Takhtajan Armen. Flowering Plants

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Superorder ROSANAE 323

Huber H. 1963. Die Verwandtschaftsverhaltnisse der Rosiﬂ oren.

Mitt. Bot. Staatssamml. Munchen 5: 1–48.

Iketani H and H Ohashi. 1991. Anatomical structure of fruits

and evolution of the tribe Sorbeae in the subfamily Maloideae

(Rosaceae). J. Jpn. Bot. 66: 319–351.

Iwatsubo Y and N Naruhashi. 1998. Chromosome study of

Exochorda racemosa (Rosaceae). J. Phytogeogr. Taxon. 46:

201–203.

Jackson G. 1934. The morphology of the ﬂ owers of Rosa and

certain closely related genera. Am. J. Bot. 21: 453–466.

Jansen S, Y Sano, B Choat, D Rabaey, F Lens, and RR Dute.

2007. Pit membranes om tracheary elements of Rosaceae

and related families: new records of tori and pseudotori. Am.

J. Bot. 94: 503–514.

Juel HO. 1918. Beiträge zur Blütenanatomie und zur Systematik

der Rosaceen. Kongl. Vetensk. Akad. Handl. 58(5): 1–32.

Stockholm.

Juel HO. 1927. Über die Blütenanatomie einiger Rosaceen.

Nova Acta Regiae Soc. Sci. Upsaliensis, vol. extr. Uppsala.

Kalkman C. 1973. The Malesian species of subfamily Maloideae

(Rosaceae). Blumea 21: 413–442.

Kalkman C. 1988. The phylogeny of the Rosaceae. Bot. J. Linn.

Soc. 98: 37–59.

Kalkman C. 2004. Rosaceae. In: K Kubitzki, ed. The families

and genera of vascular plants, vol. 6, pp. 343–386. Springer,

Berlin/Heidelberg/New York.

Kamelin RV. 2006. Rosaceae. Barnaul University, Barnaul.

Kania W. 1973. Entwicklungsgeschichtliche Untersu-chungen

an Rosaceenblüten. Bot. Jahrb. Syst. 93: 175–246.

Kubitzki K. Quillajaceae. In: K Kubitzki, ed. The families and

genera of vascular plants, vol. 9, pp. 407–408. Springer,

Berlin/Heidelberg/New York.

Lersten NR and HT Horner. 2005. Macropattern of styloid and

druse crystals in Quillaja (Quillajaceae) bark and leaves. Int.

J. Plant Sci. 166: 705–711.

Lindenhofer A and A Weber. 1999a. Polyandry in Rosaceae:

evidence for a spiral origin of the androecium in Spiraeoideae.

Bot. Jahrb. Syst. 121: 553–582.

Lindenhofer A and A Weber. 1999b. The spiraeoid androecium

of Pyroideae and Amygdaloideae (Rosaceae). Bot. Jahrb.

Syst. 121: 583–605.

Lindenhofer A and A Weber. 2000. Structural and developmen-

tal diversity in the androecium of Rosoideae (Rosaceae).

Bot. Jahrb. Syst. 122: 63–91.

Lotova LI and AC Timonin. 1998a. Anatomy of cortex and

secondary phloem of Rosaceae. 1. Spiraeoideae—Spireae; 2.

Spiraeoideae except Spiraeae and Lymno

thamneae. Bot. Zhurn. 83: 16–27 (in Russian with English

summary).

Lotova LI and AC Timonin. 1999. Anatomy of cortex and

secondary phloem in Rosaceae: 3. Quillajoideae. Bot. Zhurn.

84(2): 34–41 (in English with Russian summary).

Lotova LI and AC Timonin. 2002. Anatomy of cortex and

secondary phloem of Rosaceae. 12. Prunoideae. Bot.

Zhurn. 87(5): 68–82 (in Russian with English summary).

MacDaniels LH. 1940. The morphology of the apple and

other pome fruits. Cornell Univ. Agr. Exp. Sta., Mem.

230: 1–32.

Mai DH. 1984. Karpologischc Untersuchungen der Steinkerne

fossiler und rezenter Amygdalaceae (Rosales). Feddes

Repert. 95: 299–329.

Mauritzon J. 1939. Contributions to the embryology of the

orders Rosales and Myrtales. Lunds Universitets Arsskrift,

35(2). Gleerup, Lund, Sweden.

Melikian AP and NA Bondar. 1996a. Rosaceae. Neuradaceae.

In: A Takhtajan, ed. Comparative seed anatomy, vol. 5,

pp. 102–127. Nauka, St. Petersburg (in Russian).

Moffett AA. 1931. The chromosome constitution of the Pomoideae.

Proc. Roy Soc. London, Ser. B, Biol. Sci. 108: 423–446.

Morgan DR, DE Soltis, and KR Robertson. 1994. Systematic

and evolutionary implications of rbcL sequence variation in

Rosaceae. Am. J. Bot. 81: 890–903.

Naruhashi N and Y Toyoshima. 1979. Pollen morphology of

Japanese Rosaceae. J. Phytogeogr. Taxon. 27: 46–50.

Oh S-H and D Potter. 2002. Where does Guamatela belong?

Botany 2002 BSA/ASPT meeting, Madison (Abstract).

Oh S-H and D Potter. 2005. Molecular phylogenetic systematics

and biogeography of tribe Neillieae (Rosaceae) using DNA

sequences of cpDNA, rDNA, and LEAFY. Am. J. Bot. 92:

179–192.

Oh S-H and D Potter. 2006. Description and phylogenetic posi-

tion of a new Angiosperm family, Guamatelaceae, inferred

from chloroplast rbcL, atpB, and MatK sequences. Syst.

Bot. 31: 730–738.

Phipps JB, KR Robertson, PG Smith, and JR Rohrer. 1990.

A checklist of the subfamily Maloideae (Rosaceae). Canad.

