Takhtajan Armen. Flowering Plants

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Superorder RUTANAE 383

usually shorter than sepals, mostly keeled inside, per-

sistent, becoming more or less ﬂ eshy in fruits. Stamens

ten, in two cycles; ﬁ laments ﬁ liform, free or those

opposite the petals, adnate to petal keel; anthers large,

oblongish, tetrasporangiate, basiﬁ xed or slightly dorsi-

ﬁ xed, opening longitudinally, sometimes apiculate.

Tapetum secretory. Microsporogenesis simultaneous.

Pollen grains 3-celled, with (2)3(4)-colporate or

3(4)-pororate. Gynoecium of ﬁ ve or seldom ten

carpels; carpels free or sometimes basally united,

adnate to the shortly columnar receptacle, each with a

long slender ventral stylodium that is papillate, stig-

matic all around for most of its length, usually diver-

gent; each carpel with one pendulous ovule. Ovules

anatropous, with dorsal raphe, bitegmic, crassinucel-

late, micropyle endostomal. Female gametophyte or

Polygonum-type. Endosperm nuclear. Fruits of sepa-

rate hard-walled 5–10 achene-like mericarps, more or

less enclosed by the much accrescent, ﬂ eshy petals.

Seeds small, rather compressed, red-brown; exotesta

of cuboid, thick-walled; embryo straight, oily;

cotyledons two, large, plano-convex; endosperm scanty

or none. Contain bitter sesquiterpenoid substances and

producing kaempferol and quercetin, ellagic and often

gallic acid, strongly tanniferous. n = 10, 15.

A rather isolated taxon that is one of the most

controversial with respect to its afﬁ nities. Shows some

similarities with the Rutales, especially with the

Simaroubaceae, which is supported also by wood ana-

tomical studies (Carlquist 1985), embryology (Kamelina

1991), and chemical data (Bohm and Ornduff 1981).

1. CORIARIACEAE

A. P. de Candolle 1824. 1/15–20. Western Mediter-

ranean, temperate and subtropical Himalayas to Japan

and New Guinea, New Zealand, South Paciﬁ c islands

(including Solomons, Vanuatu, Fiji, Samoa, Society

Islands), America from Mexico to Chile.

Coriaria.

Bibliography

Bohm BA and R Ornduff. 1981. Leaf ﬂ avonoids and ordinal

afﬁ nities of Coriariaceae. Syst. Bot. 6: 15–26.

Carlquist S. 1985. Wood anatomy of Coriariaceae: Phylogenetic

and ecological implications. Syst. Bot. 10: 174–183.

Duyjes BEE. 1993. Coriariaceae. In: CGGJ van Steenis, ed.

Flora Malesiana ser. 1, 11: 385–391. Noordhoff, Groningen.

Garg M. 1981. Pollen morphology and systematic position of

Coriaria. Phytomorphology 30: 5–10.

Good RDO. 1930. The geography of the genus Coriaria. New

Phytol. 29: 170–198.

Kamelina OP. 1991. Comparative-embryological analysis as a

method of phylogenetic systematics of ﬂ owering plants.

Ph.D.. Thesis. Tashkent.

Kravtsova TI. 1996. Coriariaceae. In: A Takhtajan, ed.

Comparative seed anatomy, vol. 5, pp. 472–473. Nauka, St.

Petersburg (in Russian).

Lobreau D. 1969. Les limites de 1′ “ordre” des Celastrales

d’apres le pollen. Pollen Spores 11: 499–555.

Matthews ML and PK Endress. 2004. Comparative ﬂ oral struc-

ture and systematics in Cucurbitales (Corynocarpaceae,

Coriariaceae, Tetramelaceae, Datiscaceae, Begoniaceae,

Cucurbitaceae, Anisophylleaceae). Bot. J. Linn. Soc. 145:

129–185.

Petseon B. 1970. Coriariaceae: Taxonomy and relationships.

World Pollen Flora 1: 23–24.

Praglowski J. 1970. Coriariaceae. World Pollen Flora 1: 17–31.

Sharma VK. 1968. Floral morphology, anatomy, and embryol-

ogy of Coriaria nepalensis Wall. with a discussion of the

interrelationships of the family Coriariaceae.

Phytomorphology 18: 143–153.

Skog LE. 1972. The genus Coriaria in the Western Hemisphere.

Rhodora 74: 242–253.

Tobe H and N Suzuki. 1993. Occurrence of uni- and bitegmic

ovules in Coriaria (Coriariaceae), and evolutionary relation-

ships in the genus. Abstracts XV Intern. Bot. Congr., p. 95.

Yokohama.

Tobe H, M Suzuki, and T Fukuhara. 1992. Pericarp anatomy and

evolution in Coriaria (Coriariaceae). Bot. Mag. (Tokyo) 195:

289–302.

Yoda K and M Suzuki. 1992. Comparative wood anatomy of

Coriaria. Bot. Mag. (Tokyo) 105, 1078: 235–245.

Yokoyama J and M Hasebe. 2001. Phylogeny and biogeography

of Coriaria based on molecular data. Did Coriaria originate

in the southern hemisphere? Proc. Jpn. Soc. Plant Taxon. 16:

43–48.

