Carnivorous Plant

9/15/2015
CarnivorousplantWikipedia,thefreeencyclopedia
Carnivorousplant
FromWikipedia,thefreeencyclopedia
"InsectivorousPlants"redirectshere.ForthebookbyCharlesDarwin,seeInsectivorousPlants
(book).
Carnivorousplantsareplantsthat
derivesomeormostoftheirnutrients
(butnotenergy)fromtrappingand
consuminganimalsorprotozoans,
typicallyinsectsandotherarthropods.
Carnivorousplantshaveadaptedto
growinplaceswherethesoilisthinor
poorinnutrients,especiallynitrogen,
suchasacidicbogsandrock
outcroppings.CharlesDarwinwrote
InsectivorousPlants,thefirstwell
knowntreatiseoncarnivorousplants,in
1875.[4]
Truecarnivoryisthoughttohave
AnupperpitcherofNepentheslowii,atropicalpitcherplantthat
evolvedindependentlysixtimesinfive
supplementsitscarnivorousdietwithtreeshrewdroppings. [1][2][3]
differentordersoffloweringplants,[5][6]
andthesearenowrepresentedbymore
thanadozengenera.Theseincludeabout630speciesthatattractandtrapprey,producedigestiveenzymes,
andabsorbtheresultingavailablenutrients.[7]Additionally,over300protocarnivorousplantspeciesin
severalgenerashowsomebutnotallofthesecharacteristics.
Contents
1Trappingmechanisms
1.1Pitfalltraps
1.2Flypapertraps
1.3Snaptraps
1.4Bladdertraps
1.5Lobsterpottraps
1.6Combinationtraps
1.7Borderlinecarnivores
2Evolution
3Ecologyandmodellingofcarnivory
4Classification
4.1Dicots
4.2Monocots
5Cultivation
6Medicinaluses
7Culturaldepictions
8Seealso
https://en.wikipedia.org/wiki/Carnivorous_plant
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8Seealso
9References
10Furtherreading
Trappingmechanisms
Fivebasictrappingmechanismsarefoundincarnivorousplants.
1. Pitfalltraps(pitcherplants)trappreyinarolledleafthat
containsapoolofdigestiveenzymesorbacteria.
2. Flypapertrapsuseastickymucilage.
3. Snaptrapsutilizerapidleafmovements.
4. Bladdertrapssuckinpreywithabladderthatgeneratesan
internalvacuum.
5. Lobsterpottrapsforcepreytomovetowardsadigestive
organwithinwardpointinghairs.
Thesetrapsmaybeactiveorpassive,dependingonwhether
movementaidsthecaptureofprey.Forexample,Triphyophyllumis
apassiveflypaperthatsecretesmucilage,butwhoseleavesdonot
growormoveinresponsetopreycapture.Meanwhile,sundewsare
activeflypapertrapswhoseleavesundergorapidacidgrowth,which
isanexpansionofindividualcellsasopposedtocelldivision.The
rapidacidgrowthallowsthesundewtentaclestobend,aidinginthe
retentionanddigestionofprey.[8]
ThesundewspeciesDroseraglanduligeraemploysaunique
trappingmechanismwithfeaturesofbothflypaperandsnaptraps
thishasbeentermedacatapultflypapertrap.[9]
ThepitchersofHeliamphora
chimantensisareanexampleofpitfall
traps.
Pitfalltraps
Mainarticle:Pitcherplant
Pitfalltrapsarethoughttohaveevolvedindependentlyonatleastfouroccasions.Ingeneraltheyare
phytotelmata,waterbodiescollectedorsecretedintospecialisedcontainers,andultimatelyheldbyplants
forvariousfunctionssuchasinparticular,thetrappinganddigestionofprey.Thesimplestonesare
probablythoseofHeliamphora,themarshpitcherplant.Inthisgenus,thetrapsareclearlyderived
evolutionarilyfromasimplerolledleafwhosemarginshavesealedtogether.Theseplantsliveinareasof
highrainfallinSouthAmericasuchasMountRoraimaandconsequentlyhaveaproblemensuringtheir
pitchersdonotoverflow.Tocounteractthisproblem,naturalselectionhasfavouredtheevolutionofan
overflowsimilartothatofabathroomsinkasmallgapinthezippedupleafmarginsallowsexcesswater
toflowoutofthepitcher.
HeliamphoraisamemberoftheSarraceniaceae,aNewWorldfamilyintheorderEricales(heathersand
allies).HeliamphoraislimitedtoSouthAmerica,butthefamilycontainstwoothergenera,Sarraceniaand
Darlingtonia,whichareendemictotheSoutheasternUnitedStates(withtheexceptionofonespecies)and
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Californiarespectively.Sarraceniapurpureasubsp.purpurea(thenorthernpitcherplant)canbefoundas
farnorthasCanada.Sarraceniaisthepitcherplantgenusmostcommonlyencounteredincultivation,
becauseitisrelativelyhardyandeasytogrow.
InthegenusSarracenia,theproblemofpitcheroverflowissolved
byanoperculum,whichisessentiallyaflaredleafletthatcoversthe
openingoftherolledleaftubeandprotectsitfromrain.Possibly
becauseofthisimprovedwaterproofing,Sarraceniaspeciessecrete
enzymessuchasproteasesandphosphatasesintothedigestivefluid
atthebottomofthepitcherHeliamphorareliesonbacterial
digestionalone.Theenzymesdigesttheproteinsandnucleicacids
intheprey,releasingaminoacidsandphosphateions,whichthe
plantabsorbs.
Darlingtoniacalifornica:notethe
smallentrancetothetrapunderneath
theswollen"balloon"andthe
colourlesspatchesthatconfuseprey
trappedinside.
Darlingtoniacalifornica,thecobraplant,possessesanadaptation
alsofoundinSarraceniapsittacinaand,toalesserextent,in
Sarraceniaminor:theoperculumisballoonlikeandalmostseals
theopeningtothetube.Thisballoonlikechamberispittedwith
areolae,chlorophyllfreepatchesthroughwhichlightcanpenetrate.
Insects,mostlyants,enterthechamberviatheopeningunderneath
theballoon.Onceinside,theytirethemselvestryingtoescapefromthesefalseexits,untiltheyeventually
fallintothetube.Preyaccessisincreasedbythe"fishtails",outgrowthsoftheoperculumthatgivethe
plantitsname.SomeseedlingSarraceniaspeciesalsohavelong,overhangingopercularoutgrowths
Darlingtoniamaythereforerepresentanexampleofneoteny.
