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20 Pages·2012·2.8 MB·English

JournalofCoastal Research FortLauderdale,Florida
The Biological Flora of Coastal Dunes and Wetlands.
1. Cakile edentula (Bigel.) Hook.
M.A. Mauna, R.S. Boyd"and Lynda Olson"
'Department ofPlantSciences "Department ofBotany and
University ofWestern Microbiology
Ontario Auburn University
London,Ontario N6A 5B7, Alabama,36849-5507 USA
Canada
ABSTRACT _
..tllllllll:.· MAUN,M.A.,BOYD,R.S.,andOLSON,L.,1990.Thebiologicaltiaraofcoastaldunesandwet
lands.1.Cakileedentula (Bigel.)Hook.JournalofCoastalResearch,6(1),137-156.Fort Lau
t••!!IFJ•ll derdale(Florida),ISSN0749-0208.
'-:_~ "". Cakileedentulo(Bigel.JHook.(sea rocket)isanannualorfacultativeperennial(Br-assicaceael
thatgrows asapioneeralong the shorelines ofthe Great Lakes,theAtlanticand thePacific
...., S-- coastsofNorthAmerica,andthecoastsofJapan.theAzores,NewZealand,andAustralia.Here
wepresentasummaryofecological,physiologicalandgeomorphologicalaspectsofthebiology
ofCakileedentula.Itssuccessalong shorelines may beattributedto the production oflarge
numbersoffruits,short lifecycle,high potential forwater dispersal,resistancetosaltspray,
possessionofseeddormancy,highlevelofseedlingestablishment,rapidgrowth,andabilityto
flowerunderarangeofphotoperiods.Cakileedentulaproducesdimorphicfruits;distalandprox
imal. Distalfruits breakofffromtheparent anddispersetonewlocationswhiletheproximal
onesstay withtheparent.Thespeciesplaysasignificant roleingeomorphology.Singleor
clumped plantsaccumulatesandandaddorganicmattertothesoil,thusprovidingmoreame
nablehabitatsfor theestablishmentofsecondarycolonizers.Cakileedentulaexhibits highest
germination at alternatingtemperatures of20'e(14.5hr) and lO'e (9.5hrlunder dark con
ditions.Lightinhibitsgermination ratesbutnottotalgerminationandfruitsrequirestratifi
cationtoovercomedormancy.The plants are infested byfungi andareconsumedbyavariety
ofphytophagousinsectsandvertebrates.Thespeciesmanufacturesglucosinolates thatprotect
itfromsome insects,fungi.bacteriaand vertebrates.
ADDITIONAL INDEX WORDS:Cakile edentula,sea rocket,variation,ecology,population
biology, physiology,geomorphologicalinteractions,economicimportance.
I. INTRODUCTION mortality during storms, the outstanding fea
tures ofthespecies are resistance to salt spray,
Habitats along the shorelines of lakes and rapid growth, and abundant seed production
oceans are subject to major substrate rear during the short intervals between storms. An
rangements in space and time by dynamic elegant reviewofthesystematics and evolution
forcesofwind and wave action.In North Amer ofthe genusCakile waspresentedby RODMAN
ica, most erosion occurs in late fall and very (1974). In this review, we present information
early spring;however,throughout the year the about the ecological, physiological, and geo
coastal beach habitat is highly unpredictable, morphological aspects of the biology of Cakile
unstable and temporary.The lower part of the edentula (Bigel.) Hook.
beach is usually the most disturbed because it
isunder theconstantactionofwaves andhence II.TAXONOMY AND VARIATION
is notavailablefor colonization byplants.How
ever, the areas above the low beach (e.g., the (a) Name
mid-beach, driftline, and high beach) are free
from disturbance for short periods of time and
Cakile edentula (Bigelow) Hooker ssp. eden
some specialized plant species establish and
tula (RODMAN, 1974),Tribe Brassiceae;Fam
completetheirlifecyclesthere.One ofthemost
ily Brassicaceae (Cruciferae), Cakile edentula
characteristic genera of this habitat is Cakile.
(Bigel.) Hook ssp. edentula var. edentula, also
Although the populations suffer high rates of
referred to on the Pacific coast as C.californica
89002 received5January1989:accepted inrevision18May1989. (FERNALD,1922)or C.californica ssp.califor-
0991,1.oN,6.loV,hcraeseRlatsaoCfo lanruoJ
-ixorp eht tub dedees-eno yllausu dna delucol riap laretal ,retuo eht htiw seires tcnitsidni
eno yleslaf era stnemges htoB .emecar eht ot owt ni enrob era slapeS .gnol md 2-1 semecar
dehcatta sniamer tnemges lamixorp eht elihw etaetcarbe lanimret ,gnitagnole no enrob
suoudiced dna dekaeb si tnemges latsiD .)8491 era dna ,suomanydartet dna cihpromonitca
,YRAHOZ(eaecissarB ebirtehtfolacipytstiurf ,raluger era srewolF .ecnecserolfnI )vi(
fosnoitropravlav dnaralyts eht ot suogolomoh
eb otderedisnoc era stnemges lamixorp dna lat .)4791 ,NAMDOm ecafrus