Order 96. CORYNOCARPALES

Evergreen trees or shrubs; young stems with separate

bundles. Vessels with simple perforations, lateral pit-

ting alternate. Fibers with simple or faintly bordered

pits. Rays very broad (up to 16 cells wide). Axial

parenchyma vasicentric and in broad paratracheal

strips. Cambium storied. Sieve-element plastids of

S-type. Nodes trilacunar. Leaves alternate, condupli-

cate, coriaceous, more or less crowded towards the

ends of the shoots, shining; stipules intrapetiolar, sub-

tending the axillary bud, deciduous. Stomata paracytic.

Flowers in terminal panicles and in racemes, bisexual,

384 Subclass VI. ROSIDAE

actinomorphic, 5-merous. Plants anemophilous. Sepals

free, strongly imbricate, ﬂ eshy. Petals free, imbricate,

inserted at the margin of the disc-like receptacle.

Stamens ﬁ ve, incurved in bud, opposite and basally

slightly adnate to the petals, alternate with ﬁ ve peta-

loid staminodes attached to the outer surface of the

nectaries; ﬁ laments ﬂ attened or terete; anthers dorsi-

ﬁ xed, tetrasporangiate, introrse, opening longitudinally.

Tapetum secretory. Microsporogenesis simultaneous.

Pollen grains nearly smooth, 2- colporate, with a short

colpus at each end. Large cushionlike ﬁ ve nectaries

borne opposite and internal to petaloid scales (stamin-

odes). Gynoecium pseudomonomerous, reduced to

one fertile carpel with more or less well-expressed

trace of a second carpel and very rarely the rudiment of

the second stylodium; styles short, conduplicate,

stigma capitate; ovary superior, 1–2-locular, solitary

ovule pendulous, anatropous, bitegmic, crassinucel-

late, with micropyle directed upwards. Female

gametophyte of Polygonum-type. Endosperm nuclear.

Fruits ﬂ eshy drupes, with ﬁ brous endocarp. Seeds with

large, straight, oily, and starchy embryo, minute radi-

cle and plumule; endosperm starchy; cotyledons two,

very large. Plants containing very toxic bitter glyco-

sides, at least in the bark and seed; present kaempferol

and ellagic acid. n = 22, 23.

Both structural similarities with Coriariaceae

(Matthews and Endress 2004) and molecular studies

(Nandi et al. 1998; Savolainen et al. 2000; Soltis et al.

2000; Zhang and Renner 2003) support the afﬁ nity

with Coriariaceae

1. CORYNOCARPACEAE

Engler 1897. 1/5–7. Aru Islands, New Guinea, Bismarck

Archipelago, New Britain, New Ireland, Solomon

Islands, eastern Australia, New Caledonia, Vanuatu,

New Zealand.

Corynocarpus.

Bibliography

Carlquist S and RB Miller. 2001. Wood anatomy of

Corynocarpaceae is consistent with Cucurbitalean place-

ment. Syst. Bot. 26: 54–65.

Dawson MI. 1997. Chromosome numbers in Corynocarpus

(Corynocarpaceae). New Zealand J. Bot. 35(2): 255–258.

Hemsley WB. 1903. On the genus Corynocarpus Forst., with

descriptions of two species. Ann. Bot. 17: 179–180.

Ingle HD. 1956. A note on the wood anatomy of the genus

Corynocarpus. Trop. Woods 105: 8–12.

Matthews ML and PK Endress. 2004. Comparative ﬂ oral struc-

ture and systematics in Cucurbitales (Corynocarpaceae,

Coriariaceae, Tetramelaceae, Datiscaceae, Begoniaceae,

Cucurbitaceae, Anisophylleaceae). Bot. J. Linn. Soc. 145:

129–185.

Melikian AP and IA Savinov. 2000. Corynocarpaceae. In:

A Takhtajan, ed. Comparative seed anatomy, vol. 6, pp.

140–142. Nauka, St. Petersburg (in Russian).

Nandi O, MW Chase and PK Endress. 1998. A combined cladis-

tic analysis of angiosperms using rbsL and non-molecular

data sets. Ann. Missouri Bot. Gard. 85: 137–212.

Narayana LL, PS Narayana, and SR Vatsavaya. 1986. A contri-

bution to the numerical chemotaxonomy of Corynocarpaceae.

J. Econ. Taxon. Bot. 8: 249–254.

Nowicke JW and JJ Skvarla. 1983. Pollen morphology and the

relationships of the Corynocarpaceae. Taxon 32: 176–183.

Patel RN. 1975. Wood anatomy of the dicotyledons indigenous

to New Zealand: 9. Corynocarpus. New Zealand J. Bot. 13:

19–29.

Philipson WR. 1987. Corynocarpus J. R. and G. Forst.: An iso-

lated genus. Bot. J. Linn. Soc. 95: 9–18.

Soltis DE, PS Soltis, DR Morgan, SM Swensen, BC Mullin,

JM Dowd, and PG Martin. 1995. Chloroplast gene sequence

data suggest a single origin of the predisposition for symbi-

otic nitrogen ﬁ xation in angiosperms. Proc. Nat. Acad. Sci.

USA 92: 2647–2651.

