 8 Feb 1993,
F.
Ojeda
/39393.
SEV) and Abies azao ana (Mo occo: Jbel
Tassao , 1700
m,
IO
Feb 1993,
F.
Oj
eda J
39394
, SEV).
l,
en al side
o
he o uli e ous scales; 2,
do sal
si
de
o
he o uli e ous scales;
3,
do sal side
o
he b ac s. I and 2 ba scales 10mm. 3 ba
scale 8 mm.
Abo e I 100 m
o
ele a ion Ahies pinsapo o ms pu e o es s, bu below I 100 m i
o ms mixed communi ies. In he ee layc
o
dolomi ic soils Ahies pinsapo o ms a mixed
oak- i o es wi h species such as Que cus o undi olia and Que cus aginea, hei
p opo ions a ying ' wi h al i ude (Fig. 3). The sh ub laye , much mo e di e se,
is
composed
o
species such as Daphne lau eola subsp. la i olia, Hede a helix, Ruscus
aculea us, I is oe idissima and Ruhia pe eg ina
in
he closed o es and
o
C a aegus
monogyna subsp. h e ispina, P unus spinosa, P ilo ichum spinosum, Bupleu um
spinosum, Thymus x a undanus,
Vlex
hae icus, E inacea an hyllis, Bupleu um
gih al a icum and Cis us alhidus in he. gaps (A is a 1995). In he ee laye
o
se pen ine
soils and only occasionally on calca eous soils, Ahies pinsapo occu s wi h Que cus suhe
and wi h o he coni e s such as Pinus pinas e and Pinus halepensis. The sh ub laye
in
430
A is a & al.: Abies pinsapo Boiss.: a p o ec ed species ...
se pen ine soils is
composed
o
E ica e minalis, Genis a lanuginosa,
G.
iacan hos,
Cis us populi olius subsp. majo , Bunium alpinum subsp. macuca, Ce as ium
gib al a icum and Alyssum se pylli olium subsp. malaci anum.
San C is òbal
1400
1300
~~
1:
1200
Z
'~'
.
o
~
HO
~
w
'i
..
~B
.....
w 1000
900
8 6
o:
N-
Llanos del Re és
Fig. 3. P opo ion
o
seedlings (do ed) and ep oduc i e indi iduals (black)
01'
Abies
pinsapo
pe
plo
o
50
m2 along he al i udinal g adien
o
Sie a de G azalema
(Cidiz).
Rep oduc i e
cycIe
o
Abies pinsapo Boiss.
Abies pinsapo belongs o a e y s able genus (2n = 24), and has a ep oduc i e cycle
o
one yea (A is a & Tala e a 1994a) e y simila o hose desc ibed in o he species
o
genus Abies (Miyake 1903, Hu chinson 1915, Eis 1970, Powell 1970, Owens &
Molde
1977, Singh &
Owens
1981, 1982).
The
di e en ia ion
o
he ep oduc i e buds occu s in
la e July; b ac s
o
mic ospo ophylls a e ini ia ed h ough Augus and in o Sep embe , and
he di e en ia ion is comple ed a he end
o
he summe . Male buds a e ini ia ed
on
he
abaxial su ace
o
he elonga ing shoo s
o
he lowe b anches, and a he end
o
he
summe , hese buds
en e
a es ing phase. The do mancy is b oken a he end
o
Feb ua y
and meiosis akes pIace. Ma u e pollen g ains a e shed in he i e-celled s age be ween he
end
o
Ap iI and mid-May. The emale
buds
a e also ini ia ed
in
mid-July and a e si ua ed
on he uppe b anches, on
h
e adaxial su ace
o
he elonga ing shoo . In mid-Ap il bud
bu s akes pIace, and hen meiosis occu s coinciding wi h pollina ion.
Du ing pollen shedding, he scales
o
he emale cones sepa a e o p o ide a space
h ough which ai bo ne pollen g ains en e . The e o e, seed cone ecep i i y is
synch onised wi h pollen shedding wi hin a ee. In an indi iduaI ee lowe ing las ed o
six days (A is a & Tala e a 1994b), bu in he popula ion he lowe ing pe iod las ed one
mon h. In ecen yea s, lowe ing in ensi y and p oduc ion
o
cones
we e simila in he
h ee mai n o es s
o
Abies pinsapo.
The
mean pollen cone p oduc ion pe ee anged
om
800
o
39
200
and he emale p oduc ion anged om 62 cones
pe
ee o 198.