siD .epyt "diotnemacun-ovlav" ehtfo esac laic liosehtwolebsretemitnec wef atsujmc 51revo
eps a si,gnol mm 92-21 euqilis detnemges-owt dnetxe yam stoor latnoziroh enif ,gnol lareveS
suoecatnemacun ro ykroc-drah ,tnecsihedni .retemaid ni me 3 dna htgnel ni me 04 sdeecxe
na ,tiurF .)9591 ,TEUQCOB( eluvo suoludnep ylerartoorpattuotS ygolohproMtooR)iii(
a htiw tnemges lamixorp a dna ,eluvo tcere na
htiw tnemges latsid a ni gnitluser ,seluvo owt
.sdnarts ralucsav rojam eerht
eht neewteb sworg llaw yravo eht nehT .edis
ehtfo slanimret eht dna fael ehtfo dne eht
eno no seluvo eht htiw smrof mutpes detagnole
tatneserpyllausuerasedohtadyH .stsalboidi ro
niht a ,noitazilitref retfA .stnemges tiurf owt
sllecnisorym eb otthguoht)faelehttuohguorht
eht neewteb noitalucitra erutuf fo tniop eht ta
derettacs dna epahs ni ralugerri( sllec sniat
dehcatta era,owt yllausu ,seluvo ehT .rebmahc
noc dna llyhposem ygnops dna edasilap otni
detagnole eht ni yllacitrev desopsid atnecalp
detaitnereffid ylraelc ton si eussit amyhcneraP
raenil eht htiw ,ralucolinu yllaitini si yravO
.ecafrus eht htiw neve era dna yllaixaba suo
.mutpes ehtfo enalp eht
remun erom era atamotS .sllecdezisylsuoeneg
otralucidneprep si sebol citamgitsehtfo noital
oreteh htiw dezinituc ylniht si alutnede .rav
ucitra ehT .debol-ib ylthgils ot lacirehpsimeh
alutnede .C fo eussitfaeL .dne raloitep lasab
dna etatipac si amgits esollipap-trohS .ylesrev
eht ta fael erutam eht ni suoivbo tsom si gni
snart detalucitra yltcnitsidni dna mrofisne
boL .debolsseldnaetaloecnalbo ± ,rellamsera
,etanibrut ± ot etaloecnal si litsiP.amgits eht
sevael regnuoY .tnecsebup ro difitannip reven
woleb ro htiw level era srehtna rewol deriapnu
tub debol yletatned ro ,yletaunis ,yletanerc
eht saerehw ,amgits eht revo detresxe era
,esab eht ta etaunetta dna etalutaps ro etavobo
srehtna deriap eht ,sisehtna tA .gnol mm 4.1
otetaloitepdnaetavoyldaorb,eritnednayhself
4.0 lareneg ni era tub snemats naidem deriap
,kciht era sevael erutaM .tnebmuced netfo
eht fo esoht naht regral ylthgils era snemats
era tsomrewol eht dna ,sdrawpu dna sdraw
laretal deriapnu ehtfo srehtnA.srehtna eht ot
tuogniworgsehcnarB.thgieh ni me8 ot pudna
tnemhcatta fo tniop eht ta yltpurba repat dna
noitces-ssorc nieteret,etairtsylkaew rohtooms
tuots era stnemaliF .snemats deriap eht naht
,xeduac ydoow a gnipoleved tub tneluccus
eromyllanoitroporpetagnolesnematsderiapnu
yllaitini era smetS ygolohproM toohS )ii(
eht esuaceb sega rewolf eht sa lauqe ylraen
emoceb tub ,suomanydartet era tsrif ta snem
.tnebmucni
atS .snemats deriap eht dnetbus ,ladimaryp
yllanoisacco ot tnebmucca era snodelytoC
ro lacinoc ,hgih mm 4.0-2.0 sdnalg naidem
.)0891,ENYAP(ecafrusdnasehtevobasraeppa
owt eht ;snemats deriapnu eht yb dednetbus
taoc tiurf eht semitemos dna )1 erugiF( laegipe
era ,debolib yllausu dna yllatnoziroh desserp
si noitanimreG ygolohproM gnildeeS )i(
moc ,sdnalg laretal owt eht ,seiratcen neerg
llams ruof evah srewolF .ediw mm 5.2-5.0 dna
.)8491( YRAHOZ
gnol mm 01-1 morfyravslecideP.slateptsom ni
dna )4791( NAMDOR ,)2291( DLANREF morf
devresboera,sllecnisorym ebotthguoht,stsalb
delbmessa neeb sah tnuocca gniwollof ehT
oidi ralubut ,tneculsnart ,detagnolE .roloc
ni redneval elap ot etihw dna ediw mm 3.3
4.1 ,gnol mm 7.9-6.4 ,etavobo ± era slateP noitpircseD cimonoxaT )b(
.xepa eht ta cihpromid ylnommoc dna enil
ayhyltnetsisnoc era snigram lapeS .slapes eht .tekcor aes;)2291 ,DLANREF
fo xepa ehtta yllanoisaccodevresbo erasemohc ni 0091 ,HGUAPSLLIM( .ttuN anacirema .C
irtelpmis,esrapS.suoudicednoosdnasisehtna sa ot derrefer osla ,dlanreF sirtsucal .rav alut
ta tcere ;esab eht ta suobbig ylthgils yllausu nede .pss ,kooH ).legiB( alutnede elikaC ,acin
noslO dna dyoB,nuaM 831
Biology ofCakileedentula 139
d
a b e
9
,
l '
.
~
. ."..-::;p. '
.. ._~.... JJ.r- ""'9'f"...--~••
~ b~ -A~~.o-
..' --, -~E.:'...
Figure 1. Distal and proximalfruitsegmentsofCakileedentula var.lacustris(a), var.edentula(b),seedlings(c.d),28dayold
plant(e);matureplantwithfruits (f),andamatureplant with disperseddistalfruits(g).Barsequal1cm.
mal fruit may be multispermous or abortive. apex;while the distal fruitsegmentistruncate
Pointofarticulationofthetwo segments isusu fusiformtoturbinate.Seeds aresmooth,bronze
ally :!: constricted. Proximal fruit segment is or brown in color and oval or ellipsoid to slen
terete, :!: cylindric to expanded slightly at the derly oblong and compressed longitudinally.
