SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 58 

Classification, Evolution, 

and Phylogeny of the 

Families of Dicotyledons 

Aaron Goldberg 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1986 
ABSTRACT 

Goldberg, Aaron. Classification, Evolution, and Phylogeny of the Families of Dicotyledons. 

Smithsonzan Contributions to Botany, number 58, 314 pages, 164 figures, 2 tables, 1986. 

To some extent classification is subjective. Taxonomists differ in the relative importance 

they ascribe to particular characters and in the de ree of difference between related taxa 

they deem sufficient to constitute family or ordin3 rank. About 1000 dicot family names have been ublished Those who have attempted an overview of the s stem at the family 

level and agove in the last quarter century reco nize between 274 anJ455 dicot families 

in 39 to 82 orders. I accept 334 families and orders. In Table 1 I give my ordinal 

allocation of the families and that of 1 1 recent authors to indicate where there is agreement 

and where there are differences to be resolved. I have also constructed a dendrogram to su gest relationships and degree of advancement of the orders. f have written concise, uniform descriptions of all the families of dicots, emphasizing 

those characters that show trends between families or occur in more than one famil . Each family is illustrated by analytical drawings of the flower, fruit, seed and usual& 

inflorescence. Several species are usually used to show the range of major variation within 

families and trends toward related families. 

Angiosperms probably arose from gymnosperms, so characters or character states 

universal in gymnosperms or considered primitive in them would also be considered 

primitive in angiosperms. 

My a proach to understanding evolutionary trends in characters is to relate them to 

the ecokgical factors that might be responsible for them by their selective action. The 

dicots probably originated under warm tern erate conditions favorable for growth. A 

character states enabling them to cope with dry and hot or colaconditions and colonize 

generally unfavorable re ions. 

A second major trend kas been from wind pollination to progressive1 better adaptation 

for insect pollination. The primitive insect pollinated dicots often xave flowers with numerous spirally arranged parts; plants having flowers with few, opposite or whorled 

major evolutionary trend in them has been t K e gradual develo ment of characters and 

parts are derived. 

seDarate. The connate and adnate conditions are derived. 

The floral organs are homologous with leaves. Like leaves the parts were initially 

’General character states are primitive; specialized states are derived. In attempting to 

determine which primitive states are most primitive I considered their occurrence among 

the families. The fewer the families with a particular primative state, the more primitive 

the state. This is im ortant in deciding whether a family is low or high on the family tree 

and the position wittin its particular order. 

In accordance with the above rationale, I have constructed a table givin the primitive 

and derived states for about 100 characters. I also indicate the extent to wkch I consider 

the states reversible. 

To determine a family’s ph lo eny, it must be compared with other families considered 

to be close to it. In genera!, t%e more characters and character states in common, 

particularly uncommon ones, the more like1 are the subject families to be related. All 

parts of the plant and many characters shouyd be considered. If a famil has more than one state of a character, the state considered primitive for the particuyar taxon rather 

than the derived one should be considered in attempting to determine the extant family 

closest to its ancestor. A descendant has at least one more derived character or character 

state than its ancestor. 

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is 

recorded in the Institution’s annual report, Smithsonian Year. SERIES COVER DESIGN: Leaf 

clearing from the katsura tree Cercidiphyllum japonicum Siebold & Zuccarini. 

Library of Congress Cataloging in Publication Data 

Goldberg, Aaron. 

Classification, evolution, and phylogeny of the families of Dicotyledons. 

(Smithsonian contributions to botony ; no. 58) 

Bibliography: p. 

Includes index. 

Supt. of Docs. no.: SI 1.29:58 

1. Dicotyledons-Classification. 2. Dicotyledons-Evolution. 3. Phylogeny (Botany) I. TiQKI.S2747 no. 58 581 s [583’.0438] 85-600058 [QK495.A12] 

tle. 11. Series. 
Contents 

Introduction ............................................ 

Table 1 .-Ordinal allocation of the dicotyledon families by recent 

authors ............................................. 

Table 2.-Evolutionary trends in dicotyledons ................ Dendrogram of suggested phylogenetic relationships of the orders 

of dicotyledons ....................................... 

IncertaeSedis ........................................... References ............................................. Index to Orders and Families ............................... 

Ordinal and Family Descriptions ............................ 

Page 

1 

4 

24 

27 

29 

304 

309 

312 

... 

111 

Classification, Evolution, 

and Phylogeny of the 

Families of Dicotyledons 

Aaron Goldberg 

Introduction 

To some extent classification is subjective. 

Taxonomists differ in the relative importance 

they ascribe to particular characters and in the 

degree of difference between related taxa they 

deem sufficient to constitute family or ordinal 

rank. About 1000 dicot family names have been 

published. Those who have attempted an overview of the system at the family level and above 

in the last quarter century recognize between 

274 and 455 dicot families in 39 to 82 orders. I 

accept 334 families and 59 orders. About 76% 

of the dicot families and 76% of the orders I 

accept were recognized before 1875, the publication date of Eichler’s system, the first in which 

the concept of evolution was accepted. Of course, 

the circumscription of some families and orders 

has been changed since 1875. About 1.5% of the 

dicot families and orders were published in the 

last quarter century. 

In delimiting families and assigning them to 

orders I rely on many years of study in the 

National Herbarium and library of the Smithsonian Institution as well as examination of plants 

in the field and at botanical gardens in various 

parts of the world. I have examined numerous 

Aaron Goldberg, collaborator, Department of Botany, National 

Museum of A’atural History, Smithsonian Institution, Washington, D.C. 20560. 

1 

specimens in all families except three monotypic 

ones. 

I have written concise, uniform descriptions of 

all the families of dicots, emphasizing those characters that show trends between families or occur 

in more than one family. Each family is illustrated 

by analytical drawings of the flower, fruit, seed, 

and usually inflorescence. The illustrations have 

been reproduced as economically as possible for 

the purpose of illustrating major features; no 

attempt has been made to represent the fine 

detail that would be expected in traditional taxonomic studies published in Smithsonian Contributions to Botany. Abbreviations used in the legends are as follows: I.s. = longitudinal section; 

C.S. = cross section. 

I deeply appreciate the permission granted by 

the Oxford University Press to use many illustrations from their publication (J. Hutchinson, 

1973), and also permission of colleagues to use 

illustrations from their publications, and also 

their help in other ways. 

At present the only work in English describing 

and illustrating all the dicot families in a single 

volume is Hutchinson’s The Families of Flowering 

Plants (1 973). I believe Hutchinson’s theory of 

relationships places undue emphasis on whether 

the families are primarily woody or herbaceous. 

Also, his concept of families is narrower than 

mine. Usually he illustrates each family by a 
2 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

single species whereas I usually use several species to show the range of major variation within 

families, and sometimes the species illustrated 

show trends toward related families. He illustrates sections of seeds showing the embryo for 

half the families whereas I do so for most families. 

He illustrates leaves more frequently than I do. 

Perusal of the references dealing with chemotaxonomy indicates how little is known for many 

of the families, so I did not consider chemical 

data under the family descriptions. However, I 

have discussed under each order those chemical 

data I consider most significant in understanding 

relationships of the families, orders, and sometimes genera. I have not included data on chromosome numbers because, although they are 

significant in indicating relationships within families, it has not been determined whether this 

character is appreciably useful to show relationships between families. 

In Table 1 I give my ordinal allocation of the 

families and that of 11 authors who have attempted an overview of the system in the last 

quarter century to indicate where there is agreement and where there are differences to be resolved. In my decisions I have also considered 

the opinions of 17 earlier generalists, starting 

with Bentham and Hooker, as well as those of 

authors who have studied particular families. 

The bibliography mentions publications of several authors who have attempted to improve the 

system of classification of the dicots at the family 

level and above in the 19th and 20th centuries. 

It also mentions authors who have assembled the 

data dealing with particular characters for most 

families. Together, these publications contain 

references to the work of thousands of scientists 

who have made contributions in particular families. Stafleu and Cowan and the Kew Record of 

Taxonomic Literature can also be consulted for 

relevant literature. In the figure legends, all authors cited without publication dates are to be 

found in Stapf (1 929- 193 1). 

Jackson (1928) and Lawrence (1951) can be 

used to determine the meaning of botanical 

terms. I use the term pistil to mean a separate 

female organ. It can consist of one or more 

carpels. The gynoecium may consist of one or 

more pistils. 

Angiosperms probably arose from gymnosperms, so characters or character states occurring universally in gymnosperms or considered 

primitive in them would also be considered primitive in angiosperms. Gymnosperms are dioecious 

or monoecious. Xylem vessels are almost universally lacking. The reproductive organs are not 

subtended by a perianth. When the pollen grains 

are aperturate, they are monosulcate. 

Since gymnosperms are almost universally 

wind-pollinated, the earliest angiosperms must 

have been wind-pollinated, and since gymnosperms with this characteristic have survived, I 

see no reason why primitive wind-pollinated angiosperms could not have survived. Engler and 

his followers begin the system with the Casuarinaceae and amentiferae. However, there are 

wind-pollinated dicots that have more primitive 

character states than those groups. For example, 

Tetracentron (Tetracentraceae) and Sarcandra 

(Chloranthaceae) have vesselless wood, whereas 

Casuarina and the amentiferae have vessels. The 

carpels are connate in Casuarina and the amentiferae; they are separate in Cercidiphyllum (Cercidiphyllaceae). The amentiferae frequently have 

an inferior ovary, whereas it is superior in the 

more primitive, wind-pollinated families. The 

pollen grains are sometimes monosulcate in 

Chloranthaceae, but never in Casuarina and the 

amentiferae. In Casuarina and the amentiferae 

storage food of the seed is restricted to the embryo, whereas in the gymnosperms and more 

primitive, wind-pollinated dicots much of it is 

outside the embryo. Also, these systems give 

greater weight to the sympetalous state than I 

do, dividing the dicots into subclasses Archichlamydeae and Sympetalae. Most of the other 

systems begin with the Magnoliales, but these too 

show evidence of their anemophilous ancestry in 

having pistils with a large stigmatic surface and 

numerous stamens with abundant pollen. 

My approach to understanding evolutionary 

trends in characters is to relate them to the 
NUMBER 58 3 

ecological factors that might be responsible for 

them by their selective action. A major evolutionary trend has been from wind pollination to 

progressively better adaptation for insect pollination. Under favorable conditions where most 

families can survive and members of a species are 

often widely spaced, insect pollination is more 

proficient than wind pollination. Many plants 

developed colorful flowers or bracts, making the 

flowers easier for insects or other animals to find. 

