Takhtajan Armen. Flowering Plants

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202 Subclass V. DILLENIIDAE

Porter JM. 1998. Nomenclatural changes in Polemoniaceae.

Aliso 17: 83–85.

Porter JM and LA Johnson. 1998. Phylogenetic relationships of

Polemoniaceae: inferences from mitochondrial nad1b intron

sequences. Aliso 17: 157–188.

Porter JM and LA Johnson. 2000. A phylogenetic classiﬁ cation

of Polemoniaceae. Aliso 19: 55–91.

Prather LA and RK Jansen. 1998. Phylogeny of Cobaea

(Polemoniaceae) based on sequence data from the ITS region

of nuclear ribosomal DNA. Syst. Bot. 23: 57–72.

Prather LA, CJ Ferguson, and RK Jansen. 2000. Polemoniaceae

phylogeny and classiﬁ cation: implications of sequence data

from the chloroplast gene ndhF. Am. J. Bot. 87: 1300–1308.

Smith DM, CW Glennie, JB Harborne, and CA Williams. 1977.

Flavonoid diversiﬁ cation in the Polemoniaceae. Biochem.

Syst. Ecol. 5: 107–115.

Spencer SC and JM Porter. 1998. Evolutionary diversiﬁ cation

and adaptation to novel environments in Navarretia

(Polemoniaceae). Syst. Bot. 22: 649–668.

Steele KP and R Vilgalys. 1994. Phylogenetic analyses of

Polemoniaceae using nucleotide sequences of the plastid

gene matK. Syst. Bot. 19: 126–142.

Stuchlik L. 1967. Pollen morphology in the Polemoniaceae.

Grana Palynol. 7: 146–240.

Taylor TN and DA Levin. 1975 (1976). Pollen morphology of

Polemoniaceae in relation to systematics and pollination sys-

tems: scanning electron microscopy. Grana 15: 91–112.

Wilken DH. 2004. Polemoniaceae. In: K Kubitzki, ed. The fami-

lies and genera of vascular plants, vol. 6, pp. 300–312.

Springer, Berlin/Heidelberg/New York.

Wilson KA. 1960a. The genera of Hydrophyllaceae and

Polemoniaceae in the southeastern United States. J. Arnold

Arbor. 41: 197–212.

Order 58. LECYTHIDALES

Small to large trees or shrubs; wood siliceous and/or

with SiO

grains. Cortical vascular bundles occur in the

stem, with internal xylem (normally oriented) or external

(inverted). Secondary xylem and primary cortex often

with crystalliferous strands. Vessels typically with sim-

ple perforations, but scalariform or reticulate perforation

plates appear in some genera including Asteranthos and

Grias. Lateral pitting usually alternate, except in

Asteranthos, Grias, and Gustavia, in which it is alternate

to opposite. Fibers typically with simple pits, but bor-

dered pits in a few genera. Rays from distinctly heteroge-

neous to homogeneous (entirely heterogeneous in some

genera, including Asteranthos, Grias, Gustavia, Foetidia,

Planchonia, and others). Axial parenchyma typically

abundant and in apotracheal bands with some paratra-

cheal. Secondary phloem generally stratiﬁ ed into ﬁ brous

and soft portions. Nodes tri-, penta- or multilacunar.

Leaves alternate, simple, entire or dentate, pinnately

veined, often crowded at the ends of twigs, estipulate or

(Scytopetalaceae and Gustavia – Lecythidoideae) with

minute, caducous stipules; the structure of petiole is

mostly somewhat complex, typically containing numer-

ous vascular bundles, but with reduced number in

Napoleonaeaeae (one major bundle and two minor adax-

ial bundles). Stomata usually anisocytic. Flowers mostly

rather large and showy, solitary in the axils of leaves or

leaﬂ ike bracts, or more often in racemose or paniculate

inﬂ orescences, sometimes cauliferous, bisexual, actino-

morphic or the corolla and particularly androecium

zygomorphic, subtended by a bract and bearing two

bracteoles, typically ephemeral, pollinated by insects

and bats. Perianth forms a tube more or less adnate to the

ovary. Sepals (2)4–6(-12), imbricate or valvate, free or

more or less connate, sometimes fully connate and calyp-

trate. Petals (3)4-6 or more or absent. Stamens com-

monly numerous (up to about 1,200), in several cycles,

developing centripetally or more often centrifugally; ﬁ la-

ments mostly connate into a ring that is either actinomor-

phic or more or less expanded on one side into an

incurving ﬂ at ligule sometimes curved over the gynoe-

cium as a hood; anthers latrorse, introrse or rarely pori-

cidal, tetrasporangiate or sometimes (as in Napoleonaea)

disporangiate, opening longitudinally or sometimes (as

Gustavia) by apical pores. Tapetum secretory.

Microsporogenesis simultaneous. Pollen grains 3-celled,

3-colpate or 3-colporate, tectate- columellate, with vari-

ous types of ornamentation. Intrastaminal disc present or

absent. Gynoecium usually of 2–6(-8) united carpels,

mostly with a simple style with capitate or lobed stigma;

ovary inferior, semi-inferior or (most of Scytopetalaceae)

superior, with as many locules as carpels and with

1-many ovules per locule. Ovules anatropous to campy-

lotropous, bitegmic, tenuinucellate, with or without

endothelium, often with conspicuous funicle; micropyle

endostomal. Female gametophyte of Polygonum-type.

Endosperm nuclear. Fruits ﬁ brous berries, dry drupes, or

capsules, often very large, with one to many seeds. Seeds

small to very large, hairy, often with funicular aril; seed

coat formed primarily by the outer integument (exotestal-

mesotestal); exotestal cells variously thickened, palisade,

or low with sinuous anticlinal walls, mesotesta sclerotic

or not; endosperm almost or completely absent (well-

developed only in Scytopetalaceae); embryo large,

straight or curved, oily or proteinaceous. Generally con-

taining riterpenoid saponins and usually also producing

proanthocyanins and ellagic acid, n = 11, 13, 16–18, 21.

Superorder ERICANAE 203

The Lecythidales have been associated with the

Myrtales (Prance and Mori 1979). But many important

characters, including the secondary xylem (especially

by the absence of vestured pits) and lack of internal

phloem, and tenuinucellate (but bitegmic) ovules often

with an endothelium, they differ markedly from the

Myrtales. Therefore, Cronquist (1957, 1980, 1988)

and Dahlgren (1980, 1983) removed the family from

its traditional place, and Cronquist placed Lecythidaceae

s.l. in a separate order Lecythidales (1957) and sug-

gested placing it somewhere between Malvales and

Theales. Thorne (1968) suggested a thealean relation-

ships, and Dahlgren (1980, 1983) included this family

in Theales. In my opinion the order Lecythidales are

related to the theoid order Ochnales and could have a

common origin with it from some remote thealean

ancestor.

Key to Families

1 Seeds with endosperm. Evergreen trees or shrubs.

Leaves mostly estipulate, papery to coriaceous.

Flowers in panicles, thyrsoids, botryoids or racemes

on younger branches, often ramiﬂ orous or cauliﬂ o-

rous, rarely reduced to single ﬂ owers (Asteranthos).

Sepals united; calyx copular, thick, leathery, persis-

tent on ripe fruits, occasionally accrescent. Petals

lacking. Staminodia 6–16, or 24–28 (Asteranthos);

apex of staminodia curved inwards or not curved

inwards (Rhaptopetalum, Pierrina, Brazzeia).