Journal ofCoastalResearch,Vol.6,No.1,1990
0991 ,1.oN ,6 .loV,hcraeseRlatsaoCfolanruoJ
stnuomaecart = T"
5.1 T 6.1 0 7.48 T 2.3 0 9.7 citnaltA ireprah .pss
alutnede.C
sekaL
0 T 0 T 0.1 4.4 9.39 T 0 taerG sirtsucal .rav
alutnede.C
0 6.1 0 2.1 1.1 7.61 2.08 9.1 0 citnaltA
0 6.1 0 T 0.1 5.62 9.07 T 0 cificaP alutnede.rav
alutnede.C
0 T 0 T T 5.21 7.58 T 0 eporuE acitlab .pss
amitiram.C
3.9 8.51 0 0 T 4.05 -r 5.81 0 cificaP amitiram .pss
amitiram.C
lytub lyhporp lynetnep 2 lynetub lytub lylla lyporp 1 noitacoL axaT
-oiht -oiht -4 nwonknu -3 -ces -osi nwonknU latsaoC
-lyhtem -lyhtem
-4 -3
)%(noitisopmoC
.)0891,4791NAMDORretfa(yhpargotamorhcsagyb
segatnecrepkaepegareva eraseulav ehT .snoitacoltnereffid morfaxatelikaCfosdees ehtfonoitisopmocetalonisoculG .1elbaT
.srotcaf eseht ot snoitat sirtsucal .rav alutnede .C fo stiurf ,noitidda nI
pada tneserper otderedisnoc ton saw dna setis .)1erugiF(nigramsuoiracs a tuohtiwesabadna
sih ta erutarepmet dnas ro retaw htiw etaler kaeb trohs a htiw ,ediw mm 9-5 ,tnemges tiurf
roc ton did htworg toor ni noitairaV .yrav did latsid pmulp a sah alutnede .raV .redrob suoir
htworgtoornostceffeerutarepmettey,snoitalu aes thgils a htiw esab a dna trap suorefinimes
pop neewteb noitairav on dewohs noitanimreg eht fo htgnel eht gnideecxe ro gnillauqe kaeb
no stceffe erutarepmeT .snoitidnoc rebmahc gnol a htiw ,ediw mm 5-3 ,tnemges tiurf lat
htworg mrofinu rednu ,eniltsaoc ainrofilaC fo sidrednels a sah dna alutnede .rav naht yhself
mk0501 ot pu ybdetarapes ,snoitalupop6 morf dna tsubor ssel yllareneg si sirtsucal .raV .alut
amitiram .Cfo slaudividni werg )b0791( RUOB nede .rav dna sirtsucal .rav :alutnede .pss alut
RAB .saera owteseht morfstnalpneewtebnoit nede.Cfoseiteiravowtdebircsed)2291(DLAN
cudorp tiurf ni ecnereffid on dnuof eH .snoitid REF dna )4791( NAMDOR .seiteiraV )iI(
noc esuohneerg mrofinu rednu snoitacol htob
morfslaudividniwerg)2891( YDDEK ,alutnede .)0891 ,NAMDOR( srucco noitaz
.Cfonoitalupop afosnoitropdrawaes dnadraw idirbyh )aniloraC htroNfo sknaB retuO eht(
dnal neewtebsecnereffid cipytoce rof tset oT seicepsbus owt eht neewteb yrtapmys fo enoz
.)6791 ,NAMDOR( stsaoc eht nI .)4791 ,NAMDOR( seiteirav otni axat
htob no drawhtron sesaercni smrof hsilprup eseht gnittilps stroppus osla )1 elbaT( sdees
fo noitroporp ehT .stsaoc cificaP dna citnaltA fo sisylana etalonisoculG .)4791 ,NAMDOR(
eht htob no snoitalupop alutnede .rav alut )100.0 < P( tnacifingis ylhgih si )nrm 8.32(
nede .C ni srewolf dna stiurf gnuoy ,smets no ireprah .pssalutnede .Cdna )mm 9.71(alutnede
noitaroloc elprup fo ecnedicni eht nialpxe yam .pss alutnede .C neewteb htgnel tiurf ni ecne
noitairav cipytoce ,revewoH .)6791 ,NAMDOR( reffid ehT .hteet on dna ecafrus gnitalucitra
noitairav cipytoce naht rehtar stceffe srednuof talf a htiw ,stnemges tiurf lamixorp dna latsid
ot detubirtta neeb evah alutnede .rav alutnede debbir-thgie sah ireprah .pss alutnede elikaC
.C fo snoitalupop tsaoc cificaP dna citnaltA .drawpugnitcejorp hteetllams )6(2 dna ecafrus
neewteb htgnel latep dna emit noitarutam gnitalucitra evacnoc ot talf ± a htiw ,stnemges
,htgnel tiurf ni secnereffiD .sepytocE )iii( tiurf lamixorp dna latsid delgna-ruof sah alut
nede.pssalutnedeelikaC.alutnedednaireprah
.sekaL ,seicepsbus owt sesirpmoc alutnede .C ,)4791(
taerG eht ro tsaoc citnaltA eht rehtie morf NAMDOR ot gnidroccA seicepsbuS )i(
alutnede .ravalutnede .Cfoesohtnaht)100.0 <
P(regnolyltnacifingiserasekaLtaerGehtmorf ytilibairaV )c(
noslOdna dyoB,nuaM 041
BiologyofCakileedentula 141
(iv) Chromosome numbers. 2n = 18 has variety is alsofound alongthe Pacificcoastand
been reported by KRUCKEBERG(1948),MUL less frequently on the shores of Lakes Michi
LIGAN (1964) and TAYLOR and MULLIGAN gan, Huron, Erie, and Ontario,but is not pres
(1968). entalong LakeSuperior.Of394herbariumcol
lections studied from the Great Lakes, only
III. GEOGRAPHICAL DISTRIBUTION 7.6% were clearly C. edentula var. edentula
(RODMAN, 1974).Its sporadic occurrence sug
In eastern North America, C. edentula ssp. gests it is a recent introduction, perhaps as a
edentula var. edentula ranges from the Outer ballastplantaftercompletion ofthe ErieCanal
Banks of North Carolina to Labrador (Figure in 1825 (RODMAN, 1974).Cakileedentula var.
