oxalidales diversity and evolution of rosids wood sorrels · nectar guides, zygomorphic •...

Diversity andDiversity andEvolution of RosidsEvolution of Rosids

. . . violets, willows, and spurges . . .. . . violets, willows, and spurges . . .

OxalidalesOxalidales•• small, heterogeneous, novel group small, heterogeneous, novel groupof 6 families - seed character?of 6 families - seed character?

CephalotaceaeCephalotaceaeAustralian pitcher plantAustralian pitcher plant

OxalidaceaeOxalidaceaeWood sorrelsWood sorrels

6 genera, 770 species in the tropics and temperate areas - 7006 genera, 770 species in the tropics and temperate areas - 700belong to belong to OxalisOxalis (wood sorrel) (wood sorrel)

Oxalidaceae - wood sorrelsOxalidaceae - wood sorrels

•• plants are herbaceous creepers or woody plants are herbaceous creepers or woodyvinesvines


Oxalis corniculataOxalis corniculata - creeping yellow wood sorrel - creeping yellow wood sorrel •• typically 3-foliate typically 3-foliateleaves (the realleaves (the realshamrock)shamrock)

•• leaves are acidic to leaves are acidic totaste due to oxalictaste due to oxalicacid in the form ofacid in the form ofcalcium oxalatecalcium oxalate

6 genera, 770 species in the tropics and temperate areas - 7006 genera, 770 species in the tropics and temperate areas - 700belong to belong to OxalisOxalis (wood sorrel) (wood sorrel)


Oxalis corniculataOxalis corniculata

•• 5 merous flowers 5 merous flowers

•• fruits are 5 locular & fruits are 5 locular &winged capsules or berrieswinged capsules or berries

CA 5 CO 5 A 5+5 CA 5 CO 5 A 5+5 GG (5) (5)

OxalisOxalis


Oxalis corniculataOxalis corniculata

•• 5 merous flowers 5 merous flowers

•• fruits are 5 locular & fruits are 5 locular &winged capsules or berrieswinged capsules or berries

•• tristylytristyly common (3 levels common (3 levelsat which 2 sets of anthersat which 2 sets of anthersand 1 set of styles position)and 1 set of styles position)

CA 5 CO 5 A 5+5 CA 5 CO 5 A 5+5 GG (5) (5)

OxalisOxalis

U U U

Oxalis strictaOxalis stricta - -tall wood-sorreltall wood-sorrel


Oxalis acetosellaOxalis acetosella - -wood-sorrelwood-sorrel

Oxalis violaceaeOxalis violaceae - -violet wood-sorrelviolet wood-sorrel

•• common native and introduced wood-sorrels common native and introduced wood-sorrels

Oxalidaceae - wood sorrelsOxalidaceae - wood sorrels•• tropical fruit - carambola or star fruit: note 5 tropical fruit - carambola or star fruit: note 5carpellate structurecarpellate structure

Averrhoa carambolaAverrhoa carambola

*Malpighiales*Malpighiales•• large and diverse group of 38 large and diverse group of 38families - many of themfamilies - many of themcontributing importantly to tropicalcontributing importantly to tropicalforest diversityforest diversity

*Malpighiales*Malpighiales•• unresolved! unresolved! ““novelnovel”” clade clade•• leaf margin teeth leaf margin teeth•• ““ParietalesParietales”” subclade (placentation) subclade (placentation)•• hosts for hosts for CymothoeCymothoe butterflies butterflies

•• unusual life forms unusual life forms

*Malpighiales*Malpighiales

RafflesiaceaeRafflesiaceaeRafflesiaRafflesia

parasiteparasite RhizophoraceaeRhizophoraceaeRhizophora mangleRhizophora mangle

mangrovemangrove

PodostemonaceaePodostemonaceaePodostemonPodostemon

stream aquaticstream aquatic

23 genera, 800 species of herbs (temperate) to vines and small23 genera, 800 species of herbs (temperate) to vines and smalltrees (tropics). 400-600 of them are violets (trees (tropics). 400-600 of them are violets (ViolaViola).).

Viola tricolor - Viola tricolor - pansy, jonny-jump-uppansy, jonny-jump-up

*Violaceae - violets*Violaceae - violets

Rinorea - Rinorea - tropical shrubtropical shrub

ViolaViola have either basal leaves or cauline have either basal leaves or caulineleaves - cordate or palmately lobed orleaves - cordate or palmately lobed ordivided.divided.

StipulesStipules are well are welldeveloped.developed.

