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The Flowering Plants Handbook · 157. FAGACEAE BEECH, CHESTNUT & OAK FAMILY Trees or shrubs. Leaves ±gland-dotted,deciduous (Castanea, some Castanopsis, Fagus, some Quercus), simple,

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Page 1: The Flowering Plants Handbook · 157. FAGACEAE BEECH, CHESTNUT & OAK FAMILY Trees or shrubs. Leaves ±gland-dotted,deciduous (Castanea, some Castanopsis, Fagus, some Quercus), simple,
Page 2: The Flowering Plants Handbook · 157. FAGACEAE BEECH, CHESTNUT & OAK FAMILY Trees or shrubs. Leaves ±gland-dotted,deciduous (Castanea, some Castanopsis, Fagus, some Quercus), simple,

The Flowering Plants HandbookA practical guide to families and genera of the world

James W. Byng

Page 3: The Flowering Plants Handbook · 157. FAGACEAE BEECH, CHESTNUT & OAK FAMILY Trees or shrubs. Leaves ±gland-dotted,deciduous (Castanea, some Castanopsis, Fagus, some Quercus), simple,

CONTENTS

Chapter 1: Introduction....................................................................... 1 Notes on how to use this book.................................................. 2 Usefuldiagnosticcharacters……………………….........…....……………3 Keytomajorplantgroups..........................................................5

Chapter 2: Basal Angiosperms............................................................ 7 Amborellales………………..............................................................8 Nymphaeales……………………….....................................................9 Austrobaileyales……….................................................................11 Chloranthales.............................................................................13 Cannellales……………………………………………………….........…....………14 Piperales………………………………………………………............……………..16 Magnoliales………………………………………………….........…….………….21 Laurales………………………………………………………….........………..…….28

Chapter 3: Monocots............................................................................ 36 Acorales……………………….............................................................37 Alismatales……………………………...................................................38 Petrosaviales…………….................................................................47 Dioscoreales...............................................................................48 Pandanales………………………………………………………...........…………..51 Liliales……………………………………………………………..….........………….56 Asparagales………………….............................................................65 Arecales……….………………………....................................................86 Commelinales……………...............................................................88 Zingiberales................................................................................93 Dasypogonaceae………………………………………..……….........………….100 Poales..……………………………………………………….……….........…………101

Chapter 4: Basal Eudicots ................................................................... 116 Ceratophyllales…………….............................................................117 Ranunculales……………………….....................................................118 Sabiaceae…………………................................................................130 Proteales....................................................................................131 Trochodendrales…………………………………………………...........……….136 Buxales………………………………………………………………..........…………137 Gunnerales…………………..............................................................139 Dilleniaceae…………………………....................................................141

Chapter 5: Rosids ……………………………………........................................... 142 Saxifragales………………….............................................................143 Vitales…….……………………………....................................................153 Zygophyllales..………….................................................................154 Fabales.......................................................................................156

Page 4: The Flowering Plants Handbook · 157. FAGACEAE BEECH, CHESTNUT & OAK FAMILY Trees or shrubs. Leaves ±gland-dotted,deciduous (Castanea, some Castanopsis, Fagus, some Quercus), simple,

CHAPTER 2: BASAL ANGIOSPERMS

Basal angiosperms are the earliest diverging lineages of flowering plants and themost commonlyencounteredarethelaurels,magnoliasandwaterlilies.Thegroupoftenexhibit‘primitive’orancestralcharacterssuchasanopenflowerorganisation,designedforlargeinsect(e.g.beetle)pollination,andlittledistinctionbetweenthesepalsandpetals.Theygenerallylack‘advanced’orderivedcharacterssuch as fused petals and zygomorphy. Basal angiosperms are generallywoody plantswith notableherbaceousexceptionssuchasmanyPiperalesandtheaquaticfamiliesoftheNymphaeales.

The leavesaresimpleandoftenaromaticcontainingetherealoils.Theflowersarespirallyarrangedor 3-merous andoftenhavemanyperianth parts, stamens and carpels. The carpels are often freeandthestylesandstigmasarepoorlydeveloped.Thestamensareusuallylaminarshapedwithpoorlydifferentiatedanthers.

