Limne ica, 26 (2): 1-10 (2007)
Limne ica, 26 (2): 307-318 (2007)
c
Asociaci´
on Ib´
e ica de Limnolog´
ıa, Mad id. Spain. ISSN: 0213-8409
Phy oplank on om NE Do˜
nana ma shland (“El Cang ejo G ande”,
Do˜
nana Na u al Pa k, Spain)
I. Reyes 1,∗,G.Ma
´
ın 1,M.Reina1, A. A echede a 1, L. Se ano 1, M.A. Casco 2and
J. Toja 1
1Dp o. Biolog´
ıa Vege al y Ecolog´
ıa. Facul ad de Biolog´
ıa. Uni e sidad de Se illa. Apdo.1095, 41080-Se illa.
Espa˜
na.
2Di isi´
on Ficolog´
ıa, Facul ad de Ciencias Na u ales y Museo. Uni e sidad Nacional de La Pla a. Paseo del
Bosque s/n. 1900. La Pla a. A gen ina.
2
∗Co esponding au ho : [email p o ec ed]
ABSTRACT
Phy oplank on om NE Do˜
nana ma shland (“El Cang ejo G ande”, Do˜
nana Na u al Pa k, Spain)
The s udy a ea (“El Cang ejo G ande”, Do˜
nana Na u al Pa k) is loca ed a he final sec ion o he Guadiama Ri e channel
and i is a pa o he NE Do˜
nana ma shland. The hyd ological cha ac e is ics o he a ea a e complex and dynamic due o he
di e en o igin o i s wa e inpu : he Guadiama Ri e basin, he Guadalqui i Ri e es ua y, and ice paddies ou le s. This
a ea is included in he hyd ologic egene a ion plan o he Do˜
nana ma shland (“Do˜
nana 2005”), hough he p esen s udy was
finished be o e any modifica ion ook place. Phy oplank on ichness was high since a o al o 224 phy oplank on axa we e
egis e ed. Dia oms and Clo ophy es we e he g oups ha had he highes ichness, Ni zschia palea,Cyclo ella a omus and
Mono aphidium con o um being he mos widely dis ibu ed and abundan species. Phy oplank on exhibi ed a s ong seasonal
a ia ion, in bo h composi ion and biomass, seg ega ed in wo pe iods (flood and desicca ion) as sugges ed by he esul s o
he PCA. On he o he hand, he spa ial di e ences we e less e iden acco ding o a mul idimensional scaling analysis (MDS).
This seasonal pa e n was s onge in he o dina ion o physico-chemical a iables (conduc i i y, nu ien concen a ion and
suspended ma e ). We conclude ha his sys em is an eco one zone wi h a high hyd ologic complexi y due o a a ie y o wa e
inpu s wi h a s ong seasonali y. These hyd ologic ea u es would be he de e mining ac o in phy oplank on composi ion and
high axon ichness.
Key wo ds: Phy oplank on, Do˜
nana, ma shland, empo al seg ega ion, we lands.
RESUMEN
Fi oplanc on del NE de las ma ismas de Do˜
nana (“El cang ejo G ande”, Pa que Nacional de Do˜
nana, Espa˜
na)
El Lucio de “El Cang ejo G ande” (Pa que Na u al de Do˜
nana) es ´
a si uado en el amo final del encauzamien o del
´
ıo Guadiama y o ma pa e de la ma isma NE de Do˜
nana. Las ca ac e ´
ıs icas hid ol´
ogicas de la zona son complejas
ydin
´
amicas debido a los di e en es apo es que ecibe de la cuenca del Guadiama , del es ua io del Guadalqui i y del
desag¨
ue de los a ozales. Es a zona es ´
a incluida den o del Plan de Regene aci´
on H´
ıd ica (“Do˜
nana 2005”) aunque el
p esen e abajo finaliz´
o an es de que se lle a an a cabo las ac uaciones p e is as. La iqueza del fi oplanc on ue ele ada
ya que se egis ´
o un o al de 224 axa. Las Dia omeas y las Clo ofi as ue on los g upos que eunie on mayo iqueza,
siendo Ni zschia palea, Cyclo ella a omus yMono aphidium con o um las especies m´
as ep esen a i as, en abundancia y
dis ibuci´
on. El fi oplanc on p esen ´
o una ue e a iaci´
on es acional, en composici´
on y biomasa, iden ific´
andose cla amen e
una din´
amica empo al epa ida en dos pe ´
ıodos (inundaci´
on y es iaje) como sugie en los esul ados del an´
alisis de
componen es p incipales. En cambio, las di e encias espaciales ue on menos e iden es como demos ´
oelan
´
alisis de
o denaci´
on mul idimensional no je ´
a quico (MDS). El pa ´
on es acional ue a´
un m´
as ma cado en el an´
alisis de las p incipales
a iables ´
ısico-qu´
ımicas (conduc i idad, concen aci´
on de nu ien es y ma e ia en suspensi´
on). Se concluye que es e sis ema
co esponde a una zona de eco ono de g an complejidad hid ol´
ogica debido a la exis encia de una a iedad de apo es
h´
ıd icos suje os a a iaciones es acionales; es a complejidad se ´
ıa el ac o de e minan e de la composici´
on y ele ada iqueza
del fi oplanc on.