*Violaceae - violets*Violaceae - violetsCA 5 COZ 5 A 5 CA 5 COZ 5 A 5 GG (3) (3)

•• Flowers insect pollinated, Flowers insect pollinated,nectar guides, nectar guides, zygomorphiczygomorphic

•• Perianth 5 merous Perianth 5 merous

•• Lower Lower petal spurredpetal spurred, 2 lower, 2 lowerstamens have spurs going backstamens have spurs going backinto the petal spurinto the petal spur

•• 3 fused carpels 3 fused carpels


Pistil forms 3 partingPistil forms 3 partingcapsule in capsule in chasmogamouschasmogamousflowers (open flowers)flowers (open flowers)from out-crossingfrom out-crossing

CleistogamousCleistogamous, or closed, or closedflowers, form small capsulesflowers, form small capsulesvia self-pollination (note thevia self-pollination (note theparietalparietal placentationplacentation in inupper capsule)upper capsule)

*Violaceae - violets*Violaceae - violetsCA 5 COZ 5 A 5 CA 5 COZ 5 A 5 GG (3) (3)

V. pedataV. pedataBirdBird’’s-foot violets-foot violet V. pedatifidaV. pedatifida

Prairie violetPrairie violet


Viola affinis -Viola affinis -sand violetsand violet

V. sororiaV. sororia - dooryard violet - dooryard violet

Wisconsin state flower!Wisconsin state flower! One Oneof many stemless purpleof many stemless purplevioletsviolets

V. macloskeyi V. macloskeyi Wild white violetWild white violetA small white flowered violetA small white flowered violet


V. pubescensV. pubescens - downy yellow violet - downy yellow violeta stemmed yellow violeta stemmed yellow violet

V. canadensisV. canadensis - Canada white violet - Canada white violeta stemmed white violeta stemmed white violet


Hybanthus concolorHybanthus concolor - green violet - green violetSpecial concern, almost extripatedSpecial concern, almost extripated

*Violaceae - violets*Violaceae - violets *Violaceae - violets*Violaceae - violets•• tropical taxa of other tropical taxa of othergenera and genera and ViolaViola are arewoodywoody•• from where did the 10 from where did the 10species in Hawaii come?species in Hawaii come?


V. langsdorffiV. langsdorffi

•• DNA places them DNA places them withinwithin a apolyploid complex, amphi-polyploid complex, amphi-Beringian tundra speciesBeringian tundra species

*Violaceae - violets*Violaceae - violets•• Beringian source is Beringian source israre, especiallyrare, especiallysurprising with asurprising with agroup oncegroup onceconsidered tropicalconsidered tropical

•• bird migration bird migrationpathway frompathway fromBeringia to Hawaii isBeringia to Hawaii iscommon andcommon andpotential mechanismpotential mechanismof dispersalof dispersal

9 genera, 560 species of temperate herbs or small shrubs9 genera, 560 species of temperate herbs or small shrubs

•• opposite leaved opposite leaved

•• leaves are leaves are punctatepunctate - -with clearly visiblewith clearly visibleclear to black dots,clear to black dots,these sometimes seenthese sometimes seenon petals as wellon petals as well

Hypericaceae - St. JohnHypericaceae - St. John’’s worts wort

Hypericum perforatumHypericum perforatum - Klamath weed [ecologically invasive] - Klamath weed [ecologically invasive]Note the perforations along edge of yellow petalsNote the perforations along edge of yellow petals

CA 5 CO 5 ACA 5 CO 5 A ∞∞ GG (3-5) (3-5) Stamens many, often groupedStamens many, often grouped

Hypericaceae - St. JohnHypericaceae - St. John’’s worts wort

•• pistil made up of 3 to 5 fused pistil made up of 3 to 5 fusedcarpelscarpels

•• placentation either axile (then 3-5 placentation either axile (then 3-5locuoles) or parietal (as here)locuoles) or parietal (as here)

Hypericum pyrimidatumHypericum pyrimidatum great St. Johngreat St. John’’s worts wort

Hypericaceae - St. JohnHypericaceae - St. John’’s worts wortCA 5 CO 5 ACA 5 CO 5 A ∞∞ GG (3-5) (3-5) •• pistil made up of 3 to 5 fused pistil made up of 3 to 5 fused

carpelscarpels

•• placentation either axile (then 3-5 placentation either axile (then 3-5locuoles) or parietallocuoles) or parietal

Hypericum pyrimidatumHypericum pyrimidatum great St. Johngreat St. John’’s worts wort

Hypericaceae - St. JohnHypericaceae - St. John’’s worts wortCA 5 CO 5 ACA 5 CO 5 A ∞∞ GG (3-5) (3-5)

•• fruit a capsule fruit a capsule

Hypericaceae - St. JohnHypericaceae - St. John’’s worts wort•• HypericumHypericum common in high common in highelevation regions of tropicselevation regions of tropics

