Diversity and Evolutionof Monocots. . . orchids and palms . . .4 main groups:• Acorales - sister to all monocots• Alismatids– inc. Aroids - jack in the pulpit• Lilioids (lilies, orchids, yams)– grade, non-monophyletic– petaloid• Commelinoids– Arecales – palms– Commelinales – spiderwort– Zingiberales –banana– Poales– pineapple– grasses & sedgesLilioids - petaloid monocotsLilioids - petaloid monocots1. Terrestrial/epiphytes: plantstypically not aquatic2. Geophytes: herbaceous aboveground with below groundmodified perennial stems: bulbs,corms, rhizomes, tubers3. Tepals: showy perianth in 2series of 3 each; usually allpetaloid, or outer series notgreen and sepal-like & with nobractsAsparagales: *Orchidaceae - orchids• finish the Asparagales bylooking at the largest family -the orchids*Orchidaceae - orchidsOrchids aremycotrophic (= fungidependent) lilioids;some are obligatemycotrophsCypripedium acauleStemless lady-slipperCorallorhiza striataStriped coral rootThe family is diverse with about 880 genera and over 22,000species, mainly of the tropicsAll orchids have a protocorm - a feature restricted to thefamily.Dactylorhiza majalis protocorm*Orchidaceae - orchids• structure formed after germination and before thedevelopment of the seedling plant• has no radicle but instead mycotrophic tissueIonopsis twigepiphyteDendrobium branchepiphyte*Orchidaceae - orchidsCosmopolitan, but the majority of species are found in thetropics and subtropics, ranging from sea level to almost 5000 min nearly all environments except open water and true desert.Habit varies from herb to vine, but morethan half of the species are epiphyticOncidium trunkepiphyteOrchid root velamen (water storage) leaf tubers (water storage)*Orchidaceae - orchidsSurvive in these epiphytic and other harsh environments viaCAM photosynthesis, velamen, and leaf tubers, in addition tomycorrhizal associationSpecialized reproductive biology:*Orchidaceae - orchidsEulaema (euglossine)Perfume industryOphrys• unusual pollination systems• labellum petal for landing platform• pollen masses, reduced stamen number• numerous, dust-like seedsBulbophyllum nocturnumOnly totally night bloomingorchidFungal midge pollinated?A 2007 paper in Naturesuggests that the orchidfamily is not recent, butof late Cretaceous origin*Orchidaceae - orchids*Orchidaceae - orchidsCA 3 COZ 2+1 A 3,2,1 G (3)—• 6 tepals withlabellum (flowerresupinate orupside down)• 3 or fewerstamens• inferiorgynoecium fusedat top withstamens to formcolumn• capsule*Orchidaceae - orchidsFive subfamilies3 stamens2 stamens1 stamenCorymborkisTropidiaSobraliaEpipactisListeraCephalantheraNerviliaXerorchisMonophyllorchisMegastylis.latissimaMegastylis.raraChiloglottisAporostylisThelymitraCryptostylisOrthocerasCaladeniaCorybasMicrotisAaSpiranthesPrescottiaGoodyeraPristiglottisPachyplectronPterostylisChloraeaGalearisPlatantheraHabenariaDisaPaphiopedilumPhragmipediumSelenipediumCypripediumVanilla.imperialisVanilla.rosceriVanilla.barbellataVanilla.aphyllaVanilla.africanaVanilla.planifoliaVanilla.inodoraErythrorchis.altissimaErythrorchis.cassythoidesPseudovanillaCyrtosiaClematepistephiumEriaxisEpistephium.sp.432Epistephium.sp.433Epistephium.parviflorumEpistephium.lucidumPogonia.japonicaPogonia.ophioglossoidesCleistes.divaricataIsotria.medeoloidesIsotria.verticillataCleistes.roseaCleistes.sp.2DuckeellaNeuwiediaHypoxisRhodohypoxisCurculigoAsteliaAPOSTASIOIDEAEEPIDENDROIDEAEORCHIDOIDEAECYPRIPEDIOIDEAEVANILLOIDEAEAerangisDiaphanantheAeranthesAngraecumNeofinetiaPhalaenopsisCleisostomaPolystachyaAplectrumEarinaEpidendrumMeiracylliumCattleyaEncycliaArpophyllumMasdevalliaPleurothallisRestrepiaBletiaChysisCalypsoCoeliaCalanthePhaiusEriaPhreatiaArethusaCalopogonBletillaArundinaCoelogyneDendrochilumGlomeraThuniaCadetiaDendrobiumEpigeneiumBulbophyllumLiparisMalaxisMaxillariaCryptocentrumLycomormiumNeomooreaXylobiumLycasteAcinetaCoryanthesHoulletiaStanhopeaBifrenariaEriopsisCryptarrhenaDichaeaZygopetalumOncidiumCatasetumCyrtopodiumEulophiaAnselliaCymbidiumGrammatophyllum*Orchidaceae - orchidsRecent DNA studies by KenCameron show that vanilloidorchids diverged before the 2-stamen slipper orchidsthus, 1 stamen orchids arosetwice independently*Orchidaceae - orchidsNeuwiedia veratrifoliaApostasioideae (2 genera)• 3 stamens (usually)• most primitive of orchids withnearly actinomorphic flowers• Austral-asian distribution*Orchidaceae - orchidsVanilloideaeThe vanilloid orchidsare a small tropicalgroup of lianas, thatincludes VanillaVanilla*Orchidaceae - orchidsCypripedioideae (5 genera)• 2 stamens• “slipper” labellum• Northern HemispheredistributionCypripedium acaule - stemless lady’s-slipperThe lower petal is elaborated intothe labellum - the landing platform*Orchidaceae - orchidsLady’s-slippers have two functionalstamens with pollen masses*Orchidaceae - orchidsCypripedium acaule - stemless lady’s-slipperDeceptive pollination system fornaïve bumblebeesOther lady’s-slippers . . .Cypripedium arietinumRam’s-head lady’s-slipperthreatenedCypripedium calceolusYellow lady’s-slipper*Orchidaceae - orchidsCypripedium reginaeshowy lady’s-slipper*Orchidaceae - orchidsCypripedium candidumwhite lady’s-slipperThreatened, fen or calcareous soils*Orchidaceae - orchidsPaphiopedilumPhragmipedium*Orchidaceae - orchidsNon-native bucket orslipper orchidsMexipediumAll other orchidshave only 1functional stamenwith one or twopolliniaThe stamen is situated on a columnformed by fusion with the top of theinferior gynoecium*Orchidaceae - orchids*Orchidaceae - orchidsStructure andposition ofpollinia andcolumn allow forintricate anddifferential pollenplacement onpollinators*Orchidaceae - orchidslabellum other 2 petals 3 sepals (onebehind)other floral parts*Orchidaceae - orchidsEpidendroideaeThe epidendroid orchids,the largest group, arepredominantly epiphytes orlithophytes and include allthe showy tropical genera*Orchidaceae - orchidsOrchidoideaeThe orchidoid orchids aremostly terrestrials withtubers or fleshy rhizomesand include most temperateorchidsArethusa bulbosa - Dragon’s mouthAplectrum hyemalePutty root, Adam and eve*Orchidaceae - orchidsCalypso bulbosa - calypso orchid [threatened]*Orchidaceae - orchidsCalopogon tuberosus - grass pinknote the labellum on top!Goodyera pubescensRattlesnake plantainGoodyera tesselataRattlesnake plantain*Orchidaceae - orchidsCorallorhiza trifida -Early coral rootCorallorhiza striata -Striped coral root*Orchidaceae