J. Bot. 68: 2209–2269.

Phipps JB, KR Robertson, JR Rohrer, and PG Smith. 1991.

Origin and evolution of subfam. Maloideae (Rosaceae). Syst.

Bot. 16: 303–332.

Potter D, F Gao, PE Bortiri, S-H Oh, and S Baggett. 2002.

Phylogenetic relationships in Rosaceae inferred from chloro-

plast matK and trnL-trnF nucleotide sequence data. Plant

Syst. Evol. 231: 77–89.

Robertson KR. 1974. The genera of Rosaceae in the southeast-

ern United States. J. Arnold Arbor. 55: 303–332, 344–401,

611, 662.

Robertson KR, JB Phillis, JR Rohrer, and PG Smith. 1991.

A synopsis of genera in Maloideae. Syst. Bot. 16:

376–394.

Rohrer JR, KR Robertson, and JB Phipps. 1991. Variation in

structure among fruits of Maloideae (Rosaceae). Am. J. Bot.

78: 1617–1635.

Rohrer JR, KR Robertson, and JB Phipps. 1994. Floral morphol-

ogy of Maloideae (Rosaceae) and its systematic relevance.

Am. J. Bot. 81: 574–581.

Sax K. 1932. The origin and relationships of the Pomoideae. J.

Arnold Arbor. 12: 3–22.

Schaeppi H and F Steindl. 1950. Vergleichend-mor-phologische

Untersuchungen am Gynoeceum der Rosoideen. Ber.

Schweiz. Bot. Ges. 60: 15–50.

Sterling C. 1963. The afﬁ nities of Prinsepia (Rosaceae). Am. J.

Bot. 50: 693–699.

Sterling C. 1964, 1965, 1966a, b, c. Comparative morphology of

the carpel in the Rosaceae: I, II. Prunoideae. Oesterr. Bot. Z.

51: 36–44, 354–360, 1964; III, IV, V, VI, VII. Pomoideae.

Oesterr. Bot. Z. 52: 705–712, 52: 47–54, 418–426, 938–946,

1965; 53: 225–231, 1966; VIII. Spiraeoideae. Oesterr. Bot. Z.

53: 521–530, 951–960,1966.

Sterling C. 1969. Comparative morphology of the carpel in the

Rosaceae: X. Evolution and summary. Oesterr. Bot. Z. 116:

46–54.

324 Subclass VI. ROSIDAE

Tumanian SA. 1950. Anatomical structure of the wood of the

Caucasian representatives of the subfamily Pomoideae of the

fam. Rosaceae. Trudy Bot. Inst. Armenian Acad. Sci. 7:

69–103 (in Russian).

Vamosi JC and TA Dickinson. 2006. Polyploidy and diversiﬁ ca-

tion: a phylogenetic investigation in Rosaceae. Int. J. Plant

Sci. 167: 349–358.

Wallaart RAM. 1980. Distribution of sorbitol in Rosaceae.

Phytochemistry 19: 2603–2610.

Webb JE. 1902. A morphological study of ﬂ ower and embryo of

Spiraea. Bot. Gaz. 33: 451–460.

Wilkes S and H Glasl. 2001. Isolation, characterization, and sys-

tematic signiﬁ cance of 2-pyrone-4, 6-dicarboxylic acid in

Rosaceae. Phytochemistry 58: 441–449.

Yu T-T. 1984. Origin and evolution of Rosaceae. Acta Phytotax.

Sin. 22: 431–444 (in Chinese with English summary).

Zhang S-Y. 1992. Wood anatomy of the Rosaceae. Blumea 37:

81–158.

Zhang S-Y and P Baas. 1992. Wood anatomy of trees and shrubs

from China III. Rosaceae. IAWA Bull., n.s. 13: 21–91.

Zhou LH, ZX Wei, and ZY Wu. 1999a. Pollen morphology of

Prunoideae of China (Rosaceae). Acta Bot. Yunn. 21:

207–211.

Zhou LH, ZX Wei, and ZY Wu. 1999b. Pollen morphology of

Spiraeoideae in China (Rosaceae). Acta Bot. Yunn. 21:

303–308.

Zhou LH, ZX Wei, and ZY Wu. 1999c. Pollen morphology of

Rosoideae (Rosaceae) of China. Acta Bot. Yunn. 21:

455–460.

Zhou LH, ZX Wei, and ZY Wu. 2000a. Pollen morphology and

systematic position of the Chinese endemic genus

Dichotomanthes (Rosaceae). Acta Bot. Yunn. 22:

143–147.

Zhou LH, X Gong, and ZY Wu. 2000b. The karyomorphology

and systematic position of the Chinese endemic genus

Dichotomanthes. Acta Bot. Yunn. 22: 282–285.

Order 85. CROSSOSOMATALES

Xerophytic, glabrous, sometimes spinescent shrubs or

small trees, sometimes with hyaline to black trichomes.

Vessel elements small and short, with simple perfora-

tions; lateral pitting alternate. Fibers short, with bor-

dered pits. Rays heterogeneous, mixed uniseriate and

pluriseriate. Axial parenchyma scanty, mostly apotra-

cheal and diffuse. Sieve-element plastids of S-type.

Nodes unilacunar with one trace, or trilacunar with

three traces. Yellow acicular crystals common in all

organs. Leaves deciduous, alternate or (Apacheria)

opposite, small, simple, entire or some of them apically

tridentate, with minute, subulate, linear stipules or

estipulate. Stomata anomocytic. Flowers solitary, axil-

lary or terminal, bisexual or some of them unisexual

(plant polygamo- monoecious), actinomorphic, with

double perianth. Sepals (3-)4-5(-6), imbricate, free,

persistent. Petals (3-)4-5(-6), free, imbricate, deciduous

or persistent, usually longer than sepals, often short-

clawed. Stamens attached to a thickened nectary disc or

seated around an annular nectary disc at its base, numer-

ous (up to 50–100) in three or four cycles associated

with about ten trunk bundles (arising centrifugally or

centripetally), but in two cycles in Apacheria and some

spp. of Glossopetalon, and more or less reduced to one

cycle (antepetalous cycle lost) in rest of Glossopetalon.

