Takhtajan Armen. Flowering Plants

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Superorder HAMAMELIDANAE 121

Conti E, KJ Sytsma, and WS Alverson. 1994. Neither oak nor

alder, but nearly: The relationships of Ticodendron based on

rbcL data. Am. J. Bot. 81(6): 149 (Abstract).

Erdogan V and SA Mehlenbacher. 2000. Phylogenetic relation-

ships of Corylus species (Betulaceae) based on nuclear ribo-

somal DNA ITS region and chloroplast matK gene sequences.

Syst. Bot. 25: 727–737.

Feuer S. 1991. Pollen morphology and the systematic relation-

ships of Ticodendron incognitum. Ann. Missouri Bot. Gard.

78: 143–151.

Furlow JJ. 1990. The genera of Betulaceae in the southeastern

United States. J. Arnold Arbor. 71: 1–67.

Gomez-Laurito J and PL Gomez. 1989. Ticodendron: A new tree

from Central America. Ann. Missouri Bot. Gard. 76:

1148–1151.

Gomez-Laurito J and PL Gomez. 1991. Ticodendraceae: A new

family of ﬂ owering plants. Ann. Missouri Bot. Gard. 78: 87–88.

Hall JW. 1952. The comparative anatomy and phylogeny of the

Betulaceae. Bot. Gaz. 113: 235–270.

Hammel B and W Burger. 1991. Neither oak nor alder, but

nearly: The history of Ticodendraceae. Ann. Missouri Bot.

Gard. 78: 89–95.

Hardin JW and JM Bell. 1986. Atlas of foliar surface features in

woody plants: IX. Betulaceae of eastern United States.

Brittonia 38: 133–144.

Hickey LJ and DW Taylor. 1991. The leaf architecture of

Ticodendron and the application of foliar characters in discern-

ing its relationships. Ann. Missouri Bot. Gard. 78: 105–130.

Jager EJ. 1980. Progressionen im Synﬂ oreszenzbau und in der

Verbreitung bei den Betulaceae. Flora 170: 91–113.

Kato H, K Oginuma, Z Gu, B Hammel and H Tobe. 1999.

Phylogenetic relationships of Betulaceae based on matK

sequences with particular reference to the position of

Ostryopsis. Acta Phytotax. Geobot. 49: 89–97.

Khramova EL. 1996. Palynomorphology of the genus Alnus

(Betulaceae). Bot. Zhurn. 81(3): 42–53 (in Russian with

English summary).

Korchagina IA. 1991. Betulaceae. In: A Takhtajan, ed.

Comparative seed anatomy, vol. 3, pp. 134–140. Nauka,

Leningrad (in Russian).

Kubitzki K. 1993a. Betulaceae. In: K Kubitzki, ed. The families

and genera of vascular plants, vol. 2, pp. 152–157. Springer,

Berlin/Heidelberg/New York.

Kubitzki K. 1993b. Ticodendraceae. In: K Kubitzki, ed. The

families and genera of vascular plants, vol. 2, pp. 594–596.

Springer, Berlin/Heidelberg/New York.

Oginuma K, Kato H, Tobe H. 1999. Chromosome base number

of Corylus (Betulaceae): correction, and evolution. Acta

Phytotax. Geobot. 49(2): 99–104.

Snow N and P Goldblatt. 1992. Chromosome number in

Ticodendron (Fagales, Ticodendraceae). Ann. Missouri Bot.

Gard. 79: 906–907.

Schneider H. 1994. Sprossorganisation und Blattarchitektur bei

Betulaceae und extratropischen Fagaceae. Cramer, Berlin.

Tobe H. 1991. Reproductive morphology, anatomy, and relation-

ships of Ticodendron. Ann. Missouri Bot. Gard. 78: 135–142.

Whitcher IN, Wen J. 2001. Phylogeny and biogeography of

Corylus (Betulaceae): inferences from ITS sequences. Syst.

Bot. 26(2): 283–298.

Yoo K-O and J Wen. 2002. Phylogeny and biogeography of

Carpinus and subfamily Coryloideae (Betulaceae). Int. J.

Plant Sci. 163: 641–650.

Xing S-P, Z-D Chen, and A-M Lu. 1998. Development of ovule

and embryo sac in Ostrya virginiana (Betulaceae) and its

systematic signiﬁ cance. Acta Phytotax. Sinica 36: 428–435

(in Chinese with English summary).

Order 39. CASUARINALES

Trees and shrubs with slender, green, wiry, articulate

branchlets. Roots usually with nodules harboring nitro-

gen-ﬁ xing ﬁ lamentous soil bacteria of the genus

Frankia. Solitary or clustered crystals of calcium

oxalate usually present in some of the cells of parenchy-

matous tissues. Vessels with mainly simple or some-

times scalariform perforations with up to 30 bars; lateral

pitting opposite or alternate, very rarely intermediate.

Fibers with distinctly bordered pits. Rays heteroge-

neous. Axial parenchyma apotracheal, diffuse and in

short bands. Sieve-element plastids of S-type with up to

ﬁ ve starch grains; they also incorporate nondispersive

P-protein bodies. Nodes unilacunar with one trace.

Leaves small, scalelike, verticillate, more or less con-

nate to form a toothed sheath at each node; stipules

wanting. Stomata generally tetracytic, sometimes penta-

or hexacytic. Inﬂ orescences consist of alternating

whorls of toothlike, basally connate bracts; within each

bract two lateral, keeled, scalelike bracteoles are found.

