Takhtajan Armen. Flowering Plants

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Superorder ROSANAE 313

Paleodicraeia, Sphaerothylax, Saxicolla, Saxicolella,

Zehnderia, Stonesia, Winklerella, Thelethylax, Diplo-

bryum, Grifﬁ thella, Hanseniella, Polypleurum, Thawat-

chaia, Willisia; mourereae: Mourera, Tulasneantha,

Lonchostephus.

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Superorder ROSANAE 315

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Order 83. VITALES

Small trees, erect shrubs, and herbs or more often woody

lianas. Roots sometimes adventitious. Cambium storied.

Vessel elements usually rather large, with simple perfo-

rations; lateral pitting scalariform or sometimes opposite

to alternate. Fibers with simple or sometimes bordered

pits, septate. Rays heterogeneous. Axial parenchyma

mainly paratracheal. Sieve-element plastids of Pcs-type

that contain several polygonal protein crystalloids and

similar to the plastids of Gunnera (Behnke 1981). Nodes

3–7-lacunar. Leaves alternate or very rarely opposite,

simple or compound, of diverse shapes and structures,

stipulate or estipulate, often bearing specialized, multi-

cellular, stalked, deciduous “pearl glands” and some-

times also peltate scales. Stomata of diverse kinds.

Inﬂ orescences leaf-opposed or terminal, rarely axillary,

cymose or racemose, sometimes with 1–2 tendril-like

branches; bract and bracteoles small, sometimes nectar

secreting, the bract subtending each branch. Flowers

rather small, bisexual or (some members of haploste-

monous Vitaceae) polygamo-monoecious or dioecious,

actinomorphic, (3)4–5(6–7)-merous. Calyx mostly

weakly developed, lobed, dentate or truncate, often

much reduced. Petals 4–5 (rarely 3 or 6–7), valvate, free

or basally connate or sometimes coherent distally and

calyptra-like (Vitis), rarely persistent. Stamens 5–4

(3–7), opposite the petals; ﬁ laments free, or connate into

a tube. Anthers dorsiﬁ xed, tetrasporangiate or rarely dis-

porangiate, introrse, opening longitudinally. Tapetum

secretory. Microsporogenesis simultaneous. Pollen

grains 2-celled or 3-celled, 3- colporate, with reticulate

ornamentation. Nectary disc usually well developed,

annular or cupular or sometimes of ﬁ ve free glands, usu-

ally more or less adnate to the ovary. Gynoecium of

2-6(-8) united carpels; stylodia connate into a style with

capitate, seldom (Tetrastigma) quadriﬁ d, rarely sessile

stigma; ovary superior, but sometimes partly sunken in

the nectary disc, completely or incompletely divided

into locules, with one or two ovules per locule. Ovules

erect, axile or basal, anatropous, apotropous, with ven-

tral raphe, bitegmic, crassinucellate. Female gameto-

phyte of Polygonum-type. Endosperm nuclear. Haustoria

of the female gametophyte present or absent. Fruits

berries ( sometimes rather dry). Seeds endotestal (with

ligniﬁ ed, several-layered endotesta and persistent thin

tracheidal tegmen); embryo small to minute, straight,

surrounded by copious, horny, generally ruminate, oily

endosperm with two main folds, one from each side of

the raphe. Alkaloids present (rarely), or absent, ellagic

acid present (Vitis vinifera), or absent (Parthenocissus,

Rhoicissus), present myricetin.

Two rather closely related families Vitaceae and

Leeaceae are usually included in the order Rhamnales.

However, from the Rhamnales they differ in the pres-

ence of raphide sacs in the parenchymatous tissues,

berrylike fruits, seed anatomy (Corner 1976) and P-type

sieve-element plastids (Behnke 1974). According to

Corner (1976: 36), seed structure of Vitaceae “is

scarcely improved on that of Magnolia and may, even,

be more primitive. Certainly there is no afﬁ nity in this

regard with Rhamnales.” However, any direct links

between Magnoliales or Magnoliidae in general and

Vitales is very doubtful. Endlicher (1841) placed his

Ampelideae between Araliaceae and Corneae. Lindley

(1853: 439) placed Vitaceae s.l. between Berberidaceae

and Pittosporaceae, but indicated the connection,

although no direct, with the Araliaceae. He stated: “If

the Vine is compared with Aralia racemosa, the rela-

tionship of the present Order to it will be too obvious to

be mistaken. Suppose that Aralia racemosa had an

adherent calyx, erect ovules, with stamens opposite the

petals, and it would be a Vitis”.