Some flowers became fragrant or malodorous, 

indicating the presence of food to potential pollinators. Some produced nectar that may act as a 

substitute reward in place of reproductive organs. Access to the reward became more and 

more restricted, favoring insects with long sucking mouth parts. Some flowers became more 

specially adapted to the most proficient pollinators, the bees. 

The primitive, insect-pollinated dicots often 

have large flowers with numerous spirally arranged parts. They gave rise to plants whose 

flowers have few, opposite, or whorled parts. 

The proficiency of insects in pollination allows 

survival with fewer parts. Under harsher conditions, where fewer taxa can survive, some insectpollinated plants became gregarious and secondarily wind-pollinated. They also have small flowers with few parts. 

The dicots probably originated under warm 

temperate conditions favorable for growth. Another major evolutionary trend in them has been 

the gradual development of characters and character states enabling them to cope with dry and 

hot or cold conditions, allowing them to colonize 

generally unfavorable regions. 

By reduction of internodes and modification 

of leaves, the flowers developed a calyx that 

provides some protection of the reproductive 

organs against unfavorable climatic conditions, 

as well as enabling them to avoid some predators. 

Development of perigyny and epigyny also conserved moisture in the reproductive organs. The 

leaves evolved various conditions of the cuticle, 

epidermis, and vasculature that enable them to 

resist moisture loss and prevent collapse. Also, in 

the xylem, tracheids were converted to vessels 

and scalariform perforation plates became simple 

and the vessels wider to facilitate rapid movement of water, which is only periodically available under unfavorable conditions. 

The floral organs are homologous with leaves. 

Like leaves, the parts were initially separate. The 

connate and adnate conditions are derived. 

General character states are primitive; specialized states are derived. In attempting to determine which primitive states are most primitive I 

considered their occurrence among the families. 

The fewer the families showing a particular primitive state the more primitive the state. For example, very few families still have vesselless 

wood, so I consider that condition one of the 

most primitive of the primitive states. This is 

important in deciding whether a family is low or 

high on the family tree and the position within 

its particular order. 

In accordance with the above rationale, I have 

constructed Table 2, giving the primitive and 

derived states for about 100 characters. I also 

indicate the extent to which I consider the states 

reversible. In general, character states that are 

most constant within families are least readily 

reversed. For example, flower color is usually 

variable within families and commonly reversible, whereas cotyledon number is almost invariably constant within families and therefore difficult to reverse. The category “n” includes those 

character states that are practically irreversible. 

To determine a family’s phylogeny, it must be 

compared with other families considered to be 

close to it. In general, the more characters and 

character states in common, particularly uncommon ones, the more likely are the subject families 

to be related. All parts of the plant and many 

characters should be considered. 

I agree with those who make a distinction 

between parallelism and convergence. The same 

character or character state may arise more than 

once in a single family or independently in related families. That is parallelism. If the same 

character or character state arises in unrelated 

families it is convergence. Unrelated taxa with 
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 1 .-Ordinal allocation of dicotyledon families by recent authors. 

Familv 

Cercidiphyll 

Tetracentr 

Euptele 

Trochodendr 

"tan 

hlyrothamn 

BUX 

Hamamelid 

Daphniphyll 

Didvmel 

Magnoli 

Winrer 

Annoti 

Degeneri 

Austrobaile) 

Himantandr 

Eupomati 

Myristic 

<:anell 

Schisandr 

Illici 

Amborell 

Trimeni 

I.lonim i 

Idiosperni 

Calycanth 

Goniorteg 

Laur 

Hernandi 

Aristolochi 

Menisperm 

Cronquist Takhtajan Thorne Dahlgren Emberger 

Goldberg 1981 1983 1983 1983 1960 

Trochodendr 

Trochodendr 

Trochodendr 

Trochodendr 

Hainamelid 

Haniamelid 

Hamamelid 

Ha ma ni el id 

Hamamelid 

Hamamelid 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

hlagnoli 

Magnoli 

Magnoli 

Laur 

LdUr 

LdUr 

4ristolochi 

Ra ti u ncu I 

Hamamelid 

Trochodendr 

Hamamelid 

Trochodendr 

Hainamelid 

Hamamelid 

Euphorbi 

Hamamelid 

Daphniphyll 

Didymel 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

lllici 

lllici 

Laur 

Laur 

Laur 

Laur 

Laur 

Laur 

Laur 

Laur 

Aristolochi 

Ranuncul 

Cercidiphyll 

Trochodendr 

Euptele 

Trochodendr 

Hamamelid 

Hamamelidineae 

Hamainelid 

Hamamelidineae 

Hamamelid 

Buxineae 

Hamamelid 

Hamamelidineae 

Hamamelid 

Hamamelidineae 

Didyniel 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Laur 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Illici 

Magnoliineae 

Winterineae 

Annonineae 

Magnoliineae 

Monimiineae 

Magnoliineae 

Magnoliineae 

Annonineae 

Annonineae 

lllici 

Laur 

Monimiineae 

Laur 

Monimiineae 

Laur 

Monimiineae 

in Calycanthac 

Laur 

Monimiineae 

Laur 

Monimiineae 

Laur 

Laurineae 

Laur 

Laurineae 

Aristolochi 

Ranuncul 

Hamamelid 

Trochodendrineae 

Hamamelid 

Trochodendrineae 

Hamamelid 

Trochodendrineae 

Hamamelid 

Trochodendrineae 

Hamamelid 

Haniamelidiiieae 

Pittospor 

Brunineae 

Pittospor 

Buxineae 

Hamamelid 

Hamamelidineae 

Pi ttospor 

Buxineae 

Pi ttospor 

Buxineae 

Annon 

Annonineae 

Annon 

Winterineae 

Annon 

Annonineae 

Annon 

Annonineae 

Annon 

Annonineae 

Annon 

Annonineae 

Annon 

Annonineae 

Annon 

Annonineae 

Annon 

Annonineae 

Annon 

llliciineae 

Annon 

llliciineae 

Annon 

Laurineae 

Annon 

Laurineae 

Annon 

Laurineae 

in Calycanthaceae 

Annon 

Annon 

Annon 

Annon 

Annon 

Berberid 

La u ri neae 

Laurineae 

Laurineae 

Laurineae 

Aristolochiineae 

Berberidineae 

Cercidiphyll 

Trochodendr 

Cercidiphyll 

Trochodendr 

Hamamelid 

Hamamelid 

Bux 

Hamamelid 

Bux 

Bux 

Magnoli 

Magnoli 

Annon 

Magnoli 

Annon 

Magnoli 

Annon 

Annon 

Annon 

lllici 

Illici 

Laur 

Laur 

Laur 

in Calycanthaceae 

Laur 

Laur 

Laur 

in Lauraceae 

Aristolochi 

Ran u nc u I 

Polycarpicae 

Pol ycarpicae 

Polycarpicae 

Polycarpicae 

Ros 

Ros 

Terebinth 

Ros 

Euphorbi 

Leitneri 

Polycarpicae 

Polycarpicae 

Pol ycarpicae 

Pol ycarpicae 

Polycarpicae 

Polycarpicae 

Polycarpicae 

Pol ycarpicae 

Pol ycarpicae 

Pol ycarpicae 

in Winteraceae 

in Iblonimiaceae 

in Monimiaceae 

Pol ycarpicae 

- 

Pol ycarpicae 

Polycarpicae 

Polycarpicae 

Pol ycarpicae 

Aristolochi 

Pol ycarpicae 
NUMBER 58 5 

TABLE 1 .-continued. 

Family 

Cercidiphyll 

Tetracentr 

Euptele 

Trochodendr 

Platan 

Myrothanin 

Bus 

Hamamelid 

Daphniphyll 

Didvmel 

Magnoli 

Winter 

Annon 

Degeneri 

Austrobailey 

Hiniantandr 

Eupomati 

M>i-istic 

Cane I I 

Schisandr 

Illici 

Am horel I 

Trirneni 

Moliini 

Idiosperni 

Cal)canth 

Gomorteg 

Laur 

Hernandi 

Aristolochi 

Menisperm 

Hutchinson Melchior Stebbins Rouleau Young 

1973 1964 1974 1981 1981 

Magnoli 

Hamamelid 

in Trochodendra 

Magnoli 

Hamamelid 

Hamamelid 

Hamamelid 

Hamamelid 

Hamamelid 

Bix 

hlagnoli 

Magnoli 

Annoii 

in M'interaceae 

Laur 

Magnoli 

Annon 

Laur 

Magnoli 

Magnoli 

Magnoli 

in Monimiaceae 

Laur 

Laur 

Ros 

Ros 

Laur 

Laur 

Laur 

Aristolochi 

Berberid 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Ros 

Ros 

Celastr 

Ros 

Gerani 

Leitneri 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Hamamelidineae 

Hamamelidineae 

Buxineae 

Hamamelidineae 

Euphorbiineae 

- 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Aristolochi 

Ranuncul 

Hamamelid 

Trochodendr 

Hamamelid 

Trochodendr 

Hamamelid 

Hamamelid 

Euphorbi 

Hamamelid 

Euphorbi 

Hamamelid 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Laur 

Magnoli 

Laur 

Magnoli 

Magnoli 

Illici 

Illici 

Laur 

Laur 

Laur 

- 

Laur 

Laur 

Laur 

Laur 

Aristolochi 

Ranuncul 

Cercidiphyll 

Trochodendr 

Euptele 

Trochodendr 

Hamamelid 

Hamamelid 

Euphorbi 

Hamamelid 

Euphorbi 

Didymel 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Laur 

Magnoli 

Magnoli 

Magnoli 

Magnoli 

Illici 

lllici 

Laur 

Laur 

Laur 

Laur 

Laur 

Laur 

Laur 

Laur 

Aristolochi 

Ranuncul 

Trochodendr 

Trochodendr 

Trochodendr 

Trochodendr 

Hamamelid 

Myrothamn 

Bux 

Hamamelid 

Bux 

Euphorbi 

Magnoli 

Magnoli 

Magnoli 

Degeneri 

Illici 

Aristolochi 

Mdgnoli 

Aristolochi 

Magnoli 

Illici 

lllici 

Laur 

Laur 

Laur 

Laur 

Laur 

Laur 

Laur 

Laur 

Aristolochi 

Ranuncul 

Benson 

1979 

Hamamelid 

Hamamelid 

Ha rn a m e I i d 

Hamamelid 

Hamamelid 

Ros 

Trochodendrineae 

Trochodendrineae 

Trochodendrineae 

Trochodendrineae 

Hamanielidineae 

Euphorbi 

Hamamelid 

Hamamelidineae 

Ran 

Ran 

Ran 

Ran 

Magnoliineae 

Magnoliineae 

Magnoliineae 

Magnoliineae 

Laurineae 

Magnoliineae 

Magnoliineae 

Magnoliineae 

Laurineae 

Magnoliineae 

Magnoliineae 

Laurineae 

Laurineae 

Laurineae 

Rd n 

Ran 

Ran 

Ran 

Ran 

Ran 

Ran 

Ran 

Ran 

Ran 

- 

Ran 

Ran 

Ran 

Ran 

Aristolochi 

Ran 

Laurineae 

Laurineae 

Laurineae 

Laurineae 

Ranunculineae 
6 

Family 

Sargen t odox 

Lardizabal 

Nandin 

Berberid 

Podophyll 

Paeoni 

Ranuncul 

Circaeaster 

Halorag 

Gunner 

Hippurid 

Callitrich 

Sarraceni 

Nepenth 

Saurur 

Piper 

Lactorid 

Chloranth 

Ceratophyll 

Ca boni b 

Nyniphae 

Nel u m bon 

Prote 

Balanop 

Sininiondsi 

Leitneri 

Myric 

Jugland 

Rhoiptele 

Fag 

Betul 

Salic 

Casuarin 

F.uconinii 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 1 .-continued. 