Page 5: The Flowering Plants Handbook · 157. FAGACEAE BEECH, CHESTNUT & OAK FAMILY Trees or shrubs. Leaves ±gland-dotted,deciduous (Castanea, some Castanopsis, Fagus, some Quercus), simple,

130. DAPHNIPHYLLACEAETrees or shrubs; branchlets with leaf scars and lenticels. Leaves simple, alternate or rarely opposite, oftenclusteredatbranchends;marginsentire;petiolespresent;stipulesabsent. Inflorescences axillaryracemes;bractsconspicuous.Flowersunisexual (plantsdioecious),actinomorphic.Sepals free,±imbricateor sometimesabsent.Petals absent. Male flowers: stamen filaments often shorter than anthers; anthers basifixed. Female flowers: ovariessuperior;carpelsfused; locules2;ovules(1-)2perlocule;placentationaxileorapical;stigmasdecurrent; sometimesstaminodes. Fruitsdrupes.

Genera 1/species ca.30;Daphniphyllum.

Distribution: IndiatoAustraliaandEastAsia.

Floral formula: K(0-)3-6C0A5-14G2(-4)

References:Endress&Igersheim1999;Fishbeinet al.2001,2004;Huang1965,1966;Kubitzki2007;Min&Kubitzki2004.

131. ITEACEAE Treesorshrubs,sometimesclimbing(Itea).Leavessimple,alternate;marginsoftenspinytoothedtoentire;petiolespresent; stipulestinyor absent. Inflorescences racemesor panicles (Itea), or cymes to corymbs (Pterostemon).Flowers bisexualorrarelypolygamous,actinomorphic,hypanthium.Sepalsfreeorbasallyfused,valvate,persistent.Petalsfree,persistent,clawed(Pterostemon).Stamensalternatingwithpetals(Itea)orsepals(Pterostemon);anthersdorsifixed,introrse;staminodes5(Pterostemon).Ovariessuperiortopart-inferior(Itea)orinferior(Pterostemon);carpels fused;locules2or5;ovules4-6(Pterostemon)ormany(Itea)perlocule;placentationaxile.Fruits capsules.

Genera 2/species ca.18.

Distribution:Tropicaltonortherntemperateregions.

Floral formula: Itea K5C5A5Ĝ2 Pterostemon K5C5A5+5°Ĝ5

Notes: BothgenerawerepreviouslyassociatedwithEscalloniaceae.

References: Bohm et al.1999;Kubitzki2007a,2007b;Fishbeinet al. 2001,2004;

Generic synopsis•Itea (southeastAsiatowesternMalesia,easternNorthAmerica,East

toSouthAfrica;ca.16spp.).•Pterostemon (much-branchedshrubs;Oaxaca[Mexico];ca.2spp.).

132. GROSSULARIACEAE GOOSEBERRY FAMILYShrubs, sometimes climbing, often spiny; often glandular hairs. Leaves aromatic, usually deciduous, simple,alternate;venationpinnateto3-palmate;marginslobedortoothed;petiolespresent;stipulesusuallypresentorabsent. Inflorescences racemes,usuallyon short-shoots; sometimesbracteate, hairy.Flowers bisexual or rarelyunisexual (plants dioecious, e.g. R. diacanthum), actinomorphic, hypanthium well-developed (and lobed) andpetaloid;green,white,yelloworred;sometimesbracteolate.Sepalsfused,persistent. Petalsrarelyabsentorfree,imbricate, sometimes interpretated as staminodes, smaller than sepals. Stamens opposite the sepals; anthersbasifixed.Ovaries inferior topart inferior; carpels fused; locule1;ovules4-many;placentation parietal; style 2. Fruits berries,withpersistentperianth.

Genus 1/species ca. 150; Ribes.

Distribution:TemperatenorthernhemisphereandAndestosouthernSouthAmerica.