Anthers small, tetrasporangiate, slightly extrorse,

tetrasporangiate, basiﬁ xed to slightly ventriﬁ xed,

versatile, opening longitudinally. Tapetum secretory.

Pollen grains 2-celled, (2)3-colporate, reticulate.

Nectary disc adnate to the ﬂ oral tube. Gynoecium of

1-5(-9) free carpels, each with a short, stout stylodium

and terminal, expanded stigma, or (Apocheria) the

stigma decurrent on the stylodium; ovary superior;

ovules (1)2 to many per carpel, campylotropous, biteg-

mic, crassinucellate; micropyle zig-zag. Female game-

tophyte of Polygonum-type. Endosperm nuclear. Fruits

asymmetric follicles, ventrally dehiscent, surface

smooth or rugose. Seeds testal, provided with well-

developed, large, whitish or yellowish, and ﬁ mbriate or

small and entire aril arising around the funicle; embryo

slender, slightly curved; endosperm scanty or copious,

oily. Containing inulin, ellagic and gallic acids, and

cyanidin-3- glucoside; Glossopetalon leaves contain

acacetin and syringin; n = 6 (Crossosoma).

The afﬁ nities of the Crossosomataceae are not very

clear. They are most probably related to the archaic

members of the Rosales and especially to the

Saxifragales although in the presence of the aril they

also resemble the Dilleniaceae.

1. CROSSOSOMATACEAE

Engler 1897. 4/8 or 9. Arid parts of western United

States and adjacent Mexico.

Crossosoma, Glossopetalon (Forsellesia), Apacheria,

Velascoa.

Bibliography

Calderon de Rzedowski G and J Rzedowski. 1997. Velascoa

(Crossosomataceae), un género nuevo de la Sierra Madre

Oriental de México. Acta Bot. Mex. 39: 53–59.

Superorder ROSANAE 325

DeBuhr LE. 1978. Wood anatomy of Forsellesia (Glossopetalon)

and Crossosoma (Crossosomataceae, Rosales). Aliso 9:

179–184.

Ensign M. 1942. A revision of the celastraceous genus Forsellesia

(Glossopetalon). Am. Midl. Nat. 27: 501–511.

Holmgren NH. 1988. Glossopetalon (Crossosomataceae) a new

variety of G. spinescens from the Great Basin, U.S.A.

Brittonia 40: 269–274.

Kapil RN and RS Vani. 1963. Embryology and systematic

position of Crossosoma californicum. Curr. Sci. 32:

493–495.

Lemésle R. 1948. Position phylogénétique de l’Hydrastis

canadensis L. et du Crossosoma californicum Nutt., d’a-pres

particularités histologiques du xyléme. C. R. Acad. Sci. Paris

227: 221–223.

Mason CT. 1975. Apacheria chiracahuensis: A new genus and

species from Arizona. Madroño 23: 105–108.

Masson CT. 1992. Crossosomataceae: Crossosoma family. J.

Arizona Nevada Acad. Sci. 26: 7–9.

Melikian AP and NA Bondar. 1996. Crossosomataceae. In:

A Takhtajan, ed. Comparative seed anatomy, vol. 5, pp. 126–

127. Nauka, St. Petersburg (in Russian).

Richardson PE. 1970. The morphology of the Crossosomataceae:

I. Leaf, stem, and node. Bull. Torrey Bot. Club 97: 34–39.

Sosa V and MW Chase. 2003. Phylogenetics of Crossosomataceae

based on rbcL sequence data. Syst. Bot. 28: 96–105.

Tatsuno A and R Scogin. 1978. Biochemical proﬁ le of

Crossosomataceae. Aliso 9: 185–188.

Thorne RF and R Scogin. 1978. Forsellesia Greene

(Glossopetalum Gray): A third genus in the Crossosomataceae,

Rosineae, Rosales. Aliso 9: 171–178

Order 86. CHRYSOBALANALES

Trees or shrubs, rarely subshrubs, or woody lianas.

Vessels with simple perforations, rarely (in some

species of Trigonia) with scalariform perforation, or

mixed simple and scalariform perforations; lateral

pitting usually alternate; vestured pits present. Fibers

with bordered pits. Rays mainly heterogeneous or

rarely almost homogeneous. Axial parenchyma

apotracheal or paratrachel. Sieve-element plastids of

S-, or Ss-type. Nodes uni-, or pentalacunar. Leaves

alternate or opposite, simple, entire, pinnately veined,

sometimes conduplicate, stipulate. Stomata paracytic.

Flowers mostly rather small, in various kinds of cymose

racemose or racemose, terminal or axillary inﬂ ores-

cence or rarely solitary, bisexual or rarely polygamous

or unisexual, actinomorphic or more or less zygomor-

phic. Sepals ﬁ ve, mostly imbricate. Petals ﬁ ve, free,

usually imbricate, rarely absent. Stamens (2-)8-20 or

numerous (up to 300), all fertile or some of them sta-

minodial; ﬁ laments free or basally connate; anthers

small or thickened, tetrasporangiate, introrse, basiﬁ xed

or dorsiﬁ xed, opening longitudinally. Nectaries present

or rarely wanting. Tapetum secretory. Microsporogenesis

simultaneous. Pollen grains 2-celled, 3(4)-colpate or

-colporate. Gynoecium of 2–3 carpels; ovary superior

to quite inferior, 2–3-locular; ovules usually 1–2, rarely

numerous in each fertile locule, anatropous, bitegmic,

mostly tenuinucellate, sometimes with endothelium

and micropyle zig-zag. Female gametophyte mono-

sporic, bisporic, or tetrasporic. Endosperm nuclear.