Thesecondmajorgroupofpitcherplantsarethemonkeycupsor
tropicalpitcherplantsofthegenusNepenthes.Inthehundredorso
speciesofthisgenus,thepitcherisborneattheendofatendril,
whichgrowsasanextensiontothemidriboftheleaf.Mostspecies
catchinsects,althoughthelargerones,suchasNepenthesrajah,
alsooccasionallytakesmallmammalsandreptiles.Nepenthes
bicalcaratapossessestwosharpthornsthatprojectfromthebaseof
theoperculumovertheentrancetothepitcher.Theselikelyserveto
lureinsectsintoaprecariouspositionoverthepitchermouth,where
theymaylosetheirfootingandfallintothefluidwithin.[10]
Thepitfalltraphasevolvedindependentlyinatleasttwoother
groups.TheAlbanypitcherplantCephalotusfollicularisisasmall
pitcherplantfromWesternAustralia,withmoccasinlikepitchers.
Therimofitspitcher'sopening(theperistome)isparticularly
pronounced(bothsecretenectar)andprovidesathornyoverhangto
theopening,preventingtrappedinsectsfromclimbingout.The
liningofmostpitcherplantsiscoveredinaloosecoatingofwaxy
flakes,whichareslipperyforinsects,preythatareoftenattractedby
Brocchiniareducta:acarnivorous
nectarbribessecretedbytheperistomeandbybrightflowerlike
bromeliad.
anthocyaninpatterning.Inatleastonespecies,Sarraceniaflava,the
nectarbribeislacedwithconiine,atoxicalkaloidalsofoundin
hemlock,whichprobablyincreasestheefficiencyofthetrapsbyintoxicatingprey.[11]
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ThefinalcarnivorewithapitfallliketrapisthebromeliadBrocchiniareducta.Likemostrelativesofthe
pineapple,thetightlypacked,waxyleafbasesofthestraplikeleavesofthisspeciesformanurn.Inmost
bromeliads,watercollectsreadilyinthisurnandmayprovidehabitatsforfrogs,insectsand,moreuseful
fortheplant,diazotrophic(nitrogenfixing)bacteria.InBrocchinia,theurnisaspecialisedinsecttrap,with
aloose,waxyliningandapopulationofdigestivebacteria.
Flypapertraps
Pinguiculagiganteawithprey.The
insectwastoolargeandwasstrong
enoughtoescapefromthesticky
surface.
Theflypapertrapisbasedon
astickymucilage,orglue.
Theleafofflypapertrapsis
studdedwithmucilage
secretingglands,whichmay
beshort(likethoseofthe
butterworts),orlongand
mobile(likethoseofmany
sundews).Flypapershave
evolvedindependentlyat
leastfivetimes.
InthegenusPinguicula,the
mucilageglandsarequite
short(sessile),andtheleaf,whileshiny(givingthegenusits
commonnameof'butterwort'),doesnotappearcarnivorous.
However,thisbeliesthefactthattheleafisanextremelyeffective
trapofsmallflyinginsects(suchasfungusgnats),anditssurface
respondstopreybyrelativelyrapidgrowth.Thisthigmotropic
growthmayinvolverollingoftheleafblade(topreventrainfrom
splashingthepreyofftheleafsurface)ordishingofthesurface
underthepreytoformashallowdigestivepit.
TheleafofaDroseracapensis
bendinginresponsetothetrappingof
aninsect.
Thesundewgenus(Drosera)consistsofover100speciesofactiveflypaperswhosemucilageglandsare
borneattheendoflongtentacles,whichfrequentlygrowfastenoughinresponsetoprey(thigmotropism)
toaidthetrappingprocess.ThetentaclesofD.burmaniicanbend180inaminuteorso.Sundewsare
extremelycosmopolitanandarefoundonallthecontinentsexcepttheAntarcticmainland.Theyaremost
diverseinAustralia,thehometothelargesubgroupofpygmysundewssuchasD.pygmaeaandtoa
numberoftuberoussundewssuchasD.peltata,whichformtubersthataestivateduringthedrysummer
months.Thesespeciesaresodependentoninsectsourcesofnitrogenthattheygenerallylacktheenzyme
nitratereductase,whichmostplantsrequiretoassimilatesoilbornenitrateintoorganicforms.
CloselyrelatedtoDroseraisthePortuguesedewypine,Drosophyllum,whichdiffersfromthesundewsin
beingpassive.Itsleavesareincapableofrapidmovementorgrowth.Unrelated,butsimilarinhabit,arethe
Australianrainbowplants(Byblis).Drosophyllumisunusualinthatitgrowsunderneardesertconditions
almostallothercarnivoresareeitherbogplantsorgrowinmoisttropicalareas.
Recentmoleculardata(particularlytheproductionofplumbagin)indicatethattheremainingflypaper,
Triphyophyllumpeltatum,amemberoftheDioncophyllaceae,iscloselyrelatedtoDrosophyllumandforms
partofalargercladeofcarnivorousandnoncarnivorousplantswiththeDroseraceae,Nepenthaceae,
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AncistrocladaceaeandPlumbaginaceae.Thisplantisusually
encounteredasaliana,butinitsjuvenilephase,theplantis
carnivorous.Thismayberelatedtoarequirementforspecific
nutrientsforflowering.
Snaptraps
Theonlytwoactivesnap
trapstheVenusflytrap
(Dionaeamuscipula)andthe
waterwheelplant
(Aldrovandavesiculosa)
arebelievedtohavehadacommonancestorwithsimilar
adaptations.Theirtrappingmechanismhasalsobeendescribedasa
"mousetrap","beartrap"or"mantrap",basedontheirshapeand
rapidmovement.However,thetermsnaptrapispreferredasother
designationsaremisleading,particularlywithrespecttotheintended
prey.Aldrovandaisaquaticandspecialisedincatchingsmall
invertebratesDionaeaisterrestrialandcatchesavarietyof
arthropods,includingspiders.[12]
Droseracapensisrespondingto
capturedprey.Thissceneisabout4
hoursinrealtime.
ThesnaptrapsofDionaeamuscipula
Thetrapsareverysimilar,withleaveswhoseterminalsectionis
closerapidlywhenthesensitivehairs
dividedintotwolobes,hingedalongthemidrib.Triggerhairs(three
ontheleaflobesaretriggered.
oneachlobeinDionaeamuscipula,manymoreinthecaseof
Aldrovanda)insidethetraplobesaresensitivetotouch.Whena
triggerhairisbent,stretchgatedionchannelsinthemembranesofcellsatthebaseofthetriggerhairopen,
generatinganactionpotentialthatpropagatestocellsinthemidrib.[13]Thesecellsrespondbypumpingout
ions,whichmayeithercausewatertofollowbyosmosis(collapsingthecellsinthemidrib)orcauserapid
acidgrowth.[14]Themechanismisstilldebated,butinanycase,changesintheshapeofcellsinthemidrib
allowthelobes,heldundertension,tosnapshut,[13]flippingrapidlyfromconvextoconcave[15]and
interringtheprey.Thiswholeprocesstakeslessthanasecond.IntheVenusflytrap,closureinresponseto
raindropsandblownindebrisispreventedbytheleaveshavingasimplememory:forthelobestoshut,two
stimuliarerequired,0.5to30secondsapart.[16][17]
Thesnappingoftheleavesisacaseofthigmonasty(undirectedmovementinresponsetotouch).Further
stimulationofthelobe'sinternalsurfacesbythestrugglinginsectscausesthelobestocloseeventighter
(thigmotropism),sealingthelobeshermeticallyandformingastomachinwhichdigestionoccursovera
periodofonetotwoweeks.Leavescanbereusedthreeorfourtimesbeforetheybecomeunresponsiveto
stimulation,dependingonthegrowingconditions.