Goldberg 

Ranuncul 

Ranuncul 

Ranuncul 

Ranuncul 

Ranuncul 

Ranuncul 

Ranuncul 

Ranuncul 

Halorag 

Halorag 

Halorag 

Halorag 

Sarraceni 

Nepenth 

Piper 

Piper 

Piper 

Piper 

Nyniphae 

Nyniphae 

Nyniphae 

Nyniphae 

Prote 

Balanop 

Fag 

Fd g 

Fa g 

Fag 

Fag 

Fag 

Fag 

Salic 

Casuarin 

Urtic 

Cronquist 

1981 

Ranuncul 

Ranuncul 

in Berberidaceae 

Ranuncul 

in Berberidaceae 

Dilleni 

Ranuncul 

Ranuncul 

Halorag 

Halorag 

Callitrich 

Callitrich 

Nepenth 

Nepenth 

Piper 

Piper 

Magnoli 

Piper 

Nymphae 

Nymphae 

Nymphae 

Nymphae 

Prote 

Fag 

Euphorbi 

Leitneri 

Myric 

Jugland 

Jugland 

Fag 

Fag 

Salic 

Casuarin 

Euconimi 

Ta k h tajan Thorne Dahlgren Eniberger 

1983 1983 1983 1960 

Ranuncul 

Ranuncul 

in Berberidaceae 

Ranuncul 

in Berberidaceae 

Paeoni 

Ranuncul 

Ranuncul 

Myrt 

Saxifrag? 

Scrophulari 

Lami 

Sarraceni 

Haloragineae 

Saxifragineae 

Hippurineae 

Nepenth 

Piper 

Piper 

Laur 

Laur 

Nymphae 

Nymphae 

Nymphae 

Nelumbon 

Prote 

Balanop 

Hamamelid 

Buxineae 

Leitneri 

Myric 

Jugland 

Jugland 

Lactoridineae 

Chloranthineae 

Ceratophyllineae 

N ymphaeineae 

N ymphaeineae 

Fag 

Fag 

Salic 

Casuarin 

Eucommi 

Berberid 

Berberidineae 

Berberid 

Berberidineae 

Berberid 

Berberidineae 

Berberid 

Berberidineae 

in Berberidaceae 

Paeoni 

Berberid 

Berberid 

Corn 

Corn 

Corn 

Lami 

The 

The 

Annon 

Annon 

Annon 

Annon 

Ne I u m bon 

Nymphae 

Nymphae 

Nelumbon 

Prote 

Pittospor 

Euphorbi 

Rut 

Rut 

Rut 

Rut 

Berberidineae 

Berberidineae 

Haloragineae 

Haloragineae 

Haloragineae 

Sarraceniineae 

Nepenthineae 

Piperineae 

Piperineae 

Laurineae 

Laurineae 

Buxineae 

Rutineae 

Myricineae 

Juglandineae 

Juglandineae 

Fag 

Fag 

Viol 

Salicineae 

Casuarin 

Hamamelid 

Eucommiineae 

Ranuncul 

Ranuncul 

in Berberidaceae 

Ranuncul 

in Berberidaceae 

Paeoni 

Ranuncul 

Ranuncul 

Halorag 

Gunner 

Hippurid 

Lami 

Sarraceni 

The 

Piper 

Piper 

Lactorid 

Chloranth 

Nymphae 

Nymphae 

Nymphae 

Nelurnbon 

Prote 

Balanop 

Euphorbi 

Sapind or 

Rut 

Myric 

Jugland 

Jugland 

Fag 

Fag 

Salic 

Casuarin 

Eucommi 

Polycarpicae 

Polycarpicae 

in Berberidaceae 

Pol ycarpicae 

in Berberidaceae 

in Berberidaceae 

Pol ycarpicae 

Poi ycarpicae 

Myrt 

Myrt 

Myrt? 

Myrt? 

Sarraceni 

Sarraceni 

Piper 

Piper 

Polycarpicae 

Piper 

Polycarpicae 

Polycarpicae 

Pol ycarpicae 

in Nymphaeaceae 

Prote 

Balanop 

in Buxaceae 

Leitneri 

Myric 

Jugland 

Urtic or 

Jugland 

Fag 

Fag 

Salic 

Casuarin 

Urtic 
NUMBER 58 7 

Family 

Sargentodox 

Lardirabal 

Nandin 

Berberid 

Podoph\ll 

Paeoni 

Ranikncul 

Circaeaster 

Halorag 

Gunner 

Hippurid 

Callitrich 

Sarraceni 

Nepenth 

Saurur 

Piper 

Lactorid 

Chloranth 

Ceratophyll 

Ca bo ni b 

Nymphae 

Nel unibon 

Prote 

Balanop 

Siniiiiondsi 

Leitneri 

Myric 

Jugland 

Rhoiptele 

Fag 

Betul 

Salic 

Casuarin 

Eucommi 

Table 1 .-continued. 

Hutchinson Melchior Stebbins Rouleau Young 

1973 1964 1974 1981 1981 

Berberid 

Berberid 

Berberid 

Berberid 

Ran 

Ran 

Ran 

Berberid 

Onagr 

in Haloragaceae 

in Haloragaceae 

Onagr 

Sarraceni 

Aristolochi 

Piper 

Piper 

Magnoli 

Piper 

Ran 

Ran 

Ran 

in Nyniphaeaceae 

Prote 

Balanop 

in Buxaceae 

Leitneri 

Myric 

Jugland 

Jugland 

Fa g 

Fag 

Salic 

Casuarin 

Urtic 

Ranuncul 

Ranuncul 

in Berberidac 

Ranuncul 

in Berberidac 

Guttifer 

Ranuncul 

in Ranunculac 

Myrt 

Myrtineae 

in Haloragaceae 

Myrt 

Tubiflorae 

Sarraceni 

Sarraceni 

Piper 

Piper 

Piper 

Piper 

Ranuncul 

in Nymphaeaceae 

Ranuncul 

in Nymphaeaceae 

Prote 

Balanop 

in Buxaceae 

Leitneri 

Jugland 

Jugland 

Urtic 

H ippuridineae 

Verbenineae 

Fag 

Fag 

Salic 

Casuarin 

Urtic 

- 

Ranuncul 

- 

Ranuncul 

- 

Dilleni 

Ranuncul 

Ranuncul 

Halorag 

Halorag 

Halorag 

Lami 

Sarraceni 

Aristolochi? 

Piper 

Piper 

Laur 

Laur? 

Nymphae 

- 

Nymphae 

Nymphae 

Prote 

Fag 

- 

Leitneri 

Myric 

Jugland 

Jugland 

Fag 

Fag 

Salic 

Casuarin 

Eucommi 

Ranuncul 

Ranuncul 

Ranuncul 

Ranuncul 

Ranuncul 

Paeoni 

Ranuncul 

Ranuncul 

H ippurid 

Hippurid 

Hippurid 

Lami 

Sarraceni 

Nepenth 

Piper 

Piper 

Laur 

Laur 

Nymphae 

Nymphae 

Nymphae 

Nelumbon 

Prote 

Balanop 

Euphorbi 

Leitneri 

Myric 

Jugland 

Jugland 

Fag 

Betul 

Salk 

Casuarin 

Eucommi 

Ranuncul 

Ranuncul 

Ranuncul 

Ranuncul 

Ranuncul 

Ranuncul 

Ranuncul 

Ranuncul 

Halorag 

Halorag 

Hippurid 

Lami 

Sarraceni 

Nepenth 

Aristolochi 

Aristolochi 

Laur 

Laur 

Ranuncul 

Nymphae 

Nymphae 

lllici 

Prote 

Balanop 

Euphorbi 

Rut 

Myric 

Jugland 

Jugland 

Fag 

Fag 

Salk 

Casuarin 

Eucommi 

Benson 

1979 

- 

Ran 

Ranuncuheae 

- 

Ran 

Ranunculineae 

- 

Dilleni 

Ran 

Ranunculineae 

- 

Myrt 

in Haloragaceae 

Mvrt 

Lami 

Sarraceni 

The 

Piper 

Piper 

Ran 

Ran 

Nymphae 

Laurineae 

Laurineae 

Nymphae 

in Nymphaeaceae 

Prote 

Euphorbi 

Leitneri 

Myric 

Jugland 

Jugland 

Fag 

Fag 

Salic 

Casuarin 

Hamamelid 

Eucommiineae 
8 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 1 .-continued. 