Order Vitales probably derived from some Rosanae

ancestor.

Key to Families

1 Woody lianas usually with leaf-opposite tendrils,

rarely succulent small trees or erect herbs, branches

often swollen or articulated at nodes. Tendrils

316 Subclass VI. ROSIDAE

simple, biﬁ d, or 2–3-, or 4–12-branched, usually

leaf- opposite. Leaves evergreen, or deciduous,

gland-dotted, simple, lobed or unlobed, or com-

pound; when compound ternate, or pinnate, or pal-

mate. Stipules if present petiolar and usually

caduceus, rarely absent. Inﬂ orescences leaf-opposed

or terminal, infrequently axillary, panicles, corymbs,

or rarely spikes. Flowers small, pedicellate, with

prophyls, actinomorphic, 4–5(-7)-merous. Calyx

small, often indistinctly lobed or dentate or much

reduced and truncate. Petals free or basally connate,

valvate, sometimes coherent distally and calyptrate.

Stamens free, equal in number to and opposite the

petals; anthers tetrasporaginate or disporaginate.

Nectary disc intrastaminal, ring-shaped, copular, or

gland-shaped. Ovary 2(-6)-locular, the ovules two

per locule, anatropous or apotropous. Style long or

short, stigma inconspicuous or capitate or 4-lobed.

Female gametophyte with a haustorium. Endostome

forming the micropyle. Fruits 1–4-seeded berries.

Seeds endotestal with abaxial chalazal knot and an

adaxial raphe; embryo small, straight; endosperm

oily and proteinaceous, copious, ruminate. Present

proanthocyanins, and catechin, prodelphinisin, gal-

lic, ellagic, coffeic, chlorogenic and tartaric acids; n

= 11–16, 19–20. . . . . . . . . . . . . . . . . .1. vitaceae.

1 Small trees or erect, creeping or scrambling shrubs

without tendrils, or herbaceous plants with a woody

base; stema unarmed or with rows of prickles.

Leaves distichous, uni-trifoliolate, or 1– 4-pinnate.

Stipules attached to base of petiole and sheath-

ing, often early caducous. Stomata cyclocytic, acti-

nocytic, or rarely anomocytic. The mesophyll

contains mucilage cells sometimes with raphides

and usually with calcium oxalate crystals.

Inﬂ orescences leaf-opposed cyme, lax or condensed

by reduction of branches or peduncle. Calyx cam-

panulate or cupuliform. Petals distally cohering in

bud by ventrally apical keels, reﬂ exed at anthesis,

the basal portions connate to one another and adnate

to androecium forming a shortly tubular structure

free from calyx. Stamens connate distally (above

the common stricture with the corolla) for some

distance into a tube that may bear lobes alternate

with the anthers. Nectary disc absent. Pollen grains

3-celled or rarely 2-celled, 3-colporate. Ovary

4-6(-10)-locular (sometimes incompletely), with

one ovule per locule. Style short, entire, the stigma

slightly thickened. Female gametophyte without a

haustorium. Exostome forming the micropyle.

Large numbers of raphids are commonly in the tis-

sues of corolla, anthers, staminodial tube. Fruits

4–6-seeded berries. Seeds with ruminate endosperm;

seed coat has a thick ligniﬁ ed endotesta and incon-

spicuous tegmen; raphides in seed coat absent;

n = 10–12. . . . . . . . . . . . . . . . . . . . . . 2. leeaceae.

1. VITACEAE

A. L. de Jussieu 1789 (including Ampelopsidaceae

Kosteletzky 1835, Cissaceae Horaninow 1847,

Pterisanthaceae J. G. Agardh 1858). 15/750. Tropical

and subtropical regions, with relatively few species in

warm-temperate regions (mostly species of Vitis,

Ampelopsis, and Parthenocissus).

Cissus, Nothocissus, Cyphostemma, Acareosperma,

Cayratia, Vitis, Ampeloclssus, Parthenocissus, Yua,

Ampelopsis, Pterisanthes, Clematicissus, Tetrastigma,

Rhoicissus, Pterocissus.

2. LEEACEAE

Dumortier 1829. 1/34. Paleotropics, mainly southern

and southeastern Asia.

Leea.