Steenis van CGGJ. 1952. Corynocarpaceae. In: CGGJ van

Steenis, ed. Flora Malesiana, ser.1, vol.4, pt.3: 262–264.

Noordhoff, Groningen.

Swensen SM. 1996. The evolution of actinorhizal symbioses:

evidence for multiple origins of the symbiotic association.

Am. J. Bot. 83: 1503–1512.

Wagstaff SJ and MI Dawson. 2000. Classiﬁ cation, origin, and

patterns of diversiﬁ cation of Corynocarpus (Corynocarpaceae)

inferred from DNA sequences. Syst. Bot. 25: 134–149.

Order 97. GERANIALES

Mostly herbs, less often deciduous and evergreen trees,

shrubs, subshrubs. Vessels typically with simple

perforations; lateral pitting alternate, scalariform or

transitional. Fibers with simple pores, often septate.

Rays heterogeneous or seldom nearly homogeneous,

often wanting. Axial parenchyma paratracheal and

also with a few diffusely scattered cells. Sieve-element

plastids of S-type or Plc-type. Nodes trilacunar or

rarely unilacunar, pentalacunar or multilacunar

(Melianthaceae). Leaves alternate or less often

opposite, usually compound or dissected or more or

less lobed, rarely simple and entire, stipulate or

estipulate. Stomata paracytic or more often anomo-

Superorder RUTANAE 385

cytic, or diacytic (Francoa). Flowers usually in cymose

inﬂ orescences or in terminal racemes or panicles, less

often solitary and axillary, bisexual or rarely unisexual,

actinomorphic or rarely zygomorphic, 5-merous or

seldom 4-merous, rarely 3-merous. Sepals free or

basally connate, imbricate, valvate, or contorted,

sometimes persistent. Petals free or basally connate,

imbricate or contorted, rarely wanting. Nectary disc

represented by 5 extrastaminal glands or wanting; in

Francoaceae nectary disc interstaminal. Stamens

1–2(3) times as many as the sepals or petals, free or

more often with basally connate; ﬁ laments sometimes

subulate; anthers slightly introrse or latrorse, basiﬁ xed

or dorsiﬁ xed, tetrasporangiate, opening longitudinally.

Tapetum secretory or plasmodial. Microsporogenesis

simultaneous. Pollen grains 2-celled or 3-celled,

3- colpate or 3- colporate, sometimes (Vivianiaceae and

Rhynchothecoideae) pantoporate or (Ledocarpoideae)

inaperturate. Gynoecium of (2-)5(6) carpels; stylodia

free or more or less connate; stigma small, capitate,

shortly 5-lobed, or commisural (Francoaceae). Ovary

superior, or (Greyiaceae and Francoaceae), with

(1)2-several to numerous, more or less pendulous

ovules in each locule. Ovules anatropous to campy-

lotropous, sometimes hemitropous, epitropous, with

the micropyle directed upward, bitegmic, crassinucel-

late. Female gametophyte of Polygonum-type or very

rarely of Allium-type. Endosperm nuclear. Fruits of

various types. Seeds usually exotegmic, rarely endote-

gmic (Ledocarpaceae) or seed coat derived from both

integuments and not specialized (Rhynchothecoideae

and Vivianiaceae, except seeds of Caesarea which are

exotegmic – Boesewinkel, Bouman 2000); embryo

straight or curved; endosperm copious or scanty, some-

times wanting.

Have a common origin from a rutalean ancestor.

Key to Families

1 Nodes trilacunar or rately unilacunar.

2 Extrastaminal nectary glands usually present.

3 Fruits loculicidal capsules. Leaves estipulate.

4 Androecium of 15 stamens in two cycles

(10 short antepetalous stamens and 5 long

antesepalous ones); stamens with persistent

subulate ﬁ laments and dorsiﬁ xed, introrse

anthers. Pollen grains 3-colporate.

Acaulescent perennial herbs from thick tap-

root. Leaves rosulate, alternate, simple, pin-

natiﬁ d. Sieve-element plastids of S-type.

Inﬂ orescences radical, 1–9-ﬂ owered cymes.

Flowers bisexual, actinomorphic, 5-mer-

ous. Calyx strongly imbricate, persistent.

Corolla contorted. Pollen grains tricolpate.

Extrastaminal lobed nectary disk well

developed. Gynoecium of ﬁ ve carpels

attached to central column; stylodia united

into a simple ﬁ liform style with capitate

stigma; ovary 5-locular and 5-lobed, with

about 12–50 axile, biseriate ovules in each

locule; ovules anatropous to campylotro-

pous (becoming campylotropous after fer-

tilization, via a bulge from the inner

integument on the raphal side); micropyle

zig-zag. Fruits tardily and irregularly locu-

licidal capsules or rarely (H. tridentata)

schizocarp but the mericarps are not beaked.

Seeds with cochlear embryo with spirally

folded cotyledons and scanty endosperm;

seed coat with an exotegmen consisting of

stellately lobed cells and a crystalliferous

endotesta. . . . . . . 1. hypseocharitaceae.