2).Cakile edentula ssp.harperi occurs from the lacustrisisveryabundanton thesandybeaches
Outer Banks of North Carolina to southern ofall the GreatLakes ofNorthAmerica, except
Florida (AL-SHEHBAZ, 1985). Rare occur LakeSuperior,wherepopulationsarerare(Fig
rences ofCakile in an inland habitat have been ure 2). Occurrence is very sporadic on the St.
documented by GUIRE and VOSS (1963). Lawrence River east of Quebec. This variety
GEHUand GRANDTNER (1982)defined a sub does not occur along Atlantic or Pacific coasts.
association of Cakiletosum edentulae along the Cakileedentulavar.edentulawasnaturalized
shoreline of Magdalen Island, Quebec. The around 1880on thePacificcoastofNorthArner-
G V
I
H-
'\
I ;
I I
, I
/ \ I
r
I
. '. _.-----L-_~
/I \ I y--l ------
I I ---_1 ;
. I ;
I ! \ .
- - _J I I .0
, , I
I _
30
,
- .
e.dwtuLa
• var.
...var.Lacuihii
'0
Figure2. DistributionofCakileedentula varietiesedentulaandlacustrisin NorthAmerica. Fromspecimensin theherbariaof
BiosystematicResearch Institute,Ottawa;National MuseumofCanada;UniversityofWesternOntario, London;Universityof
California,Davis;personalobservationsandfromRodman (1974)andMortonandVenn(984).
Journal ofCoastalResearch,Vol.6, No.1,1990
0991,1.oN,6.loV,hcraeseRlatsaoCfo lanruoJ
ecnadnubaelikaCotlufgninaemeromerewsrot -aderp ,)2891 ,YDDEK( lasrepsid yb detaluger
caf latnemnorivneorcim taht dedulcnoc )6791 si alutnede elikaC fo tnetxe drawdnal ehT
,.la te RUOBRAB ;6791 ,XUAHCIBOR dna .hcaebehtfodrihtelddim eht nidekaepyllausu
RUOBRAB( seiduts lareveS .egnar sti dnetxe revocstitahtdewohsyehT.)2elbaT(noitategev
yam senilerohs gnola retaw fo ecneulfni gnita fo egde drawaes eht fo tnenopmoc tnacifingis a
redom ehT.)6791 ,.la te RUOBRAB( ecneulfni ti deredisnoc dna setisriehtfo%72 noalutnede
ni lanidutital naht rehtar lanoza sa deifissalc .C derevocsid )6791( .la te RUOBRAB ,)N°33
eb yam taht statibah ni)2 erugiF(citnaltA eht 74( setis hcaeb cificaP 43 no noitategev hcaeb
gnola 05 dna tsaoc cificaP eht gnola edutital fo yduts lacigolocenys a nI .)0891 ,EEL dna
N0
N°35tuobaot sdnetxealutnedeelikaC.noitarud KUICANGI( eporuE tsewhtron ni ytinummoc
shtnom lareves fo revoc wons a dna nosaes enil dnarts eht fo rebmem largetni na si hcihw
yrd on ,sretniw dimuh dna sremmus looc htiw ,amitiram.Cfo taht ot suogolana si noitisop sti
latnenitnoc dimuh sa deziretcarahc ,noiger hcus sA .senilerohs )sekaL taerG( retawhserf
citamilc bfD )1391( s'NEPPOK nihtiw seil dna eniram htob no seiceps dnarts reenoip
noitubirtsid s'alutnede .C fo trap rojam ehT a sa srucco alutnede .pss alutnede elikaC
stnemeriuqeR citamilC )c( ecnerruccOfo enoZ )a(
.noitubirtsidorcim
STATIBAH FO EGNAR .VI
elikaCecneulfniyamsuhtdna)0891 ,NOTLOH(
negortin fo ecruos tnatropmi na eb yam sevaw
yb sutirted fo noitisopeD .ylppus trohs ni erew
.)5891
)P ,N ,+K ,++gM( stneirtunorcam ruoF .yti
,ZABHEHS-LA( serozA eht dna ,)5891 ,ZAB
capacegnahcxenoitacroop sallew sanoitneter
HEHS-LA( dnalaeZ weN ,)2891 ,IASA( napaJ
erutsiom roop ni gnitluser ,thgiew yb %02.0
morf detroper neeb osla sah alutnede elikaC
dna 40.0 newteb yllareneg saw rettam cinagro
.)6891,NAM
etartsbuS .sllehs etarbetrevni retaw hserf dna
DOR(selaWhtuoSweN niyendyS otailartsuA
enotsemil fo stnemgarf ot eud etanobrac muic
tseW ,htreP morfsdnetxe yltnerructI .airotciV
lacfoslevelhgih dnuof)3891(NUAMdna NIW
dnaailartsuAhtuoS nialutnede.Cdecalpersah
DLAB,enilerohs noruHekaL ehtgnolA.)2591
amitiram elikaC fo noitcudortni retal A .ailart
,YRUBSILAS( slios yalc ni deniatbo saw amiti
suA nretseW otni dna ailartsuA nretsae ni
ram .C fo htworg roop dna lios deniard llew a
dnalsneeuQ otni daerps dah s'0791 eht yb dna