Stamens 60–240; ﬁ laments adnate to pseudocorolla;

anthers basiﬁ xed, tetrasporangiate, dehiscing by lon-

gitudinal slits or apical and pore-like. Pollen grains

3-colpate or 3-colporoidate (Rhaptopetaloideae).

Carpels 3–8. Ovary superior or semi-inferior. Ovules

2–4-many in each locule, pendulous. Fruits mono-

spermous or polyspermous. Seeds glabrous or

embedded in a shiny white haircoat, arising from epi-

dermis cells along the raphe (Rhaptopetalum,

Pierrina, Brazzeia). Endosperm horny, mostly mark-

edly ruminate (Kravtsova 1991); embryo well differ-

entiated; cotyledons large, ﬂ at, cordate, rarely

reduced (Asteranthos). . . . . . .1. scytopetalaceae.

1 Seeds without endosperm.

2 Petals free, 3-6(-18); stamens numerous, usually

connate at base and often forming a strip-like

structure; nectary disc absent; seeds arillate or

not; embryos undifferentiated or with ﬂ eshy coty-

ledons. Small to large trees, rarely shrubs. Stipules

absent or minute and caduceus. Inﬂ orescences ter-

minal, axillary or cauline, simple racemes, or spi-

cate branches or fascicles. Sepals 2–6 or rarely

unlobed. Petals imbricate, free; anthers bilocular,

latrorse, introrse or rarely poricidal. Ovary inferior

or semi-inferior, 2–4- or 6-locular; ovules 1–115

in each locule. Fruits indehiscent, then dry, ﬂ eshy

or woody or dehiscent. Seeds exotestal-mesot-

estal, winged or without wings; endosperm lack-

ing or very scanty. Some species accumulate

selenium. . . . . . . . . . . . . . . . .2. lecythida ceae.

2 Pseudocorolla of 30–35 fused staminodes.

Nectary disc well developed, annular, decagonal,

supported by a very complex vascular system.

Trees, shrubs or subshrubs. Leaves estipulate.

Flowers usually solitary and axillary, occasion-

ally in paniculate, axillary inﬂ orescences, actino-

morphic, usually with 4–5 bracts below the

ﬂ owers. Calyx lobes 3 or 5–6, imbricate or val-

vate, each with two large glands. Petals absent.

Stamens numerous; there are two outer cycles of

staminodia, and an inner cycle of 20 fertile sta-

mens, which are aligned into ﬁ ve groups of four

stamens in each group, the outer two have very

large, disporangiate, extrorse or latrorse anthers;

ﬁ lament ribbonlike, incurved. Pollen grains

3-colporoidate or 3-colporate. Ovary semi-infe-

rior or inferior. Style short, with ﬂ at, pentagonal

or hexagonal stigma; the entire structure is table-

like. Ovules mainly campylotropous. Fruits bac-

cate; seeds exarillate; embryos well-differentiated;

cotyledons plano-convex with short radicle and

large plumules, n = 16. . . 3 napoleonaeaceae.

1. SCYTOPETALACEAE

Engler 1897 (including Asteranthaceae Knuth 1939,

Rhaptopetalaceae van Thiegem 1908). 6/28. Tropical

West Africa and South America.

1.1 ASTERANTHOIDEAE

Flowers solitary in leaf axils. Calyx accrescent. The

margin of corolla-like structure is ciliate and curved

inward in bud and unfolds like an umbrella at anthesis.

Staminodes 24–28. Stamens 200–300. Ovary semi-

inferior. Carpels 5–8. Ovules four in each locule.

Fruits monospermous. Seeds glabrous. Cotyledons

small and strongly reduced. Pollen grains 3-colpate,

n = 21. – Asteranthos.

204 Subclass V. DILLENIIDAE

1.2 SCYTOPETALOIDEAE

Flowers in few to many-ﬂ owered inﬂ orescences in leaf

axils of youngest branches. Staminodes 6-12(-16).

Stamens 100–240. Carpels 3–5 or (6)7-8. Ovary supe-

rior. Ovules 2–4 in each locule. Fruits capsules or

drupaceous. – Scytopetalum, Oubanguias.

1.3 RHAPTOPETALOIDEAE

Inﬂ orescences ramiﬂ orous or cauliﬂ orous. Staminodes

completely united into a thick, leathery pseudocorolla.

Stamens 60–100. Anthers dehiscent by short, pore-like

longitudinal slits. Ovules many in each locule.

Fruits many-seeded. Pollen grains 3-colporoidate. –

Rhaptopetalum, Pierrina, Brazzeia.

Related to the Lecythidaceae.

2. LECYTHIDACEAE

A. Richard 1825 (including Barringtoniaceae F.

Rudolphi 1830, Foetidiaceae Airy Shaw 1965,

Gustaviaceae Burnett 1835). 17/210. Tropical, espe-

cially America and West Africa.

2.1 PLANCHONIOIDEAE

Flowers in racemose or paniculate inﬂ orescences, acti-

nomorphic. Sepals (3)4(5), free or connate, imbricate

(rarely closed). Petals (3)4(5), free, imbricate. Nectary

disc usually well developed, annular. Stamens numer-

ous, in several cycles, ﬁ laments linear, very long, free or

mostly connate at the base, with a basal staminal ring;

anthers basiﬁ xed, latrorse or less often introrse. Pollen

grains syntricolporate, with thick ectexine and the spe-

cialized ectexinous ridge along the colpus margin (mar-

ginal ridge). Style ﬁ liform, often very long, with small

stigma; ovary inferior or rarely semi-inferior, 2–4- -

locular, with two to many anatropous or campylotropous

ovules per locule. Fruits one to many-seeded drupes

with persistent calyx, sometimes broadly 4-winged.

Seeds sometimes very large (up to 6 cm in Abdulmajidia),

with undifferentiated or differentiated embryo. Cortical

bundles in stem reversely oriented. Crystalliferous

parenchyma strands absent in secondary xylem. Nodes

3-lacunar with three traces, n = 13. – Petersianthus,

Barringtonia, Chydenanthus, Abdulmajidia, Careya,

Planchonia (Combretodendron).

2.2 LECYTHIDOIDEAE

Flowers actinomorphic or zygomorphic. Sepals (4)6, free

or more or less connate, valvate or slightly imbricate.

Petals 4-6(8), free, imbricate. Stamens numerous, in sev-

eral cycles, connate at the base into a staminal ring, the

ring actinomorphic or more often prolonged on one side

and forming a hood over the fertile stamens and gynoe-

cium (the stamens on this sterile); ﬁ laments of fertile sta-

mens usually short; anthers basiﬁ xed. Pollen grains

3-colporoidate or 3-colporate, smaller, and without mar-

ginal ridge. Nectary disc absent. Style usually short,

mostly rodlike, with capitate to lobed stigma; ovary infe-

rior or less often semi-inferior, 2–6- or rarely or more-

locular. Ovules one to many. Fruits indehiscent

(drupaceous or berrylike) or dehiscent with a circumscis-

sile operculum (pyxidia), then often very large and woody.