Floral formula: K(3-)5(-9)C(0-3-)5(-9)A4-5Ĝ2

Useful species: R. nigrum(blackcurrants);R. rubrum(redcurrants);R. uva-crispa(gooseberry).

Notes: Ribes speciosumis4-merous.

References: Fishbein et al.2001,2004; Messingeret al.1999;Morin2009;Senters&Soltis2003;Schultheis&Donoghue2004;Weigend2006;Weigendet al.2002.

MaleflowersofDaphniphyllum teijsmannii

©KENPEI/CC-BY-SA-3.0

Daphniphyllum macropodum©SteveLaw/CC-BY-2.0

Itea virginica©WouterHagens/CC-BY-SA-3.0

Itea ilicifolia©Imc/CC-BY-SA-3.0

Ribes glutinosum Ribes tenue

Page 6: The Flowering Plants Handbook · 157. FAGACEAE BEECH, CHESTNUT & OAK FAMILY Trees or shrubs. Leaves ±gland-dotted,deciduous (Castanea, some Castanopsis, Fagus, some Quercus), simple,

157. FAGACEAE BEECH, CHESTNUT & OAK FAMILY Treesorshrubs.Leaves±gland-dotted,deciduous(Castanea,someCastanopsis, Fagus, some Quercus), simple, alternate or whorled in 3’s (some Trigonobalanus); marginsentire,toothedordeeplylobed(e.g.Quercus);petiolespresent,swollenatbase;stipules oftentriangular,deciduous.Flowers unisexual(plantsmonoeciousorrarelydioecious),actinomorphic. Perianth fused or free, bract-like.Male flowers in heads or catkins;stamenfilamentsfreeorrarelybasallyfused;sometimespistillode.Female flowers in spikesorsolitary,eachflowersurroundedbybracts(=cupule);staminodesabsentor6-12;ovaries inferior;carpels fused; locules2-6(-9);ovules2per locule;placentationaxiletoapical;styles3-6.Fruits nutswithsubtendingbracts(=cupule).

Genera 8/species ca.925.

Distribution: Widelydistributed.

Floral formula: P6(-9)A(4-)6-12(-many)Ĝ(2-)3-6(-15)

Useful species: Quercus suber(barkusedforcork);Castanea(chestnuts).

Notes:Theflowersareusuallywindpollinatedorlessofteninsectpollinated(Castanea).

References:Huanget al.1999;Kubitzki1993;Manoset al.1993,2002,2006;Manos&Stanford2001;Nixon1993;Nixon&Crepet1989;Oh&Manos2006;WCSP2013.

Generic synopsisFruit a solitary nut (= acorns) and round in cross-section•Lithocarpus (maleinflorescenceserect;IndiatoChinaandNewGuinea;ca.335spp.).•Notholithocarpus (cupulescalesreflexedandhookedattip;westernUSA;1sp.,N. densiflorus).•Quercus(oaks;maleinflorescencespendulous;cupulescalesrarelyreflexed;temperatenorthern

hemispheretoMalesiaandColombia;ca.430spp.).

Fruit 1-several nuts and angled (often 3-angled) in cross-section •Castanea(chestnuts;plantdeciduous,secondaryveinsprominent,ovary6(-9)-loculed,styles6+;

temperatenorthernhemisphere;ca.8spp.).•Castanopsis (cupulesusually6-lobed;tropicalandsubtropicalAsia;ca.135spp.).•Chrysolepis (plantevergreen,secondaryveinsobscure,styles3;westernUSA).•Fagus(beeches;plantsdeciduous,maleflowersindense,pendulousheads,femaleflowers(1-)2;

temperatenorthernhemispheretoMexico;ca.10spp.).•Trigonobalanus (cupulesusually3-or5-lobed;Colombia,northernThailandtoYunnan[China],

Malesia).

