k.y :phy;,,in. ponulal 'ot '!>r,,...
TRANSCRIPT
1'01..c.-., )4; 97-114, /999
RELATIONSHIPS BETWEEN SYNTAXONOMY OF THERD-SALlCORNIETEA AND TAXONOMY OF THE GENERA
SALlCORNIA AND SUAEDA IN THE IBERIAN PENINSULA
J~,"er L<>idl, M.~es Hem"" Idola Hlurrun &: Itziar GaK'a-MijangOlS
1kp"<fIMN of PlQnJ /I;"wn aNi EroIon (B""",y), "",,"try of.5d"""", U",'mll'j of,i'w lJ",q", C"""'fY.Ap. _. E·41J()8fJ 8i1000, 5""",. ,d. >;4 4 ~s 1J7, fa> + J4 44~m, E·....il f,pIo«,jOlI~.""'."
K.y_: Phy"""";,,I"n. PonulaL Sa~ .......~. Spa..
"bslnd: Th, P"J'<f <!<.ab wi"' .... 'Y"''''-'''''''''y <:>f tho: <I.,. 1J,,,,..Soliro..u.,.. '" .... lbe",," F'<n1ll..1..I>lhnl p>rt"ul.-fy "P"" tho ';Inifi<llll«' "'ot wooomy <:>f the p"'" s"!Ko",,,, >Dd s.--da Iw ",,;t lJIre<all""",. wer< <I;"-...,.,t",""" 'w<> "'!>R,, ..-.d one Modi_m. Th< A'.... " "n,,, w= ,<;"J,r"""""
Jo/i.:;/oo,,,,,,ny<>,fr"'"fI, wIth roo, :OOSOC"',,"". 1""'r""1 ,'ill< ">m....n.". ",." I<lropk>od .los,,,""(SoIKo",,,,) '1'«"'>' _ SalK"""" "''''1''''",,,,,,,,,,,,,,..... WI'" "'I« .",,< ...i<>n, <:>f och<>m wIth i<"",.lIy~iplo"IJ.,,""'".ped••. Th< Mo;/;""'....'" slhuo:< Soliro",,,,,, "",uiiu~ '" ..""',...,.,. d;<lribotod b<>o~
;n c.,.~>I ("""....m .... oa>I<m <:<10.") and inland salt .--£11<•. A 'omplo:1< .ynu""""",!, of .... cl.... m>p;<of .... di_,ion of it> assoc.- ..-.d t>bIcs ..rronanv"J: tho:rr l1"m'" """""'""'" a", >Is<> provrOe<l
INTRODUCTION
Europe'. Atl3IIhc roaslS are 'Ubjecl to a mnch rn<>re pr<",ou""ed ,idal ",g;me than ;lSMe<lilelT3Ilean ones. 11u, me.n, that the range of the AlI.nli< coa,tal ..It_,,,,,,,he.;, muchlarge., and 'hat niche. and h;OIopo. are rn<>re abund3IIt, in these .a1t_mar>he. we can basicallydi.lingui,h lwo zone., that arc u....lly named by...,.t of the authors WIth the flemISh teno>,hili and s<i>orre. lbe former teno indicate, the area flooded daily during high udes, ,,'hileschorre i. used for the levei, "hich i. only occa.ionally flooded during .pring tide. (BUF.NO
1997). Of the pl.1It commun;ues that m.ke up sail-marsh "egetation, this paper studi.. theon.. included in the 1k<ro_Salicomietea <I.." of holanic di.tribution
]),em-Salicom;e!ea communihes live in coastal mar,he. SUbjeclto udal flooding as well.. in inland marshe!; or play... where flooding and salinity depend on the ","fall ",gime 3Ildon the gCOlIlOfPhoiogic and lithologlcal condition, of the .urrounding•. Soil, can be formedby more or ies. fine materiai. depending on ""dimentation condition., which vory from being"'ndy 10 loamy or <lay, in any case. they alway. h.ve a high salinity ievel and are essen,iallymu",••I. in spile of the facl that in SOme ca"", they may be .lighlly cnriched w;th <:><g.mcm.ller,
Soch communities arc <learly pioneers within the ..It_marsh vegetation complex. 1l>eyoccupy positions in the ftfSl-line bell of the zonalton detenmned by flooding ,nten"ty, I,e. lnthe neareSt Iow tidal line, on the borden of marsh drainage system channels or In potJds lefton marsh surface, during low lide. 1l>ey are .Iso found as inillal stages of the secondorysuccession after disturba"".. have led to the dISappearance of other ,'egetation lypeS, givingw.y to mosaic vegelauon palle= ,n a blend t1,at mIXes these commun;tle, WIth patche> ofother lypes beionging 10 lwu:tltn mm-itimi, Snrrocomielta!'Ulico"".> Sparli~'len ma,itimae
" J. Loi<Ii lit ....
{)r 7..Q'u"",,,. 11Ie pllenok>gical opI;mum of th""" communities is lale summer {)r autumn,when they nOWtr and fruil; thi' occurs later in mOTe ,oulherly lemlOl"ies due to lhe earlyani,.1 of winler in territories further to the north.
Specie, nch"." is low, rarely reachIng 6 or 7 species in one relc,-e; monospec,fk'ly i,quile frequent The height of the vegelalion is variable. but it is ge""",lly lower than thal oftl>c sUlTOUnding ""rennial communuies, Cover is also u",ally low. as is binma<s. wllh a highproportion of the ground UIICO"Cred by ,'cget.lion. The>;e commun'tles ...., fOfTlled bythemphy'ic succulent Clwwpo<!iace"" (often with n",hy aniculated 'tems), moslly l>eIOllgingIQ lhe gene'" 5"lico/11I<>. Sua.d,,_ Mjcnxnemum. and Halo~pli.. The first \wo genera "'"ratller complicated from the laxooomic [lOinl of view, pro"'''';"g sever.l microspecics. w'hicharc difficult to <!etcnnioalc. even more if we consider (ha! only living material can be usc<Jfor <1eterrmnati"",", {)Ccors with Saliwm;o tau.~ fact that there are tWO pl<Jidy level> Inthe genu, Salicomia has been abundantly documented (BAIL 1964, BAIL & TUTIN 1959,CASTROVtE!O & CoEuo 1980, ComANL>l<IOf'OULOS 1968. VAL~ & CAS"'OVIEJO 1990,HERRERA eI al. 1999), and the morphological consequences and ecologlcal-phyt{)Coonoticimpllcati""s of this fact have been dealt "'Ith on a number of '",ca,i"", fG81U 1992, GtHUet a1. 1979. GI1IIU & GtuU-F!tANcK 1992, U,lfONDI1RE et .1. 1992, RtVAS-MAR'I1NEz &HERRERA 1996), 1l>e correct delenninatioo of these specie, i, essential for synlaxoOOm}' ofn..m·Salicamin.a commonilies, and vice ve""" phYlOSOCiology and dislIibulion ofwmm"niti., give us informalioo aboul lheir taxooomic ",levance. VAt.Df.s & CASTROVtF,JO(1990) accept only two 1elnlploid lan (Salicomia ~muici, S, doliclw,uuhJa) aoo "nc dIploidlawn (Saliwmio mmo.iJs;ma) wLthln the genu, ID the Iberian Peninsula. Therefore. we h.vefollo"'ed the COIICept by RtvAS_MAKI1N(;z & lIERRJ'RA (1996). since in th" worl .11 thespecies, which. in our opinion, are present In the studied area, are ",cognized.
1l>e present study provide, a complete re view of th is veget.t ioo Iype in the 1ben an Peni nsui.,laking special care of taxonomy of Salkarma '00 S"aeda ,pecie,. All the a".,l.bie dala In
the hler.tore h., been collected.oo used in the synthesis. except for those whose Sa/;,"''';otaxa determination i, not very reli.ble. A total of 330 ",loves are represented in Tabs. 4. 5and 6. Acc"Pted typology is presenttd In the ,yntaxooomic.1 scheme, which .1so givesnomencl'lUrallyp"'. 'yoonym" character ,pecies and short descriptions of a.soclallon,.oosubassociation'.
1l>e nomendatur<: of plants follow' CASTROVtElO et .1, (19S6---1997) and TlJ11N <l .1(1964-1980,1993). The only taxon for "'hlch we hO"e not followed ,hese flora' i' Se"!,,,,""'11"""'-' vor, cOmp<lC'w (HOPFM.) G. MEY.. in addiliOll to S"".da and Sal",,,m,,, tax.,"'hose nomenclatur<: is explallled in Tab, 2.