Bladdertraps
BladdertrapsareexclusivetothegenusUtricularia,orbladderworts.Thebladders(vesicula)pumpions
outoftheirinteriors.Waterfollowsbyosmosis,generatingapartialvacuuminsidethebladder.Thebladder
hasasmallopening,sealedbyahingeddoor.Inaquaticspecies,thedoorhasapairoflongtriggerhairs.
AquaticinvertebratessuchasDaphniatouchthesehairsanddeformthedoorbyleveraction,releasingthe
vacuum.Theinvertebrateissuckedintothebladder,whereitisdigested.ManyspeciesofUtricularia(such
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asU.sandersonii)areterrestrial,growingonwaterloggedsoil,andtheirtrappingmechanismistriggeredin
aslightlydifferentmanner.Bladderwortslackroots,butterrestrialspecieshaveanchoringstemsthat
resembleroots.Temperateaquaticbladderwortsgenerallydiebacktoarestingturionduringthewinter
months,andU.macrorhizaappearstoregulatethenumberofbladdersitbearsinresponsetotheprevailing
nutrientcontentofitshabitat.
Lobsterpottraps
Alobsterpottrapisa
chamberthatiseasytoenter,
andwhoseexitiseither
difficulttofindorobstructed
byinwardpointingbristles.
Lobsterpotsarethetrapping
mechanisminGenlisea,the
corkscrewplants.These
plantsappeartospecialisein
ThetipofonestolonofUtricularia
aquaticprotozoa.AY
vulgaris,showingstolon,branching
shapedmodifiedleafallows
leafshoots,andtransparentbladder
preytoenterbutnotexit.
traps
Inwardpointinghairsforce
Genliseaviolaceatrapsandleaves
thepreytomoveina
particulardirection.Preyenteringthespiralentrancethatcoils
aroundtheuppertwoarmsoftheYareforcedtomoveinexorablytowardsastomachinthelowerarmof
theY,wheretheyaredigested.Preymovementisalsothoughttobeencouragedbywatermovement
throughthetrap,producedinasimilarwaytothevacuuminbladdertraps,andprobablyevolutionarily
relatedtoit.
OutsideofGenlisea,featuresreminiscentoflobsterpottrapscanbeseeninSarraceniapsittacina,
Darlingtoniacalifornica,and,somehorticulturalistsargue,Nepenthesaristolochioides.
Combinationtraps
ThetrappingmechanismofthesundewDroseraglanduligeracombinesfeaturesofbothflypaperandsnap
trapsithasbeentermedacatapultflypapertrap.[18]
Borderlinecarnivores
Mainarticle:Protocarnivorousplant
Tobeafullyfledgedcarnivore,aplantmustattract,kill,anddigestprey[5][19]anditmustbenefitfrom
absorbingtheproductsofthedigestion(mostlyaminoacidsandammoniumions).[20]Tomany
horticulturalists,thesedistinctionsareamatteroftaste.Thereisaspectrumofcarnivoryfoundinplants:
fromcompletelynoncarnivorousplantslikecabbages,toborderlinecarnivores,tounspecialisedand
simpletraps,likeHeliamphora,toextremelyspecialisedandcomplextraps,likethatoftheVenusflytrap.
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TheborderlinecarnivoresincludeRoridulaandCatopsisberteroniana.Catopsisisaborderlinecarnivorous
bromeliad,likeBrocchiniareducta.However,unlikeB.reducta,which
producestheenzymephosphatase,C.berteronianahasnotbeenshown
toproduceanydigestiveenzymesatall.[21]Inthesepitfalltraps,prey
simplyfallintotheurn,assistedbythewaxyscaleslocatedontherim.
Roridulahasamoreintricaterelationshipwithitsprey.Theplantsin
thisgenusproducestickyleaveswithresintippedglandsandlook
extremelysimilartosomeofthelargersundews.However,theydonot
directlybenefitfromtheinsectstheycatch.Instead,theyforma
mutualisticsymbiosiswithspeciesofassassinbug(genusPameridea),
Roridulagorgonias:aborderline
whicheatthetrappedinsects.Theplantbenefitsfromthenutrientsin
carnivorethatgainsnutrientsfrom
thebugs'faeces.[22]
its"prey"viathedroppingsofa
predatorybug
AnumberofspeciesintheMartyniaceae(previouslyPedaliaceae),
suchasIbicellalutea,havestickyleavesthattrapinsects.However,
theseplantshavenotbeenshownconclusivelytobecarnivorous.[23]Likewise,theseedsofShepherd's
Purse,[23]urnsofPaepalanthusbromelioides,[24]bractsofPassiflorafoetida,[25]andflowerstalksand
sepalsoftriggerplants(Stylidium)[26]appeartotrapandkillinsects,buttheirclassificationascarnivoresis
contentious.
Theproductionofspecificpreydigestingenzymes(proteases,ribonucleases,phosphatases,etc.)is
sometimesusedasacriterionforcarnivory.However,thiswouldprobablydiscountHeliamphora[27]and
Darlingtonia,[28]bothofwhichappeartorelyontheenzymesofsymbioticbacteriatobreakdowntheir
preybutaregenerallyconsideredascarnivores.However,discountingtheenzymebaseddefinitionleaves
openthequestionofRoridula.
Evolution
Theevolutionofcarnivorousplantsisobscuredbythepaucityoftheirfossilrecord.Veryfewfossilshave
beenfound,andthenusuallyonlyasseedorpollen.Carnivorousplantsaregenerallyherbs,andtheirtraps
areproducedbyprimarygrowth.Theygenerallydonotformreadilyfossilisablestructuressuchasthick
barkorwood.Thetrapsthemselveswouldprobablynotbepreservedinanycase.
Still,muchcanbededucedfromthestructureofcurrenttraps.Pitfalltrapsarequiteclearlyderivedfrom
rolledleaves.ThevasculartissuesofSarraceniaisacaseinpoint.Thekeelalongthefrontofthetrap
containsamixtureofleftwardandrightwardfacingvascularbundles,aswouldbepredictedfromthe
fusionoftheedgesofanadaxial(stemfacing)leafsurface.Flypapersalsoshowasimpleevolutionary
gradientfromsticky,noncarnivorousleaves,throughpassiveflypaperstoactiveforms.Moleculardata
showtheDionaeaAldrovandacladeiscloselyrelatedtoDrosera,[29]butthetrapsaresodissimilarthatthe
theoryoftheiroriginveryfastmovingflypapersbecamelessreliantonglueremainsratherspeculative.