HypericumHypericum in Costa Rican paramo in Costa Rican paramo

Triadenum virginicumTriadenum virginicum marsh St. Johnmarsh St. John’’s worts wort

•• Triadenum Triadenum restrictedrestrictedto wetlandsto wetlands

Dovyalis hebecarpaDovyalis hebecarpa Oncoba spinosaOncoba spinosa

Chemically defined by Chemically defined by salicinssalicins (salicylic acid). Many (salicylic acid). Manymembers of the tropical members of the tropical ““FlacourtiaceaeFlacourtiaceae”” with showy flowers with showy flowersalso have salicins and are now part of the Salicaceaealso have salicins and are now part of the Salicaceae

*Salicaceae - willows, poplars*Salicaceae - willows, poplars

55 genera, 1000+ species of shrubs/trees - 450 are willows55 genera, 1000+ species of shrubs/trees - 450 are willows((SalixSalix), less numerous are poplars, aspens (), less numerous are poplars, aspens (PopulusPopulus).).

Populus deltoidesPopulus deltoides - -American cottonwoodAmerican cottonwood


Salix babylonica Salix babylonica --weeping willowweeping willow

*Salicaceae - willows, poplars*Salicaceae - willows, poplarsWillows (Willows (SalixSalix) are ) are dioeciousdioecious trees of temperate regions with trees of temperate regions withreduced flowers in aments - both insect and wind pollinatedreduced flowers in aments - both insect and wind pollinated

Salix babylonica Salix babylonica --weeping willowweeping willow

malemalefemalefemale

malemalefemalefemale •• nectar glandsnectar glands at base of bract allows at base of bract allowsinsectinsect as well as as well as wind pollinationwind pollination•• fruit is a capsule with cottony seedsfruit is a capsule with cottony seedsfor wind dispersalfor wind dispersal


•• many species are many species are ““precociousprecocious”” - -flower before leaves flush in springflower before leaves flush in spring Salix discolorSalix discolor - -

pussy willowpussy willow


•• species vary from large trees, shrubs, to tiny tundra species vary from large trees, shrubs, to tiny tundrasubshrubssubshrubs


Salix fragilisSalix fragilis - -crack willowcrack willow

Salix pedicellarisSalix pedicellaris - -bog willowbog willow

Salix herbacea -Salix herbacea -dwarf willowdwarf willow

Populus deltoidesPopulus deltoidesAmerican cottonwoodAmerican cottonwood

PopulusPopulus - poplars, cottonwood, aspens - poplars, cottonwood, aspensmalemale

femalefemale


•• flowers possess a flowers possess a diskdisk•• cottony seeds in capsule cottony seeds in capsule

P. grandidentataP. grandidentatabigtooth aspenbigtooth aspen

P. tremuloidesP. tremuloidestrrembling aspentrrembling aspen

•• aspens are clonal from root aspens are clonal from rootsprouts, fast growing, lightsprouts, fast growing, lightwooded, and important forwooded, and important forpulp in the paper industrypulp in the paper industry

*Salicaceae - willows, poplars*Salicaceae - willows, poplarsPopulus balsamiferaPopulus balsamiferaBalsam poplar, balm-of-gileadBalsam poplar, balm-of-gilead

Populus albaPopulus albaWhite poplarWhite poplar

Introduced from EuropeIntroduced from Europe


Populus deltoidesPopulus deltoides - -American cottonwoodAmerican cottonwood

*Salicaceae - willows, poplars*Salicaceae - willows, poplars *Euphorbiaceae - spurges*Euphorbiaceae - spurgesEuphorbiaceae s.l. isEuphorbiaceae s.l. ispolyphyleticpolyphyletic and now and nowbroken into 3 familiesbroken into 3 families

EuphorbiaceaeEuphorbiaceae

PutranjivaceaePutranjivaceae

PhyllanthaceaePhyllanthaceae

•• large cosmopolitan family of trees, large cosmopolitan family of trees,shrubs, and herbs of 222 genera andshrubs, and herbs of 222 genera and6000 species6000 species

•• leaves alternate, simple (often leaves alternate, simple (oftenpalmately lobed) or palmatelypalmately lobed) or palmatelycompoundcompound

*Euphorbiaceae - spurges*Euphorbiaceae - spurges

Ricinus Ricinus - castor oil bean- castor oil bean

•• latexlatex bearing and filled with bearing and filled withnasty chemicals (source ofnasty chemicals (source ofrubber, castor oil, tapioca,rubber, castor oil, tapioca,poinsettia)poinsettia)

Majority of the family has unisexualMajority of the family has unisexualflowers, 5 sepals, no petals, numerousflowers, 5 sepals, no petals, numerousstamens, 3 fused carpels, and capsulesstamens, 3 fused carpels, and capsules

CA 5 CO 0 ACA 5 CO 0 A ∞∞ G G 00CA 5 CO 0 A 0 CA 5 CO 0 A 0 GG (3) (3)


malemalefemalefemale

CodianumCodianum

A quite different arrangementA quite different arrangementof unisexual flowers is seen inof unisexual flowers is seen inmany of our spurges of themany of our spurges of thegenera genera EuphorbiaEuphorbia and andChamaesyceChamaesyce..