Family 

Barbey 

Urtic 

Cannab 

Sfor 

Ulm 

Caesalpini 

Fa b 

Mimos 

Papaver 

Fumari 

l'ovari 

Cappar 

Pentadiplandr 

Resed 

Brassic 

Moriirg 

Bretschneider 

Bat 

Dilleni 

Actinidi 

Sa u ra u i 

The 

Pentaphylac 

Bonneti 

Pellicier 

Medusagyn 

Eucryphi 

Paracryphi 

Symploc 

Tetranierist 

Clusi 

Quiin 

Hyperic 

Elatin 

Dipterocarp 

Huniiri 

Ancistroclad 

Cronquist 

Goldberg 1981 

Urtic 

Urtic 

Urtic 

Crtic 

Urtic 

Fab 

Fab 

Papaver 

Papaver 

Papaver 

Papaber 

Papaver 

Papaver 

Papaver 

Papaver 

Papaver 

Bat 

Dilleni 

Dilleni 

Dillerii 

The 

The 

The 

The 

The 

The 

The 

The 

The 

The 

The 

The 

The 

The 

The 

The 

Fa b 

Urtic 

Urtic 

Crtic 

Crtic 

Urtic 

Fa b 

Fa b 

Fab 

Papaver 

Papaver 

Cappar 

Cappar 

in Capparaceae 

Cappar 

Cappar 

Cappar 

Sapind 

Bat 

Dilleni 

The 

in Actinidiaceae 

The 

The 

in Theaceae 

The 

The 

Ros 

The 

Eben 

The 

The 

The 

in Clusiaceae 

The 

The 

Lin 

Viol 

Tak h tajan Thorne 

1983 1983 

Barbey 

Urtic 

Urtic 

Crtic 

Urtic 

in Fabaceae 

Fab 

in Fabaceae 

Papaver 

in Papaveraceae 

Cappar 

Capparineae 

Cappar 

Capparineae 

in Capparaceae 

Cappar 

Resedineae 

Cappar 

Capparineae 

Cappar 

Moringineae 

Sapind 

Sapind 

Dilleni 

Eric 

in Actinidiaceae 

The 

The 

The 

The 

The 

Saxifrag 

Celastr 

Eben 

The 

The 

The 

in Clusiaceae 

The 

Malv 

Cerani 

The 

C u noni i neae 

lcacinineae 

Styracineae 

Linineae 

- 

Urtic 

Crtic 

in Urticaceae 

Urtic 

in Fabaceae 

Rut 

Fabineae 

in Fabaceae 

Berberid 

Papaverineae 

in Papaveraceae 

in Capparaceae 

Cappar 

in Capparaceae 

Cappar 

Cappar 

Cappar 

Rut 

Rut 

The 

The 

in Actinidiaceae 

The 

The 

in Clusiaceae 

in Theaceae 

The 

Scytopetalineae 

Ros 

Cunoniineae 

The 

Theineae 

The 

Theineae 

in Theaceae 

The 

The 

in Clusiaceae 

The 

Malv 

Cerani 

The 

Sapindineae 

Sapindineae 

Dilleniineae 

Theineae 

Theineae 

Clethrineae 

H ypericineae 

Scytopetalineae 

Hypericineae 

Linineae 

Scytopetalineae 

Dahlgren Emberger 

1983 1960 

Urtic 

Urtic 

Urtic 

Urtic 

Urtic 

Fa b 

Fab 

Fab 

Papaver 

Papaver 

Cappar 

Cappar 

in Capparaceae 

Cappar 

Cappar 

Cappar 

Sapind? 

Cappar 

Dilleni 

Eric 

in Actinidiaceae 

The 

The? 

in Theaceae 

in Theaceae 

The? 

Cunoni 

Corn 

Corn 

in Theaceae 

The 

The 

in Clusiaceae 

The 

Malv 

Cerani 

The 

in Ulmaceae 

Urtic 

in Moraceae 

Urtic 

Crtic 

Ros 

Ros 

Ros 

Rhoead 

in Papaveraceae 

Rhoead 

Rhoead 

Olac 

Rhoead 

Rhoead 

Rhoead 

Rhoead 

Batid 

Pariet 

Pariet 

Pariet 

Pariet 

Terebinth 

in Theaceae 

in Theaceae 

Pariet 

Pariet 

in Eucryphiaceae 

Eben 

in Theaceae 

Pariet 

Pariet 

in Guttiferae 

Pariet 

Pariet 

Cerani 

Pariet 
NUMBER 58 

TABLE 1 .-continued. 

9 

Family 

Barbe) 

Urtic 

(:annab 

Mor 

Uln1 

C;,iesalpini 

Fal, 

Mimos 

Papaver 

Fumari 

Tovari 

Cdppdr 

Pentadiplandr 

Resed 

Brassic 

hloring 

Brrtschneider 

Ed 1 

Dilleni 

Actinidi 

Sauraui 

The 

Pentaph\ lac 

Bonneti 

Pellicier 

Medusaagyn 

Eurryhi 

Pat-acryphi 

Syn1ploc 

Tetrameribt 

Chi 

Quiiii 

H) peric 

Ela~in 

Dipterocarp 

H umi ri 

Anc istroclad 

Hutchinson 

1973 

Urtic 

Urtic 

Urtic 

Lt-tic 

Lrtic 

Leguniin 

Legumin 

I.egumin 

Rhoead 

Rhoedd 

CappalCappar 

Celastr 

Rered 

Brassic 

Cappar 

in Sapindacede 

Chenopodi 

1)illeni 

The 

The 

The 

The 

I 

~ 1he 

The 

1 The 

i Styrac 

1-he 

Guttifer 

I Guttifer 

(httifer 

Car~ophyll 

i Ochn 

I Malpighi 

Ochn 

Melchior Stebbins Rouledu Young 

1964 1974 1981 1981 

in Clniaceae 

Urtic 

in Moraceae 

Urtic 

Urtic 

in Leguminosae 

Ros 

in Leguminosae 

Papaver 

Papaverineae 

in Papaveraceae 

Papaver 

Capparineae 

Papaver 

Capparineae 

in Capparaceae 

Papaver 

Resedineae 

Papaver 

Capparineae 

Papaver 

Moringineae 

Sapind 

Sapindineae 

Bat 

Leguminosineae 

Guttifer 

Guttifer 

in Actinidiac 

Guttifer 

Theineae 

Celastr 

Celastrineae 

in Theaceae 

in Theaceae 

Guttifer 

Guttifer 

Dilleniineae 

Dilleniineae 

Dilleniineae 

Dilleniineae 

Eben 

Ebenineae 

in Theaceae 

Guttifer 

Theineae 

Guttifer 

Theineae 

in Guttiferae 

Viol 

Guttifer 

in Linaceae 

Tamaricineae 

Ochnineae 

Guttifer 

Ancistrocladin 

Urtic 

Urtic 

Urtic 

Urtic 

Urtic 

in Leguminosae 

Fab 

in Leguminosae 

Papaver 

Papaver 

Capper 

Cappar 

- 

Cappar 

Cappar 

Cappar 

Bat 

Dilleni 

Eric? 

- 

The 

- 

- The 

Ros 

Eben 

- 

The 

The 

in Guttiferae? 

The 

The 

Gerani 

Viol? 

Barbey 

Urtic 

Urtic 

Urtic 

Urtic 

Fa b 

Fab 

Fa b 

Papaver 

Papaver 

Capper 

Cappar 

Cappar 

Cappar 

Cappar 

Cappar 

Sapind 

Caryophyll 

Dilleni 

Eric 

Eric 

The 

The 

The 

The 

The 

Saxifrag 

Saxifrag 

Eben 

The 

The 

The 

The 

The 

The 

Gerani 

The 

Ole 

Urtic 

Urtic 

Urtic 

Urtic 

Fa b 

Fab 

Fa b 

Ranuncul 

Ranuncul 

Capper 

Cappar 

- 

Cappar 

Cappar 

Cappar 

Sapind 

Cappar 

Dilleni 

The 

- 

The 

The 

- 

- 

Scytopetal 

Cunoni 

The 

Eben 

Scytopetal 

Hyperic 

Hyperic 

Malv 

Lin 

Lin 

Benson 

1979 

- 

Urtic 

Urtic 

Urtic 

Urtic 

in Leguminosae 

Fa b 

in Leguminosae 

Papaver 

Papaverineae 

Papaver 

Papaverineae 

Papaver 

Cdpparidinede 

Papaver 

Capparidineae - 

Papaver 

Papaver 

Papaver 

Sapind 

Bdtid 

Capparidineae 

Capparidineae 

Capparidineae 

Sapindineae 

Dilleni 

The 

- 

The 

The 

- 

The 

Ros 

Eben 

- 

The 

The 

in Guttiferae 

The 

The 
10 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 1 .-continued. 

Fdnlil~ 

Marcgravi 

Car yocar 

Ochn 

Strasburgeri 

Diegodendr 

Sphaerosepal 

Scvtopetal 

Sarcolaen 

Eben 

SdpOt 

Stlrac 

Lissocarp 

Alangi 

Flacourti 

Lacistemat 

Passiflor 

Malesherbi 

Turner 

Achari 

Caric 

Viol 

Stachyur 

Scyphostegi 

Peridisc 

Hoplestigniat 

Loas 

Cucurbit 

Cist 

Cochlosperm 

Bix 

Eric 

Clethr 

Goldberg 

The 

The 

The 

The 

The 

The 

The 

The 

Eben 

Eben 

Styrac 

Styrac 

Styrac 

Viol 

Viol 

Viol 

Viol 

Viol 

Viol 

\'iol 

Viol 

L'iol 

Viol 

Viol 

Viol 

Viol 

Viol 

Cist 

Cist 

Gist 

Eric 

Eric 

Cronquist Takh tajan Thorne Dahlgren Emberger 

1981 1983 1983 1983 1960 

The 

The 

The 

in Ochnaceae 

in Ochnaceae 

The 

The 

The 

Eben 

Eben 

Eben 

Eben 

Corn 

Viol 

Viol 

Viol 

Viol 

Viol 

Viol 

Viol 

Viol 

Viol 

Viol 

Viol 

Viol 

Viol 

viol 

Viol 

in Bixaceae 

Viol 

Eric 

Eric 

The 

The 

The 

The 

The 

Malv 

Malv 

Mah 

Eben 

Eben 

Eben 

Eben 

Corn 

Ebenineae 

Ebenineae 

Styracineae 

Styracineae 

Viol 

Violineae 

in Flacourtiac 

Viol 

Violineae 

Viol 

Violineae 

Viol 

Violineae 

Viol 

Violineae 

Viol 

Violineae 

Viol 

Violineae 

Viol 

Violineae 

Viol 

Violineae 

Viol 

Violineae 

Polemoni 

Boraginineae 

Loas 

Viol 

Cucurbitineae 

Viol 

Violineae 

in Bixaceae? 