Bocconea 7 - 1997
431
Pollen e ili y in Abies pinsapo
is
e y high, nea ly 100
%,
and
in
i o, he pollen
needs only dis illed wa e and a empe a u e om
20
o 24°C o ge mina ion. Pollen
dispe sal capaci y in Abies pinsapo was s udied in Sie a de G azalema a di e en s and
densi ies. Ai pollen densi y (measu ed using pollen aps)
o
pinsapo in open o es was
ma kedly lowe han ha in closed o es (A is a & Tala e a 1994b). This e lec s a poo
suspension
o
pinsapo pollen in he ai , and in consequence a sho -dis ance pollen
dispe sal. Al hough he pollen g ains ha e wo wings, hei la ge size and he high
humidi y
o
he a ea could be impo an ac o s in educing suspension.
Al hough Abies pinsapo is a monoecious species,
in
ali he popula ions a p edominance
o
emale plan s ( ees ha de elop only emale cones) was obse ed, wi h a ew male and
monoecious indi iduaI s, hus indica ing a unc ionally subdioecious b eeding sys em
(A is a & Tala e a 1996b). Va ia ion
in
gende exp ession is obse ed be ween di e en
densi ies, and could be explained as a consequence
o
ee a chi ec u e ( emale and male
cone posi ion)
in
combina ion wi h he g ow h
o mo
A low densi y, ees a e sho -b oad
wi h b anches om g ound le el, and end o exp ess hemsel es sexually by p oducing
male cones (Fig. 4). In con as , ees g owing
a
high densi y a e ali wi h b anches limi ed
o he op hal
o
he ee, and end o emaleness. A simila si ua ion has been ound
in
o he coni e s such as Picea (Ca on & al. 1990) o Pinus (Smi h & al. 1988).
A e pollina ion, he o uli e ous scales enla ge and o e g ow he b ac s, and he
megaspo e begins ee nuclea di isions. A he end
o
lune
a chegonium o ma ion occu s
·and e ilisa ion akes pIace ( wo mon hs a e pollina ion). The emale cones achie e hei
maximum size
in
ea ly lune, and a he beginning
o
Sep embe , he emb yo
is
appa en ly
ully de eloped. Seeds begin o shed om Oc obe o No embe . Seeds eaching he so
il
en e do mancy un il Feb ua y
o
he ollowing yea , when ge mina ion begins. Seeds ha
ail o ge mina e his yea die, so ha a pe sis en seed bank
is
no o med.
Abies pinsapo indi iduals lowe
in
al e na e yea s and occasionally
in
wo consecu i e
yea s. In he las ou een yea s, A. pinsapo popula ions lowe ed
in
1982, 1983, 1985,
1987, 1989, 1991, 1992 and 1994 (A is a & Ta1a e a 1995). Cone c op cycles
o
wo o
h ee yea s a e common
in
o he i species (Singh & Owens 1981, Shea 1989a, 1989b,
Houle & Paye e 1991). Synch onisa ion
o
he ees
in
lowe ing
is
p obably due o
an
°
endogenous hy hm
o
he species con olled by in ins
ic
ac o s bu ha could be
modi ied by ex insic condi ions (Powell 1977).
Cone p oduc ion and ui
~e
om 1990 o 1993
in
Abies pinsapo o es s
o
Sie a de
G azalema, Sie a de las Nie es and Sie a Be meja we e e y di e en . The ui se was
always highe among pinsapo ees
o
he Sie a de las Nie es and Sie a Be meja ela i e
o Sie a de G azalema ees. This si ua ion e lec s he clima e suppo ed by he eas e n-
mos popula ions
o
A. pinsapo. In hese popula ions ain all du ing he lowe ing pe i od
is
hal ha
o
he Sie a de G azalema. This could in ol e a lowe pollina ion index
in
Sie a
de G azalema ees, and hus a lowe ui se (A is a & Tala e a 1995). Va ia ion
in
ui
se
was also ound be ween di e en densi ies (A is a & Tala e
a
1 996a). Low densi y A.
pinsapo popula ions had ma kedly highe cone c op p oduc ion and ui -se pe plan han
hose a high densi ies. Densi y e en a ec ed he cone size and he numbe
o
seeds pe
cone: cones
om he
high densi y zone we e signi ican ly la ge and had a highe numbe
o
seeds han hose om low densi y zone.
In he ully de eloped cone, all he o ules each he ull size al hough some
o
hem
ha e no emb yo.

432 A is a & al.: Abies pinsapo Boiss.: a p o ec ed species
...
