INTRODUCTION
A la ge po ion o he Doñana p o ec ed a ea
(Fig. 1) is composed o we lands. In hese ecosys-
ems, aqua ic mac ophy es a e esponsible o
mos p ima y p oduc ion and also play an impo -
an ole in inc easing ecosys em s uc u es o
ecycling nu ien s and elemen s. Aqua ic mac o-
phy es a e, he e o e, key elemen s in his pa a-
digma ic na u al a ea. Mo eo e , lo a is one o
he bes na u al sou ces o in o ma ion ega ding
cu en and po en ial conse a ion in any na u al
place. The sca ce numbe o s udies on his cons-
picuous g oup o o ganisms is hus su p ising.
When he las Doñana Flo is ic Ca alogue
was published wen y- i e yea s ago (Cas o iejo
e al., 1980), i was qui e ho ough a ha ime;
howe e , he bulk o new lo is ic eco ds since
hen ha e been aqua ic plan s (as shown in his
Aqua ic mac ophy es in Doñana p o ec ed a ea (SW Spain):
An o e iew
P. Ga cía Mu illo1, R. Fe nández Zamudio1, S. Ci ujano2& A. Sousa1
1Depa amen o de Biología Vege al y Ecología. Facul ad de Fa macia. Uni e sidad de Se illa. Apdo. 874.
E-41080 Se illa. Spain. pg[email p o ec ed], [email p o ec ed] & [email p o ec ed]
2Real Ja dín Bo ánico, CSIC. Plaza de Mu illo 2. E-28014-Mad id. Spain. san os@ma- jb.csic.es
ABSTRACT
A big po ion o he Doñana p o ec ed a eas co esponds o we lands; in hem aqua ic mac ophy es a e he main p ima y p o-
duce s and play also o he impo an ecological unc ions. Ne e heless, hey a e inconspicuous o ganisms and hei impo an-
ce in hese ecosys ems does no seem o be well e lec ed in he bibliog aphy abou his na u al a ea. This pape e iews he
mos signi ican in o ma ion ga he ed abou his g oup o o ganisms in his p o ec ed a ea, p o ides an upda ed ca alogue o
his g oup o plan s, and o e s some conside a ions ela ed wi h his opic.
Key wo ds: Doñana, aqua ic mac ophy es, aqua ic ege a ion, SW Eu ope.
RESUMEN
Una g an pa e de los espacios p o egidos de Doñana co esponde a humedales, en ellos los mac ó i os acuá icos son los
p incipales p oduc o es p ima ios, ealizando además o as impo an es unciones ecológicas. Sin emba go, son o ganismos
poco conspicuos y su impo ancia en es os ecosis emas no pa ece es a e lejada en las publicaciones exis en es ela i as a
es e espacio na u al. Es e a ículo ecopila la in o mación más signi ica i a sob e es e g upo de o ganismos en es e espacio
p o egido, p opo ciona el ca álogo ac ualizado de es e g upo de ege ales y o ece algunas conside aciones ela i as al ema.
Palab as cla e: Doñana, mac ó i os acuá icos, ege ación acuá ica, SW Eu opa.
Limne ica, 25(1-2): 71-80 (2006)
The ecology o he Ibe ian inland wa e s: Homage o Ramon Ma gale
© Asociación Española de Limnología, Mad id. Spain. ISSN: 0213-8409
Figu e 1. Loca ion o Doñana p o ec ed a eas. Localización
de los espacios p o egidos de Doñana.
Limne ica 25(1-2)01 12/6/06 13:54 Página 71
pape ). This indica es how li le is known abou
aqua ic plan s in his a ea.
This is coupled wi h he ac ha some signi-
ican en i onmen al e en s ha e occu ed in
his same ime pe iod, an inc ease in in ensi e
ag icul u e in he a ea, inc eased ou ism in bo -
de ing a eas, o e exploi a ion o g ound wa e s,
in asions o exo ic species, global wa ming, e c.
All o hese issues poin o he need o upda-
e he in o ma ion ela ed o he aqua ic
mac ophy es in he Doñana p o ec ed a ea,
which is he aim o his pape .
FIRST STEP: CATALOGUING
Unlike he cases o o he places o g ea na u al
alue in he sou he n Ibe ian Peninsula, such as he
Sie a Ne ada o Sie a de G azalema, which we e
p ospec ed by bo anis s du ing he nine een h cen-
u y o be o e, he i s s udies on Doñana´s lo a
appea much la e in he mid- wen ie h cen u y. In
1945 C. Vicioso, an A agonese bo anis , published
a lis o axa collec ed in he sou h pa o he
Huel a P o ince, bu among hese e e ences he e
was no da a on mac ophy es. I was no un il 1967,
when e e ences o aqua ic mac ophy e we e
ound, ha se en aqua ic mac ophy es we e
ci ed in an in e eb a e ca alogue (Maza ano ,
1967) o Guadalqui i Ma shes (belonging o
Doñana). Cabezudo la e began he sys ema ic
s udy o lo a in his p ese ed a ea, including
18 species o aqua ic mac ophy es in his s udies
(Cabezudo, 1974; 1975; 1978, and Galiano &
Cabezudo, 1976). In 1981, some yea s la e , he
b illian wo k o Cas o iejo e al. (1980) comple-
ed he in o ma ion compiled by Cabezudo, nea ly
comple ing he lis o ascula plan s in Doñana
Na ional Pa k. In subsequen yea s his Ca alogue
has changed e y li le, wi h he excep ion o
aqua ic plan s ha ha e con ibu ed some impo -
an new eco ds o he Doñana Ca alogue:
Al henia o ien alis (Ga cía Mu illo & Tala e a,
1986), Calli iche lusi anica (Piza o, 1990),
Zannichellia ob usi olia (Tala e a e al., 1986,
Lemna isulca, and Spi odella poly hyza (Ga cía
Mu illo e al., 1991), e c. (Table. 1).