Viol 

Eric 

Violineae 

Eric 

The 

Theineae 

The 

Theineae 

The 

Scytopetalineae 

The 

Scytopetalineae 

in Sphaerosepalac 

The 

Scytopetalineae 

The 

Scytopetalineae 

Malv 

Eben 

Eben 

Eben 

Eben 

Corn 

Viol 

in Flacourtiaceae 

Viol 

Viol 

Viol 

Viol 

Viol 

Viol 

The 

Viol 

Viol 

Ebenineae 

Ebenineae 

Styracineae 

Ebenineae 

Cornineae 

Violineae 

Violineae 

Violineae 

Violineae 

Violineae 

Violineae 

Violineae 

Theineae 

Violineae 

Violineae 

- 

Loas 

Viol 

Malv 

Cucurbitineae 

Malv 

Malv 

Eric 

The 

Clethrineae 

The 

The 

The 

The 

- 

Malv? 

The? 

Malv 

Eben 

Eben 

Eben 

Eben 

Corn 

Viol 

in Flacourtiaceae 

Viol 

Viol 

Viol 

Viol 

Viol 

Viol 

The 

Viol 

Viol? 

Boragin? 

Loas 

Cucurbit 

Malv 

Malv 

Malv 

Eric 

Eric 

Pariet 

Pariet 

Pariet 

Pariet 

- 

Pariet 

Malv 

Malv 

Eben 

Eben 

Eben 

Eben? 

Urnbell 

Pariet 

Pariet 

Pariet 

Pariet 

Pariet 

Pariet 

Pariet 

Pariet 

Pariet 

Urtic? 

- 

Tubiflor 

Pariet 

Cucurbit 

Pariet 

Pariet 

Pariet 

Eric 

Eric 
NUMBER 58 11 

Table 1 .-continued. 

______ 

Family 

Marcgravi 

Caryocar 

Ochn 

Strasburgeri 

Diegodendr 

Sphaeroscpal 

Scytopetal 

Sarcolaen 

Eben 

Sapot 

St) lac 

Lissocarp 

Alangi 

Flac-ourti 

Lacistemat 

Passiflor 

Malesherbi 

Turner 

.4chari 

Caric 

Viol 

Stachyur 

Scyphostegi 

Peridisc 

Hoplestigniat 

Loas 

Cucurbit 

Cist 

Cochlosperni 

Bix 

Eric 

Clethr 

Hutchinson Melchior Stebbins Rouleau Young Benson 

1973 1964 1974 1981 1981 1979 

The 

The 

Ochn 

Ochn 

Ochn 

Ochn 

Tili 

Ochn 

Eben 

Eben 

Styrac 

StVrdC 

Arali 

Bix 

Bix 

Passiflor 

Passiflor 

Loas 

Passiflor 

Cucurbit 

Viol 

Haniamelid 

Celastr 

Bix 

Bix 

Loas 

Cucurbit 

Bix 

Bix 

Bix 

Eric 

Eric 

Guttifer 

Guttifer 

Guttifer 

Guttifer 

Theineae 

Theineae 

Ochnineae 

Ochnineae 

- 

Viol 

Malv 

Malv 

Eben 

Eben 

Eben 

Eben 

Urnbell 

Cistineae 

Scytopetalin 

Sarcolaenineae 

Ebenineae 

Sapotineae 

Ebenineae 

Ebenineae 

Viol 

in Flacourtiac 

Viol 

Viol 

Viol 

Viol 

Viol 

Viol 

Viol 

Viol 

Viol 

Eben 

Viol 

Cucurbit 

Flacourtiineae 

Flacourtiineae 

Flacourtiineae 

Flacourtiineae 

Flacourtiineae 

Caricineae 

Flacourtiineae 

Flacourtiineae 

Flacourtiineae 

Flacourtiineae 

Ebenineae 

Loasineae 

Viol 

Viol 

Viol 

Eric 

Cistineae 

Cistineae 

Cistineae 

Eric 

The 

The 

The 

Malv? 

Malv 

Malv? 

Eben 

Eben 

Eben 

Eben 

Corn 

Viol 

- Viol 

Viol 

Viol 

Viol 

Viol 

Viol 

The 

Viol 

Viol 

Polemoni? 

Viol? 

Viol 

Viol 

- 

Viol 

Eric 

Eric 

The 

The 

The 

The 

The 

Malv 

Malv 

Malv 

Eben 

Eben 

Eben 

Eben 

Corn 

Viol 

Viol 

Passiflor 

Passiflor' 

Passiflor 

Passiflor 

Passiflor 

Viol 

Viol 

Viol 

Viol 

Boragin 

Loas 

Cucurbit 

Viol 

Viol 

Viol 

Eric 

Eric 

The 

The 

Scytopetal 

Scytopetal 

- 

Scytopetal 

Scytopetal 

Malv 

Eben 

Eben 

Eben 

Eben 

Corn 

Viol 

in Flacourtiac 

Viol 

Viol 

Viol 

Viol 

Viol 

Viol 

The 

Viol 

Viol 

Boragin 

Loas 

Viol 

Malv 

Malv 

Malv 

Eric 

The 

The 

The 

The 

The 

- 

Malv 

Malv 

Eben 

Eben 

Eben 

Eben 

Corn 

Viol 

- 

Viol 

Viol 

Viol 

Viol 

Viol 

Viol 

The 

Viol 

Viol 

Loas 

Loas 

Viol 

Viol 

Viol 

Eric 

Ericineae 

in Ericaceae 
12 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 1 .-continued. 

Family 

~ 

Pyrol 

Monotrop 

Em pet r 

Epacrid 

Diapensi 

Cvrill 

Lenno 

Hydnor 

Rafflesi 

Balanophor 

Ctenolophon 

lxonanth 

Irvingi 

Dichaperal 

Celastr 

Goupi 

Siphonodon t 

\‘it 

Lee 

Rhanin 

Er) thropal 

Aqulfoll 

Icacin 

Aiio 

Portulac 

Theligon 

Didiere 

Gvrostemon 

Phvtolacc 

Barbeui 

Achatocarp 

Petiveri 

Agdestid 

Nyctagin 

Stegnosperniat 

Caryophyll 

Cronquist Takhtajan Thorne Dahlgren Emberger 

Goldberg 1981 1983 1983 1983 1960 

Eric 

Eric 

Eric 

Eric 

Eric 

Eric 

Eric? 

Ramesi 

Rafflesi 

Balanophor 

Celastr 

Celastr 

Celastr 

Celastr 

Celastr 

Celastr 

Celastr 

Rhamn 

Rhanin 

Rhanin 

Rhanin 

Rhamn 

Rhanin 

Caryophyll 

Caryophyll 

Caryophyll 

Caryophyll 

Caryophyll 

Cdryophyil 

Caryophyll 

Caryoph yll 

Caryophyll 

Caryophyll 

Caryophyll 

Caryophyll 

Caryophyll 

Caryophyll 

Eric 

Eric 

Eric 

Eric 

Diapensi 

Eric 

Lami 

Rafflesi 

Rafflesi 

Santal 

in Hugoniaceae 

Lin 

in Simaroubaceae 

Celastr 

Celastr 

in Celastraceae 

in Celastraceae 

Rhanin 

Rhamn 

Rhainn 

in Olacaceae 

Celastr 

Celastr 

Caryophyll 

Caryophyll 

Caryophyll 

Rubi 

Caryophyll 

Bat 

Caryophyll 

in Phytolaccac 

Caryophyll 

in Phytolaccac 

in Phytolaccac 

Caryophyll 

in Phytolaccac 

Caryophyll 

in Ericaceae 

in Ericaceae 

Eric 

Eric 

Eric 

Eric 

Polemoni 

Rafflesi 

Rafflesi 

Balanophor 

in Linaceae 

Boraginineae 

in Linaceae 

in Simaroubaceae 

Euphorbi 

Celastr 

Celastr 

Celastr 

Rhanin 

Rhamn 

Rhamn 

in Olacaceae? 

Celastr 

lcacinineae 

Celastr 

Icacinineae 

Caryophyll 

Phytolaccineae 

Caryophyll 

Phytolaccineae 

Caryophyll 

Phytolaccineae 

Gentian 

Caryophyll 

Sapind 

Caryophyll 

Ph ytolaccineae 

in Phytolaccaceae 

Caryophyll 

Phytolaccineae 

in Phytolaccaceae 

in Phytolaccaceae 

Caryophyll 

Phytolaccineae 

Caryophyll 

Phytolaccineae 

Caryophyll 

Caryophyllineae 

Celastrineae 

Celastrineae 

Celastrineae 

Phytolaccineae 

in Ericaceae 

in Ericaceae 

Eric 

Eric 

Ros 

The 

Solan 

Rafflesi 

Rafflesi 

Balanophor 

Gerani 

Linineae 

in Linaceae 

in Rutaceae 

Euphorbi 

Celastr 

Saxifragineae 

Clethrineae 

Boraginineae 

in Celastraceae 

in Celastraceae 

Corn 

Vitineae 

in Vitaceae 

Rhamn 

in Olacaceae 

The 

Theineae 

The 

Theineae 

Chenopodi 

Portulacineae 

Chenopodi 

Chenopodiineae 

Chenopodi 

Portulacineae 

in Rubiaceae 

Chenopodi 

Rut 

Chenopodi 

Chenopodi 

Chenopodi 

in Phytolaccaceae 

in Phytolaccaceae 

Chenopodi 

Chenopodi 

Chenopodi 

Portulacineae 

Sapindineae 

Chenopodiineae 

Chenopodiineae 

Chenopodiineae 

Chenopodiineae 

Chenopodiineae 

Chenopodiineae 

Eric 

Eric 

Eric 

Eric 

Eric 

Eric 

Boragin? 

Rafflesi 

Rafflesi 

Balanophor 

Gerani 

Gerani 

in Simaroubaceae 

Euphorbi 

Celastr 

in Celastraceae 

in Celastraceae 

\‘it 

in Vitaceae 

Rhamn 

Corn 

- 

Corn 

Caryophyll 

Caryophyll 

Caryophyll 

Gentian 

Caryophyll 

Cappar 

Caryophyll 

in Phytolaccaceae 

in Phytolaccaceae 

Caryophyll 

Caryophyll 

Caryophyll 

Eric 

in Pyrolaceae 

Eric 

Eric 

Eric? 

Terebinth 

Tubiflor 

Aristolochi 

Aristolochi 

Santal 

in Linaceae 

in Linaceae 

in Simaroubaceae 

Euphorbi 

Celastr 

in Celastraceae 

- 

Rhamn 

Rhamn 

Rhamn 

Celastr 

Celastr 

Celastr 

Centrosperni 

Centrosperm 

Centrosperm 

Centrosperm? 