Palab as cla e: Fi oplanc on, Do˜
nana, ma isma, seg egaci´
on empo al, humedal.
308 Reyes e al.
INTRODUCTION
Coas al ecosys ems ( idal ma sh, es ua ies, del-
as, lagoons, e c) a e na u ally e y p oduc i e
sys ems as he esul o physico-chemical p oces-
ses aking place in shallow wa e , wi h equen
and in ense winds, and high nu ien loads (Po-
me oy, 1977; Odum, 1988; Muylae & Raine,
1999). This kind o sys ems is di ficul o s udy
because hey a e e y dynamic and subjec o
a iable s ages (Com´
ın e al., 1999). Phy o-
plank on in coas al ecosys ems unde Medi e-
anean clima e is la gely influenced by hyd o-
logic ea u es, nu ien a ailabili y and seasona-
li y (Com´
ın & Valiella, 1993; Gilabe , 2001;
Quin ana & Mo eno-Amich, 2002; Nuccio e al.,
2003; O ega-Mayagoi ia e al., 2003; Hlaili e
al., 2006). The e a e ew publica ions on phy-
oplank on assemblages in his kind o sys ems
(L´
opez, 1987; Saba e & Mu˜
noz, 1990; Romo
& Mi acle, 1994; Gilabe , 2001; Puigse e e
al., 2002; Rod igo e al., 2003; Villena & Romo,
2003; L´
opez-Flo es e al., 2006). The p esen
s udy is he fi s publica ion ela ing phy oplank-
on composi ion and ichness o seasonali y in
he Do˜
nana ma shland. The s udy a ea (“El Can-
g ejo G ande”, Do˜
nana Na u al Pa k) is loca ed
on he NE Do˜
nana ma shland. The hyd ology o
he a ea is complex and dynamic due o di e-
en wa e inpu s: Guadiama Ri e basin, Gua-
dalqui i Ri e es ua y and ou le o ice paddies
(Se ano e al., 2006). Al hough mos pa o he
Do˜
nana inland ma shland has no idal influence,
he s udy a ea is influenced by A lan ic ides om
he es ua y o he Guadalqui i Ri e . Bo h he
es ua y and he eas e n Do˜
nana ma shland ha e
been impac ed by eu ophica ion and hea y me-
al pollu ion o decades (Cab e a e al., 1984,
1987; A amba i e al., 1984, 1996; Zu e a e al.,
1987; Se ano e al., 2006). In 1998, 5 hm3o
mud and acid wa e wi h high concen a ions o
hea y me als we e dumped o he Guadiama Ri-
e , om an open-cas mine in Azanalc´
olla , and
flooded an ex ension o 2600 ha downs eam. An
ex ensi e cleaning ac i i y ook place in he i-
e floodplain ha was e en ually p o ec ed as
a bu e a ea (“Paisaje P o egido-Co edo Ve de
del Guadiama ”). Addi ionally, a hyd ologic e-
gene a ion plan (“Do˜
nana 2005”) has been de e-
loped o eco e he o iginal wa e inpu in o de
o coun e balance he endency owa ds desic-
ca ion and sil ing-up o he Do˜
nana ma shland
du ing he pas 50 yea s. The hyd ology o he
s udy a ea, in pa icula , is planned o be al e-
ed by h ee di e en modifica ions, bu none o
hem ook place be o e he end o 2004. The e-
o e, he p esen s udy on phy oplank on com-
posi ion, ichness, and dynamics was ca ied ou
p io o any modifica ion.