Very close to Vitaceae and sometimes (e.g., Thorne

1992, 2000, 2006) are considered a subfamily

Leeoideae. From Vitaceae they differ in the erect habit,

absence of tendrils, absence of nectary disc, the

presence of secondary septa in the ovary and absence

of raphides in seed coat.

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Order 84. ROSALES

Evergreen or deciduous trees and shrubs or often herbs

(usually perennial). Vessels mostly with simple

perforations. Axial parenchyma scanty to abundant, dif-

ferently patterned but mainly diffuse and scanty paratra-

cheal. Sieve-element plastids of S-type. Nodes usually

trilacunar, or unilacunar, or pentalacunar to multilacu-

nar. Leaves alternate or rarely opposite, mostly serrate or

less often entire, simple or variously compound, persis-

tent or caducous, mostly with stipules that are sometimes

adnate to the petiole; leaf mesophyll cells contain oil

bodies. Stomata anomocytic. Flowers in various kinds of

inﬂ orescences (mostly cymose), less often solitary,

bisexual or less often unisexual, actinomorphic or more

or less zygomorphic, 5-merous, cyclic, mostly with dou-

ble perianth. Hypanthium usually well-developed (not

evident in some male ﬂ owers). Sepals, petals and sta-

mens inserted on its rim; disk sometimes distinct,

intrastaminal. Sepals and petals free, usually imbricate.

Stamens few to numerous; ﬁ laments free; anthers

2- locular, tetrasporangiate, usually introrse, dorsiﬁ xed,

versatile, opening longitudinally or rarely by apical

pores. Pollen grains 2-celled, 3-colporate or rarely

3- porate, pantoporate, or 3- zonocolporate. Tapetum

secretory. Microsporogenesis simultaneous. Gynoecium

of 1 to many (up to 50), free or more or less united car-

pels, with free stylodia and usually small capitate

stigma; ovary superior to inferior, with one to many, but

mostly 1–2 ovules per carpel or per locule. Ovules pen-

dulous to subbasal, anatropous or less often hemitrop-

ous or campylotropous, bitegmic or less often unitegmic,

crassinucellate. Female gametophyte of Polygonum-

type. Endosperm nuclear, rarely helobial (Lyonothamnus).

Fruits various, ﬂ eshy or dry, dehiscent or not. Seeds

small to fairly large, wingless or rarely winged; embryo

small to large, straight or bent, linear or folded, some-

times the radicle invested; cotyledons moderately to

markedly thick, rarely convolute; endosperm mostly

present, varying between 1–20 cells thick in different

genera, sometimes abundant (as in Physocarpus and

Rhodotypos); perisperm often present. Present triter-

penes, ellagic acid, cyanogenic glycosides, ﬂ avones.

Linked to the Saxifragales through the most archaic

members of Rosaceae-Spiraeoideae.

Key to Families

1 Sepals imbricate. Evergreen or deciduous trees and

shrubs or often herbs (usually perennial). Lateral

pitting alternate or less often opposite. Parenchyma

scanty to abundant, differently patterned but mainly

diffuse and scanty paratracheal. Lysigenous muci-

lage ducts in the pith wanting. Nodes usually trilacu-

nar, or unilacunar, or pentalacunar to multilacunar.

Leaves alternate or rarely opposite, mostly serrate or

less often entire, simple or variously compound,

persistent or caducous, mostly with stipules that are

sometimes adnate to the petiole. Flowers in various

kinds of inﬂ orescences (mostly cymose), less often

solitary, bisexual or less often unisexual, actinomor-

phic or more or less zygomorphic. Sepals (3-)5(-10),

free or adnate to the ovary, imbricate, sometimes

with an epicalyx of bracteoles. Petals inserted below

the margin of the disk, as many as the calyx-lobes,

free, equal or rarely unequal, imbricate, deciduous,

rarely absent. Stamens two, three, or four times as

many as petals, sometimes more numerous (up to

400) or rarely only ﬁ ve or even (Aphanes) only one,

bent inward in bud; ﬁ laments free or more or less

connate with the ﬂ oral tube or between themselves;

anthers usually introrse or rarely (Potentilla) latrorse,

dorsiﬁ xed, versatile, opening longitudinally or rarely

by apical pores. Pollen grains 2-celled, 3-colporate

or rarely 3-porate or (Sanguisorba) pantoporate.