Male inﬂ orescences from short to elongated catkinlike

spikes; female inﬂ orescences from small globular to

ovoid heads. Flowers small and inconspicuous, unisex-

ual, monoecious or rarely dioecious, apetalous, anemo-

philous. Male ﬂ owers consist of one or two

anteroposterior, hooded, scalelike sepals (deciduous at

anthesis) and a single (sometimes more or less bifurcate)

stamen. Anthers tetrasporangiate, basiﬁ xed, opening

longitudinally. Tapetum secretory. Microsporogenesis

simultaneous(?). Pollen grains 2-celled, 3-porate, tec-

tate-granular, with thick tectum traversed by minute

channels; exine ornamentation consists of small spinules

or scabrae and sometimes appears rugulate. Female

ﬂ owers without perianth; gynoecium pseudomonomer-

ous, consists of two median (anteroposterior) carpels

with very long, threadlike, reddish stylodia united at the

base. Stigma very long, decurrent; ovary superior, later-

ally ﬂ attened, initially 2-locular, but only the anterior

locule is fertile and contains two (rarely 3 or 4) ascend-

ing ovules; the posterior one is usually empty or more

or less reduced or obsolete. Ovules orthotropous to

hemitropous(?), epitropous, bitegmic, crassinucellate,

122 Subclass III. HAMAMELIDIDAE

micropyle endostomal. Female gametophytes several to

numerous, but only one fully developed and fertile,

of Polygonum-type. Fertilization chalazogamous.

Endosperm nuclear. Female inﬂ orescence develops into

a more or less woody infrutescence with the two enlarged

bracteoles of each ﬂ ower forming two tightly appressed

lateral valves, which separate when the fruit is mature.

Fruits 1-seeded, compressed, terminally winged,

samaroid nuts. Seeds small with seed coats adnate to the

pericarp, without endosperm, and often with more than

one embryo. Embryo large, straight, oily, with two large

cotyledons and a superior radicle. Plants tanniniferous,

producing proanthocyanins, ellagic acid, and ﬂ avonoid

monomers consisting of a number of glycosidic

variations based on a conservative aglycone component

of myricetin, quercetin, and kaempferol. n = 8

(Gymnostoma) – 14. There is a remarkable differentia-

tion in chromosome size (Morawetz and Samuel 1989).

Exhibits many features in common with

Hamamelidales and to a lesser degree also with Fagales

and Betulales (especially pollen morphology, which

resembles betulaceous pollen, Erdtman 1965;

Kuprianova 1965; Zavada and Dilcher 1986). Bessey

(1915) derived Casuarinaceae from Hamamelidaceae,

which is corroborated by the totality of morphological

data (Moseley 1948) including seed anatomy (Melikian

1973). The wood anatomy of the Casuarinaceae is

more advanced than that of the Hamamelidaceae, and

their ﬂ owers originated as a result of reduction and

simpliﬁ cation of the ﬂ owers of a hamamelidaceous

type. The similarities between the ﬂ owers of

Casuarinaceae and those of Betulaceae (Eames 1961;

Flores and Moseley 1982) are a result of the evolution-

ary parallelism. In both cases the lines of ﬂ oral reduc-

tion seen in the Hamamelidales have been carried to

extremes. All of the most important morphological fea-

tures of the Casuarinaceae are found in one or another

representative of Hamamelidales. But at the same time

Casuarinaceae diverged so much from the hamamel-

idalean stock, that “there is a considerable gap between

any known living or fossil Casuarinaceae and any other

known family” (Johnson and Wilson 1989: 185).

1. CASUARINACEAE

R. Brown 1814. 4/96. Australia, New Caledonia, Fiji,

New Guinea, Malesia; concentrated in Australia and to

a lesser extent in Malesia (especially the eastern

regions) and New Caledonia. One species (Casuarina

equisetifolia) has wide littoral distribution from the

Bay of Bengal to northeastern Australia and Polynesia

(but the occurrence of the species in the Mascarenes is

regarded as a probable introduction, Johnson and

Wilson 1989, 1993).

Gymnostoma, Ceuthostoma, Casuarina,

Allocasuarina.

Bibliography

Barlow BA. 1959. Chromosome numbers in the Casuarinaceae.

Austral. J. Bot. 7: 230–237.

Boodle LA and WC Worsdell. 1894. On the comparative anat-

omy of the Casuarinaceae, with special reference to the

Gnetaceae and Cupuliferae. Ann. Bot. 8: 231–264.

Dilcher DL, DC Christophel, HO Bhagwandin, Jr., and LJ

Scriven. 1990. Evolution of the Casuarinaceae: Morphological

comparisons of some extant species. Am. J. Bot. 77:

338–355.

Flores EM. 1980. Shoot vascular system and phyllotaxis of

Casuarina (Casuarinaceae). Am. J. Bot. 67: 131–140.

Flores EM and MF Moseley. 1982. The anatomy of the pistillate

inﬂ orescence and ﬂ ower of Casuarina verticillata Lamarck

(Casuarinaceae). Am. J. Bot. 69: 1673–1684.

Flores EM and MF Moseley 1990. Anatomy and aspects of

development of the staminate inﬂ orescences and ﬂ orets of

seven species of Allocasuarina (Casuarinaceae). Am. J. Bot.

77: 795–808.

Hwang R and JG Conran. 2000. Seedling characteristics in the

Casuarinaceae. Telopea 8: 429–439.

Johnson LAS and KL Wilson. 1989. Casuarinaceae: A synopsis.