4 Androecium of 10 or 8, sometimes 5 sta-

mens, equal or unequal. Pollen grains peri-

porate. Perennial herbs, sometimes with

extensive secondary growth, very rarely

annuals. Hairs glandular. Nodes unilacunar

with one trace. Leaves opposite, entire, cre-

nate to coarsely dentate, often more or less

white-tomentose below. Flowers in termi-

nal or apparently lateral, one to many-ﬂ ow-

ered cymes, bisexual, actinomorphic,

mostly 5-merous. Sepals prominently

3-veined, more or less connate, lobes val-

vate. Petals free, imbricate. Nectary glands

as many as and alternate with the petals,

entire or 2-lobed. Gynoecium of 3(2) car-

pels. Stylodia almost free or connate, the

stigmatic surface papillose, often extended

to cover all or part of the adaxial surface;

ovary 3(2)-lobed, with two superposed,

anatropous, axile ovules (1 ascending, 1

pendulous) in each carpel. Fruits 3(2)-lobed

loculicidal capsules. Seeds with much

curved or circinate green embryo and abun-

dant ﬂ eshy endosperm, seed coat derived

from both integuments and not specialized

(except seeds of Caesarea which are exote-

gmic), n = 7. . . . . . . . . . . 3. vivianiaceae.

386 Subclass VI. ROSIDAE

3 Fruits schizocarps. Stipules present. Herbs or

less often subshrubs or shrublets, sometimes

(Sarcocaulon) with ﬂ eshy stems and bearing

essential oils. Hairs glandular. Sieve-element

plastids of S-type. Nodes trilacunar with three

traces. Leaves alternate (the upper, often) or

opposite (the lower, usually), mostly lobed,

dissected or compound, petiolate, often gland-

dotted and aromatic; stipules well developed,

interpetiolar or intrapetiolar, usually twinned

at the base of the petiole. Stomata anomacytic.

Inﬂ orescences terminal or axillary, or leaf-

opposite, often pedunculate, usually consisting

of paired ﬂ owers or umbels, commonly with

involucral bracts. Flowers bisexual, actinomor-

phic or slightly zygomorphic (Pelargonium),

5-merous. Sepals free or connate to the mid-

dle, imbricate with valvate tips, persistent.

Petals 2–5, free, imbricate or rarely contorted,

caduceus, rarely absent. Stamens 5, or 10, or

15, sometimes a few sterile, mostly more or

less connate at the base, obdiplostemonous

when more than one cycle. Anthers dorsiﬁ xed,

usually versatile, introrse. Pollen grains 3-col-

pate (Sarcocaulon and Monsoa) or more often

3-colporate, tectate- columellate, reticulate to

striate. Gynoecium of 5 carpels; style with 5–3

stigmatic branches; ovary 5-locular, 5–3-

lobed, with 1–2 pendulous or ascending, anat-

ropous to often campylotropous (becoming

campylotropous after fertilization) ovules in

each locule; micropyle zig-zag. Fruits of ﬁ ve

1-seeded mericarps that separate elastically

from a central beak (the persistent style), the

mericarps often opening to discharge the seed.

Seeds with usually or more or less curved

embryo with green cotyledons; endosperm

much reduced or absent; seed coat with

exotegmen consisting of thick-walled cells.

Containing hydrolysable tannins, ﬂ avonols

(kaempferol, quercetin, and myricetin), com-

monly alkaloids, and sometimes ellagic acid,

n = mostly 10 (Erodium), mostly 11 (Monsonia,

Sarcocaulon, Pelargonium), or mostly 14

(Geranium).. . . . . . . . . . . . . 2. geraniaceae.

2 Extrastaminal nectary glands absent. Small

shrubs or shrublets. Leaves usually opposite,

simple or compound, entire or deeply lobed or

dissected, estipulate. Flowers solitary or in few-

ﬂ owered corymbs or terminal racemes, bisexual,

actinomorphic, 5-merous. Sepals free, imbricate.

Calyx with an epicalyx of a whorl of bracteoles

(Ledocarpoideae), or epicalyx wanting

(Rhynchotheca). Petals imbricate (Wendtia) or

contorted (Balbisia), or absent (Rhynchotheca).

Stamens ten, free, alternisepalous; anthers

extrorse or introrse. Pollen grains inaperturate,

periporate (Rhynchotheca). Gynoecium of ﬁ ve

(Balbisia and Rhynchotheca) or three (Wendtia)

carpels. Stylodia free, short (Wendtia), or stig-

mas sessile (Balbisia); ovary 3- or 5-locular,

ovules 1–2 (Wendtia, Rhynchotheca), or 5–50

(Balbisia) per locule, pendulous, anatropous to

campylotropous, with ventral raphe. Fruits cap-

sules, valvular (‘septifragal’) or loculicidal.

Seeds endotegmic, embryo straight, with thin

endosperm, n = 9. . . . . . . . . 4. ledocarpaceae.

1 Nodes pentalacunar or multilacunar.

5 Stipules intrapetiolar, relatively small, papery

and caduceus (Bersama) or often large, showy

and persistent. Capsules loculicidal. Flowers

more or less zygomorphic, bisexual or polyga-

mous, in axillary or terminal racemes, often

large, resupinate by twisting of the pedicel.