Gynoecium of one to many (up to 50), free or more

or less united carpels, with free stylodia and usually

small capitate stigma; ovary with one to many, but

mostly 1–2 ovules per locule. Ovules pendulous to

subbasal, anatropous or less often hemitropous or

campylotropous, bitegmic or less often unitegmic,

crassinucellate. Fruits multifollicles, achenes, or

pomes, or drupes, rarely capsules (Lindleyieae).

Superorder ROSANAE 319

Seeds small to fairly large, wingless or rarely (e.g.

Exochorda) winged; exotestal cells periclinally

elongated, radial walls thickened, or palisade or

tubular; outer wall often becoming mucilaginous,

endotegmic cells slightly thickened, or seed coat

undistinguished; embryo small to large, straight or

bent, linear or folded, sometimes the radicle invested;

cotyledons moderately to markedly thick, rarely

convolute; endosperm mostly present, varying

between 1–20 cells thick in different genera, some-

times abundant (as in Physocarpus and Rhodotypos);

perisperm often present. Present triterpenes, ellagic

acid, cyanogenic glycosides, ﬂ avones, n = 7–10,

15–17, 27. . . . . . . . . . . . . . . . . . . . . . . 1. rosaceae.

1 Sepals valvate. Evergreen, small glabrous trees (up

to 10 m tall and 1 m in diameter) or shrubs, some-

times with saponaceous bark. Vessels with simple

perforations. Axial parenchyma abundant, apotra-

cheally diffuse, diffuse-in-aggregates and scanty

paratracheal Rays heterogenous (Zhang 1992).

Extremely numerous styloids present in the phloem

of stems, petioles and in the leaf lamina (Lersten

and Horner 2005). Nodes unilacunar with 3 traces.

Leaves alternate, simple, leathery, almost entire or

serrate, shortly petioled, margins toothed; stipules

small, caduceus, petiolar. Inﬂ orescences axillary or

terminal, 3–5- ﬂ owered, pedicels 2-bracteolate.

Flowers polygamo- dioecious (terminal ﬂ owers

bisexual, lateral ones staminate), 5-merous, rather

large, tomentose. Calyx-tube persistent, leathery,

small, subcupular, lobes broadly ovate. Petals

white, sessile, spathulate. Disk thick, ﬂ eshy, lining

the calyx-tube, lobes 5, thick, depressed,

emarginated, adnate to the calyx-lobes. Stamens

ten, ﬁ ve opposite the petals and inserted at the base

of the disk, ﬁ ve alternate and inserted at apex of the

disk-lobes; ﬁ laments subulate, free; anthers

2- locular, introrse. Pollen grains tri-zonocolporate

(Claxton et al. 2005). Gynoecium of ﬁ ve carpels,

tomentose, partly connate; styles terminal, free,

stigmatic zone elongated along styles; ovules

numerous, pleurotropous, ﬂ attened, subhorizontal,

densely 2- seriate. Fruits multifollicles, strongly

asymmetrically lobed, spreading stellately, open-

ing down both surfaces of the lobes. Seeds exot-

estal, long- and broadly winged in the upper part,

outer testa layers thickened; endosperm thin;

cotyledons convolute. Present ﬂ avone C-glycosides;

n = 14. . . . . . . . . . . . . . . . . . . . . 2. quillajaceae.

1. ROSACEAE

A. L. de Jussieu 1789 (including Agrimoniaceae

Gray 1822, Alchemillaceae Martynov 1820,

Amygdalaceae Marquis 1820, Cercocarpaceae

J. G. Agardh 1858, Chamaemoraceae Lilja 1870,

Coleogynaceae J. Agardh 1858, Cydoniaceae

Schnizlein 1856, Dryadaceae Gray 1822, Fragariaceae

Richard ex Nestler 1816, Guamatelaceae S. Oh et

D. Potter 2006, Lindleyaceae J. Agardh 1858,

Malaceae J. K. Small ex Britton 1903, Mespilaceae

Schultz-Schultz enstein 1832, Neilliaceae Miquel

1855, Potentillaceae Berchtold et J. Presl 1820,

Poteriaceae Raﬁ nesque 1815, Prunaceae Martynov

1820, Pyraceae Vest 1818, Rhodotypaceae J.Agardh

1858, Sanguis orbaceae Berchtold et J. Presl 1820,

Sorbaceae Brenner 1886, Spiraeaceae Bertuch 1801,

Tormentillaceae Martynov 1820, Ulmariaceae Gray

1822). 111/3000–3500. Subcosmopolitan, but espe-

cially in subtropical and temperate regions of the

Northern Hemisphere.

