O sis,
8:
121-126 (1993)
Response o Medi e anean g assland species o
changing ain all.
A
eply o Figue oa and Da y
R.
Fe nández
Alés
Depa amen o de Biologia Vege al y Ecologia. Uni e sidad de Se illa. Apdo. 1095. 41080 Se illa
Key wo ds:
Medi e anean annuals, empo al changes, dehesa.
Resumen.
Respues a de especies p a enses medi e áneas a cambios en la p ecipi aci6n. Una épli-
ca a Figue oa
y
Da y.
Se han e isado 10s esul ados publicados po Figue oa
&
Da y en el
Jou nal
o Ecology
( ol 79: 925-941, 1991) sob e la elación en e las a iaciones en ecuencia de las espe-
'cies de pas os medi e áneos del W de España
y
la p ecipi ación, debido a que sus esul ados con a-
dicen 10s de o os au o es que ambién han es udiado es as elaciones en o os pas os medi e áneos.
La causa de es a con adicción es iba en que 10s da os de p ecipi ación u ilizados po Figue oa
&
Da y con ienen muchos e o es. Cuando se u ilizan 10s da os co ec os 10s esul ados son conco -
dan es con 10s de 10s o os au o es.
Abs ac .
The esul s o a pape o Figue oa
&
Da y in he
Jou nal o Ecology
( ol 79: 925-941,
1991) on he ela ionships be ween changes in species equency and ain all in a Medi e anean g ass-
land o W Spain ha e been e ised, because hey do no ag ee wi h he esul s o o he au ho s ha
ha e s udied his ela ionships in o he Medi e anean g asslands. The lack o ag eemen is he esul
o he inaccu acies con ained in he ain all da a used by Figue oa
&
Da y. When he co ec ain all
da a a e used, he esul s ag ee wi h hose o he o he au ho s.
In oduc ion
Recen ly, M.E. Figue oa and A.J. Da y ha e published a pape in he
Jou nal
o
Ecology
(1991) on he ela ionships be ween changes in species abundance and ain-
all in a Medi e anean g assland in W Spain. Using a long se ies o da a (1
1
yea s)
on species equency in a dehesa g assland in Alia (Cáce es), and ain all da a om
he me eo ological s a ion o Guadalupe (Cáce es), he au ho s show ha win e
ain all and he ain all o e he whole g owing season a e good p edic o s o he
abundance o a conside able numbe o species (25%), be e han au umn and sp ing
ain all. These esul s do no i wi h o he s udies on Medi e anean g asslands o
Cali o nia (Duncan
&
Woodmansee, 1975; Pi
&
Heady, 1978), Aus alia (Rossi e ,
1966), Is ael (Na eh, 1982) and Spain (O ega
&
Fe nández Alés, 1987). All hese
au ho s ind be e ela ionships be ween species abundance and seasonal ain all
(specially wi h au umn and sp ing ain all) han wi h annual ain all.
Hobbs
&
Mooney (1985), Ma añón (1985) and Fe nández Alés (1991) ha e
shown ha , in Medi e anean annual g asslands, only a small pa o he seed bank
eme ge as seedlings, and almos all o hem eme ge du ing he i s mon h a e
he i s au umn ains, wi h many o hem dying in his ime o he yea . Mo ali y
is e y low du ing win e and sp ing, and almos all su i o s die a he end o he
sp ing, a e p oducing seeds. This means ha he bo anical composi ion and spe-
cies abundance in he g assland a e de e mined du ing he ge mina ion and seed-
ling eme gence pe iod, emaining cons an du ing he es o he g owing season
(Heady, 1958; O ega
&
Fe nández Alés, 1987). Thus, he dis ibu ion and abun-
dance o ain all du ing his c i ica1 pe iod (au umn) will be a pa amoun ac o
in de e mining he abundance o he species in he g assland (Ba olome, 1979;
O ega
&
Fe nández Alés, 1987). Since au umn ain all is no co ela ed wi h win-
e o o al ain all (Figue oa
&
Da y, 1991), he se o p ocesses desc ibed can-
no explain he ela ionships ound by Figue oa
&
Da y.
The objec i e o his s udy is o e ise he da a o Figue oa
&
Da y, in o de
o in es iga e he causes o he lack o ag eemen be ween he esul s o hese
au ho s and he esul s o he o he s udies ci ed abo e.
Me hods and esul s
The i s ma e in es iga ed is he e ec o da a ans o ma ion on he esul s o
he co ela ion be ween plan equency and ain all, since he co ela ion coe i-
cien is no in a ian o loga i hmic ans o ma ions. In o de o es o he e -
ec o he ans o ma ion used, he non ans o med equency da a o he mos
abundan species ( he only plan da a a ailable om Figu e
2
o Figue oa
&
Da y,
1991) ha e been co ela ed wi h he o al ain all o e he whole g owing season
(Figu e
1
in Figue oa
&
Da y, 1991), and he esul s compa ed wi h he ones pu-
blished by Figue oa
&
Da y using log- ans o med plan equency da a ( ables
1,
2,
3,
4 and
5
in Figue oa
&
Da y, 1991). The esul s (Table 1) show ha he
numbe o species showing signi ican co ela ion inc eases om 7 o 10 when
using non ans o med da a. Thus, i can be concluded ha da a ans o ma ions
a e no he esponsible o he disc epancies de ec ed.