In e ms o he “o he ” plan g oups in-
cluded in aqua ic mac ophy es, i.e., Cha-
ophy a and B yophy a, da a on hese a e mo e
sca ce and inconsis en .
The i s eco ds o Cha ophy a we e no ed by
Co illion (1961), who included wo o Doñana’s
Cha ophy a species in his wo k on sou he n
Spain and No h A ica cha ophy es: Cha a con-
ni ens and Ni ella lexilis. Some yea s la e ,
Comelles (1982) and Sánchez (1984) added wo
mo e axa o he lis : Tolypella hispanica and
Cha a agi e a, espec i ely. Almos one decade
la e Ga cía Mu illo e al. (1993) added nine
new eco ds o Doñana’s cha ophy e ca alogue.
Finally, he mos ecen eco ds a e on Cha a
ulga is a . oedophylla and Tolypella salina,
e e enced by Espina e al. (1997).
In he B yophy es g oup, he e a e wo
pape s on he Riella genus (Ci ujano e al.,
1988 and 1992); Riccia lui ans and Riccio-
ca pos na ans we e men ioned by Ri as
Ma ínez e al. (1980) and Sphagnum inunda-
um by Ga cía Mu illo e al. (1995).
Table 1 shows he comple e and cu en ca a-
logue o Doñana’s subme ged mac ophy es. I
includes 74 axa (21 mo e han hose ela ed in
1993 by Ga cía Mu illo e al.) o which 46 a e
Spe ma ophy a (62 %), 3 P e idophy a (4 %), 6
B yophy a (8 %) and 19 Chlo ophy a (26 %).
Besides, his able poin s o he i s lo is ic
eco d o each axon.
72 Ga cía Mu illo e al.
Table 1. Ca alogue o aqua ic mac ophy es o Doñana p o ec ed a eas. Ca álogo de los mac ó i os acuá icos de los espacios p o egidos de Doñana.
TAXA* FIRST RECORD
CHLOROPHYTA
Cha aceae Cha a aspe a De h. ex Willd. a . aspe a Ga cía Mu illo, Be nués & Mon es, 1993
Cha aceae Cha a canescens Des . & Lois. Ga cía Mu illo, Be nués & Mon es, 1993
Cha aceae Cha a conni ens Salmz. ex A. B aun Co illion, 1961
Limne ica 25(1-2)01 12/6/06 13:54 Página 72
Aqua ic mac ophy es in Doñana 73
Table 1. Con inued. Con inuación.
CHLOROPHYTA
Cha aceae Cha a agi e a Du ieu Sánchez, 1984
Cha aceae Cha a agilis Des . Fe nández Zamudio e al. (2006)
Cha aceae Cha a galioides DC. Ga cía Mu illo, Be nués & Mon es, 1993
Cha aceae Cha a hispida L. Van Vie ssen e al., 1982
Cha aceae Cha a ulga is L. a . ulga is Ga cía Mu illo, Be nués & Mon es, 1993
Cha aceae Cha a ulga is L. a . con a ia Fe nández Zamudio e al. (2006)
(A. B aun ex Kü z.) J. A. Moo e
Cha aceae Cha a ulga is L. a . oedophylla Espina e al., 1997
(Feldman) R. D. Wood
Cha aceae Cha a ulga is L. a . longib ac ea a (Kü z.) Fe nández Zamudio e al. (2006)
J. G o es & Bullock-Webs e
Cha aceae Lamp o hamnium papulosum (Wall .) J. G o es Ga cía Mu illo, Be nués & Mon es, 1993
Cha aceae Ni ella lexilis (L.) C. Aga dh Co illion, 1961
Cha aceae Ni ella hyalina (DC.) C. Aga dh Ga cía Mu illo, Be nués & Mon es, 1993
Cha aceae Ni ella enuisissima (Des .) Kü z. Ga cía Mu illo, Be nués & Mon es, 1993
Cha aceae Ni ella anslucens (Pe s.) C. Aga dh Ga cía Mu illo, Be nués & Mon es, 1993
Cha aceae Tollypella glome a a (Des .) Leonh. Ga cía Mu illo, Be nués & Mon es, 1993
Cha aceae Tollypella hispanica No ds . ex T.F. Allen Comelles, 1982
Cha aceae Tollypella salina Co illion Espina e al., 1997
BRYOPHYTA
Ricciaceae Riccia lui ans L. Ri as-Ma ínez e al., 1980
Ricciaceae Riccioca pos na ans L. Ri as-Ma ínez e al., 1980
Riellaceae Riella cossoniana T abu Ci ujano e al., 1992a
Riellaceae Riella helicophylla (Bo y & Mon .) Mon . Ci ujano e al., 1988
Riellaceae Riella no a isii (Mon .) Mon . Ci ujano e al., 1992a
Sphagnaceae Sphagnum inunda um Ga cía Mu illo e al., 1995
PTERIDOPHYTA
Azollaceae Azolla iliculides Lam. Cobo e al., 2003
Isoe aceae Isoe es ela um A. B aun subsp. ela um Galiano & Cabezudo, 1976
Ma sileaceae Ma silea s igosa Willd. Fe nández Zamudio e al. (2006)
SPERMATOPHYTA
Apiaceae Apium inunda um L. Allie & B esse , 1975
Apiaceae Oenan he is ulosa **L. Cabezudo, 1975
Apiaceae Tho ella e icilla inunda a** (Tho e) B iq. Cabezudo, 1974
Apiaceae Oenan he globulosa** L. Cabezudo, 1975