Trinomi.1 names of subspecies h.ve been abbreviated into h,nom,al one;; in table,. We i"tthem bere: A""eria p"bigem ,obsp, dep,lala (BER.N'S) "[RN. PR,i!TO et LoID!, Puccindlia/esliu:ijon",. ,ub,p, Ien"l/olla (B01SS. et RHJT.) WE. HUGHES, San.'ocomia p",.,nni, sUh>p"lpllll (UG.) CASTROV .. Suaedo .'era ,ubsp, bm"n·hlanquelii (CASlllOV. et i'EDROL) R,VASMART, et al,
MORPHOl.OGY, ECOl.OGY ANO OISTRIBUTION OF SALJCORNIAANO SUAEOATAXA
A clear tendency toward, the segregatioa ofletraploid (2n=36) ,pecies from dIploId (20=1 glone, can be obsel'\'td in genn' Salicom;o, TetraploId ,pede, possess ~ylindncal fenilesegment> ,n the lower pari of the sp,le, and s"!>equ.l flo,,'et"J, "'hile diploid ones develop
n.= Sf: ," I ",,, ..._P..-
_, .,-__",._,,_.,__)01110 f! ....... -..1 .._015 I .._
.. Of I ,....._ 5 I .._.,..e-
"
I. f' _Ill I; #wPu
51 ..__ 1'-' •5' ..__......
5' .. ••
0,5.,,_ .• , .'s. ,..--u.s.lit....._ •• _5 .._ .........
s,.."...,.;,,~... ..IJI~"""'"
s-. _rili""",-50I,,,,,,,,,_ p"".,,,fol,,,,,,"""'" """i<is-d<J b",."·i>I<Ptqw"i-SaJi"""i"",,, "..,~5~ ",I...JMIi'-SaJ"""''''.... ""'.....HI<__~'
114/.opfpJ_ ""'Ik~'
5...." .... 2Io_J6l,S: .......... 2Io_.l6.S-....!t000J6U'I' '._210_J6,S 2Io_11lS _.,_-.JI>IS .......... 2Io_J6
S, "' .... 210_11S __.2Io_11
5 ...-.2...11
S ,...1a.1I>oo1'S ""'''''. lnoJllS, 1I""01a. 201>011S, lI""iJol, lrwoll (5 """",i"...... 2n_18)S.l'I'iJol, :!n_lls:~. 2>0011
C<ltlveJl·,aded fcnile ...gment> Md latenIl f1o'o.ws usually d,,,,,,,,dy wnallcr lhan the cenll:t.l.....,. The CJOUpof~ ,ncho;b lite Al....tic Sdi<:<>ntM<Job<~Ir".. _SoI~Ihal do II(1II _ rcddi<II.. Md the Me<h_ S. ~n." Ihal bewa... ptllk .. tftl '"
fnut. 1be !WO AI1Mlbc IrInflbd$ d1ffn (rom am odIrr '" the ......m of ferule ...p'''''''''" .......na1 If'IlLt. H)--JO for S.~".. _ 6--UI for S ""'""ru. ..-todl~~
yel""""" '" ffUtL nor P"'P of d1plood """"'"' IX"""" IIwa: of Adaaoc disttI_S.Jon>nIMI ............. S. _",,_ Md S. ......,........ _.., IttOlltr "'Wk>I-' Io:ldI~ Md roattAnllal. s.we- "..,..w...1WtI al.., "W"an '" lite """""'"~ _of lbman Pau.....la (Hue"'" Iltld Alar-el Sd....",.. -m__ Md S. "...,.u. alOl'2)"
becom< ,..;I'D (11110, ..-h,1e S. t<hs<:..", ..._ does. >Dd S- ~II~~ I ....ghlllald '" Il,
In AllWJeyJl-~ lI>rte" aI.., an~a1~ of lhese 1_ ploidy ....<I.Tlw:!I, the 1eInpIotdll are """" abuni,lanl,n shUc~ the diplood, In ><:1lorTe. l1te """""n_o( the Ietraplood 5peC1eS ID the InO$I h,~ly >tre<sed (flooded) 1u.l>l1a15 of ,he: Ad:"mc...It·m....~ .uggest> lhal the ....... La.., onl, ....1cd by poIyplold""'I'on, h",·. <Olonllcd the....w an:al nf lhe: .hUe from the .d'''' '""''''01'' whc:,. diplo,ds h"e ~l00ding oondu,o",m the Mcdit......IIC." ..II-m....he' "",,Iar to Iho>e of >o:1lom: aBd lhu, l>oIh bear lOO>llydiplo,d Salk,m"" 'pc<ie.,
The mo.l abundant So!u:omw spcl:1" presenl ,n each of lhe: a""",i'"00' of11IN...SaIKY>m;~lro ..., .bu...'n In Thb I 0Ihe:, Ics••b,mdam Of I... flnl""nl Sf'C",e. an:>/tovo-.. In Incketl;. l1te ploidy level .. SI&"lrM:anlly t:<ltKlalt:d "'ilh aJl,ltI<'Cl: all,"",," I """nlypt>UJIIi IClnplood 5peCIeS, ...·hlle d,pIood spcae> mosIly occw ,n 11 and 111 The: <bromol.omenumber cbIa~ Iakea from Tun.. Cl al. (\993)
The ......w s.prcoes of S-.M IhM Lale pari in TM-s.J",_,", comml1llllle> beloQg'" cbe s-do -..n-.- plOIp. ..-bKfI doff.... from the Dl1lOfb1oul s-tM spkWfu ID ,b....__ Ihal Il<tIber • byahx IIonIi:r ... a , .la- (Jllj IJ'U T1ltu ..,.ae. can be~ .., n da< P"'P s-do _nl.-r IJ'P"'I" 0ItIy Id cbe~ coaolS of_them SpIon; s-.Ir<~ ..... "1cd:IIe,'_'" rnauaI arKIOOIWJInIllII dtsln~ MdS. .JINxnu~ ia !be A'-': -=- of cbe lb<:ruo> I'l:aursula. s-.Ir< -mu- arKI
'00
s......."" '.."" ",!»p, b",,,,,_~_,,;(CA~OV etPEDltco.) RIVAl MART, .. al" Ju"".. G<oboI. $, «rJ,,~,
S""",," >pkNJ.", (I'OcRR.) G.EN. et Gooo .. AF"""" 3. .10. 18SS
S"",Jo """"""" """"Swuda """;,,,,,,, (L.) DuMOll1".• R B<lg22. 18-10
S_da .UH",,", LAzARO IBIZA, Aw<_ F~
1'roV- G. C""l!7_ Se>illa, Se«, 4. J' PatI,74,1910
s.-.J,;, ''''«}la (WnH'.) MOO., Ann. Sd, N...(1'0,;,) 23' )17, 1831
&lkomuulolid,,,=cnya MOSS, New. l'l1)'101.11 4O'l,1912
&Iicnmw ""'nei DuVAL·jOUVK IIull. S<>< 11".."'an« IS, li6, 181>8
&~cn,"w '"ropaea L.. Sp. Pt. J, I1SJ
&kcomw I.,,,,,,,'IS pw. BALL" TImN, ",,,""';,4, ID). I'IW
~O""""""'" PW, B;,.u., TlmN, W.._i. 42lJol. 1959
Soli<wn", oWe"," PW BA.LL eo 1lm~. _ .. 4:IDI. 1959
SOU",,,.," pat_la D!JvAL-kJuV", Bull, Soc. BoLF,..nc<-Il, 115. IIj68
Sal.co"",o ",.,,11. Wooo<, 1J.<;o. Gal (l<Jr><loo) 3.10.1851
Sal,co",," "'mM"....., WOOOS, EkJo. Ou, (London)),:l9. 'SS,
Sy_ym
""",<la Jh",roso <Ub>p b"""f_ _ I. r>oo
MOO·· Il.Au~-Bl.A.>;Q «0. EIOl.6s 1958
S-Ja pro,mM p;,.u. SioMM ""''''''''''.or. "",,"""""p<> Mco;I,
S-da "",n""", , .... ""I,.", MW
_ man'""" 1llKt. """ (L.) DIJMOf<T.BRAUN·!""'''O.'' 0 IIOLOs 1958
Sabm"",, "nd<> "'"',. ..,.. D!J),f()R1"G. MEY. IS24. pop
S"Ik""ti<> jmtUjs PW B"'-L" TunN, Sal.",,,,,,,,""<Id ""'" non DuMORT 0 MEY. ISl-t, p,~,
S"liw""" "'riJa<", -.cl, non (1...) LBRAl'N_Bt-"NQ .. O. BOO/ls 19'~
Saliwmw d;m"i,"Ia", Mnss
S.albucms both occur ,n the Coa'ls of the Canlabrian fringe, where lhe former occup,e,organic m.ttor-<:nnched biOlOpes in upper shUe. wbile lbe l.uer grows in upper schorre
T.b, 2 summanz., the compl.xily in Ihese lwo g.ne"" of lhe It>.ri.n Peninsul., showingcorrecl name, and ,)'"",nyms.
eCOLOGY AND DISTRIBUTION OF THE SYNTAXA
A, a conseq""nce of the gre.'llcr diversily of biowpes in AlI.nll~ sail_marshe' w. ~an
dislinguis.h lwo .lIianc,,, in lhe French_Iberian Atlantic area: Salicornir!n dolicluwachy<>-FagiIi.J (,I iUecommun'tles) and Salicorn ion e""'pa€O_rammi".,iffUU (schorre ~ommull'lIe,).as opposed 10 only""" ,n the Medllerranean; Sali,orniOfl pal"la•. These lhree alliances. andlh<: thinel'n as""",iauOJls lhey include in the Iberian Peninsula, are represented logolh<r ,nTab. 3,
'"Table 1 SynopI" _ of .... 11It....s..l"'om....... , ...... the It><,... ~,_Ia s..__,__",_ 1_3; 506",,,,,,,,, ~w>-frn.,Ii; 4-1. Sod",,,,,,,,,,, ".....w ~IJ I
s....---. I"""""~__ '-"",_, 2 - :5,?<~"" .........._Salicomic..... ,,,mp."'" J-S-d<'"," "'... , 4 - __ do/;cltol_~)<j"S _ s..Ju~,,""""frald.. ;6 _ SahN>",m"", 0loi<:._7 - S-J< "",,;,,,,,,,,,: 8 - HaIopoplid<",," anpl,,';"",,1i>. 9 - Mit:><KM_'"," ro,aI.Ioodo" 10 _ SMOhlob_.~;_SaIi«".,., ... ""row, 11 _ S......., ...",_·Sol"""",","", """,la<: 12 - ~......_;,-Sa~<o"".....P"'~ I J _ s.~",,,,,,_ _ "',.