Fruits drupes or septicidal capsules. Seeds exotestal,

testal-tegmic, or endotegmic; embryo straight, with or

without endosperm.

Related to the Rosaceae and were by many authors

included in the Rosales, but they differ from them

markedly in tenuinucellate ovules, morphology of the

gynoecium, 4-celled female gametophyte.

Key to Families

1 Fruits septicidal capsules.

2 Fertile stamens 4–8. Evergreen trees, shrubs, or

woody lianas with T-shaped hairs. Intraxylary

phloem absent. Vessels with simple perforations,

but in some species of Trigonia occur vestigial

scalariform perforations; lateral pitting extremely

rare. Fibers with distinctly bordered pits, in some

species septate. Rays heterogeneous. Axial paren-

chyma apotracheal or paratracheal. Leaves usu-

ally opposite or rarely alternate (Trigoniastrum),

simple, entire, densely whitish hairy below;

stipules interpetiolar, often connate, deciduous.

Stomata paracytic. Flowers in terminal or axillary

cymes, racemes, or panicles, or thyrses, strongly

zygomorphic. Sepals ﬁ ve, unequal, imbricate,

united into a cup below. Petals three, or ﬁ ve,

imbricate or contorted, or valvate, the two ante-

rior (lower) ones forming a keel, the posterior

(upper) ones forming a basally saccate or spurred

standard, the laterals spatulate. Androecium uni-

lateral on anterior (lower) side of ﬂ ower, of 4–13

stamens, 4–8 (often 6) of them fertile (up to 4 sta-

minodial); ﬁ laments more or less united. Pollen

grains 3–5-porate. Nectary glands sometimes

lobed or cleft, borne in front of standard or want-

ing (Isidodendron). Gynoecium of 3(4) carpels;

style terminal, with a capitate or slightly trilobed

stigma; ovules 1–2 to rather numerous in each

locule, pendulous, anatropous, crassinucellate

(Mauritzon 1936; Corner 1976) or tenuinucellate

326 Subclass VI. ROSIDAE

(Boesewinkel 1987), with endothelium; placenta-

tion parietal; micropyle zig-zag. Fruits septicidal

capsules (Trigonia) or more often three winged

samaras. Seeds winged, hairy ( usually long-pilose),

seed coat testal-tegmic; exotesta with thickened

outer walls, endotegmic cells tanniniferous; embryo

straight, green (Trigonia); cotyledons large, ﬂ at

with very short radicle; endosperm copious and

ﬂ esh (Trigonia) or scarce (Trigoniastrum) (Plisko

2000); n = ca. 10.. . . . . . . . . . . . . 3. trigoniaceae

2 Fertile stamens three. Shrubs or subshrubs

(Euphronia acuminatissima) or trees; young

growth densely covered with white or gray

pubescent, the branchlets, petioles and blades

may become glabrous with age; mucilaginous

hypodermis lacking. Parenchyma more or less

aliform-conﬂ uent. Leaves alternate, simple,

entire, white tomentose below; stipules small.

Flowers in terminal or axillary, pubescent

racemes, bracteate, bisexual, zygomorphic.

Sepals three (5 – according A.Litt 2004), unequal,

pubescent abaxially, connate et the base. Petals

three, spatulate, clawed, purplish, generally silky

pubescent. Stamens four and generally one sta-

minodial; stamens and staminode united at the

base into tube; the split opposite staminode; ﬁ la-

ments basally connate; anthers reddish brown,

basiﬁ xed, introrse, opening by a single central

slit. Gynoecium of three carpels; the style genic-

ulate near apex, pilose, persistent in fruit, the

stigma trilobed; placentation axile. Ovary supe-

rior, 3-locular; ovules two in each locule, anatro-

pous. Fruits cylindrical septicidal, trivalvate

capsule, exocarp thin, ﬂ eshy, pubescent. Seeds

one per locule, slightly winged, glabrous, red-

dish brown. . . . . . . . . . . . . . 4. euphroniaceae.

1 Fruits drupes.

3 Stylodia free or united into simple styles.

1 Ovules tenuinucellate. Small trees, shrubs,

or woody lianas; bark of lianas often smooth,

pale, with scattered, dark, lenticellate pustules.

Mucilage cells sometimes present in the par-

enchymatous tissues and in the leaf epidermis

and hypodermis. Vessels with simple or mixed

simple and scalariform perforations; lateral

pitting alternate, rarely (Tapura) scalariform

or intermediate. Fibers with small bordered

pits. Axial parenchyma mostly paratracheal.

Indumentum consists of unicellular hairs with

warty papillae. Leaves alternate, simple, entire,

pinnately veined, stipulate; stipules usually

caducous. Flowers small, in axillary to petiolar

or epiphyllous cymes, bisexual or rarely

unisexual (the plants monoecious), actinomor-

phic or (Tapura) more or less zygomorphic,

often with articulated pedicels. Sepals ﬁ ve,

imbricate, free or rarely basally connate. Petals

ﬁ ve, usually 2-lobed or biﬁ d, imbricate, free or

basally connate into a tube. Stamens three or

ﬁ ve, free or adnate to the corolla tube, in Tapura

only three fertile and two staminodia. Anthers

thickened, opening by longitudinal slits; the con-

nective often dorsally thickened. A basal nectary

gland borne opposite each petal (conﬂ uent to

form a ring when the corolla is sympetalous).

Gynoecium of 2–3 carpels; stylodia free or

united into simple or lobed styles; stigma capi-

tate or simple; ovary superior to quite inferior;

placentation apical. Ovules two in each locule,

pendulous at apex of each locule, anatropous;

micropyle endostomal. Fruits drupes, dry or

rarely ﬂ eshy, with a unilocular or sometimes

2–3-locular pyrene that generally has only one

seed per locule; exocarp sometimes splitting.