Terebinth? 

Centrosperm 

Centrosperm 

in Phytolaccaceae 

Centrosperm 

in Phytolaccaceae 

in Phytolaccaceae 

Centrosperm 

- 

Centrosperm 
NUMBER 58 

TABLE 1 .-continued. 

13 

Family 

Pyrol 

Monotrop 

Enipet r 

Epdcrid 

Diapensi 

Cyril1 

Leniio 

Hydnor 

Rafflesi 

Bdlanophor 

Ctenolophon 

Ixonanth 

Irvingi 

Dichapetal 

Celastr 

Goupi 

Siphonodont 

\‘it 

Lee 

Rhamn 

Erythropal 

Aquifoli 

Icacin 

Cact 

Aizo 

Portulac 

Theligon 

Didiere 

Gyrostenion 

Phytolacc 

Barb4 

Achatocarp 

Petiveri 

Agdestid 

Nyctagin 

Stegnosperniat 

Cdryophyli 

Hutchinson 

1973 

Eric 

Eric 

Celastr 

Eric 

Eric 

Celastr 

Eric 

Aristolochi 

Aristolochi 

Malpighi 

Malpighi 

Malpighi 

Ros 

Celastr 

Celastr 

Celastr 

Rhanin 

in Vitaceae 

Rhanin 

Celastr 

Celastr 

Celastr 

Cact 

Caryophyll 

Car5ophyll 

Chenopodi 

Sapind 

Chenopodi 

Chenopodi 

Chenopodi 

Bix 

Chenopodi 

Chenopodi 

Thvmelae 

Pittospor 

Caryophyll 

SdnYdl 

Melchior Stebbins Rouleau Young Benson 

1964 1974 1981 1981 1979 

Eric 

in Pyrolaceae 

Eric 

Eric 

Diapensi 

Celastr 

Tubiflorae 

Aristolochi 

Aristolochi 

Balanophor 

in Linaceae 

Celastrineae 

Boraginineae 

in Linaceae 

in Simaroubac 

Thymelae 

Celastr 

Celastrineae 

in Celastraceae 

in Celastraceae 

Rhamn 

Rhamn 

Rhamn 

in Olacaceae 

Celastr 

Celastr 

Cact 

Celastrineae 

lcacinineae 

Centrospermae 

Centrospermae 

Myrt 

Myrtineae 

Centrosperrnae? 

Centrospermae 

Centrosperrnae 

in Phytolacc 

Centrospermae 

in Phvtolacc 

in Phytolacc 

Centrospermae 

in Phytolacc 

Centrospermae 

Eric 

Eric 

Eric 

Eric 

Diapensi 

Eric 

i’olemoni 

Rafflesi 

Rafflesi 

Santal - 

- 

- 

Celastr? 

Celastr 

- 

Celastr 

Rhamn 

Rhamn 

Rhamn 

Celastr 

Celastr 

Caryophyll 

Caryophyll 

Caryophyll 

Halorag? 

Caryoph yll 

- 

- 

Caryophyll 

- 

- 

- 

- 

Caryophyll 

- 

Caryophyll 

Eric 

Eric 

Eric 

Eric 

Diapensi 

Eric 

Boragin 

Rarflesi 

Rafflesi 

Balanophor 

Gerani 

Gerani 

Euphorbi 

Celastr 

Celastr 

Celastr 

Rharnn 

Rhamn 

Rhamn 

Santal 

Celastr 

Celastr 

Caryophyll 

Caryophyll 

Caryophyll 

Theligon 

Caryophyll 

Caryophyll 

Caryophyll 

Caryophyll 

Caryophyll 

- 

- 

Caryophyll 

Caryophyll 

Caryophyll 

Caryophyll 

- 

- 

Eric 

Eric 

Diapensi 

The 

Boragin 

Rafflesi 

Rafflesi 

Balanophor 

- 

- 

- 

Euphorbi 

Celastr 

- 

- 

\‘it 

- 

Rhamn 

The 

The 

Caryophyll 

Caryophyll 

Caryophyll 

in Rubiaceae 

Chenopodi 

Cappar 

Chenopodi 

Chenopodi 

Chenopodi 

Chenopodi 

Chenopodi 

Chenopodi 

Chenopodi 

Caryophyll 

- 

in Ericaceae 

in Ericaceae 

Eric 

Eric 

Eric 

Eric 

Polenioni 

Rafflesi 

Rafflesi 

Santal 

Empetrineae 

Ericineae 

Diapensiinede 

Ericineae 

Polemoniin 

- 

- 

- 

Sapind 

Celastrineae 

- 

Rharnn 

- 

Rhamn 

Sapind 

Sapind 

Cact 

Caryophyll 

Caryoph yll 

- 

Celastrineae 

Celastrineae 

Caryophyll 

Caryophyll 

Caryophyll 

in Phytolaccaceae 

in Phytolaccaceae 

Caryophyll 

Caryophyll 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 1 .-continued. 

Family 

Mollugin 

Illecebr 

Basell 

Amaranth 

Chenopodi 

Primul 

Plum bagin 

Taniaric 

Frankeni 

Myrsin 

Theophrast 

Polygon 

Plantagin 

Euphorbi 

Aextoxic 

Pand 

Sterculi 

Elaeocarp 

Tili 

Mah 

Bonibac 

Ros 

Corynocarp 

Crossosoniat 

Neurad 

Coriari 

Myrt 

Lecythid 

Barringtoni 

Asteranth 

Dialypetalanth 

Sonnerati 

Punic 

Rhizophor 

Cronquist Takhtajan Thorne Dahlgren Emberger 

Goldberg 1981 1983 1983 1983 1960 

Caryophyll 

Cat-yophyll 

Caryophyll 

Caryophyll 

Caryophyll 

Priniul 

Prim u I 

Priniul 

Primul 

Priniul 

Priniul 

Polygon 

Plantagin 

Euphorbi 

Euphorbi 

Euphorbi 

Malv 

Malv 

Malv 

Malv 

Malv 

Ros 

Ros 

Ros 

Ros 

Ros? 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Caryophyll 

in Caryophyllac 

Caryophyll 

Caryophyll 

Caryophyll 

Priniul 

Plumbagin 

Viol 

Viol 

Priniul 

Primul 

Polygon 

Plantagin 

Euphorbi 

Celastr 

Euphorbi 

Malv 

Malv 

Malv 

Malv 

Malv 

Ros 

Celastr 

Ros 

Ros 

Ranuncul 

Myrt 

Lecythid 

in Lecythidaceae 

in Lecythidaceae 

Ros 

Myrt 

Myrt 

Rhizophor 

Caryophyll 

Caryoph yllineae 

in Caryophyllacede 

Caryophyll 

Phytolaccineae 

Caryophyll 

Chenopodiinede 

Caryophyll 

Chenopodiineae 

Primul 

Plumbagin 

Tarnaric 

Tamaric 

Primul 

Primul 

Polygon 

Scrophulari 

Euphorbi 

Euphorbi 

Euphorbi 

Malv 

Scrophulariineae 

Malv 

Malv 

Malv 

Malv 

Ros 

Celastr 

Dilleni 

Celastrineae 

Ros 

Rut 

Coriariineae 

Myrt 

Myrtineae 

Myrt 

Lecythidineae 

in Lecythidaceae 

in Lecythidaceae 

Gentian 

Myrt 

Myrt 

Myrt 

M yrtineae 

Myrtineae 

Rhizophorineae 

in Aizoaceae 

in Caryophyllac 

Chenopodi 

Portulacineae 

Chenopodi 

Chenopodiineae 

Chenopodi 

Chenopodiineae 

Primul 

Primulineae 

Primul 

Plumbaginineae 

Viol 

Tamaricineae 

Viol 

Tamaricineae 

Prirnul 

Prirnulineae 

in Primulaceae 

Polygon 

Bignoni 

Euphorbi 

Euphorbi 

Euphorbi 

Malv 

Malv 

Malv 

Malv 

Malv 

Ros 

Rosineae 

Ros 

Cunoniineae 

Ros 

Rosineae 

in Rosaceae 

Rut 

Myrt 

The 

Coriariineae 

Myrtineae 

Lecythidineae 

in Lecythidaceae 

Gentian 

in Lythraceae 

in Lythraceae 

- 

Corn 

Rhirophorineae 

Caryophyll 

in Caryophyllac 

Caryophyll 

Caryophyll 

Caryophyll 

Primul 

Plumbagin 

Tamaric 

Tamaric 

Primul 

Primul 

Polygon 

Scrophulari 

Euphorbi 

Euphorbi? 

Euphorbi 

Malv 

Malv 

Malv 

Malv 

Malv 

Ros 

Celastr? 

Ros 

Ros 

Sapind 

Myrt 

The 

in Lecythidaceae 

in Lecythidaceae 

Gentian 

in Lythraceae 

in Lythraceae 

Rhizophor 

in Aizoaceae 

- 

Centrosperm 

Centrosperm 

Centrosperm 

Primul 

Plumbagin 

Pariet 

Pariet 

Primul 

Primul 

Polygon 

Tubiflor 

Euphorbi 

Terebinth 

Pand 

Malv 

Malv 

Malv 

Malv 

Malv 

Ros 

Terebinth? 

Ros 

in Rosaceae 

Terebinth 

Myrt 

Myrt 

in Myrtaceae 

in Myrtaceae 

Myrt 

Myrt 

Myrt 

Myrt 
NUMBER 58 

TABLE 1 .-continued. 