No. of «>l.....NG of ",..,ok
, ,26 .l6
,, · ,Il 16 •, ,'"
•, 9 10 11 12 Il.... 18 JS .1 14
Cbon<t<r -"" oIlJ1< _._ 0II1a..,.. .001 """"~..-j,,_ V 1
SaJ"o"",,, '""'P"'" Its-m,ol/r<".... 11 J~.JoIU:""'Iac~,.. It 11~ h<r<S"...., I It~oh<.... 11 I Vs-da ....._ 11 III It
H~""'f'I-J""DJ"M~ co",I/<>od"~pImI/<>
Sal,,,,,,,,,,, .....,rlci
DiIr tiaI <ptcl<o ol.~_Io.'_'p.kiluSpan /1... + 11 JI III
'"
•"" •
o;rr 1iaI >t><cios of s..Ii<-u- ••"'~.._ 010<1 ~i<>. _'.._',...frnt;m"" ~.,....p"""~",, ...,,"- ID 11 2 I IJ III"''''''rlp<'l'"''' I 11 n III III It
Illff ' ..1•.-ifs of S41i<.",i<Ho pan.1<u""Itn,." """'=__ Is-da'P"_
~"'~s-dob"'~J
JI."'_b~'F~~....u"",
•
•",
"",",
"'"
•
" "
O!h<' .ptri<oJ_", _"to...,H4I_ ,-r..Ja<md"Sp<"MIa"" _d",PI<uw.1" -=F..w-...J.on.,
.sa..- 1"""""Sa""""""", frwlC_r"lloclwl ""''''"''''"'mp/<. _,ro",u....,.._ .-0/8"'"
Po/vpor<»o ...",""","--p~~-Sp<r.rolono """""'
•
••"",,,",,
'"•"'",
,"
"m••
"
•
•,"
•
"•
'"•
"••
••
•
•
•
• ••
•
•
•
",
•
'" J. Loi<li et aI
P"""..,11id !=i<c"lata " "p~ ;11<."" , •J"""", ,ob"Ia,., • • •c?/,n.. ="1"",, • •l.oId C'i,"-"'k. • •ScUf"" _,., • •Sp/v""fI'U do >Wi,..,,,., " •Sa","",,,,",,, *.i • • "~"""ria d,pd<>taC<>e/okarw ""..,rid • •EJynw.< """""""'osLi-.ium bU.."",,,,,,, •hut<", t'ron)i • •flurk"""" -...rompo"""" •I'wc,on,lI;" ,,".,;jclla •P""""d/w. '"I"'''ri, • •l.mw.i_ co"'" • •C...-,i... 'P'c"",," • •PM.~mi1" """m1" •e"",,, ,,,,it,a • •l'!aJttato "''''''''''''' • •A"" sq..-at'" • •
TobIe 4. SY""I"" toI:>k of' he Saliro",j"" dol"'ho>""'hY"j",~iliJa1h.nee. 1-2 - S<J,,~,"," ooli<Itc_~,~,
J-\ _ Sdi;"",,,,,,,,,,, frogdi'; ~ - &10""",,,,,,,,, oh",,,,",,,. 1>-1 _ S..,.""'''''' ""'riJ.....
No. of <01""", , , , • , • ,No. of ",10.<" , • '" • , • •C............. <p«'" '" tbt _at""" on<! ,h••Ilian""Sahmmm doIjcloo>""'~Y<' • • '"&liromia "","'"' " • • "&Uromia "I"c,,<o '"
, •S-", ...",'i_ W " W • • •CI>aroo<'", 'p«;.s of the <I"",~'.ror-Q "01.......p«ioo4"" tnpol_ ,
" '" '" '" •1'"",••/1", """''''''''
, ," '"Span"", ""'nt""" • .. "Jp',!:"."'" ...do. W " "Sa "'o<v"';" 1''''"'''' , , •
Tn,lochi" ..,rlti.., W ,)WU:'" ..,,,,",,,,,, "
,1'''''1'''1_ "",,",",om ,11"1".__ p.>rz..Jac..<k, "l.<_".. ""rpm
Salirontl"".. <lo';"M'Ia<~"...,I _1l'.RU'A (1'195)-, <>1>. 2'J. J ",I•.. LaID! .... (l'/91): tab. 17, Id, Ill-II.,- Bn"O (Im); tab, I., %rd•. s..Jit",,.,,,...,,.j"'riw: J - H.......~ {199'i p. ,B, I "'I.. Loll>!" 01(1991)' I>b. 71. ",I,. 1-9; 4 _ BlJFN<) (1\I"n I>l>. 15, 6 "'"..~~''''''oI>~: 5 - HE."""" (1995),'ob 28, 7 ",I> S.....u,. .. ...,.m-: 6 - ONA.JSDlA (1986): tal> 25, 4 "'b.; 7 _ BIJEJ'O (I9'Tn 'oil. 21. 6 ..1>
'"
--"' 1'-- 01 .. _":4=,,:_::'::-.:: ":'":':-:::':':;":':-':_:';';':.;'.'n'. .'~'__•••••.$'.__""__ -__,as '=_
ThI1 a1llaroce gJ1)01pS A,l...uc ulHnMlJ> .."",i.,ton. tIIll 1l,~ under daJl~ fIoo<hngcondlllODt In sail mar<bes, i.e. !hose "hleh 11,'. ,n !be fi~ belt, Of 'hUe ".,. 'hUe i. flooded',",'ICe I dly. even dunng nup ncks, and JQII. arc L1Su:olly lorrntd by fine m.~ri.ls, Ifl<>'llyk>lom, ,,-hleh lCCOOnlS for Wit ,,,,,,,rally muddy leUU~.Tht dnr.,,,,,gc network is art>or=ent
and ,n some .. It-marshes lbe .hkkc «1"00; • w,de area. FOIl' "'>o<,all"'O, of this alli""". conbe i<Jenuficd ,n the lbenan Ptn,.,,,,I. Sul,wm;<!um d<Jliclw.'{l("hw" """p> olig<>Sj>CCificglOSSwon communities in A~...uc <:Oa;l.Il ,hUe. on ~)' 5011>. $u1i<:t1mi.,_jmlili, I.often fouDd 10 be in CQnta<t with lIle (alia but ,"",upi« ........-litre !be SOII"..-e """'1*'11 • shJblly hJ.&ber Ie"d ,,"""" floodilli 1$ $/Wklwn ODd don llOl laI.l .. long SolU'Ont••,,,,,,oMc""" has br:m =ordo:d 11 only 0Ilt loooiJon on lbe nlJmI Canubnan coasl (HEJI.aEaJl1~). Illllu&her Ievd. In !be 'hkU-schom!~ ....,.. T1w fOlllth aMOClauou IS S..""u,__ ",_.diwi"'-l.. lbe ...~~ ....O(_eKberplw:aoloolhanlhe.....~~_ ....... 11 <lCI:In iD !be upper Ievd oIlhe <line ......". otfWII __ 0( o<pDIC drtn....
_laloe5.. ",., Iberian <hlnbuuoD 0/ Ibeso:~ IS oh::lwa III Fi,;. I.