Seeds with large, straight, oily, green or orange

embryo and without endosperm, often with car-

uncle. Containing ﬂ uoracetic acid and pyridine

alkaloids, n = 10, 12.. . . 2. dichapetalaceae.

3 Style more or less gynobasic, with simple or

3-lobed stigmas. Trees or shrubs, rarely sub-

shrubs; woods siliceous and with SiO

grains.

Vessels with simple perforations; lateral pitting

alternate. Fibers with numerous bordered pits.

Rays mainly or wholly uniseriate heterogeneous

or rarely almost homogeneous. Axial paren-

chyma apotracheal, in numerous narrow bands.

Sieve-element plastids of Ss-type. parenchyma

in apotracheal bands; Nodes pentalacunar with

ﬁ ve traces. Leaves alternate, simple, entire, pin-

nately veined, conduplicate, often with abaxial

glands, stipulate. Flowers mostly rather small, in

various kinds of cymose racemose or racemose,

terminal or axillary inﬂ orescence or rarely soli-

tary, bisexual or rarely polygamous actinomor-

phic or more or less zygomorphic, with

well-developed ﬂ oral tube that is lined with an

annular nectary disc. Sepals ﬁ ve, imbricate.

Petals ﬁ ve, free, imbricate, sometimes absent.

Stamens (2-)8–20 or numerous (up to 300), all

fertile or some of them staminodial; ﬁ laments

Superorder ROSANAE 327

elongate and curved toward the ﬂ oral center in

bud, free or all connate or connate into groups; in

the more zygomorphic ﬂ owers all on one side of

the ﬂ oral tube; anthers small, tetrasporangiate,

introrse, dorsiﬁ xed, opening longitudinally.

Tapetum secretory. Microsporogenesis simulta-

neous. Pollen grains 2-celled, 3(4)-colpate or

-colporate. Gynoecium of three carpels united

only by the common gynobasic style, but usually

two of them more or less reduced, so that the

gynoecium may appear to be monomerous with a

basal stylodium; two ovules in each fertile loc-

ule, or sometimes the single fertile locule divided

by a median partition to appear as two uniovulate

locules; style more or less gynobasic, with a sim-

ple or 3-lobed stigma; ovary superior. Ovules

ascending from the base, anatropous, bitegmic,

tenuinucellate, with a small nucellus with the tis-

sue soon disintegrating, with endothelium;

micropyle zig-zag. Female gametophyte consists

of four cells: the egg, two synergids, and the cen-

tral cell (Tobe and Raven 1984). Endosperm

nuclear. Fruits 1-seeded drupes, often hairy

inside, medium-sized to large. Seeds exotestal,

ruminate, with large embryo and without

endosperm. Plants contain trihydroxyﬂ avonoids,

n = 10, 11. . . . . . . . . . . . 1 chrysobalanaceae.

1. CHRYSOBALANACEAE

R. Brown 1818 (including Hirtellaceae Horaninow

1847, Licaniaceae Martynov 1820). 17/495. Lowlands

of tropical and partly subtropical regions of both hemi-

spheres, but mainly Central and South America and

West Indies; reaches subtropics of North America,

South Africa, and eastern Asia.

chrysobalaneae: Chrysobalanus, Grangeria,

Licania, Parastemon; parinarieae: Bafodeya,

Exellodendron, Hunga, Neocarya, Parinari; gouepieae:

Acioa, Couepia, Maranthes; hirtelleae: Atuna,

Dactyladenia, Hirtella, Kostermanthus, Magnistipula.

2. DICHAPETALACEAE

Baillon 1886 (including Chailletiaceae R. Brown

1818). 3/180. Pantropical, but Dichapetalum reaches

southeastern Africa.

Dichapetalum, Stephanopodium, Tapura.

Traditionally are included in the Euphorbiales.

Recent molecular data indicates a relationship to the

Chrysobalanaceae and Trigoniaceae, a placement

suggested as early as 1921 by H.Hallier (Prance

2004).

3. TRIGONIACEAE

Endlicher 1841. 5/35. Madagascar (Humbertiodendron,

1), western Malesia (Trigoniastrum, 1) and central and

tropical South America: Trigonia (24) from Mexico to

Paraguay, Trigoniodendron – southeastern Brazil.

Trigoniastrum, Trigonia, Trigoniodendron, Humber-

tiodendron, Isidodendron.

Trigoniaceae traditionally placed in the Polygalales,

and related to the Vochisiaceae. Molecular data sug-

gested afﬁ nities with Chrysobalanaceae and Dicha-

petalaceae (Soltis et al. 2006).

4. EUPHRONIACEAE

Marcano-Berti 1989. 1/3. Colombia, Venezuela,

Guayana, and northwestern Brazil.

Euphronia.

Closely related to the Trigoniaceae.

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46: 265–350.

Superorder MYRTANAE

Order 87. MYRTALES

Trees, shrubs, or herbs. Intraxylary (internal) phloem

in woody stems always present. Vessels mostly with

simple or rarely (Geissolomataceae and some

Myrtaceae) with scalariform perforations and with

vestured, bordered pits; lateral pitting almost always

alternate. Fibers with simple or bordered pits, septate

or nonseptate. Rays heterogeneous to homogeneous.

Axial parenchyma most typically scanty paratracheal,

but in addition it may be apotracheally diffuse. Sieve-

element plastids of Ss-type. Nodes unilacunar or some-

times (Alzateaceae) trilacunar. Leaves opposite or less

often alternate or verticillate, simple, mostly entire,

usually with vestigial stipules, rarely estipulate.