Ca um e icilla um** (L.) Koch Galiano & Cabezudo, 1976
Apiaceae E yngium co nicula um** L. Cabezudo, 1974
Apiaceae E yngium galiodes** Lam Cabezudo, 1978
Calli ichaceae Calli iche unca a Guss. subsp. occiden alis Cas o iejo e al., 1980
(Rouy) Scho sman
Calli ichaceae Calli iche lusi anica Scho sman Piza o, 1990
Calli ichaceae Calli iche s agnalis Scop. Ga cía Mu illo, Be nués & Mon es, 1993
Calli ichaceae Calli iche b u ia Pe agna Cas o iejo e al., 1980
Calli ichaceae Calli iche ob usangula Le Gall Fe nández Zamudio e al. (2006)
Ce a ophyllaceae Ce a ophyllum deme sum L. Maza ano , 1967
Cype aceae Sci pus lui ans L. Ri as-Ma ínez e al., 1980
Limne ica 25(1-2)01 12/6/06 13:54 Página 73
SECOND STEP:AQUATIC MACROPHYTES
AND ENVIRONMENTAL FACTORS
A he end o he 1970s, González Be náldez
di ec ed a se ies o s udies on he ela ionship be -
ween he plan s in Donaña and he en i onmen
(see Ga cía No o, 1997). In his con ex , he e
was p ac ically no men ion o aqua ic plan s, wi h
jus one s udy ound on he ma sh’s ege a ion
(Allie & B esse , 1977). In 1980, Ri as Ma ínez
e al. published an excellen wo k on he ege a-
ion in he Doñana Na ional Pa k, in which hey
ca ied ou a de ailed phy osociological analysis
o he di e en communi ies o plan s in his p o-
ec ed na u al a ea. Ne e heless, despi e he
supe io quali y o he esea ch done, he in o ma-
74 Ga cía Mu illo e al.
Table 1. Con inued. Con inuación.
SPERMATOPHYTA
Ela inaceae Ela ine alsinas um L. Maza ano , 1967
Ela inaceae Ela ine mac opoda Guss. Maza ano , 1967
Ela inaceae Ela ine hexand a (Lapie e) DC. Cabezudo, 1975
Haloga aceae My iophyllum al e ni lo um DC. Maza ano , 1967
Haloga aceae My iophyllum spica um L. Van Vie ssen e al., 1982
Hyd ocha i aceae Hyd ocha is mo sus- anae L. Cabezudo, 1978
Juncaceae Juncus he e ophyllus Du ou Galiano & Cabezudo, 1976
Lemnaceae Lemna gibba L. Galiano & Cabezudo, 1976
Lemnaceae Lemna isulca L. Ga cía Mu illo e al., 1991
Lemnaceae Lemna mino L. Maza ano , 1967
Lemnaceae Spi odella poly hiza (L.) Schleiden Ga cía Mu illo e al., 1991
Lemnaceae Wol ia a hiza (L.) Ho kel ex Wimme Ga cía Mu illo, 2000
Len ibula iaceae U icula ia exole a R. B . Cas o iejo e al., 1980
Len ibula iaceae U icula ia aus alis R. B . Cabezudo, 1975
Nymphaeaceae Nupha lu eum L. Cas o iejo e al., 1980
Nymphaeaceae Nymphaea alba L. Cas o iejo e al., 1980
Polygonaceae Polygonum amphibium L. Cas o iejo e al., 1980
Po amoge onaceae Po amoge on na ans L. Galiano & Cabezudo, 1976
Po amoge onaceae Po amoge on polygoni olius Pou e Cas o iejo e al., 1980
Po amoge onaceae Po amoge on lucens L. Maza ano , 1967
Po amoge onaceae Po amoge on ichoides Cha m. & Schlech . Cabezudo, 1978
Po amoge onaceae Po amoge on c ispus L. Maza ano , 1967
Po amoge onaceae Po amoge on pec ina us L. Maza ano , 1967
Ranunculaceae Ranunculus ipa i us DC. Cabezudo, 1978
Ranunculaceae Ranunculus pel a us subsp baudo ii Allie & B esse , 1975
(God on) Meikle ex C. D. K. Cook
Ranunculaceae Ranunculus pel a us subsp saniculi olius Piza o, 1993
(Vi .) C. D. K. Cook
Ranunculaceae Ranunculus pel a us Sch ank subsp ucoides Ci ujano e al., 1992b
(F eyn) Muñoz Ga mendia
Ruppiaceae Ruppia ma i ima L. a . ma i ima Cabezudo, 1978
Ruppiaceae Ruppia d epanensis Tineo Cas o iejo e al., 1980
Zanichelliaceae Al henia o ien alis (Tz ele ) Ga cía Mu illo & Tala e a, 1986
Ga cía Mu illo & Tala e a
Zanichelliaceae Zannichelllia ob usi olia Tala e a, Tala e a e al., 1986
Ga cía & Smi h
Zos e aceae Zos e a nol ii Ho nem Cas o iejo e al., 1980
* Some axa e e eed o Doñana P o ec ed A eas ha e been ela ed wi h inco ec iden i ica ions (as Calli iche palus is L.,
Hippu is ulga is L., Zannichellia palus is L. o Zannichellia pel a a Be ol.) in o he cases hey co espond wi h synonyms
(as Ranunculus baudo ii God on; Ruppia ma i ima subsp. d epanensis L.(Tin.) Mai e & Weille o U icula ia gibba L.).
