Aquiatic macrophytes in Doñana protected area (SW Spain).

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. 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