Sepals 5 or 4, unequal, free or basally connate,

imbricate. Petals 4–5, free, clawed, ﬁ ve and

unequal or one abortive. Nectary disc extrastami-

nal, unilateral, well developed. Stamens 4–5,

sometimes 8, often declinate, free or sometimes

basally connate; anthers dorsiﬁ xed, slightly ver-

satile, introrse. Pollen grains 3-celled, 3-colpo-

rate. Gynoecium of 4(5) carpels; style with four

to ﬁ ve-lobed stigma; ovary superior, slightly

sunken, 4–5-locular, each locule with

1 basal (Bersama) or 2–5 axile (Melianthus),

erect to pendulous ovules; ovule of Bersama may

smell of mustard, and ovule of Melianthus –

smells nasty. Fruits capsules, opening loculicid-

ally or only at the apex, in Melianthus papery or

coriaceous, inﬂ ated and longitudinally deeply

sulcate and sometimes apically lobed, in Bersama

tough-walled, often woody, dehiscing

longitudinally by 4–5 valves. Seeds exotestal, the

tegmen is crushed and unligniﬁ ed; embryo small

or large, straight, endosperm copious, starchy or

oily, exarillate in Melianthus and subtended by a

large

yellowish aril in Bersama. Characteristic

anatomical feature is the presence of styloids in

Superorder RUTANAE 387

vegetative parts and sometimes also in the seed

coat. Nodes penta- or multilacunar. Evergreen

subshrubs (Melianthus), or gard-wooded shrubs,

or small trees. Leaves alternate, imparipinnate,

or occasionally trifoliate, conduplicate, strong

odour. Stomata anomocytic. Toxic principles,

known as bufadienolides, present in both genera.

Producing ﬂ avonols (quercetin), ellagic acid;

present traces of inulin and elongated prismatic

crystals. n = 18, 19.. . . . . . .5. melianthaceae.

5 Stipules absent, capsules septicidal.

6 Leaves simple, slightly peltate, crenately or

dentately lobed. Soft-wooded shrubs or small

trees with leaﬂ ess older branchlets covered

with a thin layer of periderm. Internodes green

and photosynthetic, glabrous to densely pubes-

cent, often with minute glandular hairs. Vessel

elements very short; lateral pitting scalariform

or transitional. Rays composed entirely of

square and upright cells containing raphides.

Leaves crowded apically on the branchlets,

alternate, petiolate, coriaceous, palmately or

subpalmately veined, either essentially gla-

brous or with a dense indumentum of long,

multicellular, uniseriately branched hairs

when young, often becoming glabrous with

age, minute glandular hairs present, producing

a shiny layer of glutinous exudate; petiole dis-

tinctly expanded basally into a decurrent

sheath fused with the preceding internode,

forming a pseudocortex; stipulelike out-

growths may occur on the sheath lobes; at the

end of the growing season the sheath, petiole,

and lamina are detached together, which is ini-

tiated by the formation of a periderm at the

junction of the pseudocortex and the real cor-

tex. Stomata anomocytic. Flowers conspicu-

ous, in terminal racemes eventually displaced

laterally by new leafy shoots emerging below

the peduncle, bracteolate, bisexual, actino-

morphic to slightly zygomorphic, 5-merous,

or occasionally 4-merous, pollinated by sun-

birds. The bracts, sepals and petals densely

covered with short trichomes with globose

glandular head cells. Sepals ﬁ ve, free or

shortly connate, imbricate, persistent. Petals

ﬁ ve, free, imbricate, much longer than the

sepals, minutely ciliate, partly reﬂ exed or

(Greyia ﬂ anaganii) loosely coherent apically,

brilliant red or crimson, rarely white, cadu-

cous. Extrastaminal nectary disc well devel-

oped, connected by ﬁ ve interstaminal ridges

with the base of the ovary, ﬂ eshy, cupular, with

the margin crowned by ten small vascularized

staminodia the distal portions of which are

subpeltate with a glandular area at the apex or

irregularly laceratedentate at the apex, produc-

ing copious nectar. Stamens ten, free, seem-

ingly placed in a single series but most

probably obdiplostemonous (Schonland 1914;

Dahl gren and van Wyk 1988), episepalous sta-

mens maturing ﬁ rst; ﬁ laments subulate; anthers

fairly large, with very narrow connective pro-

trusion, slightly introrse. Gynoecium of (4)5(6)

united carpels with stylodia connate into a

slender, subulate style; stigma small, shortly

5-lobed, pappilae on stigmatic cleft. Ovary

semi-inferior “because of growth of the periph-

eral tissue: (Ronse Decraene and Smets 1992:

389), deeply (4)5(6)-lobed, with slightly biﬁ d

parietal placentas developing from the intru-

sive margins of the carpels and eventually

meeting and forming the placental column

around a central hollow; the borderline between

the individual carpel margins remains clearly

distinguishable (Steyn 1977), and therefore the

ovary is essentially unilocular. Ovules numer-

ous, in two rows on each placenta. Fruits cap-

sules, mericarps ﬁ ve, comprising follicles, the

mature carpels separating from the central col-

umn or hollow, and opening ventrally. Seeds

numerous per locule, minute, elongate; seed

coat membranous, formed by both integu-

ments, the exotegmen and next layer is ﬁ brous,

but although the mesotegmic cells elongate;

embryo small, straight; endosperm copious,

ﬂ eshy. Producing ellagic acid and quercetin,

but not myricetin; present durses and raphids;

n = 16 or 17 . . . . . . . . . . . . . . .6. greyiaceae

6 Leaves lyrate-pinnatiﬁ d to pinnate or rounded-

cordate. Perennial rhizomatous, scapose herbs,

glandular-pilose or tomentose. Nodes trilacunar

with ﬁ ve traces. Leaves crowded or subradical,

conduplicate-plicate, pinnately veined (Francoa)

or palmately veined (Tetilla), estipulate.