In: PR Crane and S Blackmore, eds. Evolution, systematics,

and fossil history of the Hamamelidae, vol. 2, pp. 167–188.

Claredon Press, Oxford.

Johnson LAS and KL Wilson. 1993. Casuarinaceae. In:

K Kubitzki, ed. The families and genera of vascular plants,

vol. 2, pp. 237–242. Springer, Berlin/Heidelberg/New York.

Kershaw AP. 1970. Pollen morphological variation within the

Casuarinaceae. Pollen Spores 12: 145–161.

Moseley MF. 1948. Comparative anatomy and phylogeny of the

Casuarinaceae. Bot. Gaz. 110: 231–280.

Natarajan S, VVS Murty, and TR Seshadri. 1971. Chemo-

taxonomical studies of some Casuarina species. Phyto-

chemistry 10: 1083–1085.

National Research Council. 1984. Casuarinas: Nitrogenﬁ xing

trees for adverse sites. Academic Press, Washington, DC.

Okuda T, T Yoshida, T Hatano, K Yazaki, and M Ashida. 1982.

Ellagitannins of the Casuarinaceae, Stachyuraceae, and

Myrtaceae. Phytochemistry 21: 2871–2874.

Pant DD, DD Nautiyal, and S Singh. 1974. The cuticle, epider-

mis, and stomatal ontogeny of Casuarina equisetifrlia Forst.

Ann. Bot. 39: 1117–1123.

Redell P, GD Bowen, and AD Robson. 1986. Noduladon of

Casuarinaceae in relation to host species and soil properties.

Austral. J. Bot. 34: 435–444.

Sogo A, H Setoguchi, J Noguchi, T Jaffre, and H Tobe. 2001.

Molecular phylogeny of Casuarinaceae based on rbcL and

matK gene sequences. J. Plant Res. 114(116): 459–464.

Superorder JUGLANDANAE 123

Swamy BGL. 1948. A contribution to the life history of

Casuarina. Proc. Am. Acad. Arts and Sci. 77 (I): 132.

Torrey JG. 1983. Root development and root nodulation in

Casuarina. In: SJ Midgley, JW Turnbull, and RD Johnston,

eds. Casuarina ecology, management, and utilization,

pp. 180–192. CSIRO Melbourne.

Torrey JG and RH Berg. 1988. Some morphological features for

generic characterization among the Casuarinaceae. Am. J.

Bot. 75: 864–874.

Treub M. 1891. Sur les Casuarinees et leur place dans le systeme

naturel. Ann. Jard. Bot. Buitenz. 10: 145–231.

Ueno J. 1963. On the ﬁ ne structure of the pollen walls of

Angiospermae: III. Casuarina. Grana Palynol. 4:

189–193.

Superorder JUGLANDANAE

Order 40. MYRICALES

Evergreen or deciduous shrubs or small trees.

Trichomes when present are distinctive because of

their 2–3-celled bases and include regular balloon

glands, irregular bowl-shaped glands, and unicellular

trichomes. Roots usually bearing nodules with nitro-

gen ﬁ xing bacteria (except perhaps for Canacomyrica).

Vessels usually with scalariform perforations with a

range of 1–32 bars; Camptonia has simple perforation,

with a very small proportion of the plates bearing 1–3

bars. Fibers with bordered or both bordered and simple

pits. Rays heterogeneous. Axial parenchyma diffuse.

Sieve-element plastids of S-type. Nodes trilacunar or

unilacunar. Leaves alternate, simple, entire or serrate

to irregularly dentate or lobed, rarely (Comptonia)

deeply pinnatiﬁ d, pinnately veined, without stipules or

rarely (Comptonia) with foliaceous, semicordate

stipules, often with aromatic peltate resinous glands.

Stomata anomocytic. Flowers small, inconspicuous, in

axillary, dense, spikelike racemose inﬂ orescences, uni-

sexual (monoecious or more often dioecious), mostly

without perianth, usually subtended by a bract and two

transversal bracteoles, anemophilous. Male ﬂ owers

have 2–8 (very rarely up to 20) stamens, progressively

fewer in the more distal ﬂ owers. Filaments free or

sometimes connate at the base. Anthers ovoid,

tetrasporangiate, basiﬁ xed, extrorse, opening longitu-

dinally. Pistillodia often present. Tapetum secretory.

Microsporogenesis simultaneous. Pollen grains

2-celled, 3-porate, tectate-granular with stout columel-

lae often discernible in the infrastructural layer; the

tectum is traversed by minute channels, minutely

scabrate. Gynoecium of two united carpels, very rarely

of three carpels. Stylodia mostly long and slender,

nearly free or shortly connate at the base. Ovary

superior (Comptonia) or more or less inferior, 1-locu-

lar, with one basal erect ovule born in the anterior car-

pel. Ovules orthotropous, unitegmic (in Canacomyrica

with elongate, reﬂ exed micropylar tube), crassinucel-

late. Female gametophyte of Polygonum-type.

Endosperm nuclear. Fruits drupes with a hard endo-

carp and wax-secreting exocarp or almost a nut, some-

times enveloped by the persistent, accrescent bracteoles,

in Canacomyrica completely enclosed in an enlarged,

6-lobed perianth. Seeds with very reduced, exotestal

seed coat, straight, large embryo, and nearly or quite

without endosperm. Tanniniferous and accumulating

triterpenes and sesquiterpenes, also with protoantho-

cyanins, ellagic acid, and gallic acid. n = 8, 12.