I
~~
Il
I
~---_.
-
HIGH DENSITY
lOW
DENSITY
Fig. 4. Mo phology
o
Abi
es
pinsapo ees li ing in high and low densi y.
The
blaek zone indiea es
he ema
le
eo
ne posi ion and he s iped he male
eone
posi ion.
Bocconea 7 - 1997 433
Seed
c ap
iabili y be ween densi ies is ma kedly di e en , e.g. in 1992, a e age seed
iabili y
in
he low-densi y zone was
21
%,
bu
some ees had only 3 %
o
iable seeds. In
he high densi y popula ions seed
c ap
iabili y a e aged 82 % wi h a ange
o
54
%-
93.3 % (A is a & Tala e a 1996). Di e ences in seed iabili y a e also ound in esponse
o gende exp ession
o
he ees in low and high densi y popula ions. A high densi y,
ees ha p oduced only emale cones du ing he 10we ing pe iod had g ea e seed
c ap
iabili y han hose wi h male and emale cones (a e ages
o
80 % and 63.3 %
espec i ely). In con as , low densi y zone ees wi h only emale cones a 10we ing had
lowe seed
c ap
iabili y (a e age 8.3) han hose wi h bo h male and emale cones
(a e age
31
.8). This e lec s di e ences
in
quan i y and quali y
o
he a ailable pollen
du ing he pollina ion pe iodo
Ge mina ion
o
Abies pinsapo seeds akes pIace
in
Feb ua y. In he labo a o y,
ge mina ion
o
seeds exceeding
60
mg ( hose ha ha e an emb yo, A is a & al. 1992) is
nea 100 %. In he ield, seed ge mina ion depends on he pIace in which he seeds
allo
Inside he o es , he gaps a e he bes places o ge mina ion, anging om 26.6 % o
63.3 % (A is a 1993b). Unde he o es canopy, ge mina ion is e y low, nea 0%. The
highes mo ali y
o
seedlings occu s
in
he i s yea
o
li e, a e which mo ali y
is
p ac ically ze o. Howe e , su i al depends on he pIace in he o es a which ge mina ion
occu s. In o es gaps, su i al is e y high e en
in
he i s yea , wi h only
18
% ·
o
mo ali y. In con as ,
in
he in e io o es mo ali y
in
he i s yea
o
seedling li e is
52 %. Thus, mo ali y unde he o es canopy is always highe han
in
gaps.
One-mon h-old seedlings om seeds
collec ed am
ees a low densi y ha e a g ea e
igou
ha~
hose om he high densi y zone. This si ua ion oge he wi h he di e en
seed iabili y
in
ees
o
high and low densi y popula ions, could ha e impo an
consequences.
The
lowe ed seed iabili y ound in monoecious ees s. emale ees
in
high densi y s ands sugges s a possible inb eeding dep ession
in
he endogamous emb yos
o
A. pinsapo, and consequen ly, he selec i e abo ion
o
some
o
hem. The heo y ha
he e
is
inb ecding dep ession
in
A.
pinsapo
is
suppo ed i s ly by di e ences
in
seed
c ap
iabili y and secondly by he highe igou
o
seedlings om high dcnsi y popula ions
(A is a & Tala e a 1 996a). The esul s
o
his s udy a e impo an because hey ha e
se e al implica ions o he collec ion and p aduc ion
o
A. pinsapo seeds o
ea o es a ion plans. They p a ide addi ional. e idence agains he disc edi ed p ac ice
o
collec ing
cones om
open-g own semi-isola ed indi iduals. Al hough hese ees o en
p oduce abundan , easily-collec ed cones, a signi ican p opo ion
o
seeds a e no iable,
due o high a es
o
sel - e ilisa ion. Whe e he seeds a e iable a subsequen lack
o
igou
in
he seedlings esul s om sel ing.
The
si ua ion desc ibed he e sugges s ha he
na u al egene a ion
o
low densi y popula ions
o
Abi
es pinsapo is di icul .
Conse a ion
A p esen , mos
o
he pinsapo· o es s a e included
in
he so-cali ed 'Ne wo k
o
P o ec ed A eas
in
Andalusia': Sie a de G azalema Na u al Pa k (1984), Sie a de las
Nie es Na u al Pa k (1989) and Los Reales de Sie a Be meja Na u e A ea (1989). Bo h
Sie a de G azalema and Sie a de las Nie es Na u al Pa ks ha e been decla ed as
Biosphe e Rese es by UNESCO,
in
1977 and 1995 espec i ely. A Eu opean le el, he
434 A is a & al.: Abies pinsapo Boiss.: a p o ec ed species ...
pinsapo o es s ha e been included
in
he Rabi a s Di ec i e 92/43 and he species IS
p o ec ed by Dec e o 104/94 in he 'Andalusian Ca alogue
o
Th ea ened Wild Plan s'.