158. MYRICACEAESmall trees or shrubs. Leaves often aromatic (sweet smell), deciduous (Comptonia,Myrica), simple, alternate (spiral); margins entire to toothed or rarely pinnatifid;petioles present; stipules absent or present (Comptonia). Inflorescence a spike orcatkin-like;bracts1(-3)perflower.Flowers unisexual(plantsdioeciousormonoecious)orrarelybisexual(Canacomyrica),actinomorphic.Perianth absentorwith6tinylobes(Canacomyrica).Male flowers with bracteolesusually2; stamenfilamentsfreeorfused;anthersdorsifixed;staminodes inCanacomyrica. Female flowerswithbracteoles2-4;ovary superior to inferior; carpels fused; locule1; ovule1; placentationbasal; styles2,pink-red.Fruit a drupe(oftenwaxyandwarty)ornut-like(e.g.Comptonia, Myrica).

Genera 4/species 57.

Distribution:Temperatetotropicalregions.

Floral formula: P0(-6)A2-8(-many)Ĝ2(-3)

Notes:ThegenericnameMorellaisoftenincludedinMyricabutisretainedatgenericrankherefollowingHerbert(2005)andHerbertet al.(2006).

References:Herbert2005,2006;Herbertet al.2006;Huguetet al.2005;Kubitzki1993;Leroy1949.

Castanea sativa

FruitofQuercus robur©BöhringerFriedrich/CCBY-SA3.0AT

Lithocarpus edulis©KENPEI/CC-BY-SA-3.0

FruitsofMorella faya©Forest&KimStarr/CC-BY-SA-3.0

MaleflowersofMorella faya©Forest&KimStarr/CC-BY-SA-3.0

FruitofFagus sylvatica

Page 7: The Flowering Plants Handbook · 157. FAGACEAE BEECH, CHESTNUT & OAK FAMILY Trees or shrubs. Leaves ±gland-dotted,deciduous (Castanea, some Castanopsis, Fagus, some Quercus), simple,

306. SARCOBATACEAE GREASEWOOD FAMILYThornyshrubs. Stemserect,longshootssolitary,shortshootsclustered.Leaves deciduous,simple,alternate(spiral)toclustered;marginsentire;petiolesabsent;stipulesabsent.Flowers unisexual(plantsusuallymonoecious),actinomorphic.Male flowers in catkin-like inflorescences; perianth absent; stamens ±sessile;anthers long. Female flowers solitary or paired; perianth 1-whorled, fused,campanulate,greenish;ovariespart-inferior;carpels fused;placentationbasal.Fruits wingedachenes.

Genus 1/species 2; Sarcobatus.

Distribution:NorthAmericainsaltdesertsespeciallythesouthwestdesertsandtheGreatBasin.

Floral formula: P(0-)2A1-4G2

Confused with:Amaranthaceae‒differsusuallyintheprescenseofspines.

Notes:ThegenuswaspreviouslyplacedinChenopodiaceae(=Amaranthaceae).ThetwospeciesaresometimesbothtreatedassubspeciesofS. vermiculatus.

References:Behnke1997;Hilset al.2004;Kuhn1993.

Species synopsis•S. baileyi(0.5-1mtall,leaveshairy;restrictedtoNevada[USA]).•S. vermiculatus(1-5mtall,leavesusuallylackinghairs).

307. NYCTAGINACEAE BOUGAINVILLEA & FOUR O’CLOCK FAMILYTrees,shrubs,lianasorannualorperennialherbs;stemswithswollennodesinherbs,spines(e.g.Bougainvillea, Pisonia, Phaeoptilum),scalesorstellatehairsinLeucastereae;woodoftenturnsorangetoredwhencut. Leaves simple,alternate,oppositeorwhorled;petiolespresentorabsent;stipulesabsent.Inflorescencessolitaryflowers,cymes,panicles,spikesorumbels;bractsinvolucral,sometimeslarge,petaloidand3(e.g.Bougainvillea)or5(e.g.Mirabilis). Flowers bisexual or sometimes unisexual (plants dioecious), actinomorphic or zygomorphic (Alliona,some Colignonia).Perianth 1-whorled,fused,valvate,petaloid,oftentubular,oftenhairy.Stamen filaments freeorbasallyfused,filamentsfreeorattachedtoperianth,sometimesdimorphic.Ovaries superior;carpel1;locule1;ovule1;placentationbasal;stylelong.Fruitsachenes,sometimeswithdrupe-likeanthocarp.