The second ma e in es iga ed is he exis ence o e o s in he da a. The e-
ision o he ain all da a used by Figue oa
&
Da y has shown ha he e a e some
inaccu acies in hem. Thus, he da a on annual p ecipi a ion in Guadalupe me e-
o ological s a ion used in igu es 1 and
3
by Figue oa
&
Da y (Figu e la), do no
i wi h he da a p o ided by he Spanish Me eo ological Ins i u e o Guadalupe
s a ion (Figu e lb), no wi h he da a om Alia s a ion ( igu e lc,), ha lies close
o he s udy a ea.
In iew o such disc epancies, he non ans o med plan equency da a has
been co ela ed wi h he ain all da a om Alia Me eo ological s a ion. This s a-
ion has been selec ed because i has no nissing da a, as i happens wi h Guadalupe
s a ion (Figu e 1). P oduc -momen co ela ion coe icien s (n=ll) we e calcula-
REPLY TO FIGUEROA
&
DAVY
1
O00
-
500
E
E
-
-
s
.-
2
o
Yea s
1000
-
500
E
E
-
-
-
s
.-
2
o
Yea s
1
O00
-
500
E
E
- -
s
.-
2
o
Yea s
Figu e 1. To al ain all du ing he g owing season (Oc obe -May) a wo me eo olo-
gical s a ions close o he s udy a ea o Figue oa
&
Da y, o he yea s in which plan
equency was eco ded. Sepa a e seasonal componen s:
(
.
)
Au umn (Oc obe -
No e nbe ),
(
O
)
win e (Decembe -Febma y),
(
.
)
Sp ing (Ma ch-May). A. Da a
om Figue oa
&
Da y (1991) o Guadalupe s a ion.
B.
Da a p o ided by he Spanish
Me e ological Ins i u e o Guadalupe S a ion. This s a ion has missing da a (mon hs in
which ain all was no eco ded) in 1983,1984, 1985 and 1986. The nu nbe o non hs
wi h missing da a each yea a e ma ked a he op o he ba , in b acke s.
C.
Da a p o i-
ded by he Spanish Me eo ological ins i u e o Al a S a ion, close o he s udy a ea
han
Guadalupe and wi h no nissing da a.
ed be ween he o al equency o each species and o al ain all du ing he whole
g owing season and o e au u nn (Oc obe , No embe ), win e (Decembe , Janua y,
Feb ua y) and sp ing (Ma ch, Ap il, May).
The esul s (Table
2)
show ha , when conside ing he ain all da a o Alia ins-
ead o he ones p o ided by Figue oa
&
Da y, he numbe o species exhibi ing
signi ican co ela ion wi h ain all o e he whole g owing season alls om
10
o
6.
The mos s iking di e ences a e ound when compa ing he co ela ions
wi h he seasonal ain all (Table
2).
Thus, while Figue oa
&
Da y ind ha he
la ges numbe o species a e signi ican ly co ela ed wi h win e ain all, he co-
ela ions wi h Alia ain all da a show ha much mo e species a e ela ed wi h he
au umn p ecipi a ion
(7)
han wi h he win e (1) and sp ing
(0)
one. Thus, i can
be concluded ha he g asslands s udied by Figue oa
&
Da y beha e as he o he
Medi e anean g asslands, and ha he di e ences ound by hese au ho s we e
ela ed wi h he inaccu acies in he ain all da a.
Table
1.
Co ela ions o ans o med and non ans o med plan equency wi h he
accumula ed ain all o e he g owing season (Oc obe
-
May). Da a o ans o med plan
equency and ain all om Figue oa
&
Da y
(1991).
Coe icien s wi h an as e isk a e
signi ican o
p<
0.05.
Species T ans o med No ans o med
Vulpia b omoides
0.78"
O
81"
An hoxan hum a is a um
0.59 0.62"
Ai a ca iophyllea
0.63" 0.68"
Poa bulbosa
0.58
O
61"
Chae opogon ascicula us
0.23 O. 16
O ni hopus comp essus
0.90" 0.92"
T i olium glome a um
0.78" 0.81"
T i olium campes e
0.70" 0.70"
T i olium a ense
0.61
*
0.62"
T i olium gemellum
0.46 0.5
1
Tolpis ba ba a
0.72" 0.83"
Leon odon a axacoides
0.16 0.28
Hipochae is glab a
-0.20 -0.08
Chamae nelum nix um
0.10 -0.04
Log ia gallica
0.41 0.42
Tube a ia gu a a
0.52 0.69"
Plan ago bella di
0.57 0.58
Eupho bia exigua
-0.05 0.28
REPLY TO HGUEROA
&
DAVY
125
Conclusions
The lack o ag eemen be ween he esul s o Figue oa
&
Da y and he esul s o
o he s udies made in o he Medi e anean g assland is ela ed wi h he inaccu a-
cies in he ain all da a used by he o me au ho s. When he co ec ain all da a
a e used, his lack o ag eemen disappea .