1.1 SPIRAEOIDEAE

Carpels 2–5 (up to 12), rarely (Stephanandra,

Adenostoma, and spp. of Neillia) only one, free or

nearly so, less often more or less connate. Ovules 1–2

to several (about 8 in Chamaebatiaria and 11 in

Neillia) per carpel, pendulous or less often ascend-

ing, anatropous, hemitropous, or campylotropous,

mostly epitropous, bitegmic or unitegmic. Fruits mul-

tifollicles or (Holodiscus and Adenostoma) achenes.

Seeds mostly with scanty endosperm, rarely

(Physocarpus) endosperm copious. Leaves alternate,

mostly simple, stipulate or estipulate. Shrubs, sub-

shrubs or rarely herbs, n = 9. – neillieae:

Physocarpus, Neillia, Stephanandra; spiraeeae:

Spiraea, Pentactina, Sibiraea, Petrophytum, Kelseya,

Luetkea, Aruncus, Pleiosepalum; gillenieae:

Gillenia, Spiraeanthus; holodisceae: Holodiscus,

Xerospiraea; sorbarieae: Sorbaria, Chamaebatiaria;

adenostomateae: Adenostoma.

1.2 LYONOTHAMNOIDEAE

Carpels two, free, glandular, with four ovules in each

carpel. Leaves opposite, entire or deeply divided, pinnate

and fernlike, with small, caducous stipules. Fruits

woody. Seeds apiculate. Trees or shrubs with deciduous

bark. n = 27. – Lyonothamnus.

320 Subclass VI. ROSIDAE

1.3 FILIPENDULOIDEAE

Carpels 5–15, free, sessile, or stipitate. Ovules two per

carpel, pendulous, campylotropous, epitropous, uniteg-

mic. Fruitlets 1-seeded achenes. Seeds with very scanty

endosperm. Leaves alternate, the basal ones imparipin-

nately compound, cauline leaves similar, but becoming

smaller and less compound toward the stem apices;

stipules of the basal leaves adnate to the petioles, those

of the cauline leaves free, large, asymmetrically

reniform, clasping. Tall, rhizomatous perennial herbs,

n = 7, 8. – Filipendula.

1.4 KERRIOIDEAE

Carpels 2–8, mostly 4–5. Ovules two (Rhodotypeae)

or one (Kerrieae), pendulous or (Kerria) laterally

attached to the middle of the suture, anatropous,

epitropous, unitegmic. Fruits achenes (in Neviusia

achenes are drupaceous). Seeds with copious

(Rhodotypos) or scanty endosperm. Leaves alternate

or (Rhodotypos) opposite, simple, stipulate. Deciduous

shrubs, n = 9. – rhodotypeae: Rhodotypos; kerrieae:

Kerria, Neviusia.

1.5 COLEOGYNOIDEAE

Carpel solitary, sessile in the base of a leathery ﬂ oral

tube; style lateral, longhaired especially below, twisted,

persistent. Ovule one. Flowers apetalous. Fruits coria-

ceous, glabrous, more or less crescent-

shaped achenes. Seeds with large embryo and rather

scanty endosperm. Leaves opposite, small, entire, with

small persistent stipules; hairs mediﬁ xed. Much

branched spinescent shrubs. – Coleogyne.

1.6 RUBOIDEAE

Carpels mostly numerous, free, inserted on a nearly ﬂ at

to cylindrical receptacle. Ovules two per carpel,

collateral or superposed, pendulous, anatropous, epitro-

pous, unitegmic. Fruitlets 1-seeded drupelets crowned

on the dry or spongy, often elongated receptacle. Seeds

with small embryo and rather abundant endosperm.

Leaves alternate, simple, lobed, 3–5- foliolate or impar-

ipinnate, with stipules adnate to the petiole. Mostly

erect to trailing shrubs or scramblers, rarely herba-

ceous, n = 7. – Rubus (including Dalibarda ?).

1.7 ROSOIDEAE

Carpels numerous, free, borne on the inner surface of

an invaginated receptacle near the bottom of the hollow

center. Ovules one per carpel, pendulous, anatropous,

unitegmic. Fruits numerous achenes enclosed by the

enlarged, ﬂ eshy receptacle. Seeds with large embryo

and scanty endosperm. Leaves alternate, imparipinnate

or trifoliolate, rarely 1-foliolate or reduced to connate,

leaﬂ ike stipules; stipules adnate to the petioles and

forming wings. Erect, trailing, or scandent shrubs,

mostly armed with prickles, n = 7. – Rosa, Hulthemia.