Stomata of various types, mostly anomocytic. Flowers

in various cymose or racemose inﬂ orescences or some-

times solitary in the axils, bisexual or rarely unisexual,

actinomorphic or sometimes more or less zygomor-

phic (usually slightly zygomorphic), mostly 4- or

5-merous, sometimes apetalous (some Lythraceae and

all Alzateaceae and Penaeaceae). Sepals connate,

sometimes much reduced or obsolete. Petals alternise-

palous. Stamens numerous, rarely reduced to one;

ﬁ laments bent inward in the bud or rarely erect (as in

Psiloxylon); anthers 2-locular or (Heteropyxidoideae

in Myrtaceae, Rhynchocalycaceae) 4-locular at anthe-

sis, opening longitudinally or (some Myrtaceae and

the majority of Melastomataceae) by apical pores.

Tapetum secretory. Microsporogenesis simultaneous.

Pollen grains 2-celled or less often 3-celled, in

monads or sometimes (as in Onagraceae) in tetrads,

basically 3-colporate, but sometimes colpate or

porate, basically tectate and frequently characterized

by pseudocolpi. Intrastaminal annular nectary disc

often present. Gynoecium of 2-many (often 4 or 5)

united carpels with stylodia connate into a style (very

short in Psiloxylon); stigma simple or lobed. Ovary

superior to inferior, plurilocular to 1-locular, usually

with (1)2 to many ovules per carpel. Ovules anatro-

pous or in various taxa hemitropous, amphitropous or

campylotropous, usually bitegmic or very rarely (in

some Myrtaceae) unitegmic, crassinucellate.

Endothelium absent or rarely (Axinandra in the

Crypteroniaceae) present. Female gametophyte of var-

ious types. Endosperm nuclear. Fruits of various types.

Superorder MYRTANAE 329

Seeds mostly with very little or no endosperm. Contain

ﬂ avonols, myricetin, methyulated ellagic acid,

n = 5–14, 20, 24.

Evidently derived from the lower Rosidae, most

probably from the Saxifragales.

Key to Families

1 Nodes trilacunar with three traces. Rays

heterogeneous with long ends. Vessel-ray pits large

and simple, reticulate to scalariform. Evergreen

small trees or shrubs, sometimes hemiepiphytic;

branches opposite or verticillate; the young branches

scarcely 4-angled, the stems of the inﬂ orescence

distinctly 4-angled to narrowly winged; nodes

enlarged, especially knobby when branches are ver-

ticillate. Leaves mostly clustered at the ends of the

branches, opposite or less often verticillate, simple,

entire, thick, leathery, with axillary stipules. Stomata

anomocytic to nearly encyclocytic. Flowers in axil-

lary, basically paniculate inﬂ orescences, lacking

bracteoles, articulated at the base, bisexual, actino-

morphic, apetalous or possibly petals rudimentary,

with open campanulate, leathery, ﬂ oral tube,

5(6)-merous. Calyx lobes ﬂ eshy, valvate, persistent.

Nectary disc well developed, wide, lobed. Stamens

ﬁ ve, situated in the ﬂ oral tube at the sinuses of the

lobes; ﬁ laments green, stout, with large cordate

connective, as long as the anthers; anthers dorsi-

ﬁ xed, introrse, heart-shaped, pinkish white, ﬂ eshy,

at the anthesis held at right angles to the ﬁ lament

with the apiculate apex extending between the

lobes, resembling petals, opening longitudinally.

Pollen grains 3-colporate, psilate, except for the

slightly depressed middle of the mesocolpia, which

is psilate-punctate. Gynoecium of two carpels,

slightly bilaterally ﬂ attened, with a short style and

capitate stigma; ovary superior, 2-locular, with sep-

tum divided into two half-septa and with 40–60

anatropous ovules horizontally imbricate in vertical

rows, micropyle endostomal, placentation parietal.

Female gametophyte of Allium-type. Fruits bilater-

ally compressed dry, loculicidal capsules. Seeds

ﬂ attened, oblong to lunate, with fragile membra-

nous wing; exotestal cells low; embryo straight;

endosperm lacking. Ellagic acid and ﬂ avonoid

(quercetin); n = 14. . . . . . . . . . . . 1. alzateaceae.

1 Nodes unilacunar with one trace.

2 Plants woody or herbaceous, terrestrial or less

often aquatic.

3 Secretory cavities lined with epithelium lack-

ing. Calyx and corolla usually valvate.

4 Micropyle is formed only by one

integument.

5 Micropyle is formed only by inner integ-

ument Evergreen tree up to 12 m tall.

Leaves opposite, decussate, simple,

entire; stipules rudimentary. Stomata

anomocytic and cyclocytic. Flowers in

multiﬂ oral panicle, small, bisexual, acti-

nomorphic, 6-merous, with a short, sau-

cer-shaped ﬂ oral tube. Sepals six, white

and petal-like, recurved, and distinctly

beaked in bud, persistent. Petals six,

inserted on the rim of the ﬂ oral tube, con-

spicuously and narrowly clawed,

recurved, caducous, margin lacerate,

hoodlike, and covering the anthers in bud

only. Stamens six, oppositipetalous,

inserted near the inner rim of the ﬂ oral

tube, incurved in bud; ﬁ laments much

longer than the anthers; anthers sub-

basiﬁ xed, versatile, introrse, opening

longitudinally. Nectary disc wanting.

Pollen grains 3-colporate, heterocolpate,

apertures alternating with three pseudo-

colpi. Gynoecium of 2(3) carpels, with

simple style and capitate stigma covered

with minute papillae; ovary superior,

bilaterally compressed, in young ﬂ owers

1-locular, but subsequently rendered

2-locular (rarely 3-locular) by an exten-

sion of placentas from opposite sides of

the ovary wall; ovules ca. 15–20 per loc-

ule, anatropous, pleurotropous, super-

posed in a single vertical series. Female

gametophyte of Polygonum-type. Fruits

dorsiventrally compressed (rarely trigo-

nous) capsule, loculicidal at the apex.