Thereareoveraquarterofamillionspeciesoffloweringplants.Ofthese,onlyaround630areknowntobe
carnivorous.Truecarnivoryhasprobablyevolvedindependentlyatleastsixtimes[5]however,someof
these"independent"groupsprobablydescendedfromarecentcommonancestorwithapredispositionto
carnivory.Somegroups(theEricalesandCaryophyllales)seemparticularlyfertilegroundforcarnivorous
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preadaptation,althoughintheformercase,thismaybemoretodo
withtheecologyofthegroupthanitsmorphology,asmostofthe
membersofthisgroupgrowinlownutrienthabitatssuchasheath
andbog.
Ithasbeensuggestedthatalltraptypesaremodificationsofa
similarbasicstructurethehairyleaf.[30]Hairy(ormore
specifically,stalkedglandular)leavescancatchandretaindropsof
rainwater,especiallyifshieldshapedorpeltate,thuspromoting
bacteriagrowth.Insectslandontheleaf,becomemiredbythe
surfacetensionofthewater,andsuffocate.Bacteriajumpstart
decay,releasingfromthecorpsenutrientsthattheplantcanabsorb
throughitsleaves.Thisfoliarfeedingcanbeobservedinmostnon
carnivorousplants.Plantsthatwerebetteratretaininginsectsor
waterthereforehadaselectiveadvantage.Rainwatercanberetained
bycuppingtheleaf,leadingtopitfalltraps.Alternatively,insects
canberetainedbymakingtheleafstickierbytheproductionof
mucilage,leadingtoflypapertraps.
Thepitfalltrapsmayhaveevolvedsimplybyselectionpressurefor
theproductionofmoredeeplycuppedleaves,followedby"zipping
up"ofthemarginsandsubsequentlossofmostofthehairs,except
atthebottom,wheretheyhelpretainprey.
Artist'srestorationofArchaeamphora
longicervia,theearliestknown
carnivorousplant
ThelobsterpottrapsofGenliseaaredifficulttointerpret.Theymayhavedevelopedfrombifurcated
pitchersthatlaterspecialisedongrounddwellingpreyor,perhaps,thepreyguidingprotrusionsofbladder
trapsbecamemoresubstantialthanthenetlikefunnelfoundinmostaquaticbladderworts.Whatevertheir
origin,thehelicalshapeofthelobsterpotisanadaptationthatdisplaysasmuchtrappingsurfaceaspossible
inalldirectionswhenburiedinmoss.
Thetrapsofthebladderwortsmayhavederivedfrompitchersthatspecialisedinaquaticpreywhenflooded,
likeSarraceniapsittacinadoestoday.Escapingpreyinterrestrialpitchershavetoclimborflyoutofatrap,
andbothofthesecanbepreventedbywax,gravityandnarrowtubes.However,afloodedtrapcanbeswum
outof,soinUtricularia,aonewaylidmayhavedevelopedtoformthedoorofaprotobladder.Later,this
mayhavebecomeactivebytheevolutionofapartialvacuuminsidethebladder,trippedbypreybrushing
againsttriggerhairsonthedoorofthebladder.
FlypapertrapsincludethevarioustrueflypapersandthesnaptrapsofAldrovandaandDionaea.The
productionofstickymucilageisfoundinmanynoncarnivorousgenera,andthepassivegluetrapsinByblis
andDrosophyllumcouldeasilyhaveevolved.
Theactivegluetrapsuserapidplantmovementstotraptheirprey.Rapidplantmovementcanresultfrom
actualgrowth,orfromrapidchangesincellturgor,whichallowcellstoexpandorcontractbyquickly
alteringtheirwatercontent.SlowmovingflypaperslikePinguiculaexploitgrowth,buttheVenusflytrap
usessuchrapidturgorchangesthatgluebecameunnecessary.Thestalkedglandsthatoncemadeitand
whicharesoevidentinDroserahavebecometheteethandtriggerhairsanexampleofnaturalselection
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hijackingpreexistingstructuresfornewfunctions.Also,thenerve
likesensorysystemoftheVenusFlytrapevolvedabout135million
yearsagointheCretaceousperiod.Ithasanactivesteeltrap,like
thatofAldrovanda,thewaterwheelplant.[31]
Recenttaxonomicanalysis[32]oftherelationshipswithinthe
CaryophyllalesindicatethattheDroseraceae,Triphyophyllum,
NepenthaceaeandDrosophyllum,whilecloselyrelated,are
embeddedwithinalargercladethatincludesnoncarnivorous
groupssuchasthetamarisks,Ancistrocladaceae,Polygonaceaeand
Plumbaginaceae.Interestingly,thetamariskspossessspecialised
saltexcretingglandsontheirleaves,asdoseveralofthe
Plumbaginaceae(suchasthesealavender,Limonium),whichmay
havebeencooptedfortheexcretionofotherchemicals,suchas
proteasesandmucilage.SomeofthePlumbaginaceae(e.g.
Ceratostigma)alsohavestalked,vascularisedglandsthatsecrete
mucilageontheircalycesandaidinseeddispersalandpossiblyin
protectingtheflowersfromcrawlingparasiticinsects.Theseare
probablyhomologouswiththetentaclesofthecarnivorousgenera.
Perhapscarnivoryevolvedfromaprotectivefunction,ratherthana
nutritionalone.Thebalsams(suchasImpatiens),whichareclosely
relatedtotheSarraceniaceaeandRoridula,similarlypossessstalked
glands.
ThetrapsofCatopsisberteroniana
areunlikelytohavedescendedfroma
hairyleaforsepal.
Theonlytrapsthatareunlikelytohavedescendedfromahairyleaforsepalarethecarnivorousbromeliads
(BrocchiniaandCatopsis).Theseplantsusetheurnafundamentalpartofabromeliadforanew
purposeandbuildonitbytheproductionofwaxandtheotherparaphernaliaofcarnivory.
Ecologyandmodellingofcarnivory
Carnivorousplantsarewidespreadbutratherrare.Theyarealmostentirelyrestrictedtohabitatssuchas
bogs,wheresoilnutrientsareextremelylimiting,butwheresunlightandwaterarereadilyavailable.Only
undersuchextremeconditionsiscarnivoryfavouredtoanextentthatmakestheadaptationsobvious.
Thearchetypalcarnivore,theVenusflytrap,growsinsoilswithalmostimmeasurablenitrateandcalcium
levels.Plantsneednitrogenforproteinsynthesis,calciumforcellwallstiffening,phosphatefornucleicacid
synthesis,andironforchlorophyllsynthesis.Thesoilisoftenwaterlogged,whichfavourstheproductionof
toxicionssuchasammonium,anditspHisanacidic4to5.Ammoniumcanbeusedasasourceof
nitrogenbyplants,butitshightoxicitymeansthatconcentrationshighenoughtofertilisearealsohigh
enoughtocausedamage.