Genera 30/species ca.400.

Distribution: MostlyAmericasandsomeinOldWorld(Boerhavia, some Commicarpus, Phaeoptilum, Pisonia,Mirabilis jalapa and Bougainvilleaiswidelycultivatedinurbanareas).

Floral formula: P(3-)5(-10)A(1-)5(-20)G1

Notes: ThegenericnamesAmmocodon and SelinocarpusaresynonymouswithAcleisanthes,andthenamesHesperonia, Oxybaphus and Quamoclidion aresynonymouswithMirabilis.

Confused with:Phytolaccaceae‒differsnotablyinthesolitarycarpel,fusedandtubularperianth.

References: Bittrich&Kuhn1993;Cuenoudet al.2002;Douglas&Manos2007;Douglas&Spellenberg2010;Levin2002;Spellenberg2004.

Generic synopsisBoldoeae (woody or herbaceous; leaves alternate; stamens 3-5, free at base; anthocarp absent)•Boldoa (herbs,hookedhairspresent;MexicoandCaribbeantonorthern

SouthAmerica;2-3spp.).•Cryptocarpus (shrubs,flowersinconspicuous;GalapagosIslands,Ecuadorto

Bolivia).•Salpianthus (shrubs,flowersshowy;Mexico;1sp.,S. arenarius).

Bougainvilleae (trees or shrubs, sometimes spines; leaves alternate or opposite; stamens 4-12, often fused basally; often 3 petaloid bracts)•Belemia (flowerspink,stamens12;easternBrazil;1sp.,B. fucsioides).•Bougainvillea (oftenspiny,stamens4-10;Neotropics,B. peruviana, B. glabra

and B. spectabiliscultivatedinwarmregionsoftheworld;ca.18spp.).•Phaeoptilum(flowersmostlyunisexual;southwestAfrica;1sp.,P. spinosum).

FruitsofSarcobatus vermiculatus©MattLevin/CC-BY-2.0

Sarcobatus vermiculatus©MattLevin/CC-BY-2.0

Bougainvillea spectabilis

Page 8: The Flowering Plants Handbook · 157. FAGACEAE BEECH, CHESTNUT & OAK FAMILY Trees or shrubs. Leaves ±gland-dotted,deciduous (Castanea, some Castanopsis, Fagus, some Quercus), simple,

Caribeeae (subshrubs; leaves opposite; flowers solitary; stamens 2, fused to base of sepals; rare - known only from one collection!)•Caribea (Cuba;1sp.,C. litoralis).

Colignonieae (lianas to subshrubs; leaves opposite or whorled; stamens 5, fused basally, attached to sepals; petaloid bracts) •Colignonia (Andes;ca.6spp.).

Leucastereae (trees or shrubs; leaves alternate; stellate hairs; stamens 2-3 (12-20), fused at base) •Andradea (perianthcampanulate,stamens12-20;southeastBrazil;1sp.,A. flori-

bunda).•Leucaster (perianthcampanulate,shrubs;Brazil).•Ramisia (perianthcampanulate,trees;southeastBrazil;1sp.,R. brasiliensis).•Reichenbachia (perianthtubular;Bolivia,Argentina,Paraguay,Brazil;2spp.).

Pisonieae (woody, paired spines sometimes; leaves usually opposite or alternate or whorled; stamens (2-)5-10(-many), fused basally, attached to sepals in bisexual flowers; flowers often unisexual)•Cephalotomandra (stamens25-30;Colombia).•Grajalesia (axillaryspines,stamens6,3-wingedfruit;Mexico;1sp.,G. fasciculata).•Guapira (stamenslongerthanperianth,fleshyanthocarp;Neotropics;ca.70spp.).•Neea (stamens(5-)8(-10)andshorterthanperianth,plantdriesblack;Neotropics;

ca.80spp.).•Neeopsis (stamens6,plantsdryyellow;Guatemala;1sp.,N. flavifolia).•Pisonia (stamens2-manyandlongerthanperianth,woodyanthocarp,flowers(func-

tionally)unisexual;pantropical;ca.40spp.).•Pisoniella (stamens(6-)8(-11)andlongerthanperianth,flowersbisexual;Mexico,

Bolivia,Argentina).