Stomata (in Francoa) diacytic. Flowers in lax,

elongated racemes, bracteate, bisexual, actino-

morphic, sometimes zygomorphic, usually

388 Subclass VI. ROSIDAE

4-merous, but occasionally 5-merous. Sepals

(4)5, persistent, valvate, equal (Francoa) or

(Tetilla) unequal with two much smaller or

deﬁ cient; petals clawed or sessile, inserted at

the base of the calyx, imbricate or contorted.

Stamens 8 (Tetilla) or 4–5 (Francoa), alternat-

ing with as many staminodia; ﬁ laments free;

anthers oblong or subglobose, latrorse. Nectary

disc interstaminal, covered with nectarosto-

mata. Pollen grains 3-colporate. Gynoecium of

four (rarely two) carpels; stigmas four, rarely

two, sessile, commissural (stand over the “com-

missures,” lines of fusion of the adjacent car-

pels). Ovary semi-inferior, tetragonous, 4-lobed

at the apex, 4-locular or very rarely 2-locular;

stigma papillae on stigmatic cleft; ovules

numerous, biseriate. Fruits oblong, erect,

4- locular, septicidal (Tetilla) or loculicidal

(Francoa) capsules with numerous seeds. Seeds

very small, striolate; exotestal cells elongated,

thickened, endotesta elongated, with thickened

anticlinal walls, tegmen of pigmented cells;

embryo, minute, straight, in the copious, ﬂ eshy

endosperm. Present ellagic acid (strongly –

Francoa) and anthocyanin in roots, and traces

of inulin; n = 13. . . . . . . . . . 7. francoaceae.

1. HYPSEOCHARITACEAE

Weddell 1861. 1/6. Andes of South America from Peru

to Chile and northern Argentina.

Hypseocharis.

Hypseocharitaceae close related to the Geraniaceae

and their seeds have many common features

(Boesewinkel and Bauman 2000) In habit Hypseocharis

somewhat resembles Erodium, and its staminal

arrangement compares well with that of Monsonia and

Sarcocaulon (Rama Devi 1991), but it differs from the

Geraniaceae in its capsular fruits and estipulate leaves,

as well as in its cochlear embryo with spirally folded

cotyledons (Boesewinkel 1988).

2. GERANIACEAE

A. L. de Jussieu 1789 (including Erodiaceae Horaninow

1847). 4/c.830. Subcosmopolitan, but mainly in tem-

perate and subtropical regions.

Geranium, Erodium (including California),

Pelargonium, Monsonia (including Sarcocaulon).

Close to both the Hypseocharitaceae and the Vivia-

niaceae but their fruits are schizocarps.

3. VIVIANIACEAE

Klotzsch 1836. 4/6. Southern Brazil, Uruguay,

Argentina, and Chile.

Viviania, Araeoandra, Caesarea, Cissarobryon.

Usually included in the Geraniaceae from which

they differ in capsular fruits, estipulate leaves, and

abundant ﬂ eshy endosperm.

4. LEDOCARPACEAE

Meyen 1834 (including Rhynchothecaceae Endlicher

1841). 3/12. Andian South America.

4.1 LEDOCARPOIDEAE.

Calyx with an epicalyx of a whorl of bracteoles. Small

shrubs or shrublets. Leaves usually opposite, simple

or compound, entire or deeply lobed or dissected,

estipulate. Flowers solitary or in few-ﬂ owered

corymbs or terminal racemes, bisexual, actinomor-

phic, 5-merous. Sepals free, imbricate. Petals imbri-

cate (Wendtia) or contorted (Balbisia). Stamens ten,

free, alternisepalous; anthers extrorse or introrse.

Pollen grains inaperturate. Gynoecium of ﬁ ve

(Balbisia) or three (Wendtia) carpels. Stylodia free,

short (Wendtia), or stigmas sessile (Balbisia); ovary

3- or 5-locular, ovules 1–2 (Wendtia), or 5–50

(Balbisia) per locule, pendulous, anatropous to

campylotropous, with ventral raphe. Fruits capsules,

valvular (‘septifragal’) or loculicidal. Seeds

endotegmic, with thin endosperm, n = 9. – Balbisia

(

Ledocarpon), Wendtia.

4.2 RHYNCHOTHECOIDEAE

Calyx without an epicalyx. Many branched spiny

shrublets with 4-angled branches. Leaves small, entire,

3-ﬁ d or 3-partite. Flowers terminal, sometimes

fasciculate, apetalous. Sepals acuminate. Anthers

large, oblong. Pollen grains periporate. Gynoecium of

ﬁ ve carpels; style short, thick, with ﬁ ve elongate

stigmas; ovules two per locule, pendulous from the

middle of the loculi. Fruits septifragally dehiscent

Superorder RUTANAE 389

capsules, beaked (beaks not revolute). Seeds with

straight embryo and thin endosperm, seed coat derived

from both integuments and not specialized. –

Rhynchotheca.