In
any
case, i should be no ed ha he numbe o species co ela ed wi h ain-
all o e he whole g owing season is almos as high as when conside ing only au-
umn ains. Since many o he s udies ci ed abo e ha e es ablished he ela ion-
ships be ween species abundance and ain all g ouping he species in o amilies,
o ha e used sho se ies o da a, i would be e y in e es ing ha hese au ho s
econside hei esul s wi h he p ope ain all da a, in o de o check he ela i-
e impo ance o o al and au umn ain all in de e mining species abundance.
Table
2.
Co ela ions o non ans o med plan equency wi h accumula ed ain all o
Alia me eo ological s a ion. G owing season, Oc obe -May; Au umn, Oc obe -No embe ;
Win e , Decembe -Feb ua y; Sp ing, Ma ch-May. Coe icien s wi h an as e isk a e sig-
ni ican o
p<
0.05.
Accumula ed ain all
G owing
Species season Au umn Win e Sp ing
Vulpia b omoides
0.82" 0.64" 0.59 -0.08
An hoxan hum a is a um
0.63" 0.62* 0.34 0.11
Ai a ca iophyllea
0.7
1
*
0.21 0.68" -0.13
Poa bulbosa
0.48 0.66" 0.23 -0.16
Chae opogon ascicula us
0.26 -0.13 0.35 -0.10
O ni hopus comp essus
0.63* 0.55 0.35 0.22
T i olium glome a um
0.50 0.72" 0.14 0.15
T i olium campes e
0.81* 0.60" 0.50 0.25
T i olium a ense
0.78" 0.75" 0.4 1 0.20
T i olium gemellum
0.50 0.56 0.17 0.35
Tolpis ba ba a
0.35 0.5 1 0.14 -0.07
Leon odon a axacoides
0.34 -0.10 0.39 0.07
Hipochae is glab a
-0.11 -0.39 0.1 1 -0.19
Chamaemelum mix um
0.10 -0.16 0.09 0.37
Log ia gallica
0.07 0.61" 0.26 0.17
Tube a ia gu a a
0.29 -0.05 0.33 0.05
Plan ago bella di
0.58 0.05 0.52 0.30
Eupho bia exigua
0.47 0.12 0.50 -0.22

Re e ences
Ba olome, J.W. 1979. Ge nina ion and seedling eme gence in Cali o nia annual g ass-
land. J. Ecol 67: 273-28 1.
Duncan, D.A.
&
Woodmansee, R.C. 1975. Fo ecas ing o age yield om p ecipi a ion
in Cali o nia's annual angeland. J. Range Manag. 28(4): 327-329.
Fe nández Alés, R. 1991. Popula ion dyna nics in annual species o Medi e anean g ass-
lands. IV In e na ional Rangeland Cong ess. Mon pellie , 22-26 o Ap il o 1991.
Figue oa, M.E.
&
Da y, (1991). Response o Medi e anean g assland species o chan-
ging ain all. J. Ecol. 79: 925-941.
Heady, H.F. 1958. Vege a ional changes in he Cali o nia annual ype. Ecology 39: 402-
416.
Hobbs, R.
&
Mooney, H.A. 1985. Communi y and popula ion dynamics o se pen ine
g assland in ela ion o gophe dis u bance. Oecologia 67: 342-35 1.
Ma añón, T. 1985. Rese a de semillas en el suelo en una dehesa de Sie a Mo ena: e-
lación con la ege ación. Anal. Eda . y Ag obiol XLIV: 1805-1816.
Na eh,
Z.
1982. The dependence o he p oduc i i y o an semia id Medi e anean hil1
pas u e ecosys em on clima ic luc ua ions. Ag icul u e and En i onmen 7: 47-61.
O ega, F.
&
Fe nández Alés, R. 1987. T ends in lo is ic change in ime in Medi e anean
annual g asslands o Sou h Wes e n Spain. in Di Cas i, F.; Flo e , Ch.; Rambal,
S.
&
Roy,
J.
(eds.). Time scales and Wa e S ess. IUBS. Pa is. pp. 451-456.
Pi , M.D.
&
Heady, H.F. (1978). Responses o annual ege a ion o empe a u e and
ain all pa e ns in No he n Cali o nia. Ecology 59: 336-350.
Rossi e , R.C. (1966). Ecology o Medi e anean annual- ype pas u e. Ad . Ag on.
18: 1-56.
Manusc ip ecei ed
on
Decembe
1992