seriuqeralutnedeelikaC .tnemnorivneelbatsnu
,s'0681 eht ni enruobleM raen decudortni tsrif
rehtar a gnitaerc suht ,dnas eht fo gnitfihs
saw alutnede elikaC .)6891 ,4791 ,NAMDOR(
launitnoc sesuac sdniw dna sevaw hgih htob
ailartsuA ni derrucco sah,egnar tniojsuoiverp
yb ecnabrutsid etartsbuS.0.8 ot 5.6 tuoba morf
rieht fo trap ni alutnede .C fo ecnaraeppasid
segnar yllareneg Hp lioS .)1891,YDDEK;3891
dna amitiram .C yb noisavni yb dewollof ,alut
,NUAM dna NIWDLAB( elgnihs dna ,levarg
nede .C yb noisavni ylrae fo esac ralimis A
,dnas fo stsisnoc sworg elikaC erehw etarts
.)6891 ,DYOB ;0791
bus enilerohs eht fo lairetam tnerap ehT
,NAMDORdnaRUOBRAB(dnoyeb dna nogerO
lartnec ni ylno tnadnuba semoceb ti hguohtla
scitsiretcarahC etartsbuS )b( ,ainrofilaC ,akeruE fo htron amitiram .C htiw
yllacirtapmysworg otseunitnoctI.)0791 ,NAM
.retaw eht ot esolc oot gniworg DOR dna RUOBRAB( egnar sti fo mk 000,1
sgnildeesfo noitcurtsed eht detnemucod)3691( tsomnrehtuos eht morf deraeppasid sah alut
NOSNHOJ dna )2891( YDDEK .egde s'retaw nede .C ,alutnede .C fo egnar eht tuohguorht
eht ta ytilatrom gnildees hgih dna retaw eht daerps sti dna s'0391 eht ni ainrofilaC ot amit
morf yawa stiurf fo tnemevom hguorht tnem iram .C naeporuE fo noitcudortni eht gniwol
norivne lacisyhp eht yb detaluger si alutnede loF .1391 ni dnalsI kaidoK dna 1291 ni dnalsI
.C fo tnetxe )drawekal ro( drawaes ehT .)2891 revuocnaV no detroper saw dna ,htuos dna
,YDDEK ;b0791 ,RUOBRAB( noititepmoc htron yldipar daerps tI .)0791 ,NAMDOR dna
dna )b8891 ,DYOB ;b0791 ,RUOBRAB( noit RUOBRAB( ainrofilaC ,ocsicnarF naS raen aci
noslO dna dyoB,nuaM 241
Biology ofCakileedentula 143
Table2. Distributionpattern ofplantspecieson34beachesalongthePacificcoast(4r -33°N) showing thepercentageofbeaches
onwhichthespecieswas present in theleadingedgeandpercent coverofthespeciesontheseaward,middleandlandwardend
ofthebeach.AfterBARBOUR,DEJONGandJOHNSON (976).
Percentofbeaches
Percentageofbeachesshowingmaximumcoveron
onwhichthespecies
NameofSpecies wasrepresented Seaward 1/3 Mid 1/3 Landward 1/3
Abronialatifolia 14 24 24 38
Abroniamaritima 40 0 50 50
Ammophilaarenaria 31 0 16 72
A.breviligulata 40 0 100 0
Ambrosiachamissonis 21 6 53 35
Atriplexleucophylla 33 50 33 17
Camissoniacheiranthifolia 0 0 33 67
Cakileedentula 27 9 55 0
C.maritima 43 9 26 22
Calystegiasoldanella 0 11 11 67
Carexmacrocephala 0 0 100 0
Distichlisspicata 0 0 0 100
Elymusmollis 20 8 24 32
Fragariachiloensis 0 0 0 100
Honckenyapeploides 20 40 20 20
Lathyrusjaponicus 0 0 0 75
L.littoralis 0 0 0 100
Mesembryanthemumchilense 8 0 20 70
Poadouglasii 0 0 0 100
P.douglasiissp.macrantha 0 0 50 50
and distribution than latitudinal macrocli V. PLANT COMMUNITIES
matic gradients. The current southern limit of
C. edentula var. edentula on the Pacific coast AlongthePacificcoast,C.edentulawasfound
appears to be indirectly determined by climate growing with 19 other typical beach species
<BOYD,1986).InCalifornia,wherewintersare (Table 2),although at anyone beach generally
relatively mild, both C. edentula and C. mari five species comprised the vegetation.Only one
timacanliveintoa secondreproductiveseason. or two dominant species had any significant
Theproportionofplantssurvivingintoa second amount of cover, e.g., Ammophila arenaria
season in the open beach microhabitat is simi north of 380 latitude and Mesembryanthemum
larfor bothspecies,butmanymoreC. maritima chilense, Cakile maritima, Ambrosia chamis
plants flower and fruit during the second year sonis or Abronia maritima south ofthat lati
oflifein the foredune.As a result,reproductive tude (BARBOUR et al., 1976).