Seeds often arillate; embryo undifferentiated or with

ﬂ eshy cotyledons. n = 17 (18). – griadear: Gustavia,

Grias; couroupiteae: Allantoma, Cariniana,

Couroupita, Corythophora; lecythideae: Lecythis,

Eschweilera; couratarieae: Couratari; berthole-

tieae: Bertholletia.

2.3 FOETIDIOIDEAE

Flowers solitary and axillary or in small axillary

cymes, actinomorphic, bibracteolate. Calyx of (3)4(5)

valvate, thick, woody, deep purple, connate sepals,

which enclose the inner ﬂ oral organs until anthesis.

Petals absent. Stamens numerous, free, sometimes in

four oppositisepalous groups; ﬁ laments ﬁ liform, long

and upright, or slightly wavy in the bud, spread out in

the open ﬂ owers; anthers very small, introrse.

Intrastaminal disc inconspicuous. Style slender, with

usually four short slender divaricate stigmas; ovary

inferior, (3)4-locular, with 15–20 ovules per locule;

ovules usually campylotropous, arranged in vertical

ring around thick peltate placenta. Fruits drupaceous,

turbinate, 1- to 4-locular. Seeds with straight or

J-shaped embryo. – Foetidia.

3. NAPOLEONAEACEAE

A. Richard 1827. 2/10. Tropical West Africa.

Napoleonaea, Crateranthus.

Closely related to Scytopelataceae.

Bibliography

Appel O. 1996. Morphology and systematics of the

Scytopetalaceae. Bot. J. Linn. Soc. 121: 207–227.

Superorder ERICANAE 205

Appel O. 2004. Scytopetalaceae. In: K Kubitzki, ed. The fami-

lies and genera of vascular plants, vol. 6, pp. 426–430.

Springer, Berlin/Heidelberg/New York.

Breteler FJ. 2002. Scytopetalaceae are stipulate. Kew Bull. 57:

759–761.

Carlquist S. 1988. Wood anatomy of Scytopetalaceae. Aliso 12:

63–76.

Diehl GA. 1935. A study of the Lecythidaceae. Trop. Woods 43:

1–15.

Frame D and S Durou. 2001. Morphology and biology of

Napoleonaea vogelii (Lecythidaceae) ﬂ owers in relation to

the natural history of insect visitors. Biotropica 33:

458–471.

Knuth R. 1934. Über die Gattung Asteranthos. Notizbl. Bot.

Gart. Berlin-Dahlem 11: 1034–1036.

Knuth R. 1939a. Barringtoniaceae. In: A Engler, ed. Pﬂ anzenreich,

IV. 219: 1–82. W. Engelmann, Leipzig.

Knuth R. 1939b. Lecythidaceae. In: A Engler, ed. Pﬂ anzenreich,

IV. 219a: l–144. W. Engelmann, Leipzig.

Knuth R. 1939c. Asteranthaceae. In: A Engler, ed. Pﬂ anzenreich,

IV. 219b: l–3. W. Engelmann, Leipzig.

Kowal RR. 1989. Chromosome numbers of Asteranthos and the

putatively related Lecythidaceae. Brittonia 41: 131–135.

Kowal RR, SA Mori, and JA Kallunki. 1977. Chromosome num-

bers of Panamanian Lecythidaceae and their use in subfamil-

ial classiﬁ cation. Brittonia 29: 399–410.

Kravtsova TI. 1991. Scytopetalaceae. In: AL Takhtajan, ed.

Comparative seed anatomy, 3: 226–228. Nauka, Leningrad

(in Russian).

Leins P. 1972. Das zentrifugale Androeceum von Conroupita

guianensis (Lecythidaceae). Beitr. Biol. Pﬂ . 48: 313–319.

Lens F, P Baas, S Jansen, and E Smets. 2007. A search for phy-

logenetically informative wood characters within

Lecythidaceae s.l. Am. J. Bot. 94: 483–502.

Letouzey R. 1961. Notes sur les Scytopetalacees (Revision des

Scytopetalacees de 1’herbier de Paris). Adansonia, ser. 2, 1:

106–142.

Liben L. 1971. Révision de genre africain Napoleonaea P.

Beauv. (Lecythidaceae). Bull. Jard. Bot. Natl. Belg. 41:

363–382.

Masters MT. 1869. On the structure of the ﬂ ower in the genus

Napoleona. Bot. J. Linn. Soc. 10: 492–504.

Miers J. 1874. On the Lecythidaceae. Trans. Linn. Soc. Lond.,

Bot. 30: 157–318.

Miers J. 1875a. On Napoleona, Omphalocarpum, and

Asteranthos. Trans. Linn. Soc. Lond., Bot. 2d ser.,1: 17–19.

Miers J. 1875b. On the Barringtoniaceae. Trans. Linn. Soc., 2d

ser., 1: 47–118.

Monteiro-Scanavacca WR. 1974. Vascularizay o do gineceu em

Lecythidaceae. Bol. Univ. São Paulo, Bot. 2: 53–69.

Monteiro-Scanavacca WR. 1975a. Vascularizac o e natureza de

estruturas do androceu em Lecythidaceae. Bol. Bot. Univ.

São Paulo 3: 61–74.

Monteiro-Scanavacca WR. 1975b. Estudo da Placentaeo em

Lecythidaceae. Bol. Bot. Univ. São Paulo 3: 75–86.

Mori SA, JE Orchard, and GT Prance. 1980. Intraﬂ oral pollen

differentiation in the New World Lecythidaceae subfamily

Lecythidoideae. Science 209: 400–403.

Mori SA and GT Prance. 1990. Lecythidaceae: Part II. The

zygomorphic ﬂ owered New World genera (Couroupita,

Corythophora, Bertholletia, Couratari, Eschweilera, and

Lecythis). Flora Neotropica 21: 1–376.

Mori SA, GT Prance, and AB Bolten. 1978. Additional notes on

the ﬂ oral biology of Neotropical Lecythidaceae. Brittonia

30: 113–130.

Mori SA, C-H Tsou, C-C Wu, B Crponholm, and AA Anderberg.

2007. Evolution of Lecythidaceae with an emphasis on the

circumscription of neotropical genera: information from

combined ndhF and trnL-F sequence data. Am. J. Bot. 94:

289–301.

Morton CM, SA Mori, GT Prance, KG Karol and MW Chase.

1997. Phylogenetic relationships of Lecythidaceae: a

cladistic analysis using rbcL sequence and morphological

data. Am. J. Bot. 84: 530–540.

Morton CM, GT Prance, SA Mori and LG Thorburn. 1998.

Recircumscription of the Lecythidaceae. Taxon 47: 817–827.

Muller J. 1972. Pollen morphological evidence for subdivision

and afﬁ nities of Lecythidaceae. Blumea 20: 350–355.

Muller J. 1973. Pollen morphology of Barringtonia calyptroca-

lyx K. Sch. (Lecythidaceae). Grana 13: 29–44.

Murthy GVS. 1996. Pollen morphology of Barringtoniaceae.

J. Palynol. 32: 79–83.

Payens JPDW. 1967. A monograph of the genus Barringtonia

(Lecythidaceae). Blumea 15: 157–263.

Prance GT. 1976. The pollination and androphore structure of

some Amazonian Lecythidaceae. Biotropica 8: 235–241.

Prance GT. 2004. Napoleonaeaceae. In: K Kubitzki, ed. The

families and genera of vascular plants, vol. 6, pp. 282–284.

Springer, Berlin/Heidelberg/New York.