S.tkomiotl~1..1mMI (Tab. 5)
TIlls an..-.: melude. AtIaDbc~lballi~e ID pah. m-.I< ..... ..,.. o:i<:.-edby man In !he upptt level <Jllbo sail mash, orsduTe, ..~ Ooodula o<nlf" oaly ... 'l'fUIItu", IJda; IRroAl;.MAJUtJ<El 1990. Rr\Al;·M.nlSEZ 4 HEUEu 1996. Bl.'2.'iO 1997). 1k
ochon'e IS usually. n.. surf...., .. ,Ill ;em,~i1 ...Ii"" ""I' fOfllltd by d.y, loam or sand."Iuch hne ru<he<! • =uJn dog«< of nIallUl'y and are lIlu. qu'll' wmpact and "",-ere<!moslly by puelllll.tl don«: ¥<,e.."on, Dfa>nage 's n>eandnform and moo! of the gl.....'<Kt
<:ommunlll.. of thJ' alliance """'ut ,n liS chan....ls. Inland >ilt·m.ar;hes of the SuNtl.mic areabear~ S~I,co,."ia ""rnmun,"e, wlllch arc also u>eluJcd lie'"
1lH:.. are three as"",;a~ons in the lbenan Peninsula; the" localiOfls arc >how.. ,n Fig. 21lH: nlOSl w,doly distributed asooc,auOl\ i, $arcocornlO pn'nIIlJ·S<lI,corni,,~m mmosi...i"",.,.-..:00I«I from the Cantabrian COQIS CO the Algarve ("",rh Portugal), and eas,ly d'<bngu,wbl<by S<l1I€orlua "''''''''....Una. ..·tuch tuml reddish M the cDd of the lUrnm<r "'bel marunty 1>
reached It occupi<o cl<arano:es~ by dlwrballce of the p<fUlDlal ¥eJ<1aUO<> olll\c "'gill)'sal"'" __ ,n !he od>orre. ID COllIaCI ..,Ill the Sa",oconuetea fnuNOMJ' comm\IDI~MuchI<ss abundant IS Spe'!..u.,.;" _Salo<_et_ n"'f»<G<'. oaly rq><WICd ID !hesaIt-mars/lel. 01 A.UDri.a< (BuE."'O 19971. ..~ "o<nlf"" kMmy SOIl> <11 .......... 10"'"saluuly It IS III C<lOIIXt ..,111 oullhaIopIub<: '<1 Mm <11 J.-:nea 'The ..-..I><lo
...."M by Soroo<dd dlHJe... an: ...Iud<d ill 5 d<_ ...t""".• hmod~
w. J. l.oidO et al.
ToI:>.. S, SynopllC _ of ll>e &10«>"""" '"~1"""-'''''''''WU- .1Ii.>n« 1-4 _ S"m"""....."p<''"Iti.f.S.liw~l"," """",it"""", (I -3 - salkorni<'lOI"", '''''''''''''"'''''' 4 - 1'""",,,,,11;"<0,"",!wc;".Ia..,);S - Sp<'l'"lario ""ro--Saliro"""",,, '""'1"""". 6 - s-a.,_ ""~,,nt
OI....~Puc<'M!" ....io"'" • • • "s"IWW",i. ",,,,,ni. '" '" •Sp<'I.kl"" "",n.-. " "Amp/<.> 1"""'""' " ,FranJc..", !afflt • ",I""",poI,,,," " "T,,!Ia<M. _rl,."", '" ,"A"",•., &p'1aJd '"
,Plmol"8Q ...,nr_ " •S""'''' "',,,.,,... '"1101<_ pom.laroid" "I""co, """;,"",,, '"
,•
NQ. 01 rolumn I
No. of "'l<..!,.,,:cCOCC:C:CC-C---"'c-CCh........, opod<s or ,... -.:>o:.i.>Iio... ..... t'" .I~.n",
S"locc'",,, ",.""it""", V VSalloo",,,, '"m!"",,, IS"",da oIb<sc,,,, V
o;rr.,....i.1 >pt<l<s 0( ....._ ..U.. , ..,,,....IIi«o,••POCC;",U., ja>,,,,,,,,,,,,
,"
"
•, ,• •,
,
S<u"<"'o"';" ,..,.."....-SMk"'._ ",..<IoSiu"", ...Ii.c"''''rlo,_ """""....... UE1<Rf.RA (1 ....5)lab, 27, ..I<. 1-7. 1 _ BUENO (19')7); <ab, 17, S ",I, • 3 - CosTA (1991): 'oO 16.... 1,. I. J. 5-1:\. ",o.s:-ao'p/<."i,"",-Sol",,,,,,,,,,,,,,, ",,,,,,,i,,,iml><, _<iIo'l1ino,"", / ...<uolaSlu: 4 _ ItEJtREJlA (19\1S). t.>b. 1J, rei,8_10. S""'I'o/oorio _rlNu-S<lIko",W.", 'M'''" ..." S_ RHt;.-.0 (19\17je ...,_ 24, .J6 ..I,. S-u,,,,,, '''/t<UU:6-BW··O;OII991): .... 16.3"'l<
occurring '0 ",hones from ,he North Sea lO ,he Cantabrian COlI'l of lhe Bay of Biscay, Oa'aare available from only or>c locali,y in A,ruri", (BUENO 1997). bill i, i, in an probabili'y 4uilefrequem along !he AlIao,ic lherian COlIsdine,
Sslicomion pIItulae (Tab. 6)
Tidal oscillahoo i, much les, pronouoced on Medilerranean coas" ,han i, i. on Allan,iccoasH""", SO the flooding gradienl occupies a much narrower band and differemialioo he,ween,hUe and "'borre i, irrelevant In !he", circumstanc"". lhe halophilic annual oommuoilic<of succulents u,ually occupy lhe firsl posil,ons in woalioo. in ooll1aet wilh lhe Sp(JI1ine'eacommunilie•. TIte inland salt_marshes, .""ndant ,n some ",gions of !he Iberian renin,olawhe", lilhologicaL chmalic and geomorphologic oond,lIoo< are favoornble, also hear lhis kindof vegelatiOll ,n plac,"" whe,.. waler ,..mains loog..- In spring or early <ummer, The", sallydepressiOll' are f'"'lucn' in areOS wht,.. rocks rich in easily «:>lul>1<: min"",ls. like gyp or mar!.are dominanl, a, OCCori in ,,'me of ,l1c larges' ba,in, of lhe b,ggesl lbenan ri'ers. ,uch aslhe Ebro. Tagu, or Guad,ana
The di,ltibulion of tl1e associations of ,l1i, alli.tlCe is sl1olVo in Fig. 3. They can be separatedinlO IwO groops, continental and COlI.lal. The first group ha. '"ree association" S""ed"hmun-blnnqun//-SnIU:ornitlUm patulne i, lhe mosl frequcm and ex'eo,iv., occupyIng pi""",
'-'.
Fill. 2. Loc...", of Ih< r<k~ SdI~ .M."I""<"".......;,........ : • So"""""",,, ~,.,MU.S.hco"'''''''''
"""""""""'" • SwJtJ.,_ ""I,.n,; * SiH'8ula'ic "",,,,,,,,,-Sali,",',,,,;,,,,,,,, .0fOp""" B_...p";';' ""'P ofRJw.'-M""'l1~1'Z..... (19901
o
.-F'I 3 Loc...", of lho: ,d,~ s.JiroIwi... potu"": .. M,c....-.."",""" .",a11<Hd", •.~"'"",'-<_Sali,.""","", p<JIiJa<,. Salico",,.,,,,,, ....n". *5""'00 o."""-I>Ia"'l""ii..5<ll,ro,,,"'... JI",.IM:• s.....;" .p/mtUIIfi.-SaI..o",,,,,,,,, p<JIulM, • Halup<f/Aid"",,, ""'I'1<,icIlMl". Bj~P"'" "'Il' ofRJ'-A'·.'IAJt11.,1'Z" ,I. (1'1'10)
where waler remaln, long<r lhan in biOl<>Jl'e' co.-ered by perenmal vegelalion: M,OlXnemelumcor.lloidis i' much I,,, f~UCnl (reponed in lhe Ebro and Duero VaHeys and La Mancha)and occurs in place, where flooding i' ephemeral. HulufUplid.,um mnp!".,icaullS. how'",'e,.occur< ,n l<>o;alion. where flooding lam longer; d<ves of thi~ a,soci.lion arc only ,""or<ledfrom I,"" Ebro ba~in. althoogh Halap<p!i, amples;,au!i, i' also found In coo'lal ...11·mars,"",The 1..1 <wo ","""ialion' are defined by Microc:n.mum coral/video and !/uloP'"pllSvmplesimulis re<pecli.dy. boIh absem from S,,"'do bTU""-b/an~u"",-Salica",'''',,mPO'"'''''Thi' group of aswelalions is separale<J from lhe reSl because lhe}' ,uslaln ",meconlinenlal-<opllmum h.l<>phyle5 like Puccindl'a f=lC~'ara and SiUlt'd. ,'era ,ub~p,
b,."""-b/a",!"",;"
-.-.,-_ .......... -,......-, ,U', ''' __........,'-6 .••,,«___............. '" --_..-_,......, " ... - •_ ....._,.......1> )0·_-......__"11' ...._- _.:.-,,- _. , _,XI_ ....--,,, "___'n'l>· ....._ --,. " -,
.- , , , • , • , 0 •-" " " " " " " " " • " " • , , , , , , ," " ".....- , 0 ,
" • , , , • • • ._...l • • , 11 , , • • , • • , Jl , , • • • , •~~ • -_.- ,, • •-- . ., • • • • • • , ,--- , , , , • • • •-- , , •\w' - ,-• -, ,
0-- • • , • , • • • , ,--- • • • • • • • ,'-, , , , • - • ,_. - • • , , , ,e--..__.....--- • • -• • , • , • , • ,-..... • , • , • • , •-----, • • " •-- , , ,S- .. - • 0C.....,_~ • "'-- -...._ ...k_ •- •--- • • •..-,.......... " • , ,0......._ ....._.--- • , " , • , • • • • • , , , , , • , • , , , • • ,--....._-- " • , " ,.,."...,-- , •
,--- i.............- ,'"-.- 0 " •..... -_.- "
,-,- " '", , •
~.-, , • , • , • • " " " " " " " " " " • " " " , " " " " , • " " " INo ..-~_ , • " • , , , • , , , , • • , "
, , " • , • • ,"
, , • • • , ,-- '", ,.