In he pinsapo o es
o
Sie a de G azalema, he e seems o be no p oblem o na u al
egene a ion. Abies pinsapo does no seem o be a long-li ed species. The oldes pinsapo
o es
is
a Canada de los Cue os,
in
he Sie a de las Nie es, Miilaga p o ince. In his
o es , he oldes ees a e unde 200 yea s old. In Sie a de G azalema he e a e no ees
app oaching his age ( he oldes i was abou 123 yea s old), p obably because i
egene a ed a he beginning
o
he cen u y a e an in ense i e.
The Abies pinsapo o es
o
Sie a de G azalema has a balanced mul i-age condi ion
in
which ec ui men and mo ali y a es end o be cons an . The p opo ion
o
seedlings
in
Sie a de G aza1ema is e y high
(o
69%) and hey ha e p ac icaIly no signs
o
he bi o e
damage
(1e
ss
han 5%
o
he seedlings a e consumed by he bi o es). Size s uc u e
is
e y
di e en
in
he o he wo impo an o es s
o
A. pinsapo. In Sie a de las Nie es, he
p opo ion
o
young indi idua
ls
is e y small (only 4%), indica ing a low a e
o
ec ui men . This si ua ion e lec s i s ly, he s ong he bi o e damage caused by bo h
wild and domes ic li es ock and secondly, he s ong p uning on he lowe b anches
o
he
ees which dec eases he male eone c op p oduc ion and
in
consequence, seed iabili y
due o lack
o
pollina ion, and hi dly he ma kedly low le el
o
he e ozygosi y
o
his
popula ion (Pascual & al. 1993), which p obably a ce s seed igou . In Sie a Be meja,
diame e dis ibu ion esembled a no mal dis ibu ion, wi h ees
o
40 cm
DBR
as
h
e
mos equen . This could e lec he selec i e choice o which he ees g ea e han 60 cm
DBR
had been submi ed. In his popula ion he seedlings also su e he bi o e damage,
bu a lowe le els han
in
Sie a de las Nie es.
The e
is
a signi ican nega i e co ela ion be ween ee densi y (a measu emen
o
canopy co e ) and numbe
o
pinsapo seedlings, indica ing ha A. pinsapo es ablishmen
a es a e highe
in
he ligh gaps (A is a 1995). The small numbe
o
seedlings which
su i ed unde o es canopy su e s supp ession. Abies pinsapo
is
a mode a ely shade-
ole an species, which can su i e un il i
is
40 yea s old unde o es canopy. Du ing
hi
s
ime, i cons i u es a seedling bank, which p obably eoccupies he a ailable space when a
gap is o med.
The mai n pa hogen agen s
o
he pinsapo o es a e he coleop e an C yphalus
numidicus, he lepidop e an Dio ic ia aulloi and he ungi A milla ia meliea (Vahl. F .)
Kumm and He e obasidium annosum B es. (Cobos 1994). Li le
is
known abou he
incidence
o
such o es pes s; bu he e a e some da a on p eda ion pe cen age
o
seed
cones by Dio yc ia al/lioi (A is a & Tala e a 1995)
in
Sie a de G azalema, bu i seems
ha he highcs pa hogen impac occu s
in
he Sie a de la Nie es popula ion. This
is
being
s udied a p esen .
In
conclusion, na u al
e
gene a ion
in
he Sie a dc G azalema pinsapo o es is
gua an eed due o bo h he good seed quali y and hc absence
o
he bi o es (allowing
seedling ec ui men ). Ou side he c10sed o es ,
e
gene a ion is mo e di icul . and a
po en ial gene ic imp o emen would be possible wi h supplemen a y pollina ion
managemen as has been sugges ed
in
o he coni e o es s (F anklin
1981
, Askew 1992)
o
wi h a i icial seed ain using igo ous seeds. Regene a ion
in
Sie a de las Nie es a
nd
Sie a Be meja o es s equi es mainly seedling p o ec ion om he bi o es.
The si ua ions desc ibed ha e been ound
in
he absence
o
majo e en s such as i e.