Prance GT and SA Mori. 1978. Observations of the fruits and

seeds of neotropical Lecythidaceae. Brittonia 30: 21–33.

Prance GT and SA Mori. 1979. Lecythidaceae: Part I. The acti-

nomorphic-ﬂ owered New World Lecythydaceae (Asteranthos,

Gustavia, Grias, Allantoma, and Cariniana). Flora

Neotropica 21: 1–270.

Prance GT and SA Mori. 1983. Dispersal and distribution of

Lecythidaceae and Chrysobalanaceae. Sonderb. Naturwiss.

Vereins Hamburg 7: 163–186.

Prance GT and SA Mori. 2004. Lecythidaceae. In: K Kubitzki,

ed. The families and genera of vascular plants, vol. 6, pp.

221–232. Springer, Berlin/Heidelberg/New York.

Sarkar AK. 1983. Cytology of Barringtonia Forst.

(Barringtoniaceae) as an aid to their taxonomy. Indian Sci.

Congr. Assoc. Proc. 70: 77.

Tsou C-H. 1989. The ﬂ oral morphology and embryology of

Asteranthos and its systematic consideration. Am. J. Bot. 76

(Suppl.): 275–276.

Tsou C-H. 1993. Application of comparative embryology to

plant taxonomy: taking the Lecythidaceae as an example.

Recent Adv. Bot. 13: 293–302.

Tsou C-H. 1994a. The embryology, reproductive morphology,

and systematics of Lecythidaceae. Mem. N. Y. Bot. Gard. 71:

1–110.

Tsou C-H. 1994b. The classiﬁ cation and evolution of pollen

types of Planchonoideae (Lecythidaceae). Plant Syst. Evol.

189: 15–27.

Tsou C-H and SA Mori. 2007. Floral organogenesis and ﬂ oral

evolution of the Lecythidoideae (Lecythidaceae). Am. J. Bot.

94: 716–736.

Venkateswarlu J. 1952. Embryological studies in Lecythidaceae.

Part I. J. Indian Bot. Soc. 31: 103–116.

Vijayaraghavan MR and U Diar. 1976. Scytopetalum tieghemii:

embryologically unexplored taxon and afﬁ nities of the fam-

ily. Phytomorphology 26: 16–22.

206 Subclass V. DILLENIIDAE

Weberling F. 1958. Über das Vorkommen rudimentarer Stipein

bei den Lecythidaceae (s.1.) und Sonneratiaceae. Flora 116:

72–77.

Zeeuw de CH. 1990. Secondary xylem of Neotropical

Lecythidaceae. In: I Mori and GT Prance, eds.

Lecythidaceae – Part II, chap. II. The zygomorphic-ﬂ owered

New World genera. Flora Neotropica, Monograph 21(3),

pp. 4–59.

Order 59. SARRACENIALES

Perennial, rhizomatous, mostly acaulescent, carnivorous

herbs. Rhizomes with vascular bundles arranged in a

discontinuous ring interrupted by medullary rays of

unequal width. Bundles bounded internally and exter-

nally by sclerenchyma. Vessels with oblique, scalari-

form plates with 3–50 bars; lateral pitting from

scalariform to alternate. Fibers with bordered pits. Axial

parenchyma diffuse. Leaves alternate, borne in a radical

rosette or (species of Heliamphora) on an upright stem;

all or some of the leaves highly modiﬁ ed, transformed

into more or less elongate, often pitcherlike traps partly

ﬁ lled with digestive liquid and with a more or less devel-

oped ridge or ﬂ at wing on the ventral side and a laminar

and relatively small epidermis provided with special-

ized glands and hairs that serve to entrap insects and

other small organisms. Stomata anomocytic. Flowers

rather large, solitary on a scape or (Heliamphora) in few

ﬂ owered, sometimes axillary racemes, nodding, bisex-

ual, actinomorphic, subtended by 1–3 bracteoles. Sepals

(3)4–5(6), free, imbricate, persistent, often colored and

showy. Petals ﬁ ve, free, imbricate, caducous or

(Heliamphora) wanting. Stamens numerous or 10–15,

in Sarracenia several, arising from each of a limited

number (commonly 10) of primordia; ﬁ laments more or

less short; anthers basiﬁ xed or (Sarracenia) versatile,

opening longitudinally, introrse. Tapetum secretory.

Microsporogenesis simultaneous. Pollen grains 2-celled,

(3)4-polycolporate. Gynoecium of ﬁ ve or (Heliamphora)

three united carpels, with slightly 3-lobed style and

more or less truncate at the apex (Heliamphora), with

ﬁ ve short branches at the apex, each with a terminal

stigma (Darlingtonia) or apically expanded and peltate,

umbrella-shaped, with small stigma under the tip of

each of the ﬁ ve lobes (Sarracenia); ovary superior,

5-locular or (Heliamphora) 3-locular, but the partition

of the upper part is often not complete, and the placenta-

tion is therefore axile below and parietal above;

Sarracenia has 10 nectaries on the ovary wall above the

stamen fascicles. Ovules numerous, anatropous, uniteg-

mic or (Darlingtonia) bitegmic (?), tenuinucellate, with

endothelium and a well-developed hypostase. Female

gametophyte of Polygonum-type. Endosperm cellular.

Fruits loculicidal capsules with numerous seeds. Seeds

small, often winged, with exotestal seed coat, copious

endosperm rich in oil and protein, and small linear

embryo. Decarboxylated iridoids and O-methyl ﬂ a-

vonols present, n = 13 (Sarracenia), 15 (Darlingtonia),

21 (Heliamphora).

Probably related to the Theales, which is supported

by ﬂ ower morphology, embryology, and wood anat-

omy (see DeBuhr 1975, 1977).

1. SARRACENIACEAE

Dumortier 1829 (including Heliamphoraceae Chrtek,

Slavikova, et Studnicka 1992). 3/15. North America

(Darlingtonia and Sarracenia) and Guyana Highland.

Heliamphora, Darlingtonia, Sarracenia.

Bibliography

Arber A. 1941. On the morphology of the pitcher-leaves in

Heliamphora, Sarracenia, Darlingtonia, Cephalotus, and

Nepenthes. Ann. Bot. II 5: 563–578.

Bayer RJ, L Hufford, and DE Soltis. 1996. Phylogenetic rela-

tionships in Sarraceniaceae based on rbcL and ITS sequences.

Syst. Bot. 21: 121–134.

Bell CR. 1949. A cytotaxonomic study of the Sarraceniaceae of

North America. J. Elisha Mitchell Sci. Soc. 65: 137–166.

Chrtek J, Z Slavikova, and M Studnicka. 1992. Beitrag zur

Morphologie und Taxonomie der Familie Sarraceniaceae.

Preslia 64(1): 1–10.

DeBuhr LE. 1973. Distribution and reproductive biology of

Darlingtonia californica. M.A. thesis, Claremont Graduate

School, Claremont.

DeBuhr LE. 1975. Phylogenetic relationships of the

Sarraceniaceae. Taxon 24: 297–306.

DeBuhr LE. 1977. Wood anatomy of the Sarraceniaceae: eco-

logical and evolutionary implications. Plant. Syst. Evol. 128:

159–169.

Juniper BE, RL Robins, and DM Joel. 1989. The carnivorous

plants. Academic, London.