** The ju enile o m o hese species show mo phological, ana omical and physiological cha ac e s co esponding wi h aqua-
ic mac ophy es.
Limne ica 25(1-2)01 12/6/06 13:54 Página 74
ion on aqua ic plan communi ies was insu i-
cien (as can be deduced om he cho ological
changes a e 1980, included in Table 1).
In he 90s, in esponse o esea ches by
Mon es on Donaña’s aqua ic ecosys ems, new
da a appea ed on he ecology o aqua ic plan s in
he a ea: da a was published on hei biomass
(Dua e e al. 1990); he main ac o s ( looding
ime and salini y) con olling he dis ibu ion o
hese plan s in Doñana’s we lands we e iden i-
ied (Be nués, 1990; Dua e e al., 1990); and a
s udy was done on he ma sh’s seed bank (G illas
e al., 1993). San ama ía (San ama ía, 1995;
San ama ía & Hoo smans, 1998; San ama ía e
al., 1995; 1996), unde he di ec ion o Mon es
and wi h samples collec ed om Doñana, also
ca ied ou a se ies o s udies on he au oecology
o Ruppia d epanensis Tineo, one o he mos
abundan unde wa e mac ophy es in he Doñana
sal ma sh. In his same pe iod, Se ano & Toja
(1995), wo king in sand lagoons o Doñana,
ela ed he p esence o some aqua ic mac ophy-
es wi h o he ecological pa ame e s.
And inally, mo e ecen wo ks by Espina
(2004) and Espina e al. (2002) ha e con i-
bu ed aluable in o ma ion on he sal ma sh
aqua ic mac ophy es in ela ion o hei en i on-
men and o communi ies o helophy es.
S udies on Seed Dispe sal
A he u n o he cen u y, and as a esul o he
mul idisciplina y app oach aken by he Doñana
Biological S a ion o en i onmen al p ocesses
and wi h he in ol emen o San ama ía, a
se ies o s udies appea ed on he ole o bi ds in
he passi e anspo o o ganisms. To be su e, a
la ge numbe o hese s udies ocused on
he dispe sal o aqua ic mac ophy e seeds by
aqua ic bi ds (Cha alambidou e al., 2003;
G een e al., 2002; Figue ola & G een, 2002
and 2004; Figue ola e al., 2002; 2003 and
2005). These esea ches ha e been consolida ed
as a line o wo k, which is cu en ly being
ca ied ou in he Doñana Biological S a ion
unde he di ec ion o G een, wi h ou s anding
esul s. Likewise, Espina e al. (2004) ha e
ecen ly published s udies in his a ea.
Aqua ic Plan s and Clima e Change
As s a ed in he in oduc ion o his pape , a si e’s
lo a is one o he bes na u al sou ces o in o -
ma ion on ha a ea. This ac is e en mo e pe -
cep ible in aqua ic plan s since hei eac ion o
en i onmen al changes (due o hei pa icula
physiology) is much as e and p ecise. Based
on his p emise, ecen s udies ca ied ou in he
eas e n pa o Doñana Na ional Pa k and in he
pa k’s lagoons ha e shown how use ul diach onic
s udies on he p esence and dis ibu ion o aqua-
ic plan s can be in de ec ing clima ic changes
wi hin ela i ely ecen ime ames. The wo k o
Sousa (2004) and Sousa & Ga cía Mu illo (1998;
1999; 2003 and 2005) illus a e his ac and ind
an explana ion o he p ocesses o a idiza ion
and desicca ion o he coas al we lands o Huel a
P o ince by linking hese p ocesses o he end o
he Li le Ice Age, using –among o he hings–
he p esence o ce ain species o aqua ic plans,
he dis ibu ion o ege a ion in he we lands,
and hei changes o e ime.
THIRD STEP: REGARDING
CONSERVATION
The uniqueness o he di e se species o aqua ic
mac ophy es ound in sou h-wes e n Eu ope has
been poin ed ou by some au ho s (Cook, 1983;
Ga cía Mu illo, 2003; Mon es & Ma ino, 1987);
he majo i y o hese axa a e loca ed in he
Doñana a ea. The wo k o Ci ujano e al. (1992b)
is no ewo hy he e in i s anking o Spanish
we lands based on he p esence o ce ain species
o mac ophy es; he Doñana sal ma sh was an-
ked second among all we lands conside ed.1
In addi ion, in he la e 1990s, he Andalusian
Regional En i onmen al Agency (la e , he
En i onmen al Council) s a ed a line o esea ch
aimed a iden i ying he biology o he plan spe-
cies mos a isk om a conse a ion s andpoin .
Aqua ic mac ophy es in Doñana 75
1In he wo k ci ed, he au ho s did no conside he Doñana
lagoons o he changes in lo a, which occu ed a e he wo k’s
publica ion, which clea ly would ha e signi ican ly inc eased
he anking o his a ea.
Limne ica 25(1-2)01 12/6/06 13:54 Página 75

The mos salien ou comes o hese s udies we e
he “Andalusian Plan Species Red Lis (Dec e o
104/1994; Ley 8/2003)” and wo olumes compi-
ling he mos signi ican in o ma ion on he spe-
cies selec ed (Blanca e al., 1999-2000). In con-
as o p e ious Spanish “Red Lis s” (ICONA,
1987), his one included a signi ican numbe o
aqua ic species, as well as Al henia o ien alis,
Hyd ocha is mo sus- anae, U icula ia exole a,
Wol ia a hiza, Ma silea s igosa, and Tho ella
e icilla ainunda a (Ga cía Mu illo, 2000;
Sil es e, 2000), ound in he Doñana a ea.