outputofC.maritimainthe foredune is greater RODMAN (1974) stated that Ammophila
than C. edentula by approximately two orders breviligulataandHudsoniatomentosa weretwo
of magnitude. Farther north, where neither common species associated with C.edentula on
species can overwinter, this advantage of C. the Atlantic. VAN ASDALL and OLMSTEAD
maritima does not occur and the two species (1963) found that three species of summer
coexist. annual dicots: Euphorbia polygonifolia L.,
Cakileedentulais notthreatenedwithextinc Cycloloma atriplicifolium (Spreng.) Coult., and
tion on either coast ofNorth America. Cakile Corispermum hyssopifolium L.were associated
edentula and C. maritima have been sympatric withC.edentulaon opendune sites along Lake
for some time in the northwest (the earliest Michigan. PAYNE (1980) reported Corisper
record of C. maritima in British Columbia is mum hyssopifolium, Ammophila breviligulata
1951), but C. edentula is still dominant there Fern., Calamovilfa longifolia (Hook.) Scribn.,
(BARBOUR and RODMAN, 1970). On the east OenotherabiennisL.,SalsolakaliL.,Xanthium
coast, C. maritima has been introduced in a strumariumL.,EuphorbiapolygonifoliaL.,and
number of localities (AHLES, 1951; RIEFNER, Artemisia campestris Michx. as major species
1982)but appears to be non-aggressive. occurringwith C. edentulavar. lacustrison the
JournalofCoastalResearch,Vol.6,No.1,1990
0991,1.oN,6.loV,hcraeseR latsaoCfolanruoJ
neewteb ssenihself ni ecnereffid eht fo trap )2791(RUOBRAB ,revewoH.snoitiddatneirtun
tsael tA.yarpstlas ot esnopser ni)stnalp tnare fo ecruos tnatropmi na eb dluoc sutirted taht
lot-yarps tlas fo lacipyt sevael fo gninekciht dna htworgtnalp detimil stneirtun taht dedulc
eht( yhportrepyh fo seerged ralimis ogrednu noc )0891 ,NOTLOH( ainrofilaC dna )3891
ot elba erew dna etyhpocylg lacipyt a naht ,.late EEL( eporuE ni amitiram .Cfo seidutS
retteb yarps tlas dna ytinilas lios detarelot .sutirted ni gni
seiteirav htob ,revewoH .alutnede .rav ton tub worg stnalp fo noitcudorp tiurf lanif rehgih ni
sirtsucal .rav fo tuptuo evitcudorper desaerced detluser nosaes eht ni retal sutirted morf yawa
yarps tlas deilppa yllaicifitrA .yarps tlas gniworg stnalp no srallipretac fo noitatsefni
etarelot ot ytiliba rieht ni degrevid evah yeht yvaeh a ,revewoH .)2891 ,YDDEK( nosaes
taht nwohs evah )6891( RUOBRAB dna DYOB eht ni ylrae esaesid ffo-gnipmad fo ycneuqerf
,ecnegrevid lacigolohprom sediseB .sraey retaerg eht ot eud trap ni ,tuptuo evitcudorper
000,01 tsal eht nihtiw noitalosi evitcudorper dna lavivrus rewol dah sutirted eht ni gniworg
gniwollof tnereffid yllacigolohprom emoceb stnalP .xelpmoc saw rab elgnihs eht no "srot
evah sirtsucal .rav alutnede .C dna alutnede aderp" lamina/lagnuf dna sutirted neewteb
.rav alutnede .C taht detamitse )4791( NAM yalpretni eht ,revewoH.statibahenuddnasdna
DOR .)8791 ,b0791 ,RUOBRAB( knar noitanoz rab elgnihstsaoc citnaltA ni alutnede .C fo tup
sti htiw llew dednopserroc hcihw ,level ecnare tuo evitcudorper hgih dna noitca evaw yb tfel
lot hgih ylriaf a evah ot dnuofsaw alutnede .C sutirted neewteb noitaicossa evitisop a dnuof
,axathcaebrehto ot evitaleR.xedni ecnarelot a osla )2891 ,0891( YDDEK.)1891 ,YDDEK( lort
otni setamitse ssamoib dna ygolohprom ,ytilat noc eht naht stuptuo evitcudorper rehgih htiw
rom denibmoc dna ,noitadnuni retawaes dna stnalp egral dedleiy alutnede.C fo stolp nworg
yarpstlasfoslevelcitsilaer ot alutnede.Ctsaoc dleif ot etartin muinomma fo noitiddA .stnalp
tsew fo esnopser eht detset )7791( GNOJ ED nworg-dleif ni htworg desaercni wohs )5891
dna RUOBRAB.yarpstlasfotnarelot yrev saw ,RELLEW( stnemirepxe gniretaw dna )3891
alutnede .Ctsaoc citnaltA tahtdevresbo)9591( ,ESORIH dna IHCAK( tneirtun ,tluser a sA
NITRAM .slevel yarps tlas hgih fo tnarelot .tnetnoc rettam cinagro wol dna erutxet esraoc
ylevitaler si elikaC dna ,statibah hcaeb ni rot rieht oteudretawelbaliava dnastneirtunfosle
caflatnemnorivnetnatropminasiyarpstlaS vel wol ylevitaler niatnoc setartsbus hcaeB
.secnereffid elbissop eseht
etarelot nac ti taht setacidni sirtsucal .rav ygoloisyhP )a(
fo tnemnorivne retawhserf eht ni worg ot alut
nede .ravfo ytiliba eht ,aedi siht tset yltcerid YGOLOCE LACIGOLOISYHP .IV
ot stpmetta on fo wonk ew hguohtlA .tnemnor
ivne retawhserf eht ni stneirtun rehto ro gM .ylppus tneirtun wol dna snoitid
foycneicifed a ebyam alutnede .rav alutnede.C noc thguord ,noitca evaw evitcurtsed ,tatibah
fo tatibah hcaeb eniram eht dna sirtsucal .rav fo ytilibatsni ,)stsaoc aes gnola( slevel ytinilas
alutnede .C fo tatibah retawhserf eht neewteb elbairav otgniwoseicepstnalpthgienahterom
sutats tneirtun ni ecnereffid enO .)2891 yb dezinoloc ylerar era senildnarts naciremA
,.la te AMEZOR( sevael eht yb ro etartsbus htroN .seiceps gnirrucco-oc tnadnuba tsom
eht morf rehtie eb yam ekatpU .yarps tlas aiv owt eht era .L arafraf ogalissuT dna .llE ).L(
sevael tnalp ro dnas hcaeb eniram ot deilppus aloolehselytsohportS,eirEekaLgnolasnoitacol
erastlasesehT.stnuomaegralyllufmrah nitne emostA.Lelameyh .Edna,.rB .Amutagiveal.E
serp eb yam gM dna aC ,lC,aN elihw snoitart ,.hcielhcS mutageirav mutesiuqE ,.L asocitiurf
necnoc wol ylevitaler ni era K ,P ,N hguohtla allitnetoP ,.L sirtsepmac aisimetrA ,.L alimup
)2791 ,NIETSPE( stneirtun tnalp laitnesse sunurP ,.dlliW sucitlab sucnuJ ,ssoV )nneS
lla sniatnoc retawaeS .)2791 ,LLEWNAR( & ttelliG( mulihpommasp noryporgA htiw
gnitimil ylnommoc tsom smees ,muissatop noitaicossa ni dnarts eht no dnuof osla erew
ro/dna surohpsohp htiw gnola ,negortiN .elba stnalp lanoisaccO .hcaeb mrots eht no dnas
irav eb ot sraeppa slios hcaeb ro enud ni noit elbatsnu ni gnirrucco ,)noruH ekaL nrehtron(
atimil tneirtun eht fo erutan ehT .hcaeb nepo dnalsIniluotinaMfoenilerohsdesopxeeht morf
eht no amitiram .C fo noitubirtsidorcim eht alutnede .C detroper )4891( NNEV dna NOT
dna negortin lios neewteb noitalerroc on dnuof ROM .noruH ekaL fo enilerohs nretsaehtuos
noslO dnadyoB,nuaM 441
BiologyofCahileedentula 145
field-collected var. edentula and var. lacustris approximating normal field conditions. Water
plants is due to hypertrophy of var. edentula use efficiencyofall the Csspecieswas less than
induced by exposure to salt spray. Leaves of Atriplex.