Kubitzki K. Sarraceniaceae. 2004. In: K Kubitzki, ed. The fami-

lies and genera of vascular plants, vol. 6, pp. 422–425.

Springer, Berlin/Heidelberg/New York.

Lloyd FE. 1942. The carnivorous plants. Chronica Botanica,

Waltham, MA.

Maguire B. 1978. Sarraceniaceae. The botany of Guayana

Highland – part X. N. Y. Bot. Gard. 29: 36–62.

Superorder ERICANAE 207

Markgraf F. 1955. Über Laubblatt-Homologien und ver-

wandtschaftliche Zusammenhänge bei Sarraceniales. Planta

46: 414–446.

Newman T, S Ibrahim, JW Wheeler, WB McLaughlin, RL

Petersen, and RM Dufﬁ eld. 2000. Identiﬁ cation of sarracenin

in four species of Sarracenia (Sarraceniaceae). Biochem.

Syst. Ecol. 28: 193–195.

Neyland R and M Merchant. 2006. Systematic relationships of

Sarrceniaceae inferred from nuclear ribosomal DNA

sequences. Madroño 53: 223–232.

Shreve F. 1906. The development and anatomy of Sarracenia

purpurea. Bot. Gaz. 42: 107–126.

Terekhin ES. 1992. Sarraceniaceae. In: A Takhtajan, ed.

Comparative seed anatomy, vol. 4, pp. 21–23. Nauka, St.

Petersburg (in Russian).

Thanikaimoni G and G Vasanthy. 1972. Sarraceniaceae: palyn-

ology and systematics. Pollen et Spores 14: 143–155.

Wood CR. 1960. The genera of Sarraceniaceae and Droseraceae

in the southeastern United States. J. Arnold Arbor. 41:

152–163.

Order 60. RORIDULALES

Sparsely branched shrubs bearing glandular (tentacular)

hairs of various lengths and unicellular bristle hairs on

young stems, leaves, and calyx; the largest tentacular

trichomes are situated on the leaf margins; secretion of

tentacular hairs sticky, resinous, without proteolytic

enzymes. Vessel elements long and have oblique perfo-

ration plates with scalariform perforations that in

Roridula gorgonias have up to 83 bars per plate; lateral

pitting scalariform to alternate. Fibers are tracheids.

Rays mostly uniseriate (rarely biseriate) and composed

of upright cells. Axial parenchyma sparse, diffuse.

Leaves clustered toward the branch ends of adult shrubs,

sessile, alternate, linear-lanceolate, entire or with linear

lateral lobes, pinnately veined, circinate in vernation,

estipulate. Stomata anomocytic. Flowers in few-

ﬂ owered, axillary panicles terminated by a ﬂ ower, acti-

nomorphic, 5-merous as to the perianth and androecium,

bibracteolate. Sepals free, imbricate, glandular-hairy.

Petals free, imbricate, glabrous, purple or crimson to

nearly white. Stamens ﬁ ve, alternipetalous; ﬁ laments

free, slender, thickened at the base; anthers 2-locular,

elongate, tetrasporangiate, inverted in bud, with

basal thickening containing a nectariferous cavity,

irritable (swinging upward and becoming erect and

spreading pollen when touched at anthesis), extrorse,

basiﬁ xed, opening by terminal pores. Tapetum secretory.

Microsporogenesis simultaneous. Pollen grains 2-celled,

3-colporate, tectate-perforate and spinuliferous (R. dentata)

or without distinctly developed apertures and without

coherent tectum (R. gorgonias). Gynoecium of three

carpels; style tapering and with small, capitate stigma

(R. dentata) or expanding, terminal, apically obconical

and with a large stigma (R. gorgonias); ovary 3-locular,

superior, with solitary (R. dentata) or 2–4 (R. gorgon-

ias) ovules per lucule. Ovules pendulous, with endothe-

lium. Female gametophyte of Polygonum-type, with a

conspicuous hypostase at the chalazal end. Endosperm

cellular. Fruits 3-valved locultcidal capsules. Seeds

exotestal, rather large, reticulate (R. dentata) or warty to

honeycombed (R. gorgonias) with small, straight

embryo and copious, ﬂ eshy endosperm. Producing iri-

doid compounds. Tannins present in the seed coat and

endosperm cells, n = 6.

1. RORIDULACEAE

Martynov 1820. 1/2. Endemic to the Cape Province.

Roridula.

Hutchinson (1959, 1973) and Cronquist (1981,

1988) place Roridula in the Byblidaceae, but as Vani-

Hardev (1972) showed there are profound morphologi-

cal differences between Roridula and Byblis. Albert

et al. (1992) placed the Roridulaceae as sister to the

carnivorous pitcher plant family Sarraceniaceae.

Bibliography

Bruce AN. 1907. On the distribution, structure, and function of

the tentacles of Roridula, Notes Roy. Bot. Gard. Edinb. 17:

85–96.

Carlquist S. 1976. Wood anatomy of Roridulaceae: ecological

and phylogenetic implications. Am. J. Bot. 63: 1003–1008.

Conran JG. 2004. Roridulaceae. In: K Kubitzki, ed. The families

and genera of vascular plants, vol. 6, pp. 339–342. Springer,

Berlin/Heidelberg/New York.

Conran JG and JW Dowd. 1993. The phylogenetic relationships

of the Byblis – Roridula (Byblidaceae: Roridulaceae) com-

plex inferred from 18S rRNA partial sequences. Plant Syst.

Evol. 188: 73–86.

Lobova TA. 2000. Roridulaceae. In: A Takhtajan, ed.

Comparative seed anatomy, 6: 289–292. Nauka, St.

Petersburg (in Russian).

Vani-Hardev S. 1972. Systematic embryology of Roridula gor-

gonias Planch. Beitr Biol. Pﬂ . 48: 339–351.

Wilkinson HP. 1998. Roridulaceae. In: DF Cutler and M Gregory,

eds. Anatomy of the Dicotyledons, Saxifragales (sensu

Armen Takhtajan 1983), vol. 4, pp. 121–130. Clarendon,

Oxford.

208 Subclass V. DILLENIIDAE

Superorder PRIMULANAE

Order 61. STYRACALES (EBENALES)

Trees and shrubs. Vessels with scalariform or simple

perforations; lateral pitting from scalariform to alternate.

Fibers with bordered or simple pits. Rays mostly het-

erogeneous, mixed uniseriate and pluseriate, or some-

times all uniseriate. Axial parenchyma mostly

apotracheal. Nodes unilacunar. Leaves alternate or

rarely opposite, simple, entire or dentate to serrate,

estipulate. Stomata mostly anomocytic, sometimes

paracytic or encyclocytic. Flowers usually in various

racemose or less often cymose inﬂ orescences, bisexual

or less often unisexual, actinomorphic, with bracteoles

or (Styracaceae) without. Calyx synsepalous, usually

tubular or copulate, with (2)3-7-9 lobes. Corolla usually

sympetalous and 2–7-lobed or very rarely (Bruinsmia

in Styracaceae) petals free or nearly free. Stamens two

times as many as the corolla lobes or less often same

number, adnate at different levels upon the petals or

corolla tube; ﬁ laments free or more or less connate;

anthers tetrasporangiate, opening longitudinally or

rarely (some species of Diospyros) by apical pores.

Tapeturn secretory. Microsporogenesis simultaneous.