This was ollowed, as p oposed by he Minis y
o En i onmen , by nume ous s udies in ended o
upda e he “Red Lis ” (ICONA, 1987) na ionally.
These s udies we e compiled in he AFA P ojec
(Baña es e al., 2003), including he ed lis and
he mos ele an da a om he esea ch done on
he di e en species, al hough i was no possible
o s udy some o he species lis ed. The AFA
ed lis includes wo aqua ic mac ophy e spe-
cies ound in Doñana (U icula ia exole a and
Hyd ocha is mo sus- anae).
The su eys and esea ch ca ied ou o he
Red Lis s gene a ed nume ous a icles which
highligh ed he s a e o some species o aqua ic
mac ophy es. Included among such a icles
we e hose o Ci ujano e al. (1998) and Ga cía
Mu illo e al. (2000) on species ound in he
Doñana p o ec ed a ea.
Exo ic O ganisms
The ea ly 1980s b ough he de ec ion o he i s
exo ic o ganisms in Doñana (Ga cía Mu illo e
al., 2004b). The i s o such in ade s was he
Ame ican c aw ish (P ocamba us cla ckii) and
i s spec acula p oli e a ion. The abili y o
P. cla ckii o physically ans o m i s en i onmen
and al e he a ailabili y o esou ces o o he
species in he aqua ic ecosys ems in which i was
in oduced, deeply conce ned en i onmen alis s
and scien is s. I s e ec upon mac ophy e com-
muni ies was emendous, gi en ha hey a e
i s p incipal ood sou ce. A e he ini ial pe iod
o c aw ish expansion, nume ous Doñana
mac ophy e communi ies we e simply dwindling,
wi h some species wiped ou due o he ac i i y o
his animal (B a o e al., 1993; Dua e e al.,
1990; Ga cía Mu illo e al., 1993). The nume ous
s udies on P. cla kii, unde aken by he UAM
(Uni e sidad Au ónoma o Mad id) esea ch
eam headed by Mon es, ha e emphasised he ac
ha i is now a key elemen in mos o he aqua ic
ecosys ems in Doñana and a signi ican con ol
ac o when i comes o aqua ic mac ophy e
popula ions in his p o ec ed a ea (B a o e al.,
1993; Gu ié ez-Yu i a e al., 1998).
Likewise, he Azolla iliculoides species –a
loa ing p e idophy e na i e o he New Wo ld–
began o appea in he Doñana ma sh in he
ea ly pa o he 21s cen u y (Cobo e al., 2003
and Ga cía Mu illo e al., 2004a). In jus a cou-
ple o yea s, i s p esence has ex ended o e
nea ly he en i e ma sh, o ming ca pe s some i-
mes eaching 10cm hick, which can be clea ly
seen om he RBD (Doñana’s Biological
Rese e) plane used o bi d su eys. These ca -
pe s p e en he sun’s ays om eaching he
wa e below, he eby making i impossible o
subme ged mac ophy es (nea ly all p esen in
his a ea) o de elop. They also inc ease
eu ophyza ion since hey can ix ni ogen, and
hei espi a o y ac i i y consumes he oxygen
in he wa e below (Ga cía Mu illo e al. 2004a).
Jus as wi h he Ame ican ed c aw ish, he
changes in he aqua ic ecosys ems o Doñana
a ibu able o Azolla iliculoides may be d ama-
ic. The a emp s o con ol i ha e, o da e, been
u ile (Ga cía Mu illo e al., 2004b).
Finally, in Decembe o 2004, he opical
neophy e Pis ia s a io es was ound in some
i iga ion canals loca ed in he Doñana Pa k in
he a ea o Sanlúca de Ba ameda co e ing
3Km o canals (Ga cía Mu illo e al., 2005a).
Thanks o he quick in e en ion o he
Andalusian Regional Council on he En i on-
men – aced wi h he isk posed by his new
in asion o he Doñana p o ec ed a eas– and he
low empe a u es in Janua y 2005, he Pis ia
ca pe s we e elimina ed (Ga cía Mu illo e al.,
2005b). Ne e heless, he isk con inues o exis
since some o he Pis ia plan s sampled had lo-
we s and seeds, and i is well-known ha he
seeds o his species can emain unc ional o
long pe iods o ime bu ied unde he wa e .
76 Ga cía Mu illo e al.
Limne ica 25(1-2)01 12/6/06 13:54 Página 76
CONCLUSIONS
In he ime span since he publica ion o he las
lo is ic ca alogue on he Doñana a ea, nume-
ous e e ences whose p ima y o seconda y
objec i e is he s udy o Doñana’s aqua ic
mac ophy es can be ci ed, al hough hey a e ew
in compa ison o exis ing in o ma ion on o he
in he same a ea.
Wi h ega d o he ca alogue on aqua ic
mac ophy es, we belie e i is comple ed excep
o he addi ion o new exo ic plan s whose
e ec s encompass his na u al a ea (Cobo e al.,
2003), and he wi hd awal o o he s –ci ed by
20 h-cen u y esea che s– which ha e disappea-
ed due o he de e io a ion in wa e quali y and
he p essu e o he en i onmen su ounding
his p o ec ed na u al a ea.
The egional adminis a ion –and o a lesse
ex en he na ional one– has aken his si ua ion
in o accoun , by including some o he aqua ic
mac ophy es in i s ed lis o species in dange
o ex inc ion. Ne e heless, he numbe o axa
which should be included in he lis is g ea e
(see Cobo e al., 2002).