However,thehabitatiswarm,sunny,constantlymoist,andtheplantexperiencesrelativelylittle
competitionfromlowgrowingSphagnummoss.Still,carnivoresarealsofoundinveryatypicalhabitats.
DrosophyllumlusitanicumisfoundarounddesertedgesandPinguiculavalisneriifoliaonlimestone
(calciumrich)cliffs.[33]
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Inallthestudiedcases,carnivoryallowsplantstogrowandreproduceusinganimalsasasourceof
nitrogen,phosphorusandpossiblypotassium.[34][35][36]However,thereisaspectrumofdependencyon
animalprey.Pygmysundewsareunabletousenitratefromsoilbecausetheylackthenecessaryenzymes
(nitratereductaseinparticular).[37]Commonbutterworts(Pinguiculavulgaris)canuseinorganicsourcesof
nitrogenbetterthanorganicsources,butamixtureofbothispreferred.[34]Europeanbladderwortsseemto
usebothsourcesequallywell.Animalpreymakesupfor
differingdeficienciesinsoilnutrients.
Plantsusetheirleavestointerceptsunlight.Theenergyisused
toreducecarbondioxidefromtheairwithelectronsfromwater
tomakesugars(andotherbiomass)andawasteproduct,
oxygen,intheprocessofphotosynthesis.Leavesalsorespire,in
asimilarwaytoanimals,byburningtheirbiomasstogenerate
chemicalenergy.Thisenergyistemporarilystoredintheform
ofATP(adenosinetriphosphate),whichactsasanenergy
currencyformetabolisminalllivingthings.Asawaste
product,respirationproducescarbondioxide.
Drosophyllumlusitanicumisoneofthe
fewcarnivorousplantstogrowindry,
alkalinesoil.
Foraplanttogrow,itmustphotosynthesisemorethanit
respires.Otherwise,itwilleventuallyexhaustitsbiomassand
die.Thepotentialforplantgrowthisnetphotosynthesis,the
totalgrossgainofbiomassbyphotosynthesis,minusthe
biomasslostbyrespiration.Understandingcarnivoryrequiresacostbenefitanalysisofthesefactors.[20]
Incarnivorousplants,theleafisnotjustusedtophotosynthesise,butalsoasatrap.Changingtheleafshape
tomakeitabettertrapgenerallymakesitlessefficientatphotosynthesis.Forexample,pitchershavetobe
heldupright,sothatonlytheiroperculadirectlyinterceptlight.Theplantalsohastoexpendextraenergy
onnonphotosyntheticstructureslikeglands,hairs,glueanddigestiveenzymes.[38]Toproducesuch
structures,theplantrequiresATPandrespiresmoreofitsbiomass.Hence,acarnivorousplantwillhave
bothdecreasedphotosynthesisandincreasedrespiration,makingthepotentialforgrowthsmallandthecost
ofcarnivoryhigh.
Beingcarnivorousallowstheplanttogrowbetterwhenthesoilcontainslittlenitrateorphosphate.In
particular,anincreasedsupplyofnitrogenandphosphorusmakesphotosynthesismoreefficient,because
photosynthesisdependsontheplantbeingabletosynthesiseverylargeamountsofthenitrogenrich
enzymeRuBisCO(ribulose1,5bisphosphatecarboxylase/oxygenase),themostabundantproteinonEarth.
ItisintuitivelyclearthattheVenusflytrapismorecarnivorousthanTriphyophyllumpeltatum.Theformer
isafulltimemovingsnaptrapthelatterisaparttime,nonmovingflypaper.Theenergy"wasted"bythe
plantinbuildingandfuellingitstrapisasuitablemeasureofthecarnivoryofthetrap.
Usingthismeasureofinvestmentincarnivory,amodelcanbeproposed.[20]Aboveisagraphofcarbon
dioxideuptake(potentialforgrowth)againsttraprespiration(investmentincarnivory)foraleafinasunny
habitatcontainingnosoilnutrientsatall.Respirationisastraightlineslopingdownunderthehorizontal
axis(respirationproducescarbondioxide).Grossphotosynthesisisacurvedlineabovethehorizontalaxis:
asinvestmentincreases,sotoodoesthephotosynthesisofthetrap,astheleafreceivesabettersupplyof
nitrogenandphosphorus.Eventuallyanotherfactor(suchaslightintensityorcarbondioxideconcentration)
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willbecomemorelimitingtophotosynthesisthannitrogenorphosphorussupply.Asaresult,increasingthe
investmentwillnotmaketheplantgrowbetter.Thenetuptakeofcarbondioxide,andtherefore,theplant's
potentialforgrowth,mustbepositivefortheplanttosurvive.Thereisabroadspanofinvestmentwhere
thisisthecase,andthereisalsoanonzerooptimum.Plantsinvestingmoreorlessthanthisoptimumwill
takeuplesscarbondioxidethananoptimalplant,andhencegrowinglesswell.Theseplantswillbeata
selectivedisadvantage.Atzeroinvestmentthegrowthiszero,becauseanoncarnivorousplantcannot
surviveinahabitatwithabsolutelyno
soilbornenutrients.Suchhabitatsdo
notexist,soforexample,Sphagnum
absorbsthetinyamountsofnitratesand
phosphatesinrainveryefficientlyand
alsoformssymbioseswithdiazotrophic
cyanobacteria.
Inahabitatwithabundantsoilnutrients
butlittlelight(asshownabove),the
grossphotosynthesiscurvewillbe
lowerandflatter,becauselightwillbe
morelimitingthannutrients.Aplant
cangrowatzeroinvestmentin
carnivorythisisalsotheoptimum
investmentforaplant,asany
investmentintrapsreducesnet
photosynthesis(growth)tolessthanthe
netphotosynthesisofaplantthat
obtainsitsnutrientsfromsoilalone.
Modellingcarnivoryinplants:grossphotosynthesis,respirationand
netphotosynthesisasafunctionoftheplant'sinvestmentin
carnivorousadaptations.Nonzerooptimumcarnivoryoccursin
brightlylithabitatswithverylimitingsoilnutrients.
Carnivorousplantsexistbetweenthese
twoextremes:thelesslimitinglightand
waterare,andthemorelimitingsoil
nutrientsare,thehighertheoptimum
investmentincarnivory,andhencethe
moreobvioustheadaptationswillbeto
thecasualobserver.
Themostobviousevidenceforthis
modelisthatcarnivorousplantstendto
growinhabitatswherewaterandlight
areabundantandwherecompetitionis
relativelylow:thetypicalbog.Those
thatdonottendtobeevenmore
fastidiousinsomeotherway.