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

or 3-porate. Gynoecium of 2–8 united carpels with sty-

lodia more or less completely connate into a style, rarely

(some Ebenaceae) nearly free; stigma lobed or capitate;

ovary superior to inferior, usually plurilocular, with 1–4

or more ovules per locule; placentation axile

(Symplocaceae), axile to basal (Styracaceae), or apical

(Ebenaceae). Ovules anatropous to hemitropous, pen-

dulous (Ebenaceae), bitegmic (Ebenaceae) or unitegmic

(Symplocaceae and most of the Styracaceae), tenuinu-

cellate, sometimes (Ebenaceae and Pterostyrax and

Alniphyllum in Styracaceae) with an endothelium.

Female gametophyte of Polygonum-type. Endosperm

cellular. Fruits of various types. Seeds large, hard, often

shiny; seed coat thin, exotestal; embryo straight or

slightly curved; endosperm copious, mostly bony, often

ruminate. Without iridoid compounds.

Styracales evidently derived directly from some

thealean ancestor. They also resemble the Ericales, but

I agree with Cronquist (1981: 493) that “the relation-

ship is collateral rather than ancestral.” I also agree

that no one family of the Styracales is likely to be

directly ancestral to any of the others.

Key to Families

1 Flowers bisexual or seldom polygamo-dioecious.

Style with capitate or lobed stigma. Gynoecium of

2–5 carpels. Ovary superior to often inferior. Vessels

mostly or always with scalariform perforations; lat-

eral pitting from scalariform to opposite, rarely

alternate.

2 Pubescence of stellate or stellate-peltate trichomes.

Trees and shrubs, often with resiniferous inter-

cellular canals in the bark and wood but without

latex. Leaves alternate, simple, entire to toothed.

Stomata anomocytic. Inﬂ orescences axillary or

terminal, racemes or panicles, rarely the ﬂ owers

solitary. Flowers without bracteoles, bisexual or

rarely female in gynodioecious; sepals 4–5(-9),

connate, the lobes valvate or the calyx open in

bud, often truncate; petals 4–5(-9), mostly basally

connate. Stamens 4-10(-14), the ﬁ laments usu-

ally connate and often inserted basally on the

corolla, rarely hypogynous. Anthers linear to

oblong, basiﬁ xed. Pollen grains 3-colporate.

Stylodia completely united or (Huodendron)

united at lower level but free above; ovary supe-

rior to inferior, plurilocular below but usually

unilocular above; ovules bitegmic (Styrax) or

unitegmic (Halesia); micropyle endostomal.

Fruits nut-like, sometimes winged, or a loculi-

cidal capsule, samara, or a drupe, with persistent

calyx. Seeds one of few, rarely numerous; embryo

large, straight or slightly curved; endosperm

copious, oily. Contain quercetin, kaempferol,

tannins, a small amount of leucoanthocyanin,

saponins, coffeic and ferulic acids, n = 8 (Styrax)

or 12. . . . . . . . . . . . . . . . . . . . . 1. styracaceae.

2 Plants glabrous or with multicellular hairs or

stalked glands, but without stellate hairs.

Evergreen or rarely deciduous trees or shrubs,

commonly accumulating aluminum. Leaves alter-

nate, simple, serrate or entire. Stomata mostly

paracytic. Flowers provided with two bracteoles,

in racemes, or less often in spikes, panicles, or

fascicles, or solitary, bisexual, rarely unisexual.

Sepals ﬁ ve, connate, imbricate or valvate, persis-

tent. Petals 5 (3–11), imbricate, connate basally

in 1–2 series. Stamens 4–12 to numerous (to 100),

the ﬁ laments free or connate basally. Anthers

short and broad, mostly round ovate. Pollen grains

angular, spinuliferous. Ovary wholly or partly

inferior, usually 2–5-locular, usually with an

Superorder PRIMULANAE 209

apical 5-glandular, annular, cylindrical, or 5-lobed

disc; ovules 2–4 per lacule, pendulous, unitegmic

(Corner 1976). Fruits 1-seeded drupes or less

often baccate. Seeds with linear, straight or arcu-

ate embryo and copious endosperm; exotestal

cells with inner walls thickened or thin. Present

gallic and ellagic acids, tannins, leucoanthocya-

nins, iridoids, n = 11. . . . . . . .2. symplocaceae.

1 Flowers mostly unisexual and commonly dioecious,

rarely bisexual. Trees and shrubs, commonly with

hard, dark or black wood, without latex. Vessels with

simple perforations, or less greauently scalariform;

lateral pitting alternate. Nodes unilacunar with three

traces. Leaves alternate, rarely opposite or whorled,

simple, entire. Stomata usually paracytic. Inﬂ orescences

axillary or rarely cauliﬂ orous cymes, or ﬂ owers soli-

tary. Sepals 3–5(-8), connate, valvate or imbricate;

petals 3–5(-8), connate, usually contorted and imbri-

cate, rarely valvate. Stamens 3-50(-120 ?), usually

2–4 times the number of corolla lobes, rarely equal

in number and alternate with them; the ﬁ laments free

or connate in pairs; anthers elongated, often hairy,

adnate or basiﬁ xed, dehiscing longitudinally, more

rarely by apical pores. Pollen grains 3-colporate or

3-porate. Gynoecium of 2–8 carpels. Ovary superior,

2–16-locunar. Stylodia nearly free or more or less

connate below. Ovules 1–2 per locule, bitegmic;

micropyle endostomal. Fruits more or less juicy and

leathery berries rarely capsular. Seeds pachychala-

zal, often ruminate, testa multiplicative (not vascu-

larised), exotesta ﬁ briform or mucilaginous, cells

cuboid to palisade, endotesta crystalliferous or not,

walls thickened or not; embryo straight or slightly

curved; endosperm copious, hard. Producing

saponins, triterpenes, napthoquinone derivative of

seven methyljugone and plumbagin, ﬂ avonols, leu-

codelphinidin, myricetin, n = 15. . . . 3. ebenaceae.

1. STYRACACEAE

A.P. de Candolle et Sprengel 1821 (including Halesi-

aceae D. Don 1828). 11/160. Mediterranean, eastern

Himalayas (from Nepal to Arunachal-Pradesh), Assam,

eastern India, eastern and Southeast Asia to West

Malesia, America from United States to Argentina.

Styrax (including Pamphilia), Huodendron, Bruins-

mia, Alniphyllum, Changiostyrax, Sinojackia, Parastyrax,

Pterostyrax, Halesia, Rehderodendron, Melliodendron.

Styracaceae (together with the Symplocaceae) are

the most primitive members of the order and the near-

est to the Theales (see Copeland 1938).

2. SYMPLOCACEAE

Desfontaines 1820. 1/300. Tropical and subtropical

regions except Africa, Europe, and western Asia.

Symplocos.

Very close to the Styracaceae and approximately on

the same level of specialization.

3. EBENACEAE

Gürke 1891 (including Diospyraceae Durande 1782,

Guaiacanaceae Jussieu 1789, Lissocarpaceae Gilg

1924). 4–5/500. The largest genus Diospyros (up to

500) is pantropical with the greatest concentration of

species in Malesia; only a few species of Diospyros

occur in western Asia, Japan, and the southeastern

United States. Euclea (14) occurs in East and South

Africa, on the Comoros and Arabian peninsula.