1.8 POTENTILLOIDEAE

Carpels numerous, free, borne on convex, conical, or ellip-

soid receptacle. Ovule one per carpel, anatropous, biteg-

mic (Dryas and possibly some others) or more often

unitegmic. Fruits achenes. Seeds with endosperm 7–8

cells thick (Dryas) or more often endosperm 1-cell thick.

Leaves alternate or (Potaninia) opposite, simple or com-

pound, stipulate. Shrubs or more often perennial herbs,

rarely small trees, n = 7, 8, 9. – dryadeae: Dryas, Fallugia;

purshieae: Chamaebatia, Cowania, Purshia; geeae:

Geum (including Acomastylis, Oreogeum, Parageum),

Oreogeum, Orthurus, Taihangia, Novosieversia, Sieversia,

Woronowia, Waldsteinia, Coluria; cercocarpeae:

Cercocarpus; potentilleae: Potentilla (including

Pentaphylloides), Tylosperma, Comarum, Chamaerhodos,

Duchesnea, Fragaria, Horkelia, Horkelliella, Ivesia,

Sibbaldia (including Sibbaldianthe); alchemilleae:

Alchemilla, Aphanes; sanguisorbeae: Agrimonia,

Aremonia, Spenceria, Hagenia, Leucosiaea, Sanguisorba,

Poterium, Sarcopoterium, Margyricarpus, Tetraglochin,

Acaena, Polylepis, Cliffortia, Bencomia; potaninieae:

Potaninia.

1.9 DICHOTOMANTHOIDEAE

Carpel solitary at the base of the calyx tube, with

tomentose lateral stylodium. Ovary superior. Ovules

two, erect, subbasal, anatropous, bitegmic. Floral tube

free, ﬂ eshy, and enlarged in fruit. Fruits achenes, often

1-seeded, with coriaceous pericarp surrounded by

thick ﬂ eshy ﬂ oral tube. Seeds with endosperm 1 cell

thick. Leaves alternate, simple, entire. Trees, n = 17. –

Dichotomanthes.

1.10 PYROIDEAE (Maloideae)

Carpels 2–5, rarely one, more or less adnate to the ﬂ o-

ral tube and partly to the invaginated receptacle. Ovules

(1)2-many per carpel, anatropous, bitegmic or rarely

(Raphiolepis, Chamaemeles) unitegmic. Fruits usually

consist of more or less well-developed and usually

ﬂ eshy parenchymatous pericarp (with groups of

brachysclereids in Cydonia and Pyrus) and carpellar

Superorder ROSANAE 321

endocarps consisting of elongated scleroids. In Maleae

the inner walls of the carpels become chartaceous or

cartilaginous, rarely bony, and the carpels laterally

connate; in Crategeae the inner walls of the carpels

become hard and each carpel develops into a separate

pyrene. Seeds with endosperm 2–15 cells thick (up to

15 cells in Crataegus). Leaves alternate, mostly simple,

stipulate. Shrubs or trees, n = 17 or (Vauquelinia) 15. –

kageneckieae: Kageneckia; lindleyieae:

Vauquelinia, Lindleya; maleae: Photinia (including

Stranvaesia), Heteromeles, Eriobotrya, Rhaphiolepis,

Sorbus, Chamaemespilus, Aronia, Amelanchier, Pyrus,

Malus, Eriolobus, Peraphyllum, Docynia, Cydonia,

Pseudocydonia, Chaenomeles; crataegeae: Cotoneaster,

Malacomeles, Chamaemeles, Pyracantha, Crataegus,

Mespilus, Hesperomeles, Osteomeles.

1.11 AMYGDALOIDEAE

Carpel solitary or sometimes two (Maddenia) or two

(Oemleria and Exochorda) carpels, free from the ﬂ oral

tube; stylodium terminal or (Prinsepieae) lateral or sub-

basal. Ovules two per carpel, pendulous, anatropous,

more or less hemitropous, or (Prinsepieae) campylotro-

pous, bitegmic. Fruits mostly 1-seeded drupes, the meso-

carp ﬂ eshy and indehiscent to dryish and dehiscent, the

endocarp hard or (Prinsepieae) coriaceous, indehiscent.