Nyctagineae (herbs or shrubs; leaves opposite; stamens (1-)2-6(-18), basally fused, sometimes attached to sepals)•Abronia (herbs,stigmalinear,stamensusually(3-)4-5andshorterthanperianth;

southwestUSAtonorthernMexico:ca.20spp.).•Acleisanthes (herbsorsubshrubs,flowers>2cm,whiteandusuallysolitary,stamens

2-5andunequal,filamentsbasallyfused;TexasandCalifornia[USA]toMexico).•Allionia (herbs,involucralbractsfree,3-lobed,flowersfunnel-shaped,stamens4-7;

Americas).•Anulocaulis (herbs,stamens3-4,stickybandsonupperinternodes;Centraland

SouthUSA,Mexico).•Boerhavia (herbs,flowers<2cmandrarelysolitary,stamens1-4(-6);tropicaland

subtropicalregions;ca.20spp.).•Commicarpus (subshrubs,anthocarpwithwart-likeglands,flowersinumbels,

stamens2-6;tropicaltosubtropicalregions,especiallyAfrica,westernAsia;ca.30-35spp.).

•Cuscatlania (herbs,bracts3-8,ciliate,stamens7-8;ElSalvador;1sp.,C. volcanicola).•Cyphomeris (herbs,stickybandsonupperinternodes,stamens5,anthocarpassyme-

trical;Texas[USA]toMexico).•Mirabilis (fouro’clock;herbs,involucralbracts4-8-lobedandusuallyfused,stamens

2-6;Americas,M. jalapanaturalisedelsewhere,M. himalaica fromNorthernIndiatoChina;ca.60spp.).

•Nyctaginia (flowersindenseheads,perianthcampanulate,stamens5-8andlongerthanperianth;Texas[USA]tosouthernMexico;1sp.,N. capitata).

•Okenia (stickyherbs,flowerssolitary,stamens5-18;FloridatoMexico,Nicaragua).•Tripterocalyx (herbs,stigmalinear,stamens(3-)4-5andshorterthanperianth;fruits

withprominentlyveinedwings;NorthAmerica).

Abronia latifolia

Allionia incarnata

Acleisanthes nevadensis

Anulocaulis annulatus Boerhavia repens

©EricYarnell/CC-BY-SA-3.0

©Forest&KimStarr/CC-BY-SA-3.0

©StanShebs/CC-BY-SA-3.0

©StanShebs/CC-BY-SA-3.0

©StanShebs/CC-BY-SA-3.0

Guapira venosa

Mirabilis jalapa

©AlexPopovkin/CC-BY-2.0

©Thaumaturgist/CC-BY-SA-3.0

Pisonia umbellifera©DavidEickhoff/CC-BY-2.0

Page 9: The Flowering Plants Handbook · 157. FAGACEAE BEECH, CHESTNUT & OAK FAMILY Trees or shrubs. Leaves ±gland-dotted,deciduous (Castanea, some Castanopsis, Fagus, some Quercus), simple,

340. ACTINIDIACEAE KIWI FRUIT FAMILYTreestoshrubsorlianas.Leavesdeciduous(Clematocletha),simple,alternate(spiral);marginsentireortoothed;petiolespresent;stipulesabsent;hairscommon,especiallyonyounggrowth. Inflorescencesaxillarydensepanicles,cymesor solitaryflowers; few species cauliflorous.Flowers showy,bisexual orunisexual (plantspolygamousorfunctionallydioeciousormonoecious),actinomorphic;oftenbracteolate,small.Sepalsfreeorbasallyfused,usuallyimbricate,persistent.Petalsfreeorbasallyfused,imbricate,deciduous;oftenwhiteorredtoyellow.Stamens many infascicles,filamentsattachedtobaseofperianthorfree;anthersbasifixed.Ovariessuperior;carpelsfused;locules(3-)5(-many);ovulesmanyper locule;placentationaxile; stylesasmanyas carpels,persistent in fruit especiallyActinidia;stigmas(4-)5-30.Fruits berriessometimeswithstiffhairsorrarelyloculicidalcapsules(someSaurauia).