15 

Famil\ 

Mollugin 

Illecebr 

Basell 

Amaranth 

Chenopodi 

Pr i in u I 

Plumbagin 

Taniaric 

Frankeni 

Myrsin 

Theophrast 

Polygon 

Plantagin 

Euphorbi 

Aextoxic 

Pand 

Sterculi 

Elaeocarp 

Tili 

Malv 

Bombac 

Ros 

Corynocarp 

Crossosoniat 

lieurad 

Coriari 

Myrt 

Lecyt hid 

Barringtoni 

Asteranth 

Dialypetalanth 

Sonnerati 

Punic 

Rhiiophor 

Hutchinson Melchior Stebbinr 

1973 1964 1974 

Cdryophyll 

Polygon 

Chenopodi 

Chenopodi 

Chenopodi 

Pri in u I 

Primul 

Tamaric 

Tamaric 

Myrsin 

Myrsin 

Polygon 

Plantagin 

Euphorbi 

Celastr 

Celastr 

Tili 

in Tiliaceae 

Tili 

Malv 

Tili 

Ros 

Celastr 

Dilleni 

in Rosaceae 

Coriari 

Myrt 

Myrt 

Myrt 

Myrt as 

Rubi 

Myrt 

Mkrt 

Mvrt 

Napoleonaeaceae 

Centrospermae 

in Caryophyllac 

Centrospermae 

Centrospermae 

Centrospermae 

Primul 

Plumbagin 

Viol 

Viol 

Primul 

Tamaricineae 

Tamaricineae 

Primul 

Polygon 

Plantagin 

Gerani 

Euphorbiineae 

Sapind 

Sapindineae 

Celastr 

Celastrineae 

Malv 

Malvineae 

Malv 

Elaeocarpineae 

Malv 

Malvineae 

Malv 

Malvineae 

Malv 

Malvineae 

Ros 

Rosineae 

Celastr 

Celastrineae 

Guttifer 

Dilleniineae 

Ros 

Rosineae 

Sapind 

Coriariineae 

Myrt 

Myrtineae 

Myrt 

Myrtineae 

in Lecythidac 

in Lecythidac 

Myrt 

Myrt 

Myrt 

Myrt 

Myrtineae 

Myrtineae 

Myrtineae 

M yrtineae 

Caryophyll 

- 

Caryophyll 

Caryophyll 

Caryophyll 

Primul 

Plumbagin 

Viol 

Viol 

Primul 

Primul 

Polygon 

Plantagin 

Euphorbi 

Euphorbi 

Euphorbi 

Malv 

Malv 

Malv 

Malv 

Malv 

Ros 

Celastr? 

Dilleni 

Ros 

Sapind? 

Myrt 

Lecythid 

- 

- 

Myrt 

Myrt 

Myrt 

Myrt 

Rouleau Young Benson 

1981 1981 1979 

Caryophyll 

Caryophyll 

Caryophyll 

Caryophyll 

Caryophyll 

Primul 

Plumbagin 

Tamaric 

Tamaric 

Primul 

Primul 

Polygon 

Scrophulari 

Euphorbi 

Celastr 

Euphorbi 

Malv 

Malv 

Malv 

Malv 

Malv 

Ros 

Celastr 

Dilleni 

Ros 

Rut 

Myrt 

Myrt 

Myrt 

Myrt 

Gentian 

Myrt 

Myrt 

Myrt 

Caryophyll 

- 

Chenopodi 

Chenopodi 

Chenopodi 

Primul 

Plumbagin 

Tamaric 

Tamaric 

Primul 

Primul 

Plumbagin 

Bignoni 

Euphorbi 

Euphorbi 

Euphorbi 

Malv 

Malv 

Malv 

Malv 

Malv 

Ros 

Celastr 

Ros 

- 

Rut 

Myrt 

Lecythid 

- 

- 

- 

Myrt 

- 

Rhizophor 

in Aizoaceae 

in Caryophyllaceae 

Caryophyll 

Caryophyll 

Caryophyll 

Primul 

Primul 

Tamaric 

Tamaric 

Primul 

Primul 

Caryophyll 

Plantagin 

Euphorbi 

Euphorbi 

Euphorbi 

Malv 

Malv 

Malv 

Malv 

Malv 

Ros 

Ros 

in Rosaceae 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 
16 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 1 .-continued. 

Fan1 i I y 

Lythr 

Crypteroni 

Olini 

Melastomat 

Onagr 

Trap 

Coinbret 

Penae 

Saxifrag 

Crassul 

Parnassi 

Ereniosyn 

FrdllCO 

Davidsoni 

Hydrange 

Philadelph 

Pterostenion 

Ite 

Bauer 

Bruni 

\'ahli 

Donati 

Tetracarpae 

Escalloni 

Grossulari 

Brunelli 

Cunoni 

Greyi 

Cephalot 

Dioncophyll 

Droser 

Byblid 

Podosteni 

Cronquist Takhtajan Thorne Dahlgren 

Goldberg 1981 1983 1983 1983 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxi frag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Droser 

Droser 

Droser 

Droser 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Ros 

Ros 

in Saxifragaceae 

in Saxifragaceae 

in Saxifragaceae 

Ros 

Ros 

in Hydrangeaceae 

in Grossulari 

in Grossulari 

in Cunoniaceae 

Ros 

in Saxifragaceae 

Campanul 

in Grossulari 

in Grossulari 

Ros 

Ros 

Ros 

Ros 

Ros 

Viol 

N epen t h 

Ros 

Podostem 

Myrt 

Myrtineae 

Myrt 

M yrtineae 

Myrt 

Myrtineae 

Myrt 

Myrtineae 

Myrt 

Myrtineae 

Myrt 

Myrtineae 

Myrt 

M yrtineae 

Myrt 

Myrtineae 

Saxifrag 

Saxifragineae 

Saxifrag 

Saxifragineae 

Saxiphrag 

Saxifragineae 

Saxifrag 

Saxifragineae 

Saxifrag 

Saxifragineae 

Saxifrag 

Cunoniineae 

Saxifrag 

Pittosporineae 

in Hydrangeaceae 

Saxifrag 

Pittosporineae 

in Escalloniaceae 

in Cunoniaceae 

Saxifrag 

Saxifrag 

Campanul 

in Escalloniaceae 

Saxifrag 

Pittosporineae 

Saxifrag 

Saxifragineae 

Saxifrag 

Cunoniineae 

Saxifrag 

Cunoniineae 

Saxifrag 

Saxifragineae 

Saxifrag 

Saxifragineae 

The 

Pittosporineae 

Saxifragineae 

Campanulineae 

Saxifrag 

Saxifrag 

Podostem 

Saxifragineae 

Pittosporineae 

Myrt 

Lythrineae 

Myrt 

Lythrineae 

Myrt 

Lythrineae 

Myrt 

Lythrineae 

Myrt 

Onagrineae 

Myrt 

Lythrineae 

Myrt 

Lythrineae 

Myrt 

Lythrineae 

Ros 

Saxifragineae 

Ros 

Saxifragineae 

Ros 

Saxifragineae 

in Saxifragaceae 

in Saxifragaceae 

Ros 

Cunoniineae 

in Saxifragaceae 

in Saxifragaceae 

in Saxifragaceae 

in Saxifragaceae 

Ros 

Cunoniineae 

Pit tospor 

Bruniineae 

in Saxifragaceae 

in Stylidiaceae 

in Saxifragaceae 

in Saxifragaceae 

in Saxifragaceae 

Ros 

Ros 

Ros 

Ros 

The 

Ros 

Pittospor 

Ros 

Cunoniineae 

Cunoniineae 

Saxifragineae 

Saxifragineae 

Scytopetalineae 

Saxifragineae 

Pittosporineae 

Saxifragineae 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Saxifrag 

Saxifrag 

Droser 

Corn? 

in Saxifragaceae 

Cunoni 

Corn 

- 

Corn? 

Saxifrag 

Cunoni 

Cunoni 

in Saxifragaceae 

in Stylidiaceae 

Corn? 

Corn 

Saxifrag 

Cunoni 

Cunoni 

Saxifrag 

Saxifrag 

The 

Droser 

Pittospor? 

Podostem 

Emberger 

1960 

Myrt 

Myrt 

Thymelae 

Myrt 

Myrt 

Myrt 

Myrt 

Thymelae 

Ros 

Ros 

in Saxifragaceae 

- 

in Saxifragaceae 

in Cunoniaceae 

in Saxifragaceae 

in Saxifragaceae 

in Saxifragaceae 

in Saxifragaceae 

in Saxifragaceae 

Ros 

- 

in Stylidiaceae 

in Saxifragaceae 

in Saxifragaceae 

in Saxifragaceae 

Ros 

Ros 

Ros 

Sarraceni 

Pariet 

Pariet 

Ros 

Ros 
NUMBER 58 17 

TABLE 1 .-continued. 

Hutchinson Melchior Stebbins Rouleau Young Benson 

1973 1964 1974 1981 1981 1979 Famil) 

Lythr 

Crypteroni 

Olini 

Melastomat 

Onagr 

Trap 

Combret 

Penile 

Saxifrag 

CrdSSUl 

Parnassi 

Erenios)n 

Franco 

Davidsoni 

Hydrange 

Phihdelph 

Pierostemon 

Ite 

Bauer 

Bruni 

Vahli 

Donati 

Tetracarpae 

Escalloni 

Grossulari 

Brunelli 

Cunoni 

Grevi 

Cephalot 

Dioncophyll 

Droser 

Byblid 

Podostem 

Myrt 

Cunoni 

Cunoni 

Myrt 

Onagr 

Onagr 

Myrt 

Thymelae 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

in Cunoniaceae 

Cunoni 

Cunoni 

Cunoni 

in Escalloniac 

Cunoni 

Hamamelid 

Saxifrag 

Saxifrag 

in Escalloniac 

Cunoni 

Cunoni 

Dilleni 

Cunoni 

Cunoni 

Saxifrag 

in Flacourtiac 

Sarraceni 

Pittospor 

Podostem 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Thymelae 

Ros 

Ros 

in Saxifragac 

in Saxifragac 

in Saxifragac 

Ros 

in Saxifragac 

in Saxifragac 

in Saxifragac 

in Saxifragac 

in Saxifragac 

Ros 

in Saxifragac 

in Stvlidiaceae 

Myrtineae 

Myrtineae 

Myrtineae 

Myrtineae 

Myrtineae 

Myrtineae 

Myrtineae 

Saxifragineae 

Saxifragineae 

Saxifragineae 

Saxifragineae 

in Saxifragac 

in Saxifragac 

in Saxifragac 

Ros 

Ros 

in Melianthac 

Ros 

Guttifer 

Sarraceni 

Saxifragineae 

Saxifragineae 

Saxifragineae 

Ochnineae 

Ros 

Podostem 

Saxifragineae 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Ros 

Ros 

Ros 

Ros 

Ros 

in Grossulariac 

Ros 

- 

Ros 

Sapind 

Ros 

Viol? 

Ros? 