Although Myricales have some features in common

with the Betulaceae and Casuarinaceae, they are much

nearer to the Juglandales (Chevalier 1901; Kershaw

1909; Stokes 1937; Hjelmquist 1948; Takhtajan 1966,

1987; Dahlgren 1980, 1983; Cronquist 1981; Thorne

1989; Macdonald 1989; Kubitzki 1933), which is sup-

ported by the fossil record (Friis 1983). According to

Zavada and Dilcher (1986), Myrica exhibits particular

similarity to Rhoiptelea and Platycarya, but Canaco-

myrica is probably the closest to the Rhoipteleaceae

(see Friis 1983; Macdonald 1989).

1. MYRICACEAE

A. Richard ex Kunth 1817 (including Canaco myricaceae

Doweld 2000). 4/57. Widely distributed in America,

Africa, and Asia, but absent in Australia and New

Zealand. Comptonia (1) is endemic to eastern North

America, and Canacomyrica (1) to New Caledonia.

1.1 CANACOMYRICOIDEAE

Flowers with 6-lobed perianth. Male ﬂ owers with six

stamens. Female ﬂ owers with (5)6 staminodia. Ovules

with micropylar tube. – Canacomyrica.

1.2 MYRICOIDEAE

Flowers without perianth. Male ﬂ owers with 2–8 (very

rarely up to 20) stamens. Female ﬂ owers without sta-

minodia. Ovules without micropylar tube. – Myrica

(including Gale?), Morella, Comptonia.

124 Subclass III. HAMAMELIDIDAE

Bibliography

Abbe EC. 1972. The inﬂ orescence and ﬂ ower in male Myrica

esculenta var. farquhariana. Bot. Gaz. 133: 206–213.

Abbe LB and EC Abbe. 1971. The vessel member of Myrica

esculenta Buch. Ham. J. Minnesota Acad. Sci. 37: 72–76.

Carlquist S. 2002. Wood and bark anatomy of Myricaceae: rela-

tionships, generic deﬁ nitions, and ecological interpretations.

Aliso 21: 7–29.

Chen Z-D and A-M Lu. 1998. Systematic position of the

Rhoipteleaceae: evidence from nucleotide sequences of rbcL

gene. Acta Phytotax. Sinica 36: 1–7.

Chevalier A. 1901. Monographic des Myricacees: Anatomic et

histologie, organographie, classiﬁ cation, et description des

especes, distribution geographique. Mem. Soc. Sci. Nat.

Cherbourg 32: 85–340.

Collins RP and AF Halim. 1973. Chemotaxonomy of the

Myricaceae; II. Essential oil analysis of three Central

American species of Myrica. Lloydia 36: 320–325.

Davey AJ and CM Gibson 1917. Note on the distribution of

sexes in Myrica gale. New Phytol. 16: 147–151.

Diboll AG. 1959. Comparative anatomy of staminate catkins of

Myricaceae. M.A. thesis, Claremont Grad. School, Claremont.

Elias TS. 1971. The genera of Myricaceae in southeastern United

States. J. Arnold Arbor. 52: 305–318.

Ferguson DK. 1998. The contribution of micromorphology to

the taxonomy and fossil record of the Myricaceae. Taxon 47:

333–335.

Fletcher WW. 1955. The development and structure of the root-

nodules of Myrica gale with special reference to the nature

of the endophyte. Ann. Bot. 19: 501–573.

Friis EM. 1983. Upper Cretaceous (Senonian) ﬂ oral structures

of juglandalean afﬁ nity containing Normapolles pollen. Rev.

Palaeobot. Palynol. 39: 161–188.

Gladkova VN. 1962. Fragments of the history of the Myricaceae

family. Pollen Spores 4: 345.

Hakansson A and RP Collins. 1955. Endosperm formation in

Myrica gale L. Bot. Not. 108: 6.

Halim AF and RP Collins. 1973. Essential oil analysis of the

Myricaceae of the eastern United States. Phytochemistry 12:

1077–1083.

Kershaw EM. 1909. The structure and development of the ovule

of Myrica gale. Ann. Bot. 23: 353–362.

Killick DJB. 1969. The South African species of Myrica.

Bothalia 10: 517.

Kubitzki K. 1993. Myricaceae. In: K Kubitzki, ed. The families

and genera of vascular plants, vol. 2, pp. 453–457. Springer,

Berlin/Heidelberg/New York.

Leroy JF. 1949. De la morphologic ﬂ orale et de la classiﬁ cation

des Myricaceae. C. R. Acad. Sci. Paris 229: 1162–1163.

Leroy JF. 1957. Sur deux Amentiferes remarquables de la Flore

Asiatico Paciﬁ que et Paciﬁ que. Proc. 8th Paciﬁ c Science

Congress 4: 459–464.

Macdonald AD. 1974. Floral development of Comptonia pere-

grina (Myricaceae). Canad. J. Bot. 52: 2165–2169.

Macdonald AD. 1977. Myricaceae: Floral hypothesis for Gale

and Comptonia. Canad. J. Bot. 55: 2636–2651.

Macdonald AD. 1978. Organogenesis of the male inﬂ orescences

and ﬂ owers of Myrica esculenta. Canad. J. Bot. 56:

2415–2423.

Macdonald AD. 1979. Development of the female ﬂ ower and

gynecandrous partial inﬂ orescence of Myrica californica.

Canad. J. Bot. 57: 141–151.