Seeds with endosperm 1 to 15 cells thick. Leaves alter-

nate, simple, stipulate. Trees or shrubs, n = 8. – osma-

ronieae: Oemleria; exochordeae: Exochorda;

amygdaleae: Maddenia, Pygeum, Laurocerasus,

Padus, Cerasus, Padellus, Prunus, Armeniaca, Persica,

Amygdalus; prinsepieae: Prinsepia.

Doubtfully Rosaceous genus: Guamatela.

The Spiraeoideae represent the least advanced

subfamily and are nearest to the ancestral stock.

It is widely accepted that all other subfamilies are

offshoots from a basically spiraeoid-like stock. The

nearest to the Spiraeoideae are Lyonothamnoideae and

Quillajoideae. Both of them are taxonomically isolated,

especially the latter. Based on their study of the rbcL

sequence, Morgan et al. (1994) even conclude that

Quillaja is not a member of the family Rosaceae,

which has been accepted by some recent authors,

including Kalkman (2003). As regards Lyonothamnus,

I agree with Morgan et al. (1994:899) that “it may be

described most accurately as an isolated descendant of

the ancestral spiraeoid complex.”

Filipenduloideae occupy a somewhat intermediate

position between them and Rosoideae although, accord-

ing to Hegnauer (1973), because of the presence of

ellagic acid and the absence of cyanogenetic substances,

Filipendula is deﬁ nitely rosoid, not spiraeoid. Nakai

(1943), who established the subfamily Filipenduloideae,

placed it after his Potentilloideae, Ruboideae,

Sanguisorboideae, Cercocarpoideae, and Kerrioideae.

Kerrioideae and the closely related Coleogynoideae

(Golubkova 1991) with their achenous fruits and basic

chromosome number x = 9 are probably also derived

from the spiraeoid complex.

The traditional subfamily Rosoideae has been

divided by Nakai (1943) into a number of segregate

subfamilies, some of which are fairly distinct. His

Filipenduloideae, Kerrioideae, Ruboideae, Rosoideae

s.str., and Potentilloideae I ﬁ nd acceptable. However,

I accept the Potentilloideae in a somewhat broadened

sense and include in them – albeit with some hesita-

tion – also his Sanguisorboideae and Cercocarpoideae

while excluding Coleogyne. The Potentilloideae are

the most heterogeneous subfamily (even sensu Nakai),

and there are some isolated genera, such as Cercocarpus,

Purshia, and other genera with basic chromosome

number x = 9.

Dichotomanthoideae are close to the Pyroideae but

differ in ﬂ ower morphology, in which, according to

Rohrer et al. (1991), they resemble more the

Amygdaloideae.

The Pyroideae have deﬁ nite afﬁ

nities with the

Spiraeoideae, especially with the Sorbarieae, and are

closely linked to them via Vauquelinia, Kageneckia,

and Lindleya. They have been derived from spiraeoid

ancestors by autopolyploidy or allopolyploidy

(Gladkova 1972; Cronquist 1981; Takhtajan 1987,

1997; Morgan et al. 1994).

The Amygdaloideae have probably also originated

directly from the spiraeoidlike ancestor. Exochorda

belongs to the Amygdaloideae (Juel 1918; Goldblatt

1976; Takhtajan 1987, 1997; Thorne 1992, 2000;

Morgan et al. 1994) although it differs in having follicu-

lar fruits, winged seeds, and in its chemical makeup.

However, according to Challice (1981), the presence of

arbutin in Exochorda indicates that this genus should

perhaps remain in the Spiraeoideae and not be trans-

ferred to the Amygdaloideae (where arbutin is absent).

2. QUILLAJACEAE

D. Don 1831. 1/2–3. Peru, Chile, southern Brazil.

322 Subclass VI. ROSIDAE

Quillaja.

Quillaja had typically been placed in Rosaceae,

sometimes as a separate subfamily (Takhtajan 1987,

1997), because of several shared features, including

follicular fruits and winged seeds. However

Quillajaceae differ from Rosaceae in valvate sepals,

pollen grains (Claxton et al. 2005), the base chromo-

some number (n = 14), which is not found elsewhere in

Rosaceae, and in the presence of saponins and

trihydroxy-substituted ﬂ avonoids. In analyses based

on rbcL sequence data, Quillaja appeared as sister to

Polygalaceae, but these relationship was not strongly

supported (Savolainen et al. 2000).

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