f1'oroN1Iio'_ , , ,-- '" "'-- , '"1-..._ , , " "-- f---~- • ,;- " ,.-.- ,_..._,,~ , •''-Ioffl, ,
f"''''''''-....._~--~ " ,._, ......~-- • f-.......- "
•".....,..,.,... _,;.oIis, I _ 8~'" ,,"""', ,,,• ., L, , ... " __, , """" .. ,; " .." " L<
,11';_..-.._, ,- ""',·MA>'rMZ'" C"'''I""r "". ,_ " , 17. J ~"." -~'.IJMI<) 0011), .... '. " ....." _w:-:. I"""," I.' ",k.,'"lA..,.. .._"..., .... ',0 "', "ADO'",; (IW>( toO, I, '" D.' - U I'_' l>. J '..... 1_ ........... " ...'. ' .w, I ....,
.<-4, ...._ ....--.-..-...._,._ ....~ Oo<iIo"m), "". n,'Ob H.""""" _..o ..........; 10 _"_ .. ,;, Cl"" .. l,""10-". ""'Mo<~_"_, 11 _ BEl.MOI<I'EI< LAoo.a.{ '" toO. , ........ "'" s-4o_.__,., """""RODO~11 ...., .... 7. <d. 2.'"~__. 12_U... " ...'.... ,u ..,.....__·__....__; B-8.,.,...(I""" 1' '-11, 14_........., (1'*"' .... L'"". ",,,_ .. So<....... ; ,,_~ 1'_)." ".' s-.o._. ........'"_ v 'oo ,{I_,'" Ju J, '_"-"_..........u..-......._ ~_,,' -CJo""""" .. ,)- .... 1, I ~__"', ,,,,,,",...-,,,_, ,. _v~ .. ,;,,"',)' ... I.' ~,...._~ , lO_CA""" """"",n...)-, ''',' ...... ...,__~__, »_ ....-z.I'..... """,, "".. ,0""., ..._,,-,..~~ __'.' _""",,.,,,__",,,,"-__............ .-w,:1 -000.000"" ,,", ,."". ,_n, ''''''.>0---''.'' -R"",,,,-n"', "" , ... 1-1,"""' , ... _...... 1".""" Ill, ~~. '_', u __""'-"""""". l2, ...,. ,_~ l ...."""""'_,-;·""..-.-_,_......""'~__...__ ,.....,,, - ""''''.'' • ..-. ~ "0@0l· ..... lJ.,.. 1--1 ....__.----.. ,,. _<»or.
""',, ,..,. ". ""- L_" . .., ,..,...... """_, ,., _ ""'""" "" """"" ~. "M')" "" '. '"" 11-1>~_ ..,.-, ,. - .......,"....-,,~. "08lJ1' "" ", ..., H,""__""""'""'_~""_,,,,,-,,_ ..",-, ""'"'_ "" -.. ,,~ (l'")-''' " ...''"' -, ..._----"""'----...... _ ..~"_,,.-COOTA".. II ... ".~k ')_21, ... .....__ '-,,"'",
......-..... .-,,_, :lO_Coo,,, ''''''''''. \$,"" ll-l.l_...,'-""__~_,......, ~;,.." ,., l., ..... ',A, ... _ ............$0/0 ~ ~~~
-"'- """-___""_.........~_..,'_11 --.;."""""" " ..." , .... """"'" I'"'' ,. :lO.' _, "_"'--M"'.'" ',;_,_,~_ H- ......,.<l.." .... ", "... '_'.•"""'<l""" :lO,' _ ~
H' J. L.oodI et .1
n.. C<l&lIuol puup i. also formed by LhrI'<' _'aIJOn<. S..-Jo _n"'..o~·Salicomi~'"",pGllllM '1 dllolnbulCd ia !he ""eSIefII Medll,:rnnean -.:I " fTeqlloellI 011 I!le Mcdz.."."..."...C'Q<II"IC 01 dlie Iberian Pau.....lL S<lrn< pi-.. spttIfor 10 tho o:oosoaI Mcdzl<jjM •lo:aIl-~ ],U AnILuuw- __A._.~ ..... au<:ICWMI8 from doe~_
s-do~ .. -=r~ frequeM ... !hi• ......aaoo. wboch oloo llI:CW\ .. IIx ialaod~ oi ,. be:<. s.-.Io> .rpIooM1Il"-SoI~"..lOJM .. <b~ iD tho
........~~ cL Ibt I~ PmulsulL from cadiz 10 !he Alpve Tboo d1'iUDCt1.c(eallIre 01: Ihd aoooc>alJOe .. <ha"~ _ spcacs Iitc s-.M .u.r.... IS. __
~j. s.."o<"'oaa1"''''-' -.:I s,...n_ _ "IUIWJ, ""'"'"VII •ere. Al.-.c ,1IflueJo:e. 1llI'
..... asloOClIUOn ..Sal~"-""'....""'" ..... <le><:ribed by \he IUdlor 10 be '" tIx ...,.around~ and is diSU'lbulOd la "",!Mm F.-r -.:I Ul !he Thynauan ,,~. ll>lDdlCalOf spte... is !he.etnpIoid Sal"'-Qnl'" .--rici and i, OCCUI'> ,n loo1&.l~~"1 ..-ale.- pond>
A<~""'-'.>: W..._fuJ fa< JOII'PO" "..""~ PI ,*,,~2. f_ '"' 1).,.«>600 GratroI '"NI>;.C_fr"" (Ilaoquo Go ). no ,t><lqo""', ....",__.01""""_, "'I<-nlodp<d
., "' '~'hd I'"fI<" "I"if", I,
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8.u..L PW (1'l().l1- A...- at~. -._F~ 1/",.... r,..- ,.,... hp. .., 119,~
1lAU.1'''' .. 1\..... T.o. (I_~ N<.- __ ....... '" .-. _. _ ..._. 1~}-lO5_
8fuoo'<Tl'D ..~ S. (1'IlI1l- _do 10 lion) ' 010 loo __ do" a-....-H , ' ~l } lo).-ll'1
11I,....... , llWSl ,."., MO" __ 4I%;JIiloI« ...-..«... _ A'J'O'_..._ PloD """"'" Uoo_ .... No l-.
8lA~C • Mol.-. I (1'" "'__ 01_*3. _'_ .... _1 c__•__r. ''s '-, m_M
IIIIllJoO D£ll96.l, a,...,.,. ..,... ......_. t'••u _*__Bou'lsO 11l!:(l'l&1tC " __<to. =.....-.. lJcItftp
,5qtn. a_JII j-1IO
3OIlltO lIIl('99litlo~<to"'lllroa....C...."d<~I'J<$ M_ol .... -..I.-.PIoM_o,.o=)._ E-.C-'Son-.Ci.&«. 110 '~261
B"'~~·BlA'rlQl.'I!T I. (19JJ).I'...J_*r,.,,,,,, .......~ 1'_. Il~,"""'''Sobt... " .._<I· Mooq><II....
B....UN-Bi.AI<QlIl!t J.• IlOl.6s O. (1951) l... IN'III''''''''' ofptw.< ;Ju bow.. """..... ran <t .....
dy......."""......1.. £1_, £>p. A.1d lHll I lti/).IIUU ~.BI-",q.JItr J. :le 11JXEN 11. (I 'UJ), Obe..."", ...~ 1'<;<..._ .......,... M,tt< l<uropo>. C""""",,
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IIlIENO A (1'1\17). ~.""" Y"'P""'''''"''' u" 'JlO<J"'" ...'""'""". Coad Mo<110 A_. N.....1eu .l.C.....J.~Seo- PubI Prift<.......*A_<>--.
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(1 __19911 ""'_/-5.1 RaI_ .......... C5JC._C'al',,-'Ol)$ ( .... ). l.M ....~,_. .. n._-... ..._» 1$1-m0l0Ir,,-~ l 0_ 1\ ..- .. _ I .. .s.- ....~
!, • I _MOLH.... N. -...,:la O_ll.
'"Cosr_ Le, (1'191). FIoro " "'8"~;k. do 1"'''1"'' ""'"roi da Rio F0nMS4 (1'1"", aNi "'6'''''''''' JT- R",
Fo.....,,., ...,.ro.! pari). f'l,O. Tho:.i~ Un;. T«ni<. <k L,>bo«. L,"""_C\IR<XI A. (1'1%), La V<r«oc>6 <le140111 <le ['&re (1I): ... """"nO." hohllik.; IWo>-nitnlfi". (,1&>",.