Macdonald AD. 1980. Organogenesis of the female reproduc-

tive structure of Myrica pensylvanica. Canad. J. Bot. 58:

2001–2006.

Macdonald AD. 1989. The morphology and relationships of the

Myricaceae. In: PR Crane and S Blackmore, eds., Evolution,

systematics, and fossil history of the Hamamelidae, vol. 2,

pp. 147–165. Claredon Press, Oxford.

Macdonald AD and R Sattler. 1973. Floral development of

Myrica gale L. and the controversy over ﬂ oral concepts.

Canad. J. Bot. 51: 1965–1975.

Stokes J. 1937. Cytological studies in the Myricaceae. Bot. Gaz.

99: 387–399.

Sundberg MD. 1985. Pollen of the Myricaceae. Pollen Spores

27: 15–28.

Vikhireva VV. 1957. Anatomical structure and development of

the female ﬂ ower of the common bog myrtle Myrica gale

L. Trudy Komarov Bot. Inst. Acad. Sci. USSR, ser. 7, Morf.

Anat. Rast. 4: 270–287 (in Russian).

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Comparative seed anatomy, vol. 3, pp. 141–148. Nauka,

Leningrad (in Russian).

Yen TK. 1950. Structure and development of the ﬂ ower and the

fruit of Myrica rubra. Peking Natural Hist. Bull. 19: 1–20.

Order 41. JUGLANDALES

Deciduous or evergreen, aromatic trees or rarely large

shrubs with naked or scaly buds. Vessel elements

elongate, with scalariform or more often simple perfo-

rations, but in Alfaroa and Engelhardtia some of them

scalariform with a few bars; lateral pitting alternate.

Fibers with bordered pits; some gelatinous ﬁ bers

usually also present. Rays heterogeneous or sometimes

(in some Juglandaceae) almost homogeneous. Axial

parenchyma apotracheal to paratracheal. Sieve-element

plastids of S-type. Nodes trilacunar or sometimes

pentalacunar with ﬁ ve traces. Leaves alternate or less

often opposite, stipulate or estipulate, pinnately com-

pound (or trifoliolate), with leaﬂ ets entire or serrate.

Leaves provided with aromatic, resinous, several-celled,

peltate, basally sunken glandular scales; hairs mainly

nonglandular. Stomata anomocytic. Flowers small,

borne in aments, apetalous, unisexual (monoecious or

rarely dioecious) or gynomonoecious (the central ﬂ ower

of dichasium in Rhoiptelea), borne in elongate, droop-

ing, and rarely erect, catkins or spikelike inﬂ orescences,

that are sometimes grouped into panicles. Each ﬂ ower

of Rhoiptelea has two lateral bracteoles and four small,

scarious, persistent on the fruit sepals in two cycles. In

Juglandaceae ﬂ owers more or less reduced. The less

Superorder JUGLANDANAE 125

reduced ﬂ owers (e.g., Juglans) consist of two lateral

bracteoles and four sepals, but mostly there are only

three, two, or even one sepals. Platycarya lost both the

sepals and bracteoles and thus reached the end of the

reduction line. The bract and two bracteoles (when

present) are fused with the ﬂ oral receptacle and appear

as a part of the calyx, the stamens thus appearing to be

on the involucre and calyx in male ﬂ owers; modiﬁ ca-

tions in female ﬂ owers led to the reduction of the calyx

and the adnation of the bract and bracteoles with the

entire ovary to form a cupular involucre that ripens with

the fruit to form a husk. Stamens 3–105 (mostly 5–40)

in Juglandaceae, in Rhoiptelea – 6; ﬁ laments short,

anthers tetrasporangiate, basiﬁ xed, opening longitudi-

nally. Tapetum secretory. Microsporogenesis simulta-

neous, rarely successive. Pollen grains 2-celled or

3-celled, 3-porate or pantoporate, 3-colporate or sel-

dom 4-colporate, tectate-granular. Gynoecium regu-

larly of two united carpels, rarely individual ﬂ owers

with three or four carpels; stylodia free or united at the

base, seldom stigmas sessile. Ovary superior or infe-

rior, incompletely 2-locular (2-locular below and 1-loc-

ular above), in Juglandaceae often falsely 4- to 8-locular

below because of secondary partitions (sometimes

5-locular at the very apex due to intrusions from the

wall). Ovules solitary, attached to the partial partition,

campylotropous, or orthotropous, bitegmic or uniteg-

mic, crassinucellate. Female gametophyte of

Polygonum-type. Fertilization chalazogamous, very

rarely porogamous. Endosperm nuclear. Fruits are nuts

(often samaroid), or drupe-like but rather dry. Seeds

large, obovoid (Rhoiptelea) or more often 2- to 4- to

8-lobed. Seed coat reduced to thin exotestal layer

(Rhoiptelea) or outer epidermis as a layer of rather

large, thin-walled cells, highly vascularized. Embryo

massive, with 2- (Rhoiptelea) or 4-lobed oily cotyle-

dons; endosperm wanting or very scanty.

Has much in common with Myricales (including

serolological similarities between Juglandaceae and

Myricaceae – Chupov 1978; Petersen and Fairbrothers

1979). Especially many common features with Myrica-

ceae in the structure of ﬂ ower, pollen grains, ovules, fruit,

basic chromosome number and chemistry. Both orders

derived from the hamamelidalean type ancestor.