Related to the Geraniaceae, however, they clearly

differ from the Geraniaceae in absence of nectary

glands, free stamens, inaperturate pollen grains, free

stylodia, capsular fruits, and also endotegmic seeds.

5. MELIANTHACEAE

Berchtold et J. Presl 1820. 2/16 (including Bersamaceae

Doweld 2001). Tropical and South Africa; Melianthus

(8) is endemic to South Africa, often occurring in arid

areas; Bersama (8) is usually conﬁ ned to forest regions

in South and tropical Africa.

Melianthus, Bersama.

Related to the Geraniaceae.

6. GREYIACEAE

Hutchinson 1926. 1/3. South Africa, mainly the eastern

Transvaal escarpment and the Drakenberg Mountains.

Greyia.

Greyiaceae related to the Melianthaceae.

7. FRANCOACEAE

Adr. Jussieu 1832. 2/2. Mountains of middle Chile.

Francoa, Tetilla.

Very closely related to the Greyiaceae.

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Order 98. ZYGOPHYLLALES

Small trees, shrubs, or subshrubs, less often perennial

or annual herbs. Stems often swollen or jointed at the

nodes. Trichomes usually unicellular and simple, more

rarely unicellular and 2-armed, sometimes glandular.

Usually without scattered mucilage cells (except for

spp. of Nitraria) and without secretory cells. All

elements of wood are usually storied, except in

Balanites. Vessel elements very small to moderately

large, very to extremely short, exclusively with simple

perforations and with vestured pits (Carlquist 2005);

lateral pitting alternate and very small and numerous

or (Nitrariaceae) intermediate between opposite or

alternate; rarely (Balanitaceae) intervascular pits ves-

tured. Fibers with bordered or (Tetradiclidaceae) sim-

ple pits. Rays usually narrow and short, homogeneous

or rarely slightly heterogeneous. Axial parenchyma

predominantly apotracheal, almost entirely paratra-

cheal in Bulnesia and Nitraria, scanty metatracheal

and vasicentric in Tetraena. Sieve-element plastids of

S- or less often (some Zygophyllaceae) of Pcs-type.

Nodes trilacunar. Leaves opposite or less often

alternate, mostly paripinnate, often 2-foliolate, less

often simple and entire, very rarely (Pintoa) gland-

dotted, usually with well-developed and commonly

slender stipules (sometimes modiﬁ ed into spines).

Stomata anomocytic or (Balanitaceae) encyclocytic.

Flowers in inﬂ orescences (cymose or seldom race-

mose) or solitary, usually bisexual, commonly

actinomorphic, (4)5(6)-merous. Sepals free or some-

times basally connate, imbricate or valvate. Petals usu-

ally free, imbricate or contorted, rarely valvate,

sometimes wanting. Stamens in (1)2(3) cycles; ﬁ la-

ments often with basal glands; anthers small or

(Peganum) large, tetrasporangiate, dorsiﬁ xed, introrse,

opening longitudinally. Tapetum secretory (becoming

the false periplasmodium in Tetradiclidaceae).

Microsporogenesis simultaneous. Pollen grains

2-celled or rarely 3-celled, of various types, but mostly

3-colporate. Nectary disc usually well developed, more

or less distinctly intrastaminal, sometimes modiﬁ ed

into a gynophore. Gynoecium of ﬁ ve or less often four,

seldom six or only two united carpels, with a slender

(sometimes very short) simple style; stigma capitate or

sometimes lobed or cleft; ovary superior, multilocular,

with one to several (rather numerous in Peganum) pen-

dulous or seldom ascending ovules per locule. Ovules

epitropous or rarely (Nitraria) apotropous, anatro-

pous to sometimes hemitropous, campylotropous, or

orthotropous, bitegmic, crassinucellate, mostly with

an endothelium (except in Nitraria, Peganum,

Malacocarpus, and Seetzenia), sometimes

(Malacocarpus and Seetzenia) with a funicular-pla-

cental obturator. Female gametophyte of Polygonum-

type. Endosperm nuclear. Fruits often capsules or

schizocarps, rarely drupes or berries. Seeds typically

exotestal-endotestal, with endotegmic cells elongate

tangentially; embryo straight or slightly curved, in

hard oily endosperm, which is sometimes absent.

Closely related to the Linales and had a common

origin from a rutalean stock.

Key to Families

1 Leaves usually stipulate.

2 Leaves alternate.

3 Glabrous or hairy perennial herbs (Peganum)

or shrubs (Malacocarpus) with raphids.

Leaves entire or irregularly dissected, leathery

or ﬂ eshy, sessile, simple, with minute seta-

ceous stipules on some leaves (usually absent

in Malacocarpus); glands with a clavate or

spherical head are characteristic of Peganum;

hypoderm present beneath the upper epidermis

392 Subclass VI. ROSIDAE

in Peganum. Flowers solitary and axillary,

bisexual, actinomorphic, 4–5-merous. Sepals

leaﬂ ike, entire or pinnately cleft, valvate.