Mo eo e , he impo ance o his a ea o he
conse a ion o aqua ic mac ophy es is e iden
since i con ains nume ous species o aqua ic
mac ophy es, many o which a e limi ed-a ea
species (“endemic species”), ela i ely uncom-
mon in aqua ic plan s.
Along he same lines, a con lic has a isen in
he a ea’s lo a conse a ion-managemen , ully
a ec ing he g oup o aqua ic plan s: many o
he aqua ic mac ophy e species ound in Doñana
co e wide a eas o dis ibu ion (in heo y);
howe e (in eali y) hese a eas a e la gely ag-
men ed, wi h simila ly agmen ed popula ions.
These species should be included in he ed
lis s, since hei ulne abili y is qui e high, a
ac which has been igno ed by en i onmen a-
lis s when i comes o he ( heo e ical) dis ibu-
ion a eas o hese species.
As s udies a e conce ned on aqua ic
mac ophy es and how hey ela e o en i onmen-
al ac o s, while he e a e a numbe o quali y
s udies on his subjec , mo e wo k on he basic
aspec s o his ela ionship would be o alue
(e.g., how mac ophy es ela e o nu ien s, ac-
o s de e mining mac ophy e dis ibu ion, s u-
dies on succession, e c.). Finally, a en ion mus
be called o he de e io a ion o Doñana’s wa e s,
he e ec s o which ope a e on wo le els:
1. By causing he disappea ance o so-called
“di icul en i onmen ” specialis plan s ha
had aken e uge in his na u al se ing (such
as plan s om bogs o oligo ophic we lands)
2. By acili a ing he in asion o exo ic species,
some o which ha e g ea po en ial o habi a
modi ica ion and i s consequences.
A ques ion emains: Is he e s ill ime o us
o comp ehend he ull complexi y o he na i e
aqua ic sys ems o Doñana o a e he ans o -
ma ions de ec ed in ecen yea s he beginning
o an i e e sible p ocess ha will p o oundly
change his place?
ACKNOWLEDGMENTS
These s udies we e suppo ed by he P ojec
Doñana 2005 om he Spain En i on-
men al Minis y.
BIBLIOGRAPHY
ALLIER, C. e V. BRESSET. 1975. La ege a ion des
milieux sales de la Rese e Biologique de Doñana
(Bas Guadalqui i , Espagne). Colloques phy oso-
ciologiques, 4: 257-269.
ALLIER, C. e V. BRESSET. 1977. E ude phy o oso-
ciologique de la Ma isma e de sa bo du e (Rese e
Biologique de Doñana). In: Doñana. P ospección e
in en a io de ecosis emas. F. Ga cía No o, J.
Me ino, L. Ramí ez Díaz, M. Ródenas, F. Sancho
Royo, A. To es, F. González Be náldez, F. Díaz
Pineda, C. Allie , V. B esse & A. Lacos e (eds.):
59-110. ICONA. Monog a ía nº 18. Minis e io de
Ag icul u a. Mad id.
BAÑARES, A., G. BLANCA, J. GÜEMES, J.C.
MORENO SANZ & S. ORTIZ. 2003. A las y lib o
ojo de la lo a ascula amenazada de España.
Minis e io de Medio Ambien e. Mad id. 1027 pp.
BERNUÉS, M. 1990. Limnología de los sis emas
acuá icos supe iciales del Pa que Nacional de
Aqua ic mac ophy es in Doñana 77
Limne ica 25(1-2)01 12/6/06 13:54 Página 77
Doñana. Tésis Doc o al. Uni e sidad Au ónoma
de Mad id. 242 pp.
BLANCA, G., B. CABEZUDO, J. E. HERNÁNDEZ-
BERMEJO, C. M. HERRERA, J. MUÑOZ & B.
VALDÉS (eds.) 1999-2000. Lib o ojo de la Flo a
sil es e amenazada de Andalucia.Tomo I: Espe-
cies en pelig o de ex inción. Tomo II: Especies ul-
ne ables. Conseje ía de Medio Ambien e. Se illa.
Tomo I, 302 pp; omo II, 375 pp.
BRAVO, M. A., C. DUARTE & C. MONTES. 1994.
En i onmen al ac o s con olling he li e his o y o
P ocamba us cla kii (Decapoda, Camba idae) in a
empo a y ma sh o he Doñana Na ional Pa k (SW.
Spain). Ve h. In . Ve ein. Limnol., 25:2450-2453.
CABEZUDO, B. 1974. No a co ológica sob e la
lo a de Huel a. Lagascalia, 4(2): 281-284.
CABEZUDO, B. 1975. No a co ológica sob e la
lo a de Huel a II. Lagascalia, 5(1): 77-83.
CABEZUDO, B. 1978. Plan as de la Rese a
Biológica de Doñana II. Lagascalia, 8(2): 166-182.
CASTROVIEJO, S., E. VALDÉS-BERMEJO, S.
RIVAS-MARTÍNEZ & M. COSTA. 1980.
No edades lo ís icas de Doñana. Anales Ja dín
Bo ánico de Mad id, 36: 203-244.
CHARALAMBIDOU I. C., L. SANTAMARÍA & O.
LANGEVOORD. 2003. E ec o inges ion by
i e duck dispe se s on he e en ion ime, e ie-
al and ge mina ion o Ruppia ma i ima seeds.
Func ional Ecology, 17: 747-753.
CIRUJANO, S., C. MONTES, P. MARTINO, S.