Key to Families

1 Leaves with leafy, thick, papery and asymmetri-

cally caducous stipules. Ovary superior, 2-locular;

stigmas two, lamelliform; ovules one per locule,

campylotropous, bitegmic, but only one ovule

develops. Flowers borne in elongate spikes con-

sisting of 3- ﬂ owered dichasial glomerules that are

clustered into large, nodding, terminal panicles

look like a horse tail (the species is named the

“horse-tail tree” – Wu and Kubitzki 1993). Each

dichasium is suspended by a large bract. The lat-

eral ﬂ owers of each dichasium are female, but

more or less reduced and frequently abortive, each

ﬂ ower with two bracteoles. Sepals four, in two

cycles, small, scarious, brown, persistent on the

fruit. Stamens six. Pollen grains 3-colporate or

seldom 4-colporate with very short colpi appear-

ing to approximate the porate condition, scabrate.

The female ﬂ owers has no staminodes. Fruits

small, 2-winged nuts. Vessel elements elongate,

with scalariform perforations that have 1–2 (up to

20 bars). Axial parenchyma abundant, mostly

vasicentric. Produce protoanthocyanins, ellagic

acid, and ﬂ avonols (myricetin, quercetin and kae-

mpferol), n = 16. . . . . . . . . . .1. rhoipteleaceae

1 Leaves estipulate. Ovary inferior; ovules orthotro-

pous, unitegmic. Flowers unisexual, borne in

elongate, drooping, and rarely erect, catkins or

spikelike inﬂ orescences, that are sometimes grouped

into panicles. Male and female inﬂ orescences sepa-

rate or combined into an androgynous panicle; indi-

vidual ﬂ owers in the axils of entire or 3-lobed

bracts. The calyx adnate to the bract or wanting.

Stamens 3–105, mostly 5–40. Pollen grains 3-porate

or pantoporate, spinulose. Fruits are nuts enclosed

in an adherent, thin or thick, ﬁ brous indehiscent or

dehiscent husk, or nuts with a thin dry skin, or thin-

skinned nutlets with wings, which are formed by

the bract or by two bracteoles. The whole fruit is

drupelike, but the husk is derived from the involu-

cre and calyx, and the skin from the calyx alone,

neither one from the pericarp. Hence the fruit is not

a true drupe; it is sometimes called a tryma (Manning

1978); nut loculicidal in germination. Vessel ele-

ments elongate, mostly with simple perforations,

but in Alfaroa and Engelhardtia some of them sca-

lariform with a few bars. Axial parenchyma mostly

apotracheal, but sometimes intermediate between

apotracheal and paratracheal. Mostly with abundant

tannin and producing proanthocyanins, ellagic acid,

ﬂ avonol glycosides, including those of myrcetin,

quercetin, kaempferol, and napthoquinones, n = 15,

16, 28. . . . . . . . . . . . . . . . . . . . . 2. juglandaceae

126 Subclass III. HAMAMELIDIDAE

1. RHOIPTELEACEAE

Handel-Mazzetti 1932. 1/1. Southwestern China

(Yunnan, Guangxi, Guizhou) and North Vietnam.

Rhoiptelea.

In many respects this unigeneric family approaches

closely to the hypothetical intermediate between

Hamamelidales and Juglandaceae.

2. JUGLANDACEAE

A.P. de Candolle ex Perleb 1818 (including

Engelhardtiaceae Reveal et Doweld 1999, Platycary-

aceae Nakai ex Doweld 2000, Pterocaryaceae Nakai

1930). 9/60. Mainly northern temperate and subtropi-

cal regions with extensions southward to South and

Southeast Asia, New Guinea, West Indies, and Central

America and the western mountains of South America

as far as northern Argentina.

Juglandaceae are closely related to the Rhoipte-

leaceae, but are somewhat more advanced. Each ﬂ ower

of the reduced inﬂ orescence of Juglandaceae morpho-

logically corresponds to the central ﬂ ower of the

3-ﬂ owered dichasium of Rhoiptelea.

2.1 PLATYCARIOIDEAE

Bracts of female ﬂ owers persisting on the axis of inﬂ o-

rescence (not dispersed with the fruit). Pollen grains

small, 3-porate, with pseudocolpi. Vessels with simple

perforations. – Platycarya.

2.2 ENGELHARDTIOIDEAE

Bracts of the female ﬂ owers fall together with the fruit,

3-lobed. Pollen grains small, 3- to 4-porate. Vessels

with scalariform or simple perforations. Leaf paren-

chyma with druses. – Alfaroa, Alfaropsis, Oreomunnea,

Engelhardtia.

2.3 JUGLANDOIDEAE

Bracts of the female ﬂ owers fall together with the fruit,

entire. Pollen grains large, 3-polyporate. Vessels with

simple perforations. – juglandeae: Cyclocarya,

Pterocarya, Juglans; hicorieae: Carya.

Manning (1978) recognized two subfamilies

(Platycaryoideae and Juglandoideae), the former with

one monotypic east Asian genus (Platycarya) and the

latter with three tribes (Engelhardieae, Juglandeae, and

Hicorieae). Recently Iljinskaya (1990) reviewed previ-

ous classiﬁ cations and came to the conclusion that

Manning’s tribe Engelhardieae is so distinctive that it

deserves the rank of a subfamily.