MATERIAL AND METHODS
S udy a ea
Do˜
nana has a Medi e anean clima e wi h A lan ic
in luence, gene ally classi ied as d y sub-humid.
Rain all is qui e a iable, bo h wi hin a yea
and o e he yea s. I has a 580mm yea ly
a e age ain all, abou 80 % o which is dis ibu ed
h oughou a we pe iod om he end o Sep embe
o he beginning o Ap il. Summe s a e e y d y
and ho , while win e s a e sho and mild. Wa e
balance is gene ally de icien as ain all exceeds
e apo anspi a ion only du ing 3-4 mon hs o he
yea (Siljes ¨
om&Clemen e,1990).
Abou 23 000 ha o he Do˜
nana ma shland is
p o ec ed as ei he a Na ional Pa k o Na u al
Pa k (Fig. 1). The s udy si e co e s 250 ha o he
final s e ch o he Guadiama Ri e enc oached
by wo pa allel le ees buil in 1956 (Fig. 1). I in-
cludes na u al and man-made wa e ways, isola-
ed wa e -bodies, and a beachlike-floodplain lo-
cally named Lucio, which gi es name o he
whole si e (“Lucio, El Cang ejo G ande”). Mos
o he Guadiama Ri e wa e flow is di e ed
owa ds he Guadalqui i Ri e es ua y h ough
an a ificial canal which is pe manen ly connec-
ed o he s udy a ea by wo inle s (Fig. 1, si es
B and C), seasonally by a floodga e (Fig. 1, si e
D), and occasionally du ing floods by he ou -
flow o he Guadiama Ri e (Fig. 1, si e A). Wa-
e in he s udy a ea is gene ally empo a y and
shallow excep a he floodga e whe e wa e can
each up o 2 m du ing high- ide. Wa e empe-
a u e anges om 8◦C o30
◦C, conduc i i y (a
Phy oplank on om NE Do˜
nana ma shland 309
Wa e ways
A i icial canal
Pa k bounda ies
Ricepads
Figu e 1. Loca ion o he s udy si e (Lucio El Cang ejo G ande) a he end o he Guadiama Ri e channel. Sampling s a ions a he
inle si es a e ep esen ed by le e s (A, B, C, and D) while he inne si es a e numbe ed. Localizaci´
on del ´
a ea de es udio (Lucio El
Cang ejo G ande) en el amo final del encauzamien o del ´
ıo Guadiama . Se indican las es aciones de mues eo de las “en adas”
median e le as (A, B, C y D) y con n´
ume os los pun os in e io es.
20◦C) om 0.5 mS cm−1 o 15.7 mS cm−1,Na
+
is he dominan ca ion in he floodplains while
Ca2+domina es in some wa e ways. Wa e flo-
wing h ough he a ificial canal is usually u bid
due o a high load o ino ganic suspended ma e
la gely composed o CaCO3pa icles associa ed
o P (Reina e al., 2006). In con as , he sou he n
ma shland gene ally p esen s a be e wa e qua-
li y wi h highe anspa ency and ela i ely lowe
concen a ions o N and P (Espina e al., 2002).
Vege a ion in ele a ed a eas is domina ed by Sa -
coco nia u icosa and Ho deum ma inum, while
aqua ic eme gen mac ophy es (Ph agmi es aus-
alis,Sci pus ma i imus,Juncus subula us)and
subme ged mac ophy es g ow in flooded a eas
(Cha a galioides,Calli iche unca a,Ranuncu-
lus pel a us,Ruppia d epanensis).
Twen y-one sampling si es we e loca ed wi -
hin he s udy a ea and we e sampled ele en i-
mes om Sep embe 2002 o Sep embe 2004
(Fig. 1). Fou si es co esponded o inle s, o
which only h ee ac ed as ou le s du ing floods
o idal flows (si es B, C and D). Dissol ed oxy-
gen, pH, and elec ical conduc i i y we e measu-
ed in si u. Su ace wa e was collec ed wi h 1.5
L PET bo les a each si e o chlo ophyll ade-
e mina ion (me hanol ex ac ion ollowing Ma -
ke e al., 1980), suspended ma e (g a ime i-
cally wi h p e iously d y fil e s a 100◦C), and
nu ien concen a ions in 2-4 eplica es a e fil-
a ion h ough WHATMAN GF/C fil e s in he
labo a o y: ni i e (S ickland & Pa sons, 1972),
ni a e (Gol e man, 1991), ammoniun (Rodie ,
1981), and phospha e (Mu phy & Riley, 1962).