Genera 3/species ca.315.

Distribution:HimalayasandEastAsiatoAustralasiaandtheAmericas.

Floral formula: K(3-)5(-9)C(3-)5(-8)Amany G3-5(-many)

Useful species: Actinidia deliciosa(kiwifruit,chinesegooseberry).

Confused with:SladeniaceaeandTheaceae‒differsbytheusuallyfleshyfruitandmorethan10ovulesperloculeintheovary.

Notes: Clematoclethracontainsabout20speciesaccordingtoLiang(1984)orjustonespeciesaccordingtoTang&Xiang(1989)andthelattercircumscriptionisadoptedhere.

References:Anderberget al.2002;Dressler&Bayer2004;Geutenet al.2004;Liet al.2002;Liang1984;Tang&Xiang1989.

Generic synopsis•Actinidia (lianas,ovarymulti(15+)-locular,stylesfree;KurilIslandstoSakhalin[Russia]

throughChina,Taiwan,Himalayas,IndochinaandMalaysia;ca.65spp.).•Clematoclethra (lianas,ovary5-locular,stylesfused;montanewesternandCentralChina;1

sp.,C. scandens).•Saurauia (treesorshrubs,flowersbisexualorplantsfunctionallydioecious;Himalayas,East

Asia,southeastAsiatoFiji,Neotropics,Queensland[Australia];ca.250spp.).

341. CLETHRACEAE LILY-OF-THE-VALLEY FAMILYShrubsorsmalltrees(Clethra arborea).Leavesdeciduousintemperatespecies,simple,alternate(spiral),clusteredatbranchesends;margins entireor toothed;petiolespresentor absent; stipulesabsent; oftenpubescent with hairs simple, fascicledand/or stellate. Inflorescences racemes,panicles, fasciclesorumbel-like;oftenbracteate,deciduous.Flowersbisexualorfunctionallyfemale(someClethra),actinomorphicorslightlyzygomorphic(Purdiaea);bracteolesabsent. Sepalsfreeorfused,imbricate,bluntly-lobed,unequalinPurdiaea,persistent,enlarginginfruit(Purdiaea).Petals freetorarelypartlyfused,imbricate;whiteorrarelypink(Clethra)orpinktoviolet(Purdiaea).Stamens 2-whorled, filaments free or attached to petals; anthers dorsifixed. Ovaries superior; carpels fused,sometimesnectariesatbase; locules3-5;ovules1 (Purdiaea)ormany(Clethra)per locule;placentationaxile in upperportion; style 1. Fruitsloculicidalcapsules(Clethra)orindehiscentandovoidtosubglobose(Purdiaea)

Genera 2/species ca.95.

Distribution:Americas,tropicalandsubtropicalAsiaandMadeira[Portugal].

Floral formula: K5(-6)C5(-6)A5(-6)+5(-6)G3-5

Notes: PurdiaeawaspreviouslyplacedinCyrillaceaeanddiffersfromthatfamilybythepubescentleavesandinflorescencesandunequalsepals.

References:Anderberg&Zhang2002;Fioret al.2003;Schneider&Bayer2004;Thomas1960;WCSP2013.

Generic synopsis•Clethra (sepalsequal,carpels3,oftenfoundinacidsoils;Ameri-

cas,tropicalandsubtropicalAsia,Madeira;ca.80spp.).•Purdiaea (sepalsunequal,carpels3-5;BelizetoPeru,especially

Cuba;ca.12spp.).