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A. Takhtajan, Flowering Plants, 129

Mostly herbaceous plants, undershrubs, or low shrubs,

less often trees; succulents and halophytes common,

woody species generally with anomalous secondary

growth or otherwise anomalous stem structure. Vessels

with simple perforations. Sieve-element plastids of

P-type (Caryophyllales, Physenales) or of S-type

(Polygonales, and Plumbaginales). Nodes unilacunar

or less often trilacunar or multilacunar. Leaves simple,

mostly entire, sometimes with stipules. Stomata mostly

paracytic, anomocytic, or diacytic. Flowers bisexual or

unisexual, cyclic. Tapetum secretory. Microsporogenesis

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

3-colpate or of derived types. Gynoecium apocarpous

(certain Phytolaccaceae and Gisekia) or more often

syncarpous, frequently pseudomonomerous. Ovules

mostly bitegmic or rarely (in some Nyctaginaceae)

unitegmic, crassinucellate. Endosperm nuclear. Fruits

of various types. Seeds with mostly curved embryo

and usually without endosperm (except in Polygonaceae

and Plumbaginaceae), but mostly with perisperm.

Related to the Ranunculidae, especially to the lower

Ranunculales. The Phytolaccaceae in particular are

linked with the ranunculids, especially with the

Lardizabalaceae and Menispermaceac. In all probabil-

ity both subclasses share a common origin from some

Magnoliidae.

The rbcL sequences conﬁ rm the monophyly of the

Caryophyllidae. According to Olmstead et al. (1992),

the Caryophyllidae, including Polygonaceae and

Plumbaginaceae, appear as a strongly supported,

monophyletic group, whose circumscription in tradi-

tional classiﬁ cations coincides completely with the

molecular evidence.

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Subclass IV. CARYOPHYLLIDAE

130 Subclass IV. CARYOPHYLLIDAE

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Superorder CARYOPHYLLANAE

Order 42. CARYOPHYLLALES

Perennial or annual herbs, shrubs or trees, rarely climb-

ing, often more or less succulent. Anomalous second-

ary thickening of the stems fairly frequent. Vessels

with simple perforations, very rarely also reticulate.

Fibers with simple or seldom bordered pits. Axial

parenchyma diffuse (Stegnospermataceae, Barbeui-

aceae) or vasicentric (Phytolaccaceae and Achato-

carpaceae). Sieve-element plastids of P-type with a

peripheral ring of protein ﬁ laments and with or with-

out (Amaranthaceae and Chenopodiaceae) a central

protein crystal (Behnke 1994). Nodes unilacunar or

rarely trilacunar, very rarely (Rhabdodendraceae)

multi lacunar. Leaves alternate or opposite, seldom ver-

ticillate, simple and mostly entire, sometimes reduced

to spines, often showing Kranz anatomy, estipulate or

rarely with small stipules. Stomata of various types.

Flowers in various types of inﬂ orescences or solitary

in the axils, bisexual or less often unisexual, usually

actinomorphic, mostly 4–5-merous, entomophilous or

less often anemophilous. Sepals (1)2–5(-10), free or

more or less connate below, sometimes petaloid. Petals

two to many or lacking, free or more or less connate.

Stamens one to many, often as many or twice as many

as the sepals or petals, in multistaminate ﬂ owers often

developing centrifugally, especially in Aizoaceae and

Cactaceae; anthers usually opening longitudinally.

Pollen grains 3-celled or rarely 2-celled, 3-colpate to

pantocolpate and pantoporate, tectate-columellate,

with spinulose-annular perforate/punctate tectum.

Tapetum secretory. Microsporogenesis simultaneous.

Gynoecium syncarpous (eusyncarpous, paracarpous,

or lysicarpous) or less often more or less apocarpous

(but with short, basal, distinctly fused carpellary sec-

tions (see Rohweder 1965; Hofmann 1977; Hofmann

in Behnke and Mabry 1994), of two to many carpels,

with distinct stylodia. Ovary superior, half inferior, or

inferior. Nectaries present or absent. Ovules hemitrop-

ous to campylotropous, rarely nearly anatropous,

bitegmic, crassinucellate. Female gametophyte usually

of Polygonum-type. Endosperm nuclear. Fruits of vari-

ous types. Seeds with peripheral and mostly curved or

coiled embryo, bordering or encircling the more or less

copious perisperm or (Didiereaceae) nearly or quite

without perisperm. Perisperm completely absent in

Monococcus of Phytolaccaceae); endosperm lacking

or scanty. Seed-coat exotestal-endotegmic or exotestal.

Mostly producing betalains (betacyanins and betaxan-

thins) but not anthocyanins (only in the Molluginaceae

and Caryophyllaceae producing anthocyanins and not

betalains). At least some species of Phytolacca contain

glucosilonates (Daxenbichler et al. 1991).

Key to Families

1 Stylodium gynobasic. Evergreen tall shrubs or small

trees. Secretory cavities with resinous contents scat-

tered through parenchymatous tissues. Vessels with

simple perforations; lateral pitting alternate. Fibers

with many small simple or bordered pits. Rays het-

erogeneous to sometimes homogeneous. Axial

parenchyma paratracheal and very scanty diffuse or

largely in apotracheal bands. Sieve-element plastids

of Pcs-type. Leaves alternate, simple, entire, leath-

ery, with short-stalked, peltate hair, obscurely pellu-

cid-punctate; stipules minute and caducous or

obsolete, or absent. Stomata anomocytic. Flowers in

axillary or supra-axillary racemes or racemoid

cymes, with thick rachis, usually bisexual, actino-

morphic. Calyx very short, turbinate, more or less

entire or with ﬁ ve imbricate lobes. Petals ﬁ ve, oblong,

sepal-like, pellucid-punctate, slightly imbricate

below or more or less valvate at apex, caducous.