var.edentulaexposedto levelsofsaltspraysim
ilarto those found at the leading edge ofbeach (b) Phenology
vegetation were 2.3-fold as thick as water
sprayed leaves of var. lacustris (BOYD and In far northern latitudes C. edentula germi
BARBOUR, 1986). This difference decreased to natessoonafterthe groundis fullythawed, and
1.3-fold when both varieties were sprayed with occurs during a relatively short time. Both
water. Great Lakes and Atlantic populations germi
No water relations studies have been per nate over a period of 4-6 weeks (PAYNE and
formed on C. edentula, but DE JONG's (1979) MAUN, 1984; KEDDY, 1982). Observations at
study of California C. maritima revealed that PineryProvincialPark,Canadaoveraperiodof
plantssurvivedrainlesssummersby tolerating several years showed that the first emergence
higher xylem sap tensions than Ambrosia of C. edentula var. lacustris occurred in late
chamissonis or Abronia maritima seedlings. April on the upper undisturbed beach areas,
Drought, however, hasbeenconsideredto be an whereas lakeward seedlings emerged a few
important factor in C. maritima seedling mor days later. Observations by KEDDY (1982) on
tality (BARBOUR, 1970a). DE JONG (1979) the Atlantic coast also showed earlier germi
also reported lower xylem tensions in southern nation of landward seeds, which may have
California C. maritima relative to northern resulted from warmer sand temperatures.
California populations, which he attributed in Flowering begins in the end of July or begin
part to the greater amount and seasonal even ning of August in the Grand Bend region in
ness of precipitation to the north. BALDWIN southwestern Ontario, Canada. Most plants
and MAUN (1983) remarked that soil moisture continueto flower and fruit throughout August
content, especially atshallowdepths (1-10 em), and September,butoccasional flowering plants
was the most important discriminating factor may be found in October or November. Fruits
in the Great Lakes sand dune environment. mature within about three weeks.
Moisture was lowestin JuneandJuly, atwhich In the mediterranean-climate region of Cali
time a drastic reduction in the seedling popu fornia, flowers and fruits ofC.edentula may be
lations of C. edentula var. lacustris occurred. found almost any time ofyear.For example,at
PAYNE and MAUN (1984)estimated that 32% Eureka in northern California, C. edentula
of the plants in their population died from flowered from June to January (JOHNSON,
drought stress. 1963).NearSanFrancisco,somemembersofan
BARBOUR et al. (1985) compared photosyn artificially-established population in foredune
thetic rates and leafwater loss of C. maritima and open beach habitats survived into a second
under high and low soil nitrogen regimes, find reproductiveseason and flowered in late winter
ingthatplantsmaintainedthesamerate ofnet andspring(BOYD, 1986). However, thepeakof
photosynthesis under both nutrient regimes. flower and fruit production of both C. edentula
However, under low nitrogen conditions there and C. maritima was in the summer and fall.
was an increase in leafconductance and there Germination timing is also less well-defined
fore transpiration, resulting in a decline in an in California. JOHNSON (1963) observed seed
already low value for water use efficiency. lings from planted seeds in November, March
All four Cakile species investigated to date andAprilandfrom naturally-dispersedseedsin
possess the C; photosynthetic pathway (DE December. In an artificially-established open
JONG, 1978; BARBOUR et al., 1987), and beach colony of C. edentula, BOYD (1986)
therefore it seems likely that C. edentula does recorded seedling emergence from January to
aswell. DEJONG(1978)comparedC.maritima May, with a large peak in April. This germi
andtwo otherC, plantsto a C Atriplexspecies, nationpatterndifferedfromthatofC.maritima
4
finding that the C. species reached maximum at the same site, which peaked in January and
rates ofphotosynthesis at higher temperatures April with no May germination.