ENRIQUEZ & P. GARCÍA MURILLO. 1988.
Con ibución al es udio del géne o Riella Mon .
(Sphae oca pales, Riellaceae) en España.
Limne ica 4: 41-50.
CIRUJANO, S., C. FRAILE & P. GARCÍA MURI-
LLO. 1992a. No as sob e el géne o Riella Mon .
Anales Ja dín Bo ánico de Mad id, 50(1): 113-115.
CIRUJANO, S. M. VELAYOS, F. CASTILLA & M.
GIL. 1992b. C i e ios bo ánicos pa a la alo a-
ción de las lagunas y humedales españoles
(Península Ibé ica e Islas Balea es). ICONA.
Mad id. 456 pp.
CIRUJANO, S., L. MEDINA, P. GARCÍA MURILLO
& J. L. ESPINAR. 1998. Riccioca pos na ans (L.)
Co da (Ricciaceae) en la Península Ibé ica. Anales
Ja dín Bo ánico de Mad id 56: 366-368.
COBO, M. D., SÁNCHEZ GULLÓN, E. & GARCÍA
MURILLO, P. 2002. Flo a y Vege ación. In: Pa que
Nacional de Doñana. V. Ga cía Canseco .: 108-
174. Canseco Edi o es SL. Tala e a de la Reina.
COBO, M. D., E. SÁNCHEZ GULLÓN & P.
GARCÍA MURILLO. 2003. Da os ace ca de la
p esencia y ges ión de especies in aso as y xenó-
i as en un espacio p o egido eu opeo pa adigmá-
ico. In: Con ibuciones al conocimien o de las
especies in aso as en España. L. Capde illa
A güelles, B. Zille i & N. Pé ez Hidalgo, N.
(eds.): 38-41. GEI. León.
COMELLES, M. 1982. No es locali a i e isió de la
dis ibució de les espéces de ca ó i s a Espanya.
Tesis de Licencia u a. Uni e sidad Cen al de
Ba celona. 132 pp.
COOK, C. D. K. 1983. Aqua ic plan s endemic o
Eu ope and he Medi e anean. Bo anische
Jah büche Sys ., 103: 539-582
CORILLION, R. 1961. Les ege a ions p écoces des
Cha ophycées d´Espagne Me idionale e du
Ma oc Occiden al. Re ue Géné ale de Bo anique,
68: 317-330.
DECRETO 104/1994, de 10 de Mayo (BOJA de 14
de Julio): Ca álogo Andaluz de Especies de Flo a
Sil es e amenazada: 1-9.
DUARTE, C., C. MONTES, S. AGUSTI, P. MARTI-
NO, M. BERNUES & J. KALFF. 1990. Biomasa
de mac o i os acua icos en la ma isma del Pa que
Nacional de Donana (SO. España): impo ancia
and ac o es ambien ales que con olan su dis i-
bucion. Limne ica, 6: 1-12.
ESPINAR, J. L. 2000. Dis ibución espacial y em-
po al de las comunidades de mac ó i os acuá icos
de la Ma isma Salada del Pa que Nacional de
Doñana. Tesis de Licencia u a. Uni e sidad de
Se illa. 126 pp.
ESPINAR, J. L. 2004 Ecología de las comunidades
de g andes heló i os de la ma isma del Pa que
Nacional de Doñana. Tesis Doc o al. Uni e sidad
de Se illa . 214 pp.
ESPINAR, J. L., S. CIRUJANO y P. GARCÍA
MURILLO. 1997. Con ibución al conocimien o
de los ca ó i os del Pa que Nacional de Doñana.
Ac a Bo ánica Malaci ana, 22: 209-211.
ESPINAR J. L., L. V. GARCÍA,.P. GARCÍA MURI-
LLO & J. TOJA. 2002. Subme ged mac ophy e
zona ion in a Medi e anean sal ma sh: a acili a-
ion e ec om es abilished helophy es? Jou nal
o Vege a ion Science, 13: 1- 15.
ESPINAR, J. L., L. V. GARCÍA, J. FIGUEROLA, A.
J. GREEN & L. CLEMENTE. 2004. Helophy e
ge mina ion in a Medi e anean coas al ma sh:
Gu -passage by ducks changes seed esponse
o salini y. Jou nal o Vege a ion Science, 15:
315-322.
FERNÁNDEZ ZAMUDIO, R., S. CIRUJANO, I.
NIETO, A. SOUSA & P. GARCÍA MURILLO.
78 Ga cía Mu illo e al.
Limne ica 25(1-2)01 12/6/06 13:54 Página 78
2006. No edades lo ís icas en el Pa que Nacional
de Doñana. Ac a Bo ánica Malaci ana.
FIGUEROLA, J. & A. J. GREEN. 2002. How e-
quen is ex e nal anspo o seeds and in e e-
b a e eggs by wa e bi ds? A s udy in Doñana, SW
Spain. A chi ü Hyd obiologie, 155: 557-565
FIGUEROLA, J. & A. J. GREEN. 2004. E ec s o
seed inges ion and he bi o y by wa e owl on see-
dling es ablishmen : a ield expe imen wi h wige-
ong ass Ruppia ma i ima in Doñana, sou h-wes
Spain. Plan Ecology, 173: 33-38.
FIGUEROLA, J., A. J. GREEN & L. SANTAMA-
RIA. 2002. Compa a i e dispe sal e ec i eness o
wigeong ass seeds by wa e owl win e ing in
sou h-wes Spain: quan i a i e and quali a i e
aspec s. Jou nal o Ecology, 90: 989-1001
FIGUEROLA J., A. J. GREEN & L. SANTAMA-
RIA. 2003. Passi e in e nal anspo o aqua ic
o ganisms by wa e owl in Doñana, sou h-wes
Spain. Global Ecology and Biogeog aphy, 12:
427-436.