Drosophyllumlusitanicumgrowswhere
thereislittlewater,butitisevenmore
extremeinitsrequirementforbright
lightandlowdisturbancethanmost
Modellingcarnivoryinplants:grossphotosynthesis,respirationand
netphotosynthesisasafunctionoftheplant'sinvestmentin
carnivorousadaptations.Anoptimumcarnivoryofzerooccursin
poorlylithabitatswithabundantsoilnutrients.
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othercarnivores.Pinguiculavalisneriifoliagrowsinsoilswithhighlevelsofcalciumbutrequiresstrong
illuminationandlowercompetitionthanmanybutterworts.[39]
Ingeneral,carnivorousplantsarepoorcompetitors,becausetheyinvesttooheavilyinstructuresthathave
noselectiveadvantageinnutrientrichhabitats.Theysucceedonlywhereotherplantsfail.Carnivoresareto
nutrientswhatcactiaretowater.Carnivoryonlypaysoffwhenthenutrientstressishighandwherelightis
abundant.[40]Whentheseconditionsarenotmet,someplantsgiveupcarnivorytemporarily.Sarracenia
spp.produceflat,noncarnivorousleaves(phyllodes)inwinter.Lightlevelsarelowerthaninsummer,so
lightismorelimitingthannutrients,andcarnivorydoesnotpay.Thelackofinsectsinwinterexacerbates
theproblem.Damagetogrowingpitcherleavespreventthemfromformingproperpitchers,andagain,the
plantproducesaphyllodeinstead.
Manyothercarnivoresshutdowninsomeseasons.Tuberous
sundewsdiebacktotubersinthedryseason,bladderwortsto
turionsinwinter,andnoncarnivorousleavesaremadebymost
butterwortsandCephalotusinthelessfavourableseasons.
Utriculariamacrorhizavariesthenumberofbladdersit
producesbasedontheexpecteddensityofprey.[41]Parttime
carnivoryinTriphyophyllumpeltatummaybeduetoan
unusuallyhighneedforpotassiumatacertainpointinthelife
cycle,justbeforeflowering.
Themorecarnivorousaplantis,thelessconventionalits
ParttimecarnivoryinTriphyophyllum
habitatislikelytobe.Venusflytrapsliveinaveryspecialised
peltatummaybeduetoanunusuallyhigh
habitat,whereaslesscarnivorousplants(Byblis,Pinguicula)are
needforpotassiumatacertainpointinthe
foundinlessunusualhabitats(i.e.,thosetypicalfornon
lifecycle,justbeforeflowering.
carnivores).ByblisandDrosophyllumbothcomefrom
relativelyaridregionsandarebothpassiveflypapers,arguably
thelowestmaintenanceformoftrap.Venusflytrapsfiltertheirpreyusingtheteetharoundthetrap'sedge,
soasnottowasteenergyonhardtodigestprey.Inevolution,lazinesspays,becauseenergycanbeusedfor
reproduction,andshorttermbenefitsinreproductionwilloutweighlongtermbenefitsinanythingelse.
Carnivoryrarelypays,soevencarnivorousplantsavoiditwhenthereistoolittlelightoraneasiersourceof
nutrients,andtheyuseasfewcarnivorousfeaturesasarerequiredatagiventimeorforagivenpreyitem.
Thereareveryfewhabitatsstressfulenoughtomakeinvestingbiomassandenergyintriggerhairsand
enzymesworthwhile.Manyplantsoccasionallybenefitfromanimalproteinrottingontheirleaves,but
carnivorythatisobviousenoughforthecasualobservertonoticeisrare.
Bromeliadsseemverywellpreadaptedtocarnivory,butonlyoneortwospeciescanbeclassifiedastruly
carnivorous.Bytheirveryshape,bromeliadswillbenefitfromincreasedpreyderivednutrientinput.Inthis
sense,bromeliadsareprobablycarnivorous,buttheirhabitatsaretoodarkformoreextreme,recognisable
carnivorytoevolve.Mostbromeliadsareepiphytes,andmostepiphytesgrowinpartialshadeontree
branches.Brocchiniareducta,ontheotherhand,isagrounddweller.
Manycarnivorousplantsarenotstronglycompetitiveandrelyoncircumstancestosuppressdominating
vegetation.Accordingly,someofthemrelyonfireecologyfortheircontinuedsurvival.
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Forthemostpartcarnivorousplantpopulationsarenotdominant
enoughtobedramaticallysignificant,ecologicallyspeaking,but
thereisanimpressivevarietyoforganismsthatinteractwithvarious
carnivorousplantsinsundryrelationshipsofkleptoparasitism,
commensalism,andmutualism.Forexample,smallinsectivores
suchastreefrogsoftenexploitthesupplyofpreytobefoundin
pitcherplants,andthefrogMicrohylanepenthicolaactually
specialisesinsuchhabitats.Certaincrabspiderssuchas
MisumenopsnepenthicolalivelargelyonthepreyofNepenthes,and
other,lessspecialised,spidersmaybuildwebswheretheytrap
insectsattractedbythesmellorappearanceofthetrapssome
scavengers,detritivores,andalsoorganismsthatharvestorexploit
thoseinturn,suchasthemosquitoWyeomyiasmithiiarelargelyor
totallydependentonparticularcarnivorousplants.Plantssuchas
Roridulaspeciescombinewithspecialisedbugs(Pameridea
roridulae)inbenefitingfrominsectstrappedontheirleaves.
Associationswithspeciesofpitcherplantsaresomanyandvaried
thatthestudyofNepenthesinfaunaissomethingofadisciplinein
Nepenthesmirabilisinaroadcutin
itsownright.Camponotusschmitzi,thedivingant,hasanintimate
Palau.Showinghabitandhabitat.
degreeofmutualismwiththepitcherplantNepenthesbicalcaratait
notonlyretrievespreyanddetritusfrombeneaththesurfaceofthe
liquidinthepitchers,butrepelsherbivores,andcleansthepitcherperistome,maintainingitsslippery
nature.Theantshavebeenreportedtoattackstrugglingprey,hinderingtheirescape,sotheremightbean
elementofmyrmecotrophytotherelationship.Numerousspeciesofmosquitoeslaytheireggsintheliquid,
wheretheirlarvaeplayvariousroles,dependingonspeciessomeeatmicrobesanddetritus,asiscommon
amongmosquitolarvae,whereassomespeciesofToxorhynchitesalsobreedinpitchers,andtheirlarvaeare
predatorsofotherspeciesofmosquitolarvae.Apartfromthecrabspidersonpitchers,anactualsmall,red
crabGeosesarmamalayanumwillenterthefluid,robbingandscavenging,thoughreputedlyitdoessoat
someriskofbeingcapturedanddigesteditself.