and light intensities, but the C species had RODMAN (1974) found that greenhouse or
s
higher photosynthetic rates at temperatures growth chamber-grown plants of all species he
JournalofCoastalResearch,Vol. 6,No.I,1990
0991.1 .oN.6.loV.hcraeseRlatsaoCfolanruoJ
hgih ro eniltfird eht gnola enoz a ni stnalp EEL( ycneicifed tneirtunorcam ro/dna)4891
ynam )2( ;retnec eht morf ecnatsid gnisaercni ,NUAM dna ENYAP( sserts thguord rehgih
htiw yldipar denilced ytisned sretsulc hcus yb desuac eb yam hcaeb hgih eht no ytilatroM
nI.raeysuoiverpehtfostnalpegral yb deipucco .stnalp hcaebdim rof %05 naht ssel ot derap
setis ta gnirrucco retemaid nre 06 ot 8 spmulc moc sa ytilatrom %57 tuoba dereffus stnalp
ni stnalp deretsulc ylesned )1( :sgnipuorg hcaeb hgih dna eniltfirD .setar ytilatrom ni
fo sepyt eerht fo detsisnoc tsaoc citnaltA eht secnereffiddesuacoslatatibahorcim nisecneref
gnola )1891,YDDEK( alutnede.rav dna noruH fiD .ytilatrom elbaredisnoc desuac sgnildees
ekaL no)0891,ENYAP( sirtsucal .rav alutnede fo lairub peed ni detluser taht mrots gnirps
.CfogninrettaplaitapS.remmusotgnirpsmorf a tub raey gniwollof eht rewol erew setar yti
neve dna ,raey ot raey morf seirav alutnede latroM.llafniarwol ylemertxefoesuaceb8791
.Cfonoitubirtsid ehttaht deton)0891( ENYAP gnirudsirtsucal .rav alutnede.C rofsetarytilat
.tsaoc ainrofilaC eht gnola yllacol stekcor rom hgih detroper )4891( NUAM dna ENYAP
aes fo ecnaraeppasid dna ecnaraeppa gnitaut ,elpmaxe roF.)2891,YDDEK;4891,NUAMdna
culf detroper )b0791( RUOBRAB .laremehpe ENYAP(raey ot raey morfylediw yrav otytilat
ylhgih eb nac alutnede .C fo snoitalupoP rom esuac yam srotcaf citoiba dna citoib rehto
.spmulc regral ni thguordfo noit dnaezisnoitalupop,ytilibatciderpnu tatibaH
cnufasassertsytisnedretaergoteudylbaborp
tluser a ,stnalp derettacs fo gnitsisnoc slort scimanyD noitalupoP )b(
noc tnecajda naht etar ytilatrom naem rehgih
yltnacifingisadahspmulcegral ylno,revoeroM .)6891
.tnereffid yllacitsitats ton erew setar eht ,reve ,DYOB( tatibah enuderof eht ni nosaes evitcud
woh;ezispmulc laitini gnisaercnihtiwetaryti orperdnocesa otnignivivrusslaudividni wefa
latrom ni esaercni na dewohs)0891 ,ENYAP( htiw,lainnerep evitatlucaf a sastcati setamilc
alutnede .Cfospmulc rof sevrucpihsrovivruS redlim nI.sdeessasnoitidnocelbarovafnuseviv
.dne drawaes eht ta noit rus dna sedutital nrehtron ni nosaes gniworg
cudorp tiurf fo lortnoc tnedneped-ytisned yb eno nihtiw elcyc efil sti setelpmoc taht)etyhpo
detaluger saw)1891,YDDEK morf atad( hcaeb reht( launna tneluccus a si alutnede elikaC
euqinitraM no ecnadnuba elikaC taht dewohs
)5891( NOSNIKTAW yb enod gnilledoM .dne noitannereP )a(
drawaes eht ta tsetaerg erew tuptuo evitcud
orper no ytisned fo tceffe eht elihw ,tneidarg YGOLOIB NOITALUPOP .IIV
eht fo dne drawdnal eht ta tsetaerg erew pihs
rovivrus no ytisned fo stceffe )3( dna ,tuptuo .amitiram .C fo deeps evitcudorper
evitcudorper decneulfni yltnacifingis stceffe eht dehgiewtuo ,sesac emos ni ,taht srotcaf
tnedneped-ytisned dna tnednepedni-ytisned rehtoynam ybdetceffasawtuptuoevitcudorper
htob)2(;tnatsnocdeyatsytilatromtnednepedni latot taht tub ,ainrofilaC ,seyeR tnioP ta snoit
ytisned saerehw tneidarg eht gnola yltnaci alupop nworg-dleif ni alutnede.C naht reilrae
fingis degnahcpihsrovivrusfoecnedneped-ytis derewolf amitiram .C taht dnuof )6891( DYOB
ned )1( :taht dnuof eH .hcaeb ydnas a fo sdne .snoitidnoc rebmahc htworg rednu devresbo
drawdnal ot drawaes morf gnihcterts tneidarg amitiram .C fo emit noitarutam retsaf eht
latnemnorivne eht no alutnede .rav alutnede ot eud neeb evah yam ailartsuA dna ainrofilaC
.Cfo yhpargomed eht deiduts)1891( YDDEK ni amitiram .C yb alutnede .C fo tnem
.)2891 ,YDDEK( lahtel eb yam ecalper lacirotsih taht detseggus )6891( NAM
sehcaebeniramnonoitadnuniralimiS.thguord DOR .alutnede .C tsaoc cificaP rof tceffe s'red
gnirud retteb worg dna evivrus yltneuq nuof a ot eud saw ro ecnacifingis evitpada dah
esnoc dna sevaw hgih yb detadnuni eb yam ecnereffid siht fi enimreted ot elbanu saw eH
yllanoisacco stnalp hcaeb wol ,)retaw hserf{ .snoitalupoptsaoccificaP rofsyad 33 ottsartnoc
noruH ekaL gnola remmus gniruD .alutnede ni ,syad 44 fo egareva na ni derutam snoitalu
.C fo setar htworg desaerced dna ytilatrom poptsaoccitnaltA,alutnede.CroF.srehtonaht
gnildees ot ylivaeh setubirtnoc laitnetop retaw setar noitarutam retsaf ylbaeciton dah seiceps
lios wol enuderof eht no taht dewohs )a689l( rosnoitalupopemostub,noitanimregfoshtnom
.la te LLADNYT .)5891 ,KUICANGI dna owt nihtiw detiurf dna derewolf detagitsevni
noslOdnadyoB,nuaM 641