Actinidia chinensis

MaleflowersofSaurauia zahlbruckneri

©MNolf/CC-BY-SA-3.0

©StanShebs/CC-BY-SA-3.0

Clethra alnifolia©H.Zell/CC-BY-SA-3.0

Purdiaea cubanensis©ScottZona/CC-BY-2.0

Page 10: The Flowering Plants Handbook · 157. FAGACEAE BEECH, CHESTNUT & OAK FAMILY Trees or shrubs. Leaves ±gland-dotted,deciduous (Castanea, some Castanopsis, Fagus, some Quercus), simple,

APIALESTrees,shrubs,lianasorherbs.Leaves sometimesaromatic,simpleorcompound,alternateorrarelyopposite;marginsentire, toothedordissected; stipulesabsentor sometimespresent (Araliaceae, someApiaceae). Inflorescences often simple or compound umbels. Flowers small, bisexual or sometimes unisexual, actinomorphic or rarelyzygomorphic.Sepals freeorfused,oftenreducedtoabsent.Petals usuallyfreeorsometimesfused.Stamens 5or3-many (Araliaceae), filaments free fromperianthor rarelyattached.Ovaries inferioror rarelysuperior (e.g.Pittosporaceae,Pennantiaceae).Fruitsusuallydrupesorschizocarps.

Notes:Manyspeciesintheorderhavealternate,compoundleavesandumbelinflorescences.

1.Ovarysuperior..................................................................................................................................... 21.Ovaryinferiororpart-inferior............................................................................................................... 4

2.Leavessimple;flowers5-merous;widelydistributed……….................................................................. 32.Leavescompound;flowers3-5merous;Hawaii[USA]distribution…………..Araliaceae (Tetraplasandra)

3.Petalsvalvate;fruitsdrupes……………………………………………………………………………………....Pennantiaceae

3.Petalsimbricate;fruitscapsulesorberries…………………..……………………………………………...Pittosporaceae

4.Fruitsdryschizocarps;styles2;carpels2;petalsvalvate;flowersusuallyinsimpleor

compoundumbels…………………………………………………………………………..……………………….......Apiaceae

4.Fruitsdrupesor(lessoften)dryfruits;petalsvalvateorimbricate;descriptionnotasabove………......4

5.Carpels3,ovary1-locular;leavessimplealternate;flowersunisexual(plantsdioecious);

NewZealandandsouthernSouthAmericadistribution……............................................Griseliaceae

5.Carpels2tomany,ovary1-tomulti-locular;leavessimpleorcompound............................................6

6.Inflorescencesusuallysimpleumbels,spikesorheads;petals3-12;anthersdorsifixed..................... 76.Inflorescencespaniclesorracemes;petals5;anthersbasifixedordorsifixed…………….. Torricelliaceae

7.Carpels2;petalswithinflexedtips;fruitsdrupesorschizocarps..................................Myodocarpaceae

7.Carpels2-200;petalsusuallynotasabove;fruitsdrupesorberries,rarelyschizocarps……..Araliaceae

407. PENNANTIACEAETrees,shrubs(P. baylisiana)orrarelylianas(sometimesP. cunninghamii);stemsoftenlenticellate.Leaves simple,alternate (spiral); margins entire or toothed (see notes); venation pinnate, 4-6 lateral pairs; stipules absent. Inflorescences solitary flowers, cymes or panicles; ramiflorous or cauliflorous (P. baylisiana, sometimes P. endlicheri).Flowersminute(ca.2-3mmlong),functionallyunisexual(plantsdioecious). Sepalsfree,minute(ca.1mmlong).Petalsfreeorfused,valvate;whitetogreenish.Stamen filaments freeorattachedtopetals,reducedinfemaleflowers;anthersdorsifixed.Ovaries superior; carpelsfused;locule1;ovule1;placentationapical;styleveryshorttoabsent;stigma3-lobedordisc-like.Fruits blackishdrupes,(7-15x5-7mm).

Genus 1/species 4; Pennantia.

Distribution:EasternAustraliaandNewZealand.

Floral formula: K5C5A5G2-3

Notes:ThegenuswaspreviouslyincludedinIcacinaceae.

References:Gardner&deLange2002;Karehad2001,2002,2003;Plunkettet al.2004.

ImmaturefruitsofPennantia corymbosa©Kahuroa/CC01.0