Stamens 25–50 more or less in two cycles; ﬁ laments

very short, ﬂ attened, persistent and recurved; anthers

very long, basiﬁ xed, caducous. Pollen grains 3-celled,

3-colpate and rarely 4-colpate (Prance 2003). Nectary

Superorder CARYOPHYLLANAE 131

disc wanting. Gynoecium of single carpel with long,

thickened gynobasic stylodium with unilateral (dor-

sal) decurrent stigma. Ovule usually solitary (some-

times another aborted), basal, hemicampylotropous

and epitropous, bitegmic (bitegmic condition

restricted to the micropylar part). Fruits small, drupa-

ceous, with thin exocarp and slightly woody endo-

carp, shortly stipitate, within the cupular calyx. Seed

with curved, green embryo and large, ﬂ eshy cotyle-

dons, small radicle bent inward toward the hilum;

endosperm wanting; exotestal cells tangentially elon-

gated. Presence triterpenoids and O-alkylated ellagic

acid (Walter-Filho et al. 1985, 1989); alkaloids

absent, n = 10.. . . . . . . . . . . 2. rhabdodendraceae

1 Stylodium not gynobasic.

2 Sieve-element plastids with a central protein

crystal.

3 Producing betalains, very rarely anthocyanins.

4 Carpels free or united. Pollen grains 3-colpate.

Sepals ﬁ ve, imbricate. Petals absent or present.

5 Fruits of more or less drupelike fruitlets or syn-

carpous and baccate, or fruits indehiscent, by

abortion 1-seeded, with coriaceous pericarp

adherent to the seed (Agdestis). Herbs, shrubs,

woody lianas, or rarely trees, more or less suc-

culent, or non-succulent, mesophytic or xero-

phytic. Raphides present. Sieve-element

plastids of P3a-type. Nodes unilacunar. Leaves

alternate, entire, usually estipulate. Flowers in

racemes or spikes, bisexual or rarely dioecious,

actinomorphic or more or less zygomorphic,

apetalous. Perianth segments 4–5, in one whorl.

Stamens (2-)4 to many; ﬁ laments ﬁ liform, free

or basally connate, sometimes inserted on a

ﬂ eshy disc; anthers dorsiﬁ xed, introrse, tet-

rasporangiate. Pollen grains 3-colpate via

panto colpate to pantoporate, 2- or 3-celled.

Carpels 3–16, free or more or less connate;

ovary superior, rarely semi-inferior (Agdestis),

4–12-locular, ovules solitary in each locule,

campylotropous or hemitropous (Agdestis).

Seeds more or less reniform, perisperm copious

or moderate, soft or granular and whitish;

embryo well differentiated, curved; cotyledons

ﬂ at or folded; endosperm absent. Betalains

present; saponins, alkaloids and ﬂ avonols pres-

ent or absent; Gallesia smells of onions; n = 9,

rarely 17. . . . . . . . . . . . . .1. phytolaccaceae

5 Fruits achenes with membranous epicarp

densely covered by warts. Annual herbs.

Leaves opposite or subverticillate. Flowers in

dichasia, bisexual or rarely unisexual, more or

less actinomorphic. Petales ﬁ ve. Stamens 5 or

10–15, rarely 8, sometimes 20 or more, and

then they are paired in groups of three, inserted

on a ﬂ at disc, alternating with petals; ﬁ laments

dilated at the base; anthers basiﬁ xed. Carpels

ﬁ ve or less often three or ten, pseudoapocar-

poous, laterally compressed, with short stylo-

dia on their ventral side; ovule solitary in each

locule, anatropous. Fruit achenial; seeds

smooth, pitted. Present anthocyanins and

raphides; n = 9.. . . . . . . . . . . . 3. gisekiaceae

4 Carpels always united. Pollen grains of various

types. Petals present or absent.

6 Ovary mostly superior.

7 Fruits winged. Shrubs with thorny branches.

Wood rayless. Leaves opposite below, alter-

nate above, deciduous, linear, entire, ﬂ eshy,

glabrous to hairy. Inﬂ o rescence spike, cylin-

dric, densely catkin-like. Flowers unisexual.

Male ﬂ owers numerous, spirally arranged,

with peltate bracts, without perianth. Stamens

2–3 or more, with very short ﬁ laments and

long anthers; pollen grains pantoporate, dis-

tinctive by its raised pore margins produced

by elongated columellae (crassimarginate

pores – Nowicke 1994). Female ﬂ owers 1–3,

below male ﬂ owers, perianth cup-like, con-

nate with ovary; ovule solitary. Carpels two,

stigmas two, horizontally expanded. Seed

erect with transparent testa; embryo com-

pressed, green, n = 9. . . . 4. sarcobataceae.

7 Fruits not winged.

8 Petioles articulated at the base. Large, branched

woody lianas, blackening on drying. Axial

parenchyma diffuse. Leaves alternate, entire.

Protein crystal in the sieve-element plastids

globular. Flowers in short, rigid, axillary

racemes. Pedicels long and slender, thickened

below the calyx. Receptacle slightly convex.

Sepals ﬁ ve imbricate. Stamens 20–25, in sev-

eral series, inserted on an annular disc (annu-

lus of receptacle); ﬁ laments short, free; anthers

linear, 2-locular, introrse, sagittate. Pollen

grains 3(4–6)-colporoidate, the exine undu-

lated to verrucate. Gynoecium of two united

carpels; ovary 2-locular; stylodia linear-oblong,