Tetraclis (3) endemic to Madagascar, Lissocarpa

(8) tropical South America (from Guyana to Bolivia).

3.1 EBENOIDEAE

Pollen grains 3-colporate. Ovary superior, 2–16-locu-

nar, ovules one or less frequently two; indumentum

present; bracteoles alternate. – Diospyros, Royena,

Tetraclis, Euclea.

3.2 LISSOCARPOIDEAE

Pollen grains 3-porate. Ovary inferior, 4-locular;

ovules two; indumentum absent; bracteoles suboppo-

site. – Lissocarpa.

Related to the Styracaceae and Symplocaceae, but

the afﬁ nity is not very close. A very heterobathmic fam-

ily: ovules bitegmic and stylodia not completely united

or sometimes nearly free, but ﬂ owers mostly unisexual.

Bibliography

Barth OM. 1982. The sporoderma of Brazilian Symplocos pollen

types (Symplocaceae). Grana 18: 65–69.

Berry PE. 1999. A synopsis of the family Lissocarpaceae.

Brittonia 51: 214–216.

210 Subclass V. DILLENIIDAE

Berry PE, V Savolainen, KJ Sytsma, JC Hall, and MW Chase.

2001. Lissocarpa is sister to Diospyros (Ebenaceae). Kew

Bull. 56: 725–729.

Contreras LS and NR Lersten. 1984. Extraﬂ oral nectaries in

Ebenaceae: anatomy, morphology, and distribution. Am. J.

Bot. 71: 865–872.

Copeland HF. 1938. The Styrax of northern California and

the relationships of the Styracaceae. Am. J. Bot. 25:

771–780.

Caris P, LP Ronse Decraene, E Smets, and D Clinckemaillie.

2002. The uncertain systematic position of Symplocos

(Symplocaceae): evidence from a ﬂ oral ontogenetic study.

Int. J. Plant Sci. 163: 67–74.

Dickison WC. 1993. Floral anatomy of the Styracaceae, includ-

ing observations on intraovarian trichomes. Bot. J. Linn.

Soc. 112: 223–255.

Dickison WC and KD Phend. 1985. Wood anatomy of the

Styracaceae: evolutionary and ecological considerations.

IAWA Bull. 6: 3–22.

Duangiai S, B Wallnöfer, R Samuel, J Munzinger, and MW

Chase. 2006. Generic delimitation and relationships in

Ebenaceae sensu lato: evidence from six plastid DNA

regions. Am. J. Bot. 93: 1808–1827.

Franceschi, G. de. 1993. Phylogénie des Ebénales: analyse de

l’ordre et origin biogéographique des espèces indiennes.

Institut Français de Pondichery, Département de Écologie

33: 1–153.

Fritsch P. 1996. Isozyme analysis of intercontinental disjuncts

within Styrax (Styracaceae): implications for the Madrean-

Tethyan hypothesis. Am. J. Bot. 83: 342–355.

Fritsch PW. 1999. Phylogeny of Styrax based on morphological

characters, with implications for biogeography and infrage-

neric classiﬁ cation. Syst. Bot. 24: 356–378.

Fritsch PW. 2001. Phylogeny and biogeography of the ﬂ owering

plant genus Styrax (Styracaceae) based on chloroplast DNA

restriction sites and DNA sequences of the internal tran-

scribed spacer region. Molec. Phylogen. Evol. 19: 387–408.

Fritsch PW. 2004. Styracaceae. In: K Kubitzki, ed. The families

and genera of vascular plants, vol. 6, pp. 434–442. Springer,

Berlin/Heidelberg/New York.

Fritsch PW, CM Morton, T Chen, and C Meldrum. 2001.

Phylogeny and biogeography of the Styracaceae. Int. J. Plant

Sci. 162(6, Suppl.): 95–116.

Fritsch PW, BC Cruz, F Almeda, Y Wang, and S Shi. 2006.

Phylogeny of Symplocos based on DNA sequences of the

chloroplast trnC-trnD intergenic region. Syst. Bot. 31:

181–192.

Kolbe K-P and J John. 1980. Serology and systematics of the

Ebenales and the Theales. Biochem. Syst. Ecol. 8: 249–256.

Komar GA. 1992. Ebenales. In: A Takhtajan, ed. Comparative

seed anatomy, 4: 45–48. Nauka, St. Petersburg (in Russian).

Li Y-H and C-H Yu. 1985. Pollen morphology of Styracaceae

and its taxonomic signiﬁ cance. Acta Phytotax. Sinica 23:

81–90 (in Chinese with English summary).

Liang YH. 1986. Pollen morphology of the family Symplocaceae

from China. Acta Bot. Austro Sinica 2: 111–121.

Machado SR, V Angyalossy-Alfonso, and BL de Morretes.

1997. Comparative woody anatomy of root and stem in

Styrax camporum (Styracaceae). IAWA J. 18: 13–25.

Mallavadhani UV, AK Panda, and YR Rao. 1998. Pharmacology

and chemotaxonomy of Diospyros. Phytochemistry 49:

901–951.

Manshard E. 1936. Embryologische Untersuchungen an Styrax

obassia Sieb. et Zucc. Planta 25: 364–383.

Morawetz W. 1991. The karyology of some Neotropical

Styracaceae. Plant Syst. Evol. 177: 111–115.

Morton CM and WC Dickison. 1992. Comparative pollen mor-

phology of the Styracaceae. Grana 31: 1–15.

Morton CM, MW Chase, KA Kron, and SM Swensen. 1996.

A molecular evaluation of the monophyly of the order Ebenales

based upon rbcL sequence data. Syst. Bot. 21: 567–586.

Morton CM, MW Chase, KA Kron, and SA Swensen. 1997.

A molecular evaluation of the monophyly of the order Ebenales

based upon rbcL sequence data. Syst. Bot. 21: 567–586.

Ng ESP. 1971. A taxonomic study of the Ebenaceae with special

reference to Malesia. Doctoral thesis, University of Oxford,

Oxford.

Nikiticheva ZI. 1983. Ebenales. In: MS Yakovlev, ed. Com-

parative embryology of ﬂ owering plants: Phyto laccaceae-

Thymelaeaceae, pp. 221–233. Nauka, Leningrad (in

Russian).

Nooteboom HP. 1975. Revision of the Symplocaceae of the Old

World (New Caledonia excepted). Leiden University Press,

Leiden.

Nooteboom HP. 2004. Symplocaceae. In: K Kubitzki, ed. The

families and genera of vascular plants, vol. 6, pp. 443–449.

Springer, Berlin/Heidelberg/New York.

Oever L van den, P Baas, and M Zandoe. 1981. Comparative

wood anatomy of Symplocos and latitude and altitude of

provenance. IAWA Bull. 2: 3–24.

Oliver D. 1895. Lissocarpa benthamii Gürke. Hooker’s Icon.

Plant. 25: t.2413.

Periasamy K. 1966. Studies on seeds with ruminate endosperm.

VI. Rumination in the Araliaceae, Aristolochiaceae,

Caprifoliaceae, and Ebenaceae. Proc. Indian Acad. Sci. 60B:

127–134.

Schadel WE. 1978. Leaf anatomy and venation patterns of the

Styracaceae. Thesis, Master of Arts, University of North

Carolina at Chapel Hill.

Schadel WE and WC Dickison. 1979. Leaf anatomy and vena-

tion patterns of Styracaceae. J. Arnold Arbor. 60: 8–27.