Seeds narrowly winged (expanded

raphe), the wings is formed by divisions

and elongation of cells of the funiculus;

seed coat papery, rather smooth; embryo

more or less ﬂ attened, cotyledons folded

inside; endosperm lacking. Contain

flavonoids and accumulated Aluminium;

n = 10.. . . . . . 2. rhynchocalycaceae.

5 The micropyle is formed by the outer

integument. Evergreen, densely leafy

330 Subclass VI. ROSIDAE

ascending shrubs. Young branches more

or less densely covered with long

unicellular two-armed (Malpighian) hairs

that have one long and one quite short

arm. Branches with four longitudinal

ridges and with relatively inconspicuous

transverse leaf scars. Young stems some-

what tetragonal. Vessels with scalariform

perforations that have rather numerous

(12–23) bars; lateral pitting opposite or

scalariform. Fibers are tracheids. Rays

heterogeneous, mixed uniseriate and

pluriseriate. Axial parenchyma scanty and

diffuse and occurs in strands of 4–5 very

long cells. The stem cortex contains scat-

tered stone cells. Small calcium oxalate

crystals present in ray cells. Leaves oppo-

site, subsessile, leathery, simple, entire,

with the thickened, slightly recurved mar-

gins pinnately veined; epidermis contain-

ing some mucilaginous cells; stipules

small, subulate, situated on the sides of

the short, petiolelike leaf base. Stomata

anomocytic. Flowers solitary, terminal

(on short axillary branches), or axillary,

subtended by three pairs of decussate

bracts, these increasingly large and

petaloid (pink) above, often also with ves-

tigial ﬂ oral buds in axils of uppermost

bracts, on short, sharply 4-angled pedicel,

bisexual actinomorphic, apetalous, 4-mer-

ous throughout. Sepals large, imbricate,

pointed, rose-colored, persistent. Nectary

disc wanting. Stamens in two cycles of

four each, the outer cycle alternating with

the sepals; ﬁ laments slender, long erect,

free or inserted at the base of the calyx;

anthers dorsiﬁ xed, versatile, tetrasporan-

giate, with short, apical connective, open-

ing longitudinally. Tapetum secretory.

Pollen grains usually 3-colporate, but

some 6-pantoporate grains also occur; the

exine tectate-columellate, perforate.

Gynoecium of four united carpels; ovary

superior, 4-locular, 4-ridged, and

4-grooved with essentially free stylodia;

stylodia basally and centrally separate,

but their upper slender parts connivent

and twisted, separate apically and covered

with stigmatic papillae adaxially; each

locule with two pendulous ovules attached

centrally in the upper middle part of the

ovary. Female gametophyte of Polygonum-

type. Fruits hard, 4-lobed, loculicidal cap-

sules enclosed in the persistent calyx.

Seeds reniform, smooth, with a short,

whitish funicle, which is swollen and

forms a collarlike caruncle; embryo

straight, linear with ﬂ eshy, linear cotyle-

dons; endosperm present. Plants

Al-accumulators. 3. geissolomataceae.

4 Micropyle is formed by both the

integuments.

6 Female gametophyte of Penaea-type

(tetrasporic, 16-nuclear). Glabrous

small, evergreen, ericoid shrubs or

shrublets, procumbent to ascending or

erect. Leaves opposite, decussate,

entire, acuminate to retuse, often cori-

aceous; stipules wanting or small, each

divided into a small row of ﬁ ngerlike

or rarely hairlike, secretory structures

in the leaf axil. Stomata anomocytic.

Flowers solitary in the upper leaf-

axils, bisexual, actinomorphic, apeta-

lous, 4-merous, with a variably long

ﬂ oral tube. Calyx lobes valvate or

reduplicate-valvate, persistent, calyx

often coloured like a corolla. Stamens

four, inserted in throat of calyx and

alternating with calyx lobes; ﬁ laments

very short, narrow, erect; anthers

4-locular, introrse, basiﬁ xed, opening

longitudinally; connective ﬂ eshy,

much expanded, often much longer

than the frequently well separated loc-

ules. Pollen grains 3–6-colporate,

always provided with pseudocolpi

isomerous with the apertures. Nectary

disc lacking. Gynoecium of four car-

pels; style with capitate or 4-lobed

stigma; ovary superior, 4-locular, often

4-lobed, with 2–4 anatropous, pendu-

lous or ascending ovules per locule.

Placentation basal to axile, or axile, or

axile to apical. Fruits ﬁ rm, often hard

loculicidal capsules, included in the

persistent ﬂ oral tube. Seeds elongate,

Superorder MYRTANAE 331

arilate, endotestal cells much elon-

gated, endotegmen ﬁ brous, nearly or

quite without endosperm; embryo with

a large hypocotyl and small cotyle-

dons. Contain non-hydrolysable tan-

nins, n = 10–12. . . . . 4. penaeaceae.

6 Female gametophyte of Polygonum- or

Oenothera-type.

7 Female gametophyte of Polygonum-

type. Oxalate raphids wanting.

8 Each ﬂ ower is basally subtended by

a short internode ending with some

blunt teeth, a “calyculus” (which is a

stem structure). Flowers in panicu-

late inﬂ orescences with the branches

ending in 3-ﬂ owered cymules, small,

bisexual, actinomorphic, 4(5)-mer-

ous, with ﬂ oral tube prolonged

over the ovary, truncate or obscurely

and minutely 5-toothed, ultimately

deciduous. Calyx represented by a

narrow rim at the throat of the ﬂ oral

tube, usually with (4)5 more or less

linear-spatulate or obovate spatulate,

white or pinkish, valvate lobes, not

enclosing the petals in bud. Petals

4(5), inserted at the throat of the ﬂ o-

ral tube, scalelike, alternating with

end much shorter than the calyx

lobes, incurved and closing the tube

in bud, spreading and reﬂ exed at

anthesis, valvate, usually pink to

crimson, pubescent. Stamens (4)5,

inserted on the inner rim of the ﬂ oral

tube below the petals, incurved in

bud; very short, not exserted and

with the anthers almost ﬁ lling the

mouth of the ﬂ oral tube; anthers dor-

siﬁ xed, not versatile, introrse, with

thickened connective, opening longi-

tudinally. Pollen grains heteropolar

with one pole colpate and the other

heterocolpate; pseudocolpi restricted

to one polar face only. Nectary disc

lacking. Gynoecium of (2)4–5 car-

pels; style with a capitate stigma;