I'u«wllW-&JironIj.,,,, , c..J.ijn", "'n'''''' (Ve!""'",,, from Ill< Ebro Del.. (ll): ~ib< andlIak>notrophilw<_n"'" (p"<",,,,,U",-.5ah"""",'",...J (ook,......"/;,,...« ......). F01", Bet- M",10: 113-1.19
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Gll"u J,M (Im): la salkorneo .",,,,,,11es d'Eu""",. 'y<O!m< WlOllOOllq... <1 e>"" lie <I<' d< """""",-MnCoIJoq Ph;«»U<. 18 l17_241
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pIoyt<:>/<o!os;e CoIJ<>q, PkyI=x, 18; 2S-40GtHU I.M '" Gl'>llJ.1'RAr<cx I. & CARD!' B (I97l!), le, SaI",,,,,,~,,, .,..".,,, <I ,,"""'.,-,..... ,.. ""orlll
mM"'mnlen f,..",..•. ..le," &<. Molac. 4: 'N-l!1.HEIlREJU M. (19\I~), F..«oo;,,& la v<r<"""loI Y fkn ;·,,"ul>., & I>. , ......... &1 rlo Ao6n (Cont_.) Gu""'''''''
1 142\1I1EJ<REJ<A M .. I'EJ<NANOCZ CASAOO M.A. '" FEJ<NA.'DE.Z PIl'Ero j,A. (19S9)' El 1<ne'O Sal"""",, L. <0 ,I
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<I< V>I<""a. v.~;".I'1GJ;Arn S. (19~J): Inlroduzlon< 0110 stud" fi'''''''''''k>s'''' <lel" p;."",,, "",,. or""'.... coo """0;;01",,
"Ioanloo .11. '<t<U"_ I~"",""" (<001;"""'''''''') (lo,.-cd>o,iOIl '" th< pIo)'t"""1O!oI".1 'nod;' of"" u<ltf1lV,,,,,,,, 1'1..... wllh ,['«;"1 r<m.wb "" ..,.,1>1"'1<\0';"" l'on<" iOIl)). ..I"h, 80/ 5<,.J 1_2S
RKJUALA (1%8): AIf'J"""»OC'O<"'""',je Iocl_SoIk",'"""' ',,""" B•._Bc. e1Tx IW 1 on 10 1"0,""''''<I< AI"""". C..u.ct. Bc<. (&"",1_) 7: '<15-99'i
R'VA,·MA"'WEl S (l'l84)' 11<1""'''' h,_,o< No<uto IV S'iod. &7<. J: 1_16
'" J. loidi .. al.
Rrw••·MAR1fNI'Z S. (1\19(1), Smla>"""""" de 1.0 <1_ n",,,,~m,,'.... on Europ> occMl<t>tal. frdI. MMir,16, J5'J-364
Rm<S_MAln1N1'.Z S" C"-'ffll P.I'ERNANIlEz-GO/.7.Au:z F. NAWdW>C. PIl.ADOJ.M '"~_MATAD. 11\1901 BIOf"'fmlk.J< lo> P...r.u..to IbJ~ Id", JIaka"" ye......... Copy~ In XJ_d< fl""""""""la. Gnn.ta.
RIVAS_MAl<Tl!.1!Z S. " C=A M (1916): Dolo< _ 10 ...,.."''''''' hal«,la do< I. M.ne,," <E>¥or...} Co/hf.P~_.•,31-91'
RIVAS·MAKIfNEl S .• COSTA M .• l;A<n<>VII'.lO S. '" YAU!01.< E (198(l): La ",,""";00 do DOOano l.<>""""2. S-l\lO.
R"'AS-MAl<rtJ<I!z S, '" IiDul£AA M. (1996): DokJ< """" s..""""", l.(C~)... EopoN. A",w,lard. _, MwbW \.01 1_114.
SAlAZAJ. C. (1996)0 Es-s.o/iJm«~iro<k la "',""" "","""~~ 8<_) """"~<k/~M.-. PhD. n...... Ullioenio1ao:l d< JokI. J....
SA.."'" E ""'- '" ROORlGut:z.()C\I(M R. (1991): Su<1oo. 'oJ<U<.... Y "" ..I><......... rorw of~, p<W
.....idad ... I. R,oj.o medi•. lr<h<a J: 10'1-141n.m.~ TG_. IlU.GD NoA.• 0iA-.u A.O.• I'lliIoNDSOI< lR.. HEYWOOD v,H., MooIU< O,M.• VAI.EI'·n'~
Il.H.• WAL1lJtS S.M . .le WIJI. D.A. (1\19)): FlcN .._a /. F.d. l. Cart'ltrio4< UDi-.i<l' IW»,'-TUI1N T.G.• Ib!vWOOD V.H.• BUJ.GE.'I N.A.•~ D.M .. VAUN"NIlD,Il.• WAL1lJtSS.M I< W"". D.A(1%l-l'l8Oj: flt= ,_ I_l M. 1 c_,. Un,"""., Pr<...C~
T\)XEN R, '" 0Il..000.1a F. (19~); DIe Pfb.U'"."k s.-""" If T<il Eu""'ibornchePhan<ropmm-G<><II",lIoft<n Spont<", V,rof! G,oboI. In". R"",llllric~ U 1-)2;8,
UJ.<A C. (19&6), fkm' y "'''lOCi&< M Ja HiIK... Toodool<uwL PIID. Th<.;~ Un;'ttSidod de N.,."., """"""'"V '-Ulll>; B .t C...-nlOVIEJO S. (19\1(1),~ L. In, CA...-.:wn-:x> s. « al. (<d•.~ ~',,"" ilK,;.., 1, ~<aI
J_n _"'0, C.sJ.c.. Madrid, pp. 'JI-s,nV'-Ulll>;_Fu""J A .• G0007.AlEZ B s J.L .t MouNA R. (199))' florn, wJ'f<I<>6o M I<n .ooI<ado"., .k
C""*,,,IIIo,"'_ (SE M "lIta< ~ I.... E.ss.A~ E>..... D.plla<........ AR>ac0l<, AJbo<><t<Va.O,V"," M .. CARUo;as M,A .t CDo~'ANO5 (l9S'J)' !.as 10&»... <1<1 Camp> <I< c.b<.... (COodad R<aI)
801. C-rI"', I., ~SO,
Encl. AI'l"'ftdi>. pp. 111_114
n-c 5 se
APPENDIX
,."" .._,...- '"
SYJlTlUOHOlIIY 01' lME Jh1ERO-S.UCOAJaIEA ct.ASS {TIIb. I'~......._. .$'. ' (......1<-IILuoQl."ETI'J'~
.... _. __-. $ n- .... (IIaAL-~"""TCDN I,.JI ...__~ _____"' .. _ n I f _ -~' ._
lIIL·.....'lJAa ... 'J ~"; _..,'''''' .-.._.. r _- •• ·.... t9lJ ....... - ..........__-k·it.-t9». __ ", ~._" 1"1' ____ .. •• _ (1IJ're,u.~ co oL 19lIQ 1t..,-_WAll:IDu. 19901'" ... _ of
TI>i ' ...~ ......".._"'_-..w"' _f<>nnodbJ lor _r_bJ~ •.... .. r.......... by ~1'11(19'B~ "'" ptOII<IOOCI rco .... r"""". _toN (n.- ' ..__~.).-I ... ook<(n...:-r"""""'"""l. bid ..~ B•._BL 19JJ.....do • _ 1W<>ph;1., <laMC~n...:-r_ The~ <It "", olaM .... .." """,PIOll. bool .... 0<\01 "'_"'1 • h,p ....on. '0 P""P oU -.: 1~ """"'"'n..... of lI<JII.nhnrOOll hlIop/lilo< _ .... of >ai,
............... tak<""P by 'TOXEN (io Tl!XI'J< It; (la,Jl.OOOfU< t9~8),"''''' "'lObh>l>od to< rno<Iern """""'" of,0< n.."'.$<oI.",......,.., <I.... wo' haIon;~i~ v<&<,""'" <Adl"i«l
~ R,Tr.•• C1tHu 11 C1l!H1J.FIlAHCK'_
SI""~... BR_iL .. It Tll I~J P p. ("'. I):~"" h ·'L" 01 1952 P. p........n.-_ """""'1'11 ,9j;J (.. _ ..... (.... SI) ...,.1·4 '), C..._~I'lCi'<Am ,mp.p.-.,n..~_Il"J)r.R. TX CIIUJ) 1~ __ ..... \ .... JfI~
lWoo, ~, S ti ... f R."J)r .. ot>n-;~';;.;_;;;;RlA."'-"" I"'"c.. _ ..... __""'-__
1'1Ieoo $ U ,t 14 "" n _ coo.u .. G&lu--F_ I_
S,... ~ ' •• f """'""' IfSJ 1Mt..~ _ ..s I... I _Il TlL. Iln .. o.ao t~ l"'W);;C. _ • I' OW'MA.~I",_~<-<-)<1
Il I ,,,,",, • f._ "I .,." GI:Nl co~ .. 0fMv lm __ ~(•• ' .. - ' I' ',.J .."" CIIooa. ,""S .. 0t>I\; .. e->-~"'-"" ."'"• $' .. d t' '" k_it.- .. Il"J)r t9U)
5,. ._ 1 " , ..,.,.,..,...,uO GtHJ .. Rrv-.M.unMz •• GtHJ .. CI8lu--F_ I_s'" n.-~"f><='R."J)r.R"J)r.. o.oo I~__'" (... If,.),5rot<.. _ .......