Nepenthesrajahhasaremarkablemutualismwithtwounrelatedsmallmammals,thetreeshrewTupaia
montanaandtheSummitRat,Rattusbaluensis.Thetreeshrewsandtheratsdefecateintotheplant'straps
whilevisitingthemtofeedonsweet,fruitysecretionsfromglandsonthepitcherlids.[42]Thetreeshrew
alsohasasimilarrelationshipwithatleasttwoothergiantspeciesofNepenthes.Moresubtly,Hardwicke's
WoollyBat(Kerivoulahardwickii),asmallspecies,roostsbeneaththeoperculum(lid)ofNepenthes
hemsleyana.[43]Thebat'sexcretionsthatlandinthepitcherpayfortheshelter,asitwere.Totheplantthe
excretaaremorereadilyassimilablethanintactinsectswouldbe.
TherealsoisaconsiderablelistofNepenthesendophytesthesearemicrobesotherthanpathogensthatlive
inthetissuesofpitcherplants,oftenapparentlyharmlessly.
Classification
Seealso:Listofcarnivorousplants
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Theclassificationofallfloweringplantsiscurrentlyinastateofflux.IntheCronquistsystem,the
DroseraceaeandNepenthaceaewereplacedintheorderNepenthales,basedontheradialsymmetryoftheir
flowersandtheirpossessionofinsecttraps.TheSarraceniaceaewasplacedeitherintheNepenthales,orin
itsownorder,theSarraceniales.TheByblidaceae,Cephalotaceae,andRoridulaceaewereplacedinthe
SaxifragalesandtheLentibulariaceaeintheScrophulariales(nowsubsumedintotheLamiales[44]).
Inmoremodernclassification,suchasthatoftheAngiospermPhylogenyGroup,thefamilieshavebeen
retained,buttheyhavebeenredistributedamongstseveraldisparateorders.Itisalsorecommendedthat
DrosophyllumbeconsideredinamonotypicfamilyoutsidetherestoftheDroseraceae,probablymore
closelyalliedtotheDioncophyllaceae.Thecurrentrecommendationsareshownbelow(onlycarnivorous
generaarelisted):
Dicots
Asterales(sunfloweranddaisyorder)
Stylidiaceae
Stylidium(triggerplants,aborderlinecarnivore)
Caryophyllales,(carnationorder)
Dioncophyllaceae
Triphyophyllum(atropicalliana)
Drosophyllaceae
Drosophyllum(Portuguesedewypine)
Droseraceae(sundewfamily)
Aldrovanda(waterwheelplant)
Dionaea(VenusFlytrap)
Drosera(sundews)
Droserapollis
Droserapites
Droseridites
Fischeripollis
Palaeoaldrovanda
Saxonipollis
Nepenthaceae(tropicalpitcherplantfamily)
Nepenthes(tropicalpitcherplantsormonkey
cups,includingAnurosperma)
Ericales(heatherorder)
Roridulaceae
Roridula(aborderlinecarnivore)
Sarraceniaceae(trumpetpitcherfamily)
Archaeamphora
Sarracenia(NorthAmericantrumpetpitchers)
Darlingtonia(cobraplant/lily)
Heliamphora(sunormarshpitchers)
Lamiales(mintorder)
Byblidaceae
Byblis(rainbowplants)
Lentibulariaceae(bladderwortfamily)
Pinguicula(butterworts)
Genlisea(corkscrewplant)
Stylidiumturbinatum
Aldrovandavesiculosa
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Utricularia(bladderworts,includingPolypompholyx,thefairyapronsorpinkpetticoats
andBiovulariaanobsoletegenus)
Martyniaceae(allborderlinecarnivores,relatedtothesesameplant)
Ibicella
Oxalidales(woodsorrelorder)
Cephalotus(Albanypitcherplant)
Monocots
Poales(grassorder)
Bromeliaceae(bromeliadorpineapplefamily)
Brocchinia(aterrestrialbromeliad)
Catopsis(aborderlinecarnivore)
Eriocaulaceae(pipewortfamily)
Paepalanthusbromelioides(aborderline
carnivore)
Cultivation
Byblisliniflora
Inhorticulture,carnivorousplantsareconsideredacuriosityora
rarity,butarebecomingmorecommonincultivationwiththe
adventofmassproductiontissueculturepropagationtechniques.
Venusflytrapsarestillthemostcommonlygrown,usuallyavailable
atgardencentersandhardwarestores,sometimesofferedalongside
othereasytogrowvarieties.Nurseriesthatspecializeingrowing
carnivorousplantsexclusivelyalsoexist,moreuncommonor
demandingvarietiesofcarnivorousplantscanbeobtainedfrom
specialistnursies.CaliforniaCarnivoresisanotableexampleof
suchanurserythatspecializesinthecultivationofcarnivorous
plants.Itisownedandoperatedbyhorticulturalist,PeterD'Amato.
Althoughdifferentspeciesofcarnivorousplantshavedifferent
cultivationrequirementsintermsofsunlight,humidity,soil
moisture,etc.,therearecommonalities.Mostcarnivorousplants
requirerainwater,orwaterthathasbeendistilled,deionisedby
Cephalotusfollicularis
reverseosmosis,oracidifiedtoaroundpH6.5usingsulfuricacid.
Commontapordrinkingwatercontainsminerals(particularly
calciumsalts)thatwillquicklybuildupandkilltheplant.Thisisbecausemostcarnivorousplantshave
evolvedinnutrientpoor,acidicsoilsandareconsequentlyextremecalcifuges.Theyarethereforevery
sensitivetoexcessivesoilbornenutrients.Sincemostoftheseplantsarefoundinbogs,almostallarevery
intolerantofdrying.Thereareexceptions:tuberoussundewsrequireadry(summer)dormancyperiod,and
Drosophyllumrequiresmuchdrierconditionsthanmost.
Outdoorgrowncarnivorousplantsgenerallycatchmorethanenoughinsectstokeepthemselvesproperly
fed.Insectsmaybefedtotheplantsbyhandtosupplementtheirdiethowever,carnivorousplantsare
generallyunabletodigestlargenoninsectfooditemsbitsofhamburger,forexample,willsimplyrot,and
thismaycausethetrap,oreventhewholeplant,todie.
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Acarnivorousplantthatcatchesnoinsectsatallwillrarelydie,
althoughitsgrowthmaybeimpaired.Ingeneral,theseplantsare
bestlefttotheirowndevices:afterunderwateringwithtapwater,
themostcommoncauseofVenusflytrapdeathisproddingthetraps
towatchthemcloseandfeedingtheminappropriateitems.
Mostcarnivorousplantsrequirebrightlight,andmostwilllook
betterundersuchconditions,asthisencouragesthemtosynthesise
redandpurpleanthocyaninpigments.NepenthesandPinguicula
willdobetteroutoffullsun,butmostotherspeciesarehappyin
directsunlight.
ManySarraceniahybridsareeasyto
Carnivoresmostlyliveinbogs,andthosethatdonotaregenerally
grow.
tropical.Hence,mostrequirehighhumidity.Onasmallscale,this
canbeachievedbyplacingtheplantinawidesaucercontaining
pebblesthatarekeptpermanentlywet.SmallNepenthesspeciesgrowwellinlargeterraria.