Petals free, contorted and imbricate. Stamens

12, or 15, in three cycles, attached to the

nectary disc; ﬁ lament dilated at the base.

Pollen grains 3-colporate, with reticulate orna-

mentation. Gynoecium of 3–2 carpels, with

simple cylindrical style continued into long

3–2-keeled stigma papillate along the keels;

ovary superior, 3–2-lobed and 3–2-locular,

with numerous pendulous, anatropous ovules

per locule. Fruits loculicidal capsule (Peganum)

or 2-locular berry (Malacocarpus). Seeds

angular, with spongy, mucilage testa bearing

small, funicular outgrowth; embryo green,

slightly curved, in ﬂ eshy endosperm. Present

alkaloids, n = 12.. . . . . . . . . . 3. peganaceae.

3 Succulent shrubs, sometimes spiny, secretary

cavities sometimes present, with mucilage

cells. Leaves entire or sometimes 2–3 toothed

above, petiolate, simple stipules minute, intra-

petiolar, free of one another, often caduceus or

persistent; very large clusters of crystals occur

in the leaves. Stomata anomicytic. Flowers in

the axils of small caducous bracts, in scorpioid

cymose inﬂ orescences, actinomorphic, 5-mer-

ous. Calyx synsepalous, small, ﬂ eshy, toothed,

imbricate, persistent. Petals free, concave,

conduplicate-valvate. Stamens 15–10, ﬁ ve of

them antesepalous, the other ones in pairs or

solitary antepetalous; ﬁ laments subulate, with-

out basal appendages; anthers versatile. Pollen

grains 3(4)-colporate, with thick ectexine.

Gynoecium of three or rarely of six carpels,

with a continuous ﬂ eshy style; stigma of three

(rarely 6) papillate decurrent ridges; ovary

3(6)-locular, not lobed, with one pendulous, apo-

tropous ovule per carpel. Fruits drupaceous,

1-seeded, derived from single fertile locule, with

ﬂ eshy exocarp with ligniﬁ ed inner part (named

scleromesocarp) and stony thin fovealate endo-

carp (in N. sphaerocarpa the drupe is almost

dry). Seeds with a straight, green embryo, with

the radicle next to hilum; endosperm wanting,

n = 12, 30. . . . . . . . . . . . . . . . . 4. nitrariaceae.

2 Leaves opposite or sometimes alternate, usually

pinnate, rarely 3-foliolate or 2-foliolate or simple

and entire, leathery, ﬂ eshy, or modiﬁ ed into

spines, usually with stipules (sometimes spiny,

or scaly, or leafy, persistent). Stomata mostly

anomocytic. Small trees or shrubs, rarely peren-

nial or annual herbs, resinous or not resinous,

more or less succulent. Wood often ﬂ uorescing;

vessels with simple perforation and usually with

vestured pits. Axial parenchyma diffuse or

diffuse-in-aggregates; rays heterogenerous or

homogeneous; crystals one per cell or septate

portion of cell in wood or secondary phloem

(Carlquist 2005). Nodes often swollen or jointed,

trilacunar with three traces. Flowers in cymose

inﬂ orescences or solitary, terminal or axillary, or

leaf-opposed, bisexual or very rarely unisexual

and dioecious (Neoluederitzia), actinomorphic or

rarely zygomorphic, 5-merous or sometimes

4-merous. Sepals free or less often basally con-

nate, imbricate or valvate. Petals usually free,

imbricate or contorted, rarely valvate, sometimes

wanting (Seetzenia, Miltianthus). Nectary disc

usually present, extrastaminal or intrastaminal.

Stamens in (1)2(3) cycles, alternisepalous, or

oppositisepalous, when the outer whorld is sta-

minodal; ﬁ laments often with basal ligular

appendages; anthers versatile, introrse, or latrorse.

Pollen grains mostly 3-colporate, less often 3-col-

pate (Seetzenia, Sisyndite, Viscainoa) or pantopo-

rate (Tribuloideae). Gynoecium of ﬁ ve or less

often four carpels, gradually attenuated into a

simple style with lobed or capitate stigma; ovary

often angled or winged, 5–6-locular (in Tribulus

locules secondarily divided by ‘false septa’), with

one to many axile, pendulous, apotropous to

orthotropous or campylotropous ovules per loc-

ule; micropyle not zig-zag. Endothelium present

(absent in Seetzenia). Fruits usually loculicidal

and/or septicidal capsules or schizocarps, rarely

(Sericodes) succulent and berrylike. Fruit elasti-

cally dehiscent (when of cocci), or passively

dehiscent. Seeds with straight to weakly curved,

green embryo, with oily or without endosperm;

seed-coat with endotestal crystal layer and a lig-

niﬁ ed endotemgen. Present anthroquinones, ﬂ a-

vonols (kaempferol and quercetin), some species

produce steroid and triterpenoid saponins, alka-

loids (harman, harmin, and harmol); n = 6, 8–13

or (Tetraena) 14. . . . . . . . . 1. zygophyllaceae.

1 Leaves estipulate (Balanitaceae) or stipules minute

and only visible in the youngest leaves.