ovary inferior, with as many locules

as carpels, with (2)3 superposed

ovules per carpel. Ovules pendulous,

campylotropous, hypotropous. Fruits

drupaceous, marked by a circular

scar left by the deciduous ﬂ oral tube;

endocarp woody, longitudinally

ribbed. Seeds usually one per locule

and nearly always only one per fruit,

without endosperm; embryo oily,

with a short radicle and unfolded,

spirally twisted or convolute cotyle-

dons. Evergreen shrubs or trees up to

25 m tall; stem often ﬂ uted and but-

tressed; young leafy branchlets quad-

rangular. The inner bark and leaves

(when crushed) usually with a strong

almond scent. Leaves opposite or

rarely ternate, entire, coriaceous,

with rudimentary stipules. Stomata

paracytic to anomocytic, verging on

encyclocytic. Fibers with simple pits,

septate. Contain non-hydrolysable

tannins, cyanogenic glycoside pruna-

sin; n = 15, 20. . . . . 5. oliniaceae.

8 Flowers not subtended by

“calyculus.”

9 Placentation apical. Evergreen or

deciduous trees up to 50 m tall,

shrubs, subshrubs (in savannas,

often do not exceed 20 cm), or

often lianous (in forests, often

climb above 30 m high), some-

times mangroves, rarely spiny;

the bark is often ﬂ aking. Roots

rarely with pneumatophores

(Laguncularia racemosa); stems

rarely with spines. Fibers with

simple pits, frequently septate, or

(Strephonema) with distinctly

bordered pits. Intraxylary (inter-

nal) phloem commonly but not

always present, interxylary

(included) phloem often present

as well (restricted to some genera

of the tribe Com-breteae); muci-

lage ducts present in Terminalia.

Leaves alternate, opposite or less

often verticillate, entire, simple,

variously pubescent, sometimes

glandular (Combretinae), condu-

plicate or supervolute (revolute in

332 Subclass VI. ROSIDAE

Laguncularia), often with two

gland-containing ﬂ ask-shaped

cavities at base; stipules lacking

or vestigial. Stomata anomocytic

or sometimes encyclocytic, rarely

(Strephonema) paracytic. Flowers

in racemes, spikes, or heads,

bisexual or sometimes unisexual,

actinomorphic or seldom slightly

zygomorphic, with a usually

fairly short ﬂ oral tube (sometimes

absent). Sepals 4–5(-8), often

appearing as lobes of ﬂ oral tube,

valvate or imbricate or sometimes

very small. Petals 4–5(-8), mostly

rather small, imbricate or valvate,

often lacking. Stamens usually

twice as many as the sepals and

mostly in two cycles; ﬁ laments

inﬂ exed in bud; anthers usually

versatile, opening longitudinally.

Pollen grains 3(4)- colporate,

mostly heterocolpate, with pseudo-

colpi (absent in Strephonema).

Well-developed intrastaminal disc

is very often present. Gynoecium

of 2–5(-8) carpels with elongate

style ending in a punctate or sel-

dom capitate stigma; ovary infe-

rior, or rarely (Strephonema)

semi-inferior, 1-locular, with

2(-6), sometimes up to 7(20),

only one developing, anatropous

ovules pendulous on an elongate,

slender funicle from the top of

locule; placentation apical.

“Fruits 1-seeded pseudocarps,

formed from the ovary and

surrounding the lower hypan-

thium; the fruit wall ( pericarp

plus hypanthium) may be thin

and hard, or become differenti-

ated as spongy or succulent tis-

sue, or develop 2–5 wings” (Stace

2007). Seeds without endosperm;

embryo green, oily, with folded

or spirally twisted cotyledons;

rarely (Strephonema) cotyledons

massive and hemispheric or

(some African spp. of Combretum)

united, some species of Terminalia

have three or more cotyledons.

Contain tannins,

5- desoxyﬂ avonoids, ﬂ avonoid

sulphates, myricetin, only few

alkaloids and triterpene saponins;

n = 7, 11–13. 6. combretaceae.

9 Placentation not apical, only

rarely (Trapaceae) apical-axile.

10 Flowers polygamo-dioecious

or bisexual, in axillary racemes

or panicles, very small, acti-

nomorphic, 4–5-merous.

Sepals 4–5, appearing as val-

vate lobes on the cupulate or

shortly tubular ﬂ oral tube.

Petals absent (Crypteronia) or

small and connate apically and

falling off as a cup when the

ﬂ ower opens (Axinandra).

Stamens as many as and alter-

nate with the sepals, inserted

on the margin of the ﬂ oral

tube, inﬂ exed in bud, and with

a wide connective, which is

conduplicate in Axinandra.

“The anther endothecium

degenerates early, and the

anthers dehisce not via differ-

ential shrinking of endothe-

cium cells, but via rupture of

walls along their thinnest sec-

tions, caused by the shrinking

of connective cells” (Renner

2007). Pollen grains 2- or

3-colporate, with apertures

alternating with pseudocolpi.

Gynoecium of 2–6 carpels;

style with capitate stigma;

ovary superior to semi-infe-

rior, with numerous, anatro-

pous ovules on axile placentas.

Endothelium present at least

in Axinandra. Fruits charta-

ceous or woody, loculicidal

capsules, the valves often held

together apically by persistent

style. Seeds usually small, ﬂ at,