Shyamala R and N Gulati. 1981. Contribution to the embryo-

logy of Symplocos cochinchinensis. Indian J. Bot. 4:

229–235.

Shyamala R and N Gulati. 1986. Origin and structure of integu-

ments in Ebenales: I. Symplocaceae. Swamy Bot. Club 2:

113–117.

Takasi Y. 1972. Embryological studies in Ebenales, part 4.

J. Jpn. Bot. 47: 20–28.

Veillet-Bartoszewska M. 1960. Embryogenie des Styra-

cacees: Developpement de l’embryon chez le Styrax ofﬁ ci-

nalis L. Compt. Rend. Hebd. Seances Acad. Sci. 250:

905–907.

Wallnöfer B. 2001. The biology and systematics of Ebenaceae:

a review. Ann. Naturhist. Mus. Wien. 103B: 485–512.

Wallnöfer B. 2004a. Ebenaceae. In: K Kubitzki, ed. The families

and genera of vascular plants, vol. 6, pp. 125–130. Springer,

Berlin/Heidelberg/New York.

Wallnöfer B. 2004b. Lissocarpaceae. In: K Kubitzki, ed. The

families and genera of vascular plants, vol. 6, pp. 236–238.

Springer, Berlin/Heidelberg/New York.

Wallnöfer B. 2004c. A revision of Lissocarpa Benth. (Ebenaceae

subfam. Lissocarpoideeae (Gilg in Engler) B. Walln. Ann.

Naturhist. Mus. Wien, B, 105: 515–564.

Superorder PRIMULANAE 211

Wang Y, PW Fritsch, S Shi, F Almeda, BC Criz, and LM Kelly.

2004. Phylogeny and infragenetic classiﬁ cation of Symplocos

(Symplocaceae) inferred from DNA sequence data. Am. J.

Bot. 91: 1901–1914.

White EB. 1956–1963. Notes on Ebenaceae: I, II. Bull. Jard.

Bot. L’Etat 26: 237–246; 277–307, 1956. III. Bull. Jard. Bot.

L’Etat 27: 515–531, 1957. IV. Bol. Soc. Brot. 36: 97–100,

1962. V. Bull. Jard. Bot. L’Etat 33: 345–367, 1963.

Wood CE and RB Channell. 1960. The genera of the Ebenales

in the southeastern United States. J. Arnold Arbor. 41:

1–35.

Yamazaki T. 1970, 1972. Embryological studies in Ebenales: 1.

Styracaceae. 2. Symplocaceae. 4. Ebenaceae. J. Jpn. Bot. 45:

267–273; 353–358, 1970. 47: 20–28, 1972.

Order 62. SAPOTALES

Trees and shrubs with well-developed, elongate laticif-

erous sacks in the pith, phloem, cortex, and leaves that

contain a refractive white substance resembling resin

or guttapercha. Indumentum in all genera, except

Delpydora, of appressed unicellular hairs with two equal

or unequal arms commonly mixed with erect or spread-

ing simple hairs. Vessels with simple perforations; lat-

eral pitting alternate. Fibers typically with simple pits,

but pits occasionally bordered. Rays heterogeneous.

Axial parenchyma apotracheal. Sieve-element plastids

of S-type. Nodes trilacunar or seldom unilacunar. Leaves

alternate or seldom opposite, simple, nearly always

entire, pinnately veined, with small to large and some-

times persistent stipules (e.g., Madhuca spp.) or more

often estipulate. Stomata usually anomocytic. Flowers

mostly rather small, solitary or more often in small

cymes, sometimes (Sarcosperma) in large paniculate

inﬂ orescences, bisexual, actinomorphic, often nocturnal

and bat- pollinated. Sepals (4)5(-12), free or nearly so

and imbricate, or sometimes in two cycles of two, three

or four. Corolla sympetalous, with 4–8 imbricate lobes,

sometimes with paired appendages. Stamens in 1–3

cycles, opposite the corolla lobes, inserted within the

corolla tube or occasionally at the base of the corolla

lobes, rarely free, often some of them staminodial;

anthers tetrasporangiate, opening longitudinally.

Tapetum secretory. Microsporogenesis simultaneous.

Pollen grains 2-celled, 3–5(6)-colporate, with colpi pro-

vided by granular membrane; ectexine nearly always

with a thick and usually sparsely punctate or perforate

tectum and a very narrow interstitium, either granular or

with very reduced columellae. Gynoecium of 2-14(-30)

united carpels, rarely (several species of Pouteria sect.

Franchetella) of one carpel, usually hairy; style with

capitate or slightly lobed stigma; ovary superior, typi-

cally 5-plurilocular (unilocular in Diploon), with axile

or sometimes basal or axile-basal placentation and one

ovule per carpel. Ovules anatropous to hemitropous,

apotropous, unitegmic, tenuinucellate, without endothe-

lium. Female gametophyte of Polygonum-type.

Endosperm nuclear. Fruits typically 1- to many-seeded

berries with ﬂ eshy, leathery, or rarely woody outer peri-

carp, rarely (as in Pradosia) drupelike or sometimes, as

in four genera of Mimusopeae, tardily dehiscing 1-sev-

eral-seeded loculicidal capsule. Seeds medium-sized to

large, ellipsoid, light brown to black, shiny, often with

variously expanded hilum (basal, basi-ventral or adax-

ial); seed coat thick and hard, its outer part (8–25 cells

thick or more) often forming a heavily ligniﬁ ed, scle-

rotic layer of compressed, pitted cells, and its inner part

(of similar thickness) thin-walled, aerenchymatous, car-

rying the vascular bundles, eventually more or less

crushed; embryo large, usually with small radicle;

endosperm sometimes relatively copious (Manilkara and

some other genera), but mostly scanty or even wanting.

Producing saponins, C-30 oxidised triterpenes, pyrrhliz-

idine alkaloids, ﬂ avonols, leucodelphinidin, myrcetin, but

without iridoid compounds, n = (10-)13(-14).

Sapotales are related to the Styracales and share a

common origin from the thealean ancestor. However,

they differ from the Styracales in many important char-

acters including a well-developed system of latex

sacks, nuclear endosperm, absence of the endothelium,

and seed coat anatomy.

1. SAPOTACEAE

A.L. de Jussieu 1789 (including Achradaceae Vest

1818, Boerlagellaceae H.J. Lam 1925, Bumeliaceae

Barnhardt 1895, Sarcospermataceae H.J. Lam 1925).

53/1200. Pantropical, with a few species extending

into temperate regions.

mimusopeae: Mimusops, Vitellariopsis, Autranella,

Tieghemella, Baillonella, Vitellaria, Manilkara,

Labramia, Faucherea, Northia, Labourdonnaisia,

Letestua, Inhambanella, Neolemonniera, Lecomtedoxa,

Gluema, Eberhardtia; isonandreae: Palaquium,

Aulandra, Isonandra, Madhuca, Payena, Burckella,

Diploknema; sideroxyleae: Sideroxylon, Neohems-

leya, Nesoluma, Argania, Sarcosperma, Diploon;

chrysophylleae: Pouteria, Aubregrinia, Breviea,

Micropholis, Chromolucuma, Chrysophyllum, Eccli-

nusa, Delpydora, Pichonia, Sarcaulus, Elaeoluma,