Abies pinsapo
is
a mode a ely shade- ole an species, which could domina e whe e he e is
Bocconea 7 - 1997 435
no exogenous dis u bance, bu less ole an species (such as Q. o undi olia and Q.
aginea) a e capable
o
pe sis ing as mino componen in he canopy
o
o es cap u ing
space when majo e en s occu (Ba den 1979, 1981). In addi ion, Abies pinsapo seeds
nei he ge mina e a e i e no esp ou bu Que cus spp. do bo h. The e o e, i e, a
ecu en dis u bance
in
Medi e anean ecosys ems, could be an impo an ac o
in
de e mining species composi ion and dynamics, and is p obably he mos impo an dange
in
A. pinsapo o es s.
Re e ences
A is a,
M.
1993: Supe i encia de las pllin ulas de Abies pinsapo Bois
s.
en su habi a na u al. -
Anales Ja din Bo . Mad id
51:
193-198.
1993: Ge minaci6n de
la
s
se
millas y supe i encia de las plan ulas de Abies pinsapo Boiss.
-Ac a Bo . Malaci ana 18: 173-178.
1995 The s uc u e and dynamics
o
an
Abiespinsapo
o es in sou hem Spain. -Fo es
Ecol. Manag.
74:
81-89.
& Tala e a, S. I 994a. Phenology and ana omy
o
he ep oduc i e phase
o
Abies pinsapo
Boiss. (Pinaceae), -Bo . J. Linn. Soc. 116: 223-234.
& -I 994b: Pollen dispe sal capaci y and pollen iabili y
o
Abies pinsapo Boiss. -Sil ae
Gene ica
43:
155-
15
8.
& - 1995: P oducci6n de pi ias y ciclos de cosechas en Abies pinsapo Bois
s.
-
An
ales del
Ja din Bo . Mad id
53:
5-12.
& - 1996a: Densi y e ec on he ui -se , seed c op iabili y and seedling igou
o
Abies
pinsapo. -Annals
o
Bo . 77: 187-192.
&-
1996b: Gende exp ession
in
Abies pinsapo Boiss., a Medi e anean i . -Annals
01'
Bo . (
in
p ess).
- & He e a, J. 1992: Viabilidad y ge minaci6n de las semillas de Abies pinsapo Boiss. -
Ac a Bo . Malaci ana 17: 223-228.
Askew,
G.
R. 1992: Po en i al gene ic imp o emen due o supplemen al mass pollina ion
managemen
in
coni e se ed o cha d.
-Fo e
s Eco!. Manag. 47: 135-147.
Ba den,
L.
S. 1979:
T
ee
el"l
acemen
in
small canopy gaps
01'
a Tsuga canadiensis o es
in
he
sou hem Appalachian, Tenessee. -Oecologia 44: 141-142.
1981: Fo es de elopmen
in
canopy gaps
o
a di e se ha dwood o es
o
he sou hem
Moun ains. -Oikos
37:
205-209
Ca on, J. E., Wang,
B.
S. P. & Schooley, H. O. 1990: E ec
o
ee spacing, cone s o age and
p echilling on ge mina ion
o
Picea glauca seed. -Fo es Ch on. 66: 388-392.
Ceballos, L. F.
C.
& Bola ios,
M.
M. 1928: Flo a y mapa o es al. El pinsapo y el abe o de
Ma uecos.~
Ins i u o Nacional Expe . Ag on. Fo es a
l,
Mad id 1: I-57.
Cobos,
J.
M.
1994: Es ado i osani a io de los pinsapa es andaluces. -
In
: Ges i6n y conse aci6n
de los pinsapa es Andaluces. -Asociaci6n Fo es al Ahdaluza. Jun a de Andalucia.
Eis, S. 1970: Rep oduc ion and ep oduc i e i egula i ies
o
Abies lasioca pa and Abies g andis.
-Canadian J. Bo . 48:
141
-
143
.
Fa jon,
A.
1990: Pinaceae. -Koel z Scien i ic Books, Koenigs ein.
F anco, J. 1950: Abe os. -Anais Ins i u o Supe io Ag onomia 17.
F anklin,
E.
C.
1981: Pollen managemen handbook. -USDA Fo .
Se o
Ag ic. Handbook 587, 98
pp.
Houle,
G.
& Paye e, S. 1991: Seed dynamics ol' Abi
es
balsamea and
Ac
e saccha um
in
a
deciduous o es
01'
No heas em No h Ame ica. -Ame ican J. Bo . 78: 895-905.
Hu chinson, A.
H.
1915: Fe ilisa ion
in
Abies balsamea. -Bo . Gaz. 60: 457-472.