The concen a ion o o al P in he wa e was de-
e mined in duplica es as phospha e a e acid di-
ges ion wi h 0.5 M H2SO4and0.5gK
2S2O8a
120◦C o 3-4 h (De G oo & Gol e man, 1990).
Wa e samples o 125 ml we e p ese ed in
si u wi h lugol, and he de e mina ion o phy o-
plank on composi ion and abundance was pe o -
med in duplica es. Mos phy oplank on axa we e
iden ified o species wi h an op ical mic oscope
(magnifica ion: 400x and 1000x). Abundance o
310 Reyes e al.
Figu e 2. Phy oplank on composi ion in he s udy a ea.
Composici´
on flo ´
ıs ica del fi oplanc on en la zona de es udio.
phy oplank on cells was obse ed wi h an in-
e ed mic oscope ollowing U e m¨
ohl’s me hod.
S a is ical analyses we e pe o med wi h he so -
wa e SPSS 12.0; communi y analyses (MDS,
ANOSIM, PCA, SIMPER) wi h PRIMER 5.
An analogue o he uni a ia e ANOVA called
ANOSIM was used o es o di e ences be -
ween mul i a ia e samples om di e en si es o
om di e en seasons (flooding s. d ying). A
SIMPER es was used o iden i y he axa p i-
ma ily p o iding he disc imina ion be ween sea-
sons. Simila i y ma ices o he mul i a ia e sam-
ples we e calcula ed using he B ay-Cu is coe -
ficien a e a log- ans o ma ion [log (x+1)] o
he o iginal axa abundances in o de o pe o m
a non-me ic mul idimensional scaling (MDS).
The Shannon di e si y index (H) was calcula ed
as bi s pe cell using loge.
RESULTS
Phy oplank on composi ion included 224 axa:
80 dia oms, 71 chlo ophy es, 39 cyanobac e ia,
19 euglenophy es, 6 ch ysophy es, 5 c yp ophy-
es, and 4 dinoflagella es (Fig. 2). Phy oplank on
assemblages we e e y a iable in bo h ime and
space as only 13 axa had a equency o ap-
pea ance highe han 50 %, and jus h ee spe-
cies eached a equency highe han 90 % (Mo-
no aphidium con o um,Cyclo ella a omus and
Ni zschia palea). Fo y- wo % o axa appea ed
10
8
6
4
2
0
-2
-4
-6
-8
PC2
PC1
-8 -6 -4 -2 024
6810
2003
2004
lood desicca ion
PC1
PC2
15
10
5
0
-5
-10
-15 -10 -5 0 5 10 15
Figu e 3. PCA o dina ion o phy oplank on assemblages
a he di e en sampling si es g ouped in o flooding and
desicca ion pe iods o each hyd ologic cycle. O denaci´
on PCA
de la comunidad del fi oplanc on en las dis in as es aciones de
mues o ag upadas en pe iodos de inundaci´
on y es iaje pa a
cada ciclo hid ol´
ogico.
only du ing flooding, and 5 % o axa we e
es ic ed o d y imes, whe eas he majo i y o
axa appea ed a any o he season. The e o e a
ew axa we e widely dis ibu ed bu no clea
dominance was ound. The di e si y index (H)
anged om 0.37 o 2.93 in he whole o he
s udy a ea, and om 0.51 o 2.84 a he inle
si es. Minimum di e si y coincided wi h he
occasional bloom o Ch ysidalis sp.1 wi hin he
s udy a ea du ing June 2003 and a he inle si es
Phy oplank on om NE Do˜
nana ma shland 311
in June 2004 ha eached 93 % and 96 % o
ela i e abundance, espec i ely.
No spa ial dis ibu ion o phy oplank on was
obse ed acco ding o he MDS o dina ion. In
con as , he PCA o dina ion o phy oplank on
assemblages showed a empo al pa e n whe e
composi ion and abundance we e seasonally se-
g ega ed wi hin each cycle (Fig. 3). The PCA
analysis showed ha 23.4 % and 21.3 % o he
a ia ion we e explained by he fi s wo axes
in bo h he 2002/2003 and he 2003/2004 cy-
cles, espec i ely. Phy oplank on assemblages
du ing flooding we e p ima ily a anged along
he fi s PCA axis, while assemblages du ing
d y pe iods we e mo e spa sely dis ibu ed along
he second PCA axis. An ANOSIM es was
used o es whe he phy oplank on assembla-
ges we e significan ly seg ega ed du ing floo-
ding and d y pe iods. No clea seg ega ion was
ound o he whole s udy a ea du ing ei he
he 2002/03 o he 2003/04 cycles (R=0.48and
R= 0.61, p<0.01, espec i ely). The inle si-
es, howe e , we e significan ly seg ega ed in o
wo seasons du ing each cycle (R=0.99 and
R= 0.87, p<0.01, espec i ely). Few axa con-
ibu ed mo e han 5 % o his seg ega ion ac-
co ding o a SIMPER es , and only some o
hem accoun ed o he simila i y o phy oplank-
on assemblages wi hin ce ain pe iods: a) Ch oo-
coccus a . minu us and Synechocys is sp. o
2002/03 flooding ; b) Te adesmus a . c ocini
o 2003/04 flooding; c) Limno h ix a . planc-
onica o he 2003 d y pe iod; and d) Me is-
mopedia wa mingiana,Chlo ococcum sp. and
Ch oococcus dispe sus o he 2004 d y pe iod.
The es o he con ibu ing species appea ed
Table 1. Maximum abundance (cells/ml) o he main phy oplank on axa in he s udy a ea du ing each cycle in he flooding and
disseca ion pe iods. Abundancias m´
aximas (c´
elulas/ml) de los p incipales axa fi oplanc ´
onicos en la zona de es udio du an e cada
ciclo en la ´
epoca de inundaci´
onydees iaje.
Cicle 2002/03 Cicle 2003/04
Flood Dessica ion Flood Dessica ion
Ch oococcus a . minu us 8.1·1039.3·103103103
Oscilla o ia a . planc onica — 1.3·104——
Synechocys is sp. 3.9·103—10
26.1·104
Chlamydomonas sp. 3 1.6·1030.1·1034.1·103103
Chlamydomonas sp. 4 — 3.1·1040.5·1032.2·103
Chlo ella ulga is 2.5·1031030.3·103103
Coelas um mic opo um 3.8·1043.7·1033.9·103—
Mono aphidium ci cinale 3·1032.4·1030.2·1030.4·103
Mono aphidium con o um 1.9·1041.4·1042.4·1047.2·104
Mono aphidium sp. 5.1·1031033.6·1031.7·103
Py amimonas sp. — — 8.2·1030.3·103
Scenedesmus eco nis 2.1·1031030.2·1031.4·103
Sch oede ia sp. 2.1·1032.8·1041031.8·103
Te adesmus a . c ocini 4.8·104—8.3·104—
Cyclo ella a omus 7.6·1037.6·1032.1·1042.6·104
Ni zschia acicula is 0.8·1031.2·1040.2·1031.2·104
Ni zschia longissima 0.4·1031.7·1041040.4·103
Ni zschia palea 6.1·1032.8·1041.5·1043.2·104
Ch ysidalis sp.1 — 8.3·105—2.6·105
Gymnodinium sp. 3.6·1035.6·1030.6·1030.6·103

312 Reyes e al.
PC1
PC2
PC2
3
1
-1
-3
-5
-5 -3 -1 1 3
2003
2004
5
3
1
-1
-3
-5
-5 -3 -1 1 35
PC1
Figu e 4. PCA o dina ion o physico-chemical a iables
a he di e en sampling si es g ouped in o flooding and
desicca ion pe iods o each hyd ologic cycle. O denaci´
on PCA
de las a iables ´
ısico-qu´
ımicas en las dis in as es aciones de
mues eo ag upadas en pe ´
ıodos de inundaci´
on y es iaje pa a
ambos ciclos hid ol´
ogicos.
h oughou he s udy pe iod and only showed
changes in abundance, such as M. con o -
um, N. palea, and C. a omus.
Abundance o he main phy oplank on axa
anged widely wi hin each pe iod (Table 1).
Some axa eached maximum abundance du ing
he d y pe iods (e.g., Sch oede ia sp., N. acicu-
la is, N. palea,yLimno h ix a . planc ´
onica),
while o he s we e ei he mo e abundan du-
Figu e 5. E olu ion o e he s udy pe iod o he a e age
abundance o main phy oplank on axonomic g oups ( %)
and a e age o al abundance (cells/ml) a all he inle si es.
E oluci´
on a lo la go del pe ´
ıodo de es udio de la abundancia
media de los p incipales g upos axon´
omicos del fi oplanc on
( %) y abundancia media o al (cells/ml) en el conjun o de los
pun os de “en ada”.
ing flooding (Chlamydomonas sp. 3 y Coelas-
um mic opo um) o main ained a ela i ely high
abundance (1.4 ·104cells/ml) du ing all seasons
(Mono aphidium con o um). On some occasions
la ge abundances we e p eceded and/o ollowed
by negligible abundances (Table 1) sugges ing
ha episodes o coloniza ion ook place (e.g.,
Coelas um mic opo um,Te adesmus a . c o-
cini,Py amimonas sp.and Ch ysidalis sp. 1).
The concen a ions o o al P and suspen-
ded ma e we e gene ally high and eached up
o 974.7 μg/l and 529.8 mg/l, espec i ely, while
N-NO−
3concen a ion anged om negligible a-
lues o 7.1 mg/l. Suspended ma e was la gely
ino ganic as phy oplank on con ibu ed wi h only
30 % o he o al. Plank onic chlo ophyll a an-
ged om 2.0 μg/l (Feb ua y 2003) o 479.5 μg/l
(Sep embe 2003), and was posi i ely co ela ed
wi h cell abundance ( = 0.59, p<0.001).
Some o hese a iables we e c oss-co ela ed
wi h o he , such as chlo ophyll and conduc i i y
( = 0.53, p<0.01). Elec ical conduc i i y was
highe in summe ; June a he han Sep embe
a ained he highes alues a he inle si es
because o he dilu ion e ec o eshwa e
d aining om ice pads a he end o he
cul i a ion season (Sep embe -Oc obe ).
En i onmen al a iables showed a seasonal
seg ega ion wi hin each cycle (Fig. 4). The fi s
Phy oplank on om NE Do˜
nana ma shland 313
Table 2. Numbe o samplings, numbe o s a ions, o al samples (n), su ace o he s udy a ea (km2), and o al cumula i e ichness
o phy oplank on in se e al coas al sys ems s udied by di e en au ho s. N´
ume o de mues eos, n´
ume o de es aciones mues eadas,
o al de mues as (n), supe ficie del ´
a ea es udiada (km2) y iqueza o al acumulada del fi oplanc on en di e sas sis emas cos e os
es udiados po di e en es au o es.
Si es Sys em Au ho Numbe o
samplings
Samplings
si es n´
A ea To al ichness
o phy oplank on
Lucio Cang ejo G ande
(Do˜
nana) En es e es udio 011 30 <146 2.4 224
1 Guadiama (En emu os,
Do˜
nana)
Toja e al., sin publica 018 030<43 5 089
5 Es ua io Guadalqui i L´
opez, 1987 027 10 <270 50 259
Lagunas San a Olalla y Dulce
(R.B.Do˜
nana)
Lopez e al., 1991 026 06<132 0.6 117
Laguna de Ta elo (Do˜
nana) Se ano e al., 2004 020 020<40 0.16 055
4 Albu e a Ad a S´
anchez Cas illo, 1987 013 030<39 >0.4 058
6 Albu e a Valencia Romo & Mi acle, 1995 114 03<342 21 131
Albu e a de Mallo ca Puigse ´
e e al., 2002 020 04<105 0.7 074
2 La Sa o Rod igo e al., 2003 002320<64 13 128
3 Del a del Eb o Saba e & Mu˜
noz, 1990 012 040<48 — 127
wo axis o a PCA o dina ion g ouped 67.3 % and
68.3 % o 2002/03 and 2003/04, espec i ely.
The a iabili y accumula ed by he fi s PCA
axis was associa ed o d y condi ions, and
was co ela ed o he concen a ion o o al P,
suspended ma e , plank onic chlo ophyll a,and
conduc i i y. Flooding was associa ed o he
second PCA axis, pH and ni a e concen a ions
being posi i ely co ela ed, and ammonium and
ni i e nega i ely co ela ed o his axis.
The ela i e abundance o phy oplank on axa
(%) also ollowed a seasonal pa e n a he inle
si es (Fig.5). Chlo ophy es domina ed om he
end o win e un il he beginning o summe when
ch ysophy es became he dominan ones, hough
heywe e apidly o e u nedbydia oms ha domi-
na ed h ough au umn and win e . Small dia oms,
such as C. a omus, we e dominan un il hey we e
eplaced by chlo ophy es nea he end o win e .
Cumula i e phy oplank on ichness was signi-
fican ly co ela ed wi h he numbe o samplings
( = 0.77, p<0.05). The numbe o samplings
was cons ained by he a ailabili y o wa e a
each si e du ing he s udy. The e o e, he inle
si es, being pe manen , accumula ed he highes
ichness. Ou s udy si e (“Lucio El Cang ejo
G ande”) showed a ela i ely high cumula i e i-
chness compa ed o o he coas al we lands wi h
Medi e anean clima e (Table 2). Bo h he num-
be o samplings and he su ace a ea we e aken
in o accoun in o de o compa e ou esul s. Si-
es 1, 2 and 3 accumula ed a highe ichness wi h
a lowe numbe o samplings. The s udy si e had
a simila ichness han o he si es o simila su -
ace a ea (si e 4) bu highe ichness pe ha han
nea by si es (si es 5 and 6).
DISCUSSION
Phy oplank on o coas al we lands is gene ally
composed o cosmopoli an species able o cope
wi h wide a ia ions in salini y, u bidi y, and eu-
ophy. In a eas domina ed by a Medi e anean
clima e, his a iabili y is o en a anged in floo-
ding and d ying pe iods acco ding o wa e a ai-
labili y. The e o e, he unc ioning o his kind
o sys em is gene ally d i en by ac o s ela ed
o seasonali y and hyd ology (Ma ´
ın & Com´
ın,
1992; Com´
ın e al., 1999). The hyd ology o
he s udy si e is a he complex wi h bo h na-
u al and a ificial wa e bodies ecei ing wa e
inpu s om di e en o igins depending on he
314 Reyes e al.
season. Di e en wa e inpu s would allow o
he coloniza ion o o ganisms om di e en o i-
gins (Com´
ın e al., 1999). Fo his eason, hyd o-
logy is expec ed o play an impo an ole in he
composi ion o phy oplank on assemblages in he
s udy a ea, pa icula ly a he inle si es, whe e
la ge exchanges a e likely o occu . Simila ly,
Romo (1997) ound ha he phy oplank on com-
posi ion o inle channels d aining he ice pad-
dies was di e en om he composi ion a he
inne si es o he Albu e a o Valencia. In he
p esen s udy, he composi ion and abundance o
phy oplank on assemblages a he inle si es we e
seg ega ed seasonally, a he han spa ially, in o
a flooding and a d ying pe iod. Simila ly, he in-
fluence o empo al a he han spa ial a iabili y
was mo e ele an o he composi ion o phy o-
plank on in he Albu e a o Valencia (Romo &
Van To engen, 1995). Du ing d y pe iods, phy o-
plank on assemblages we e mo e he e ogeneous
han du ing flooding, p obably due o he com-
bina ion o idal hy hms and e apo a ion in iso-
la ed wa e bodies. This isola ion was la ge in
2002/03 when he floodga e emained closed du-
ing bo h he flooding and d y seasons, while in
he ollowing cycle i was open om he end o
June ill mid Oc obe 2004.
The main phy oplank on axa in ol ed in his
seg ega ion belonged o cyanobac e ia, dia oms,
chlo ophy es, and c ysophy es. Chlo ophy es and
dia oms ha e been epo ed o each a la ge
abundance in coas al we lands inging ice pad-
dies in he Eb o Del a (Fo ´
es & Com´
ın, 1992).
Cyanobac e ia a e common in ma shes du ing
he summe (Ma ´
ın & Com´
ın, 1992; Quin ana
& Mo eno-Amich, 2002; L´
opez- Flo es e al.,
2006). In he es ua y o he Guadalqui i Ri-
e , cyanobac e ia domina ed in sp ing-summe
wi h species belonging o he gene a: Limno h ix,
Ch oococcus and Me ismopedia (L´
opez, 1987).
In he p esen s udy, L.a . planc onica,C. dis-
pe sus and M. wa mingiana we e mo e abundan
du ing d y pe iods a he inle si es. The chlo o-
coccal chlo ophy e Te adesmus a . c ocini was,
in con as , mo e abundan du ing flooding pe-
iods. I is likely ha he in e mi en appea ance
o his axon was due o episodes o coloniza-
ion as his species has been eco ded in he i-
cini y, bo h in he es ua y o he Guadalqui i Ri-
e (L´
opez, 1987) and in he Guadiama Ri e
(Ma ´
ın & Ga c´
ıa-No o, 2006). Ano he axon li-
kely in ol ed in coloniza ion episodes was Ch y-
sidalis sp. 1, bu his axon has no been eco -
ded ea lie in nea by a eas. Ch ysophiceans a e a
significan ac ion o ma ine phy oplank on and
can also each high abundance in b ackish, Me-
di e anean we lands (So okin e al., 1996; C uz-
Piza o e al., 2003; Rod igo e al., 2001). The
in e mi en appea ance o his axon du ing June
2003 and 2004 con ibu ed o he seasonal seg e-
ga ion o phy oplank on assemblages in flooding
and d ying pe iods. The concen a ion e ec o
e apo a ion could explain i s high abundance du-
ing summe , which was 4 imes highe han he
mos abundan axon eco ded in he es ua y o
he Guadalqui i Ri e (L´
opez, 1987). In ac ,
phy oplank on abundance and chlo ophyll con-
cen a ion we e always highe du ing he d y pe-
iods a all si es, while dilu ion e ec s domina-
ed du ing flooding pe iods. This seasonal pa -
e n explains he c oss-co ela ion be ween chlo-
ophyll concen a ion and elec ical conduc i i y.
Al hough P concen a ion was associa ed o d y
pe iods and ni a e o flooding pe iods, nu ien
concen a ions we e high enough o ensu e p i-
ma y p oduc ion all yea long. To al P concen a-
ion was highe du ing d y pe iods because bo h
algal biomass and suspended ma e we e highe
in he summe , ino ganic suspended ma e and
o al P being posi i ely co ela ed a he inle si es
(Se ano e al., 2006). Ni a e was highe du ing
flooding because o d ainage and uno o ag i-
cul u al soils in he Guadiama Ri e wa e shed
(Cab e a e al., 1984). Consequen ly, he mos
abundan axa in he s udy pe iod co esponded o
he unc ional g oups o Reynold’s (2002) classi-
fica ion D, J and X1 which a e indica i e o sha-
llow sys ems ich in nu ien s.
The posi i e co ela ion be ween cumula i e
phy oplank on ichness and numbe o samplings
( = 0.77, p<0.05) indica e ha a longe s udy
pe iod and/o a highe equency o samplings
a e equi ed o eco d he o al numbe o
axa a he s udy a ea. Few axa had a wide
dis ibu ion wi hin he s udy a ea. Only M.
con o um,C. a omus and N. palea had a
Phy oplank on om NE Do˜
nana ma shland 315
equency o appea ance highe han 90 %. They
a e species ypical o es ua ies (L´
opez, 1987;
T igue os & O i e, 2000 and 2001), able o
cope wi h apid en i onmen changes due o
hei sho ep oduc ion ime (K ieni z e al.,
1997 in Neg o e al., 2000).
In conclusion, phy oplank on cumula i e ich-
ness was high in ela ion o o he coas al
we lands unde medi e anean clima e despi e
ha he s udy si e ecei es a highe inpu o nu-
ien s compa ed o he sou he n Do˜
nana ma sh-
land (Espina e al., 2002). The s udy a ea was
ea u ed as a fluc ua ing en i onmen influenced
by coloniza ion episodes due o i s hyd ologic he-
e ogenei y and wa e inpu a iabili y. A la ge
pa o his hyd ologic a iabili y is man-made
and u u e modifica ions a e planned acco ding o
he hyd ologic egene a ion plan o he Do˜
nana
ma shland (“Do˜
nana 2005”).
ACKNOWLEDGEMENTS
We a e e y g a e ul o Pila and Julio ( om
Hue a Tejada), o he ange s o Do˜
nana Na u al
Pa k, and o ou colleagues Gonzalo Ma ´
ın, A an
A echede a, Ma a Reina, Da id Le´
on, Vi gilio
He moso, Paco Blanco, and Pepe P enda. This
wo k was pa ly unded by he Conseje ´
ıa de
Medio Ambien e, Jun a de Andaluc´
ıa, Spain.
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