Manycarnivoresarenativetocoldtemperateregionsandcanbegrownoutsideinaboggardenyearround.
MostSarraceniacantoleratetemperatureswellbelowfreezing,despitemostspeciesbeingnativetothe
southeasternUnitedStates.SpeciesofDroseraandPinguiculaalsotoleratesubfreezingtemperatures.
Nepenthesspecies,whicharetropical,requiretemperaturesfrom20to30Ctothrive.
Carnivorousplantsrequireappropriatenutrientpoorsoil.Most
appreciatea3:1mixtureofSphagnumpeattosharphorticultural
sand(coirisanacceptable,andmoreecofriendlysubstitutefor
peat).NepentheswillgrowinorchidcompostorinpureSphagnum
moss.
Ironically,carnivorousplantsarethemselvessusceptibleto
infestationbyparasitessuchasaphidsormealybugs.Although
smallinfestationscanberemovedbyhand,largerinfestations
necessitateuseofaninsecticide.
Isopropylalcohol(rubbingalcohol)iseffectiveasatopical
insecticide,particularlyonscaleinsects.Diazinonisanexcellent
systemicinsecticidethatistoleratedbymostcarnivorousplants.
MalathionandAcephate(Orthene)havealsobeenreportedas
tolerablebycarnivorousplants.
CultivatedNepenthesrajahandafew
otherspecies.
Althoughinsectscanbeaproblem,byfarthebiggestkillerof
carnivorousplants(besideshumanmaltreatment)isgreymold
(Botrytiscinerea).Thisthrivesunderwarm,humidconditionsand
canbearealprobleminwinter.Tosomeextent,temperatecarnivorousplantscanbeprotectedfromthis
pathogenbyensuringthattheyarekeptcoolandwellventilatedinwinterandthatanydeadleavesare
removedpromptly.Ifthisfails,afungicideisinorder.
Theeasiestcarnivorousplantsforbeginnersarethosefromthecooltemperatezone.Theseplantswilldo
wellundercoolgreenhouseconditions(minimum5Cinwinter,maximum25Cinsummer)ifkeptin
widetraysofacidifiedorrainwaterduringsummerandkeptmoistduringwinter:
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Droseracapensis,theCapesundew:attractivestrapleavedsundew,pinkflowers,verytolerantof
maltreatment.
Droserabinata,theforkleavedsundew:large,Yshapedleaves.
Sarraceniaflava,theyellowtrumpetpitcher:yellow,attractivelyveinedleaves,yellowflowersin
spring.
Pinguiculagrandiflora,thecommonbutterwort:purpleflowersinspring,hibernatesasabud
(hibernaculum)inwinter.Fullyhardy.
Pinguiculamoranensis,theMexicanbutterwort:pinkflowers,noncarnivorousleavesinwinter.
Venusflytrapswilldowellundertheseconditionsbutareactuallyratherdifficulttogrow:eveniftreated
well,theywilloftensuccumbtogreymoldinwinterunlesswellventilated.SomeofthelowlandNepenthes
areveryeasytogrowaslongastheyareprovidedwithrelativelyconstant,hotandhumidconditions.
Medicinaluses
Astudypublishedin2009byresearchersfromTelAvivUniversity
indicatesthatsecretionsproducedbycarnivorousplantscontain
compoundsthathaveantifungalpropertiesandmayleadtothe
developmentofanewclassofantifungaldrugsthatwillbe
effectiveagainstinfectionsthatareresistanttocurrentantifungal
drugs.[45][46]
Culturaldepictions
Carnivorousplantshavelong
beenthesubjectofpopular
interestandexposition,
muchofithighlyinaccurate.
Fictionalplantshavebeen
featuredinanumberof
books,movies,television
series,andvideogames.
Typically,thesefictional
depictionsinclude
Depictionofanativebeingconsumed
AudreyJunior,themaneatingplant
exaggeratedcharacteristics,
byaYateveo("Iseeyou")
fromthecultfilmTheLittleShopof
suchasenormoussizeor
carnivoroustreeofCentralAmerica,
Horrors
possessionofabilities
fromLandandSeabyJ.W.Buel,
beyondtherealmofreality,
1887
andcanbeviewedasakindofartisticlicense.Twoofthemost
famousexamplesoffictionalcarnivorousplantsinpopularculture
arethe1960sblackcomedyTheLittleShopofHorrorsandthetriffidsofJohnWyndham'sTheDayofthe
Triffids.Othermovies,suchasTheHellstromChronicle(1971),andtelevisionseriesutilizeaccurate
depictionsofcarnivorousplantsforcinematicpurposes.
Theearliestknowndepictionofcarnivorousplantsinpopularculturewasacasewhereinalargeman
eatingtreewasreportedtohaveconsumedayoungwomaninMadagascarin1878,aswitnessedbyDr.
CarlLiche.LichereportedtheeventsintheSouthAustralianRegisterin1881.Thewoman,picturedinan
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accompanyingartwork,wassupposedtohavebeenamemberoftheMkodos,a"littleknownbutcruel
tribe".TheaccounthasbeendebunkedaspuremythasitappearsDr.Liche,theMkodos,andthetreewere
allfabrications.[47]
Seealso
Aggressivemimicry
Carnivorousfungus
Predatorydinoflagellate
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Furtherreading
SlackA(1986).InsecteatingPlantsandHowtoGrowThem.
WikimediaCommonshas
SherborneUK:Alphabooks.ISBN090667042X.
mediarelatedto
JuniperBE,RobinsRJ,JoelDM(1989).TheCarnivorous
Carnivorousplants.
Plants.AcademicPress,SanDiego.
CarnivorousPlantDatabase
Wikisourcehasseveral
(http://www.omnisterra.com/bot/cp_home.cgi)providesan
originaltextsrelatedto:
uptodate,searchabledatabaseofallthepublishedspeciesof
Carnivorousplants
carnivorousplants.
CarnivorousPlantFAQ(http://www.sarracenia.com/faq.html)atSarracenia.com
ListoffilmsandTVshowsthatfeaturecarnivorousplantsmostofthemfictional
(http://www.sarracenia.com/faq/faq1395.html)
BotanicalSocietyofAmericaCarnivorousPlantsOnline
(http://www.botany.org/carnivorous_plants/)
InnerWorldofCarnivorousPlantsfromtheJohnInnesCentre
(https://wikis.nbi.ac.uk/InnerWorlds/index.php/Inner_World_of_Carnivorous_Plants#.UThnmRyePA
k)
Ellison,A.M.2006.NutrientLimitationandStoichiometryofCarnivorousPlants.
(http://harvardforest.fas.harvard.edu/publications/pdfs/ellison_PlantBio_2006.pdf)PDF(334KB)Plant
Biol.8:740747.
Retrievedfrom"https://en.wikipedia.org/w/index.php?title=Carnivorous_plant&oldid=678686755"
Categories: Carnivorousplants Carnivory
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