Ros 

Podostem 

Myrt 

Saxifrag 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Campanul 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Saxifrag 

Sapind 

Sax i frag 

The 

Nepenth 

Saxifrag 

Podostem 

Myrt 

- 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Saxifrag 

Saxifrag 

Saxifrag 

Hydrange 

Saxifrag 

Cunoni 

Hydrange 

Hydrange 

Saxifrag 

Saxifrag 

Saxifrag 

Bruni 

Saxifrag 

- 

Saxifrag 

Hydrange 

- 

Cunoni 

Cunoni 

Saxifrag 

Saxifrag 

Scytopetal 

Saxifrag 

Pittospor 

Podostem 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Myrt 

Ros 

Ros 

in Saxifragaceae 

in Saxifragaceae 

in Saxifragaceae 

in Saxifragaceae 

in Saxifragaceae 

in Saxifragaceae 

in Saxifragaceae 

Ros 

- 

in Saxifragaceae 

in Saxifragaceae 

Ros 

Ros 

Ros 

Ros 

Ros 

Podostem 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 1 .-continued. 

Family 

Hldropstachy 

Begoni 

Datisc 

Arali 

Api 

Davidi 

N yss 

Carry 

Corn 

Caprifoli 

Valerian 

Adox 

Dipwc 

Rut 

Cneor 

Siniaroub 

Burser 

Anacdrd 

Meli 

Akdni 

Sapind 

Melianth 

Hippocastan 

Staphyle 

Acer 

Sabi 

Gerani 

Viviani 

Limnanth 

Oxalid 

Connar 

Tropaeol 

Ba Isa ni i n 

Stackhousi 

Cronquist Takhtajan Thorne 

Goldbere 1981 1983 1983 

Dahlgren Emberger 

1983 1960 

Droser 

Begoni 

Begoni 

Api 

Api 

Corn 

Corn 

Corn 

Corn 

Dipsac 

Dipsac 

Dipsac 

Dipsac 

Rut 

Rut 

Rut 

Rut 

Rut 

Rut 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Gerani 

Gerani 

Gerani 

Gerani 

Gerani 

Gerani 

Gerani 

Gerani 

Callitrich 

Viol 

Viol 

Api 

Api 

in Nyssaceae 

Corn 

Corn 

Corn 

Dipsac 

Dipsac 

Dipsac 

Dipsac 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Ranuncul 

Gerani 

in Geraniaceae 

Gerani 

Gerani 

Ros 

Gerani 

Gerani 

Celastr 

Scrophulari 

Begoni 

Begoni 

Arali 

Arali 

Corn 

Corn 

Scrophulariineae 

Corn 

Corn 

Dipsac 

Dipsac 

Dipsac 

Dipsac 

Rut 

Rut 

Rut 

Rut 

Rut 

Rut 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Gerani 

Geraniineae 

in Geraniaceae 

Rutineae 

Rutineae 

Rutineae 

Rut i neae 

Rutineae 

Rutineae 

Gerani 

Gerani 

Connar 

Gerani 

Gerani 

Celastr 

Lirnnanthineae 

Geraniineae 

Balsaminineae 

Balsamininede 

Celastrineae 

Pittospor 

Viol 

Viol 

Arali 

in Araliaceae 

in Nyssaceae 

Corn 

Cornineae 

Corn 

Cornineae 

Corn 

Cornineae 

Dipsac 

Dipsac 

Dipsac 

Dipsac 

Rut 

Rut 

Rut 

Rut 

Rut 

Rut 

Rut 

Rut 

Rut 

Rut 

Ros 

Rut 

Rut 

Gerani 

Gerani 

Gerani 

Gerani 

Rut 

Gerani 

Gerani 

Celastr 

Bruniineae 

Begoniineae 

Begoniineae 

Rutineae 

Rutineae 

Rutineae 

Rutineae 

Rutineae 

Rutineae 

Sapindineae 

Sapindineae 

Sapindineae 

Sapindineae 

Cunoniineae 

Sapindineae 

Sapindineae 

Geraniineae 

Geraniineae 

Geraniineae 

Geraniineae 

Fabineae 

Geraniineae 

Geraniineae 

Hydrostachy 

Cucurbit 

Cucurbit 

Arali 

Arali 

Corn 

Corn 

Corn 

Corn 

Corn 

Dipsac 

Corn 

Dipsac 

Rut 

Rut 

Rut 

Rut 

Sapind 

Rut 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Cerani 

Gerani 

Tropaeol 

Gerani 

Sapind 

Tropaeol 

Balsamin 

Celastr 

Ros 

Pariet 

Pariet 

Umbel1 

Umbell 

in Nyssaceae 

Umbell 

Carry 

Umbel1 

Rubi 

Rubi 

Rubi 

Rubi 

Terebinth 

Cneor 

Terebinth 

Terebinth 

Terebinth 

Terebinth 

Terebinth 

Terebinth 

Terebin t h? 

Terebinth 

Celastr 

Terebinth 

Terebinth? 

Gerani 

in Geraniaceae 

Gerani 

Gerani 

Ros 

Gerani 

Terebinth? 

Celastr 
NUMBER 58 

TABLE 1 .-continued. 

19 

Family 

Hydrostachy 

Begoni 

Datisc 

Arali 

Api 

Davidi 

Nyss 

Carry 

Corn 

Caprifoli 

Valerian 

Adox 

Dipsac 

Rut 

Cneor 

Siniaroub 

Burser 

Anacard 

Meli 

Akani 

Sapind 

Melian t h 

Hippocastan 

Staphyle 

Acer 

Sabi 

Gerani 

\'iviani 

Limnanth 

Oxalid 

Connar 

Tropaeol 

Balsamin 

Stackhousi 

Hutchinson Melchior Stebbins Rouleau Young Benson 

1973 1964 1974 1981 1981 1979 

Podosteni 

Cucurbit 

Cucurbit 

Arali 

Unibell 

in Kyssaceae 

Arali 

Arali 

Arali 

Arali 

Valerian 

Saxifrag 

Valerian 

Rut 

Celastr 

Rut 

Rut 

Sapind 

Meli 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Gerani 

Pittospor 

Gerani 

Gerani 

Dilleni 

Gerani 

Gerani 

Celastr 

Hydrostachy 

Viol 

Viol 

Urnbell 

Urnbell 

Urnbell 

Urnbell 

Urnbell 

Begoniineae 

Begoniineae 

Urnbell 

Dipsac 

Dipsac 

Dipsac 

Dipsac 

Rut 

Rutineae 

Rut 

Rutineae 

Rut 

Rutineae 

Rut 

Rutineae 

Sapind 

Anacardiineae 

Rut 

Rutineae 

Rut 

Rutineae 

Sapind 

Sapindineae 

Sapind 

Sapindineae 

Sapind 

Sapindineae 

Celastr 

Celastrineae 

Sapind 

Sapindineae 

Sapind 

Sapindineae 

Gerani 

Geraniineae 

in Geraniaceae 

Gerani 

Gerani 

Ros 

Gerani 

Sapind 

Celastr 

Lirnnanthineae 

Geraniineae 

Legurninosineae 

Geraniineae 

Balsamineae 

Celastrineae 

Scrophulari 

Viol 

Viol 

Urnbell 

Urnbell 

Corn 

Corn 

Corn 

Corn 

Dipsac 

Dipsac 

Dipsac 

Dipsac 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

- 

Gerani 

- 

Gerani 

Gerani 

Sapind 

Gerani 

Gerani 

Celastr 

Scrophulari 

Begoni 

Begoni 

Api 

Api 

Corn 

Corn 

Corn 

Corn 

Dipsac 

Dipsac 

Dipsac 

Dipsac 

Rut 

Rut 

Rut 

Rut 

Rut 

Rut 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Gerani 

Gerani 

Gerani 

Gerani 

Connar 

Gerani 

Gerani 

Celastr 

Hydrostachy 

Viol 

Viol 

Arali 

Arali 

Corn 

Corn 

Corn 

Corn 

Dipsac 

Dipsac 

Dipsac 

Dipsac 

Rut 

Rut 

Rut 

Rut 

Rut 

Rut 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Gerani 

Tropaeol 

Gerani 

Fa b 

Tropaeol 

Gerani 

Celastr 

Loas 

Loas 

Urnbell 

Urnbell 

Corn 

- 

Corn 

Corn 

Rubi 

Rubi 

Rubi 

Rubi 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Sapind 

Gerani 

Rutineae 

Rutineae 

Rutineae 

Rutineae 

Rutineae 

Rutineae 

Sapindineae 

Spindineae 

Sapindineae 

Sapindineae 

Sapindineae 

Sapindineae 

Sapindineae 

Gerani 

Gerani 

Ros 

Gerani 

Gerani 

Sapind 

Celastrineae 
20 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 1 .-continued 

Family 

Lin 

Erythroxyl 

Zygophyll 

Balanit 

Malpighi 

Polygal 

Krameri 

Trigoni 

t'ochysi 

Ole 

Salvador 

Logani 

Plocospermat 

Apocyn 

Asclepiad 

Convolvul 

Cuscut 

Rubi 

Colunielli 

Gentian 

Menyanth 

Olac 

Aptandr 

Octokneni 

Opili 

Medusandr 

Cardiopterid 

Santal 

Loran ti1 

Misodendr 

Grubbi 

Geissoloniat 

Gon\st,l 

Thvnielae 

Cronquist Takhtajan 

Goldbera 1981 1983 

Thorne 

1983 

Dahlgren Emberger 

1983 1960 

Gerani 

Gerani 

Gerani 

Gerani 

Gerani 

Polygal 

Po I yga I 

Polygal 

Polygal 

Ole 

Ole 

Gentian 

Gentian 

Gentian 

Gentian 

Gentian 

Gentian 

Gentian 

Gentian 

Gentian 

Gentian 

Santal 

Santal 

Santal 

Santal 

Santal 

Santal 

Santal 

Santal 

Santal 

Santal 

Thynielae 

Thynielae 

Thynielae 

Lin 

Lin 

Sapind 

in Zygophyllac 

Polygal 

Polygal 

Polygal 

POlygdl 

Polygal 

Scrophulari 

Celastr 

Gentian 

in Apocynaceae 

Gentian 

Gentian 

Solan 

Solan 

Rubi 

Ros 

Gentian 

Solan 

Santal 

in Olacaceae 

in Olacaceae 

Santal 

Santal 

Celastr 

Santal 

Santal 

Santal 

Eric 

Celastr 

in Thynielaeaceae 

Myrt 

Gerani 

&rani 

Rut 

Rut 

Polygal 

Polygal 

Polygal 

Polygal 

Polygal 

Ole 

Linineae 

Linineae 

Rut i neae 

Rutineae 

Celastr 

Gentian 

in Loganiaceae? 

Gentian 

Gentian 

Celastrineae 

Polenioni 

Polemoni 

Gentian 