FIGUEROLA, J., L. SANTAMARIA, A. J. GREEN,
I. LUQUE, R. ALVAREZ & I. CHARALAMBI-
DOU. 2005. Endozoocho ous dispe sal o aqua ic
plan s: does seed gu passage a ec plan pe o -
mance? Ame ican Jou nal o Bo any, 92: 696-699.
GALIANO, E. F. & B. CABEZUDO. 1976. Plan as
de la Rese a Biológica de Doñana (Huel a).
Lasgascalia, 6: 117-176.
GARCÍA MURILLO, P. 2000. Al henia o ien alis;
Hyd ocha is mo sus- anae; U icula ia exole a y
Wol ia a hiza. In: Lib o Rojo de la Flo a
Sil es e Amenazada de Andalucía. TomoII. G.
Blanca, B. Cabezudo, J. E. He nández-Be mejo,
C. M. He e a, J. Muñoz & B. Valdés (eds.): 31-
34; 173-177; 361-363 & 373-375. Conseje ía de
Medio Ambien e. Jun a de Andalucía. Se illa.
GARCÍA MURILLO, P. 2003. Mac ó i os acuá icos
en los humedales andaluces. Medio Ambien e, 42:
38-41
GARCÍA MURILLO, P. & S. TALAVERA. 1986. el
Géne o Al henia Pe i . Lagascalia, 14: 102-114.
GARCÍA MURILLO, P. S. CIRUJANO & M. BER-
NUES. 1991. Lemna isulca L. y Spi odella
poly hiza (L.) Scheiden, nue as pa a el su de la
Península Ibé ica. Anales Ja dín Bo ánico de
Mad id, 48: 268-270.
GARCÍA MURILLO, P., M. BERNÚES, & C.
MONTES, 1993. Los mac o i os acuá icos del
Pa que Nacional de Doñana (SW España).
Aspec os lo ís icos. Ac as VI Cong eso Español
de Limnología, 261-267.
GARCÍA MURILLO, P., A. SOUSA & E. FUERTES.
1995. Sphagnum inumda um Russ., nue o pa a
Andalucía. Anales Ja dín Bo ánico de Mad id, 53:
245-245.
GARCÍA MURILLO, P., S. CIRUJANO, L. MEDI-
NA, & A. SOUSA. 2000 ¿Se ex ingui á
Hyd oca is mo sus anae L. de la Península
Ibé ica? Po ugaliae Ac a Biológica, 19: 149-158.
GARCÍA MURILLO, P., M. D. COBO, E.
SÁNCHEZ GULLÓN & H. GARRIDO. 2004a.
Una plan a acuá ica ame icana in ade Doñana.
Que cus, 218: 46- 47.
GARCÍA MURILLO, P., M. D. COBO, E. SÁN-
CHEZ GULLÓN & H. GARRIDO. 2004b.
Plan as exó icas e in aso as en Doñana. Medio
Ambien e, 46: 45-53.
GARCÍA MURILLO, P., E. DANA SÁNCHEZ, &
C. RODRÍGUEZ. 2005a. Pis ia s a io es L.
(A aceae) una plan a acuá ica exó ica en las p oxi-
midades del Pa que Nacional de Doñana (SW
España). Ac a Bo ánica Malaci ana, 30: 235-236.
GARCÍA MURILLO, P., E. DANA SÁNCHEZ y C.
RODRÍGUEZ HIRALDO. 2005b. La lechuga de
agua amenaza con in adi Doñana. Que cus, 232:
36-37.
GARCÍA NOVO, F. 1997. The ecosys ems o
Doñana Nacional Pa k. In: The ecology and con-
se a ion o Eu opean dunes. F. Ga cía No o, R.
M. N. Ca w o d & M.C. Díaz Ba adas (eds.): 77-
116. Uni e sidad de Se illa. Se illa.
GREEN A. J., J. FIGUEROLA, & M. I. SÁNCHEZ
2002. Implica ions o wa e bi d ecology o he
dispe sal o aqua ic o ganisms. Ac a Oecologica,
23: 177-189
GRILLAS, P., P. GARCÍA MURILLO, O. GEERTZ-
HANSEN, N. MARBÁ, C. MONTES, C. M.
DUARTE, L. TAM HAM, A. GROSSMAN. 2000.
Subme ged mac ophy e seed bank in a
Medi e anean empo a y ma sh: abundance and
ela ionship wi h es ablished ege a ion.
Oecologia, 94: 1-6.
GUTIÉRREZ-YURRITA, P. J., G. SANCHO, M. A.
BRAVO-UTRERA, A. BALTANÁS & C. MON-
TES. 1998. Die o he ed swamp c ay ish P ocam-
ba us cla kii in na u al ecosys ems o he Doñana
Na ional Pa k empo a y esh-wa e ma sh (Spain)
Jou nal o C us acean Biology, 18(1): 120-127.
ICONA. 1987. Lib o ojo de las especies ege ales
amenazadas de España peninsula e islas
Balea es. ICONA-MAPA. Mad id. 676 pp.
LEY 8/2003, de 28 de Oc ub e (BOJA de 12 de
No iemb e): Ley de la lo a y auna sil es es.
Aqua ic mac ophy es in Doñana 79
Limne ica 25(1-2)01 12/6/06 13:54 Página 79