Flower or inflorescence?Flower or inflorescence?

*Euphorbiaceae - spurges*Euphorbiaceae - spurgesThe The ““flowerflower”” of our of ourflowering spurgeflowering spurge is actually is actuallya highly modifieda highly modifiedinflorescence = inflorescence = cyathiumcyathium

Shown here are 3 cyathia; theShown here are 3 cyathia; thewhole unit here is onewhole unit here is onecyathiumcyathium

*Euphorbiaceae - spurges*Euphorbiaceae - spurgesEuphorbia corollataEuphorbia corollata - flowering spurge - flowering spurge

Cyathium is composed of:Cyathium is composed of:glandsglands


appendages of glandsappendages of glands

*Euphorbiaceae - spurges*Euphorbiaceae - spurgesCyathium is composed of:Cyathium is composed of:

glandsglands

1- stamened male flowers (no perianth) 1- stamened male flowers (no perianth)




1- stamened male flowers (no perianth) 1- stamened male flowers (no perianth)




1- tricarpellate female flower (3 styles)1- tricarpellate female flower (3 styles)

Euphorbia esulaEuphorbia esula - leafy spurge - leafy spurge

•• one of several species one of several specieslabelled labelled ““obnoxiousobnoxiousweedweed”” by state law by state law

*Euphorbiaceae - spurges*Euphorbiaceae - spurges *Euphorbiaceae - spurges*Euphorbiaceae - spurges

Euphorbia dendroidesEuphorbia dendroides

Euphorbia pulcherrima - Euphorbia pulcherrima - poinsettiapoinsettia

Euphorbia canariensisEuphorbia canariensis

EuphorbiaEuphorbia is a is a ““giantgiant”” genus (> 2,000 spp.) with some genus (> 2,000 spp.) with somespectacular radiations - spectacular radiations - is the cyathium a is the cyathium a ““key innovationkey innovation””??

Hura Hura - poison arrow- poison arrow


Manihot esculenta -Manihot esculenta -manioc, cassava, tapiocamanioc, cassava, tapioca Hevea Hevea - rubber- rubber

•• economically important members economically important membersfrom Neotropicsfrom Neotropics


•• besides its use as a poison, what besides its use as a poison, whatother features does other features does Hura crepitansHura crepitanspossess that makes it a pretty nastypossess that makes it a pretty nastyplant?plant?

Passifloraceae - passion flowerPassifloraceae - passion flowerSpectacular group of vines and trees in theSpectacular group of vines and trees in thepantropics - one ingrediant of Hawaiian punchpantropics - one ingrediant of Hawaiian punch

PassifloraPassiflora

parietal placentationparietal placentation

berry fruitberry fruit

Passifloraceae - passion flowerPassifloraceae - passion flowerCo-evolutionary relationships involving leaf shapes, HeliconidCo-evolutionary relationships involving leaf shapes, Heliconidbutterfly egg layers, extrafloral nectaries, and patrolling antsbutterfly egg layers, extrafloral nectaries, and patrolling ants

A small family of herbs most noted as the source of flaxA small family of herbs most noted as the source of flax(linen), linseed oil, and base chemicals for linoleum.(linen), linseed oil, and base chemicals for linoleum.

Linaceae - flaxLinaceae - flax

Linum bienneLinum bienne

Linum narbonenseLinum narbonenseLinum sulcatumLinum sulcatum - -grooved yellow flaxgrooved yellow flax

BruguieraBruguiera

Rhizophoraceae - mangrovesRhizophoraceae - mangrovesRhizophora -Rhizophora -red mangrovered mangrove

Asian and Madagascar tropical trees andAsian and Madagascar tropical trees andone widespread mangroveone widespread mangrove

Byrsonima spicataByrsonima spicata

Malpighiaceae - malpighsMalpighiaceae - malpighsDiverse woody family of pantropical region with distinctiveDiverse woody family of pantropical region with distinctiveclawed petals and floral nectaries.clawed petals and floral nectaries.

MalpighiaMalpighia HiptageHiptage

oxalidales diversity and evolution of rosids wood sorrels · nectar guides, zygomorphic •...

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