(M + .,....11. Tx.lfl4_M (....J111.~_· A_/NIoh,Go'Ho;<lGEttO--f'IIo.toO:IMl_ .,,... (... SI' 1fm~"Gl)'l'" RPr'.u-M~ 19112 "" (...SI. SoI«<>_ dnI",~-frqik'00tt; .. Gl'''u·~ .... GQru 19'11_ ,11<1· ( J1)
HoIooypu.· $<JI"""""'_fntI;~' 00!Hu .. OtHU-I'hNCl< 1_0..0.:......0< ,..'" .1<>Itwmio <loIi,wwAy<I. S. kd",m, (. S. fra"I,,)
~__~GfJtU..~_.I_
S,.,. 5oIJ< IrJ-otIl\J .. OtHIJ-~-"""'-""1982 nom. On"" 1",,5),SaticonUs<_"""'"'K........u .. R. Tx 1"14 nom. M, <.... ]7), 500II....-._ .."... Clt'l$ll~~st;N 19j;~ nom...d (... ltl.1). $ ! ..__..,.......,.. o:..tu.JOri .. OtHe 1__ ..d.l"'. lI>, 7)
_,.... G!>tl' .. Gl!tJl;..hAI<O; (I9C:! ." n:I 11)"-~(_ .... _s.. "" N.......... ...
_b!' $ I .._ ., I •.••"";;.~M;.;;.~_; d.J_ .......u._1<>d, _""".. _ ........~ ......¥--
" 1 _ .._'..SI" :s-. _~abIlJ"Gl!Nl~"'-"" 1'If2_ ...... l.. ,~:s-....__..'_pno
___...~.u-~I_
11 I ,,. Ql!Nl; .. ~lt'1f2 ........ U)
j. loidI et 81
European A,lone'" C!loo,,,,,,,,~" ... lonn«I by ,.1I, den'" P"J"oW'Oom of S<llirontid 1I."""".. .,.j
often by s-da ""'nl..... "'n",h I".. In more ItabIe sandy '" ..oo.1oan>y >Oil' '" the uppe' ,I,m ....I
~ """'"" lllllU .. at...."""""" ,_S,n .. S<ll,ce'."'_ <>Nc."" Gt/m .. GI'>!W_PltAN<'K 1982""", ",vol. (Ml.')Ho:>k>typ",: Gl'Jm & G'.HIJ-FRArocK (1982, tab 45. ",I. 4),Europe"" A,1>..", Cl»'wor! oommun,.... form«! by the d'p1o>J S<llicvmw (>/,><"ro """b "" OIl the
" ....;';"".,1 fri"~< belween"ill<..-.:I ocb<n< !<vd,
s.-tum_,_(.. LIT_........'CUT l\127l Gt.... "01. In 11, 1'<.,.10.,.,..,.. 8<JlNo 1W1Sy". Sal"",",;,,,,,,, "'1T>f!O<O< DE LrrARDt£RE et MALCUIT 1927 fIOOl. Hlte. (0I'l. 29); SIMN<kr_
"""''''''"'PO< (DELrr1JlDItItE .. MAL<;IJrr 1927) G£HW« al. ;n K, Ix. 1974 (.... a)l; S"",d",,", """''OC"'P'"G£Hw '" OrnI}-FRA..'lCK 1'I6'J nom. mod. (111, 2b, n, M""'·S""""..... """,'OCarpn<' O!HU 1976 fIOOl, ,.,,1(111 3.); "-,,,,,, 'ripolii-S-d<'_ "",ri_ G£Hw 1976 «>r'. GtI<w .. Ornl}-FRANCK 1'184 fIOOl ,n,,1 (111
h'l«""ypo" G£Hw.t G£HW·FRA..'lCK (1'Il!2: 1>.0. aa. "I. 22); l«,u",; lIu,;.-.o (1997: 2")R<I." ..1y den..,. 'all <""""",,"i,;,, of eorll<f ph<nolocy. <lomJI1>ted by s-da ..,,,,,",,,, whdl o«u~
"'J>"'< ,,,..,..."'he<! b",._.Ioc_d On !be _ ,IUe. T1><)- arr <h'tribu<<<l .11 0"" ,be "rr~",,_AH."t"<""'" of F "I'"
SMi<:omlon ..,roPM<H~"""- GtHU el GEHIJ-FAAHCK 1984
S)'tI __ S<l1"'Qn""" Ta.,,,,I,:;,,,,,, K. n. 1974 pro•. """'. 'm"1. I .... .)/»; Sai,corn"", ,_""I Grn~ eoGi."u-FPAr;cK 19114 p. p."""
HokJIypo,. Solicemi""", !""iu.,.",,,,,,,,,,,ioo<u O£'W 1976 (. Sabromitl"'" do·"''''<I,I<''''·'''''''''''''''''''Gum 1976 OOIT. GtHU« G£HW.Fl<AHCK 1984). '-="... :GF>!lJ.i. G~HlJ_F"ANCK (1984 61)
Ch""",1<ri><i;:: la"': S<lkc"mi<t .i".,. S .....,,,,,...,. s-aa alh",:""
Sw<ocomio> ".,.nnJs.salI"cm_ _ I...... (at.... .. GtHU-AlAHC. l\179l Rlv...H'..m.u 'MOSy". "",<",,11,,, """#""""S<lli<",,,,,,,,," .-",,""",G!HU<l Grnl_I'RANCK 197\1 """'. illtl. ('" 31),
oon R. n '974; .I._do 'l'1,ndt"'i'-Snli"'m"'.... """"''''''''"' R'YA,<-MA011Ne, '" .1. 1980~. PH<>Iooypo.· GtHw '" GrnW_I'RA."C. (197\1: 3SS, 'ob. 2, rei ag)Eu_ A,lone" (C__ A,Ian,~ ..-.:I Med'l<fT>rI<>n 1b<n..·AtllII1tk) (Ia",wor! coornuM"" I""","
by SaI;""",in ,"""',;,,,... (~, ... " """",iI)), ..-b"b hI'< oo!l>e "W'" "'bone """I 00 >andy Of! ...-.:I-loamy
""I,. Ot.., ...."""y.., liko P"-cWltin ""'''''''' Of! Sn,,-""omi<t P""""'........"",'" f""" ,10< 'um,,<Miingp<r<rlnlol <omtnu";,ie~of"• .....;.;'1'1'< Th<y..., pr,,,,,.. lrom ,be C.....b"." OOtil'o ,be Alp"'" .,.j Ci<:h,.",.~I "h_m""h"
v.ri.biIity",11,,,,,,,..,,,,, ,.._"";""0 ,ub-. ""'-' boc loco>. i' ,be typocol ,ulIo""",,,,,,oo,Ho,,"ypo<: G£Hw '" GrnW_FRA."C" (197\1: 3SS, tab. 2, "I 4g),,,uiltrlli<l'I>U,,,joKk._ Grn~" I'RO\'= 197.P....""U'"fOlcoClda," " linked '0 'lay_',," ""I< ""hi<h..., ..,,,,,,Iy dned·"", it> ......,." _ ....'umn, w"h
• oon.lderable "'reas< ;. <>l' """..."",.,.. "'nog ,.... per>OO
S_m ""~Gt"U111& O<KI'. G.... e1a1. 1M3S)·". .1...,.,£""", vulgon', Gf';.. ,,« Gt"U-FkANCK 1%9"""",..,.t, (an 21>, 7); .\.""<d.t"",, p""'Ta.., GIJIIl'
1976(m, 43)Lo<""~po~ G~H" & Gf';t'U-nANCK (1'Il!2: tab 4g, ,d. 161, l.«"mo GFJ'U & G~HU_FRA""" (191l4: 62)Open C<>m1OO.,U., __ by s..n.do oJb<J<'''' (. S...,dot ...";,,,,,,,, v•. VU/gdTU) "'h",h ""'u, i" ""
b,gbe' "'b<n< Ie"" on ,,~II"""I1'Od...-.:ly Of! ..ooy-Ioomy >0,1" T1><)- "''''' A,I ...,,,, oli<rribut....
~_ .....m-s.ficomjot<on__1lufHO .. p, PMlO In 8...., 1917
1I<>1oo)'p"<; 8lJI'.-.o (I '197' 241, "to, 24. reI. 26)
GI""W<41 ,omtnun"", lo>nllOd by 'Ioe d'pIood Soloca""" '"_a. '''1IIUy redd,>I1 at """""'y. T1><)- oc<u,,n tbe "F'fI" "'b<n< 00 f"'l"""'Y I1oo<I«l~ with"..-,..leno"",, 10...- .... 'nily
'"SMlcomion """'1ft GfHU ~ GtH FIWICIC 1984
Syn .. Mic"""""""" co",11cidU RlV ·MAk1'1N',., Cl GE,fU In RIVAH,'ARTf~EZ 1984 (.yn'., 'Jo.);}M_SaI;'o,""," ""'" noo BR,·BL 19J); .0;:,1",,,,",,,,, "","ii Guw .. Gtlm.f1lA""" 19S4 p.~, (an, ~),
Saiiromi<>o """"'u,""'" RNAS_MAJ<TlNl1Z ..... 1980 prov, 1IOOl. i.,'al. (.... JlJ), """ •. Tx. 1974flok>typu>: s-Jo _.in""",_SoIironI~_pal..... BRlUO .. AJ.~A.'<> G£Hu et GeI,u-f1lA.-.c~ II/S4
oorr R1V....·MAnj""" 19'J(): L«tum: 08t" & Gb<IJ-FAA""'" (1984- 61)Chon<,m,,,,, wO, Mi<:~<<=11'>'4... Sa/iro....", ,_rid, S. paJui<..
II~ _ 1tIv........11""" n R...........-.. COO"TA 1~71
L«""YP'" .tVAS-MARTfNEl & COSTA (1976; 32, 'ob. 6. ",I. 4): l«,om- Loo'" &. B'U""ut< (1998 I f>l)W..l<m M«lil<mnean as"",,",,,,,, di>lribuled in conoinenlal lalt.n>ar>i>e, in bioIope> where l'Ioodi"ll i,
opb<.......1ond ,he",;] i. """",," b)' • thl<k ",It.. eN'" Dom",,[«J by Moc,.,.."'....... coroJIood", ~ 0c<"P'e'tht- ""' in Iho holoo<nc>.fIef 1I>OIK>'p«ifoc II./op<pli, 00..",".."",. (BAl>IA ",.1. 1992).
$_ mw/I~_IM'_ IIfluu.o .. "'-"'" •• GtHu .. G£HIJ-I'IWtCK ,_ """R1V............INU 1"0
Sy.__ .\:01""'...-"....~ ""n", B<l!U.O '" FlJRNAR' 1976 noo (SCHlU"Z 1939) CHRIS1lANSEN 19l1:A'*'x"""", 61auci-SaJjCQml.- """"ti,ti""", BRULl.O« Fuo.'ARI .. G~.HLJ '" .1. 1918 """' ",al. (art
JO; 51oס:i<'<io ""8"m-Jaiie,,,..'<I,,," ""'_ B~lJU.O .. I'!J'NA" " G&tu .. G&tu-Fl<ANCK 1934 I 4J)1.«""11"''' G&tu .. ai, (1973; 8l!. ",I. 12. W>. 2); I.«tum otHU '" otHU-Fl<AHCK (1'134' 02),Coo MoO "" <""""'-'''''1eS ~I"'oO by r<dd,sh dipO:Hd Sa!;"ornitJ "",uJ,,, "h~n
h"" in ,.inte,·f1oo:>ok,j Ilnd ,,,,,,,,,,,,_dry ""is [)I<'ribu~"' .... ,.,,,",m pan of [I>< M<d,""'....... !la"... it I>a<be<" rec<Jr<>ed ," tl>< V. i<nc,.".,.TarTaooneo", and.~I< """"" """ i...... M"", oon<>-AI"""""", 1"<"''''''"
s.,'I<..,,_~O. DO 1IOl.64 ... 8 .......0 .. F........ '~
Syn.- SaIic"",u""" ......n.,; O. 00 BOI.O:> 1961 pro'. (art. JlI), Anh"""""", ~ta.<;':;"I;<",",,'_<_ne;o Of! BoWs" ae.U <'- ai. 1973 nom, in.al. (.... l.).
1.«",>,f'U" !lOWs (1962: lab, ~1, rei. J); l='um Il(..os (1996; '(6),
W,,,,,... M<dJt<rnnean rowal (5",;1,. ~'n"" CO"",.. LanJ-. C""'"'ll.... Ca<alo,,,.) JWs"""",,,,,,,-,nnleS dorru",,,," by .... mldlsh leI1op1ood Sal...""".. ......,;.., wbo<ll 10.. COl saody_lo>amy "';1•. They
"""u~y epbemer..J p<><><lo ""110 "t~"""__ tbal dry nul in "'''''''''' (>I ...... o:nd of .... ",""i. ,.h""lead. 10 hip ....,"" """'<TIt~,
"oriof>;lijy
~i<l....," • ...,ri<:; 0, "" BoWs 1\Ill) I< 'yp;eal -"'N<oociJ'''"".1.«",,>,1"''; BOI.O:> (1962. tall. ~7...I. l); l...,tu '- 10«0.,U<Hd........ .....m;_ O. ",,!lOWs 1961Le<""l'P'''; BOI.o. 11 %2, tall. 17...I. I I; L<c1,,", .... 1<1«I
S""'''' """"" (S. ""''''.... I'''''P) """.... "'''''" """f",..,,,,, ''''''''''''''
$'-_~I-SaII«>m_PfItu_ Bfl.-BI.. et O. BoLe. 1151 """. AJ'lOS-M....,,-.... 1_
5j".: A... Sol,,,,, ...,,, Iv""", ... et SuiUdo IN-..if<>Iio BR._BL .. O. "'" IlOI.6s 19~8 ("". 4l). SwKJo,,,u,,,u"",-$aI..,,,,,,;'_ _ ,;"..........'" CI.,., ..,O 19lI1.",", RlvA'_MAIITI,,:Z <1 al. 1980; 5......do
,pk",u"".-$aI;ro",;.,,,,,, P"'uJ«. RIVA,..MAIl11~El 198-1 nom, inVlll (an, h), IIO!l K'VA,·M~u<1011980 "'n. R<VAS-MARlfNEl 19'1O
Le<1OIy1"'>: BUl·~_B'.A"QIJET &< BoI.6s (19lS: 99, tall 2l. ..1 I); I.«Ium: RJ\'A'-MA..-ri"", (199()-
."lbenoiJ ,"land oaI'·..-w<h 11>.._ «>mmunl'i<, 10",,£ to h.o""." flooded r", loo.; 1""'00<1, and ,."b Io~'
"""'I<"W~< '0<JI<nl. I.. pI1<ooiOfkal 0(11<""'''' is '" <MIy an"'"," Ilnd" d",ribut<d """nly in ....Ebro bwn (RIOj..-.oand B:ud<noo-Mon<v<» "";to,,,) orJ<J ,n u. MancM (MMJ<l><lo 0«1"'). In tl>< Gu;o:Ialqo,m
Valley (H;~", o«lOr)lh<n: b."" boerI~~I.""'on<""too",,,.. by MAiITI.~U PAIlRAS "'al. (1'1841on<! G.-CiA Fl'F.l'<"ffi5 (1996) ~b",~ "'-'0 be<o ..dud<d .. "~ .",,,,iot,,,,,.
llomi"""," by Sa~conUa paI"ld. Ih" ,,,,,"nental ."OC""IOII dlff... from ,be IWO 00lSllI1 (>ne:I .1", doJmj", "'"
by • i n11>< "",,,,nee 01 oome coo,;ne ",aI ;pe<ioe,. 0' Swuda "'''''''''''p h,,,,,"-hIaNj,,,,,,;, Puuu..II'"1=<;<0""Ilnd .4<10",1"" I;norulu.
'" J, LoicII 01 ..
_ Th<_.-......,--.~__ (Jtr.·......M_193ot) =- .. _.... • ... _ L.> _ (_~. _ OIl" _; ,? W..,. ca" ,." (1911 la,.... 1.1)\* od. lJ~ _.__ -.I" _.-... 4'/<,.._ f ' ow__ ,
Jtr..u--....z ..... 191O ... did_ ..= "pr _,_<aft<dy tr_.-_ ..... --- <#$ ... - ....... -"'__ .. ..-..to-" : ..," ..__ ,...._ .... ~tan.2j1
_ •• f ;' - 11I_.... 1__._"' ,....._
s,..._,' . of' U.~.. "'I_iort .)~
" ' »""_ .",......MAa1t'o!z 11_ ....... n. od. \ ... _ ;; d ... 5 5 _
U..u-MAIn1!;t:z 19IO.~ ...., &Iaw_'" ........ __ .- <Jl .... _ p ,
(C_ ('lowO _ Alp. _~__ .. ......, doy .... I'Iooclod .. pcnod'Ilor)I <Io1ko '-......... ' _ by Soa/;r...,.;",... ..... ,....,..,. al__
-=........--...,.,.,..s,....-_ ...s..---_.....IaI>lIk,__ ,.,...... _ ..... IM< ....... .. "'" t)'l"'oI ...bo"""'...... """iIR.od '" ....
Gail"...,.Onubon.. _ Alp<> ...... _ •.
1Io1011P''' RIVA'_MAkT1~ <1 al. (I~ ". 1>1>. 13. ",I. 3).<""IN.._..,.INot_ RlVA.$.M;,/(J'fNIlZ .............. '- 10<0I_ff"" R""J-MARTfNEz .... (1980" .~, lab n. Id. 9); 5,• .- ~,-$<>I,,,,•...,_
......,.w_ <"",;_~ R!V.s-MAI'rlI'o:z .. >I. I9$)_ ...w. (.. 211,7)c..,......-'- ...... c..... <....... __ .. pb<...._ -..... ~..."-.:1...- ......,- '-,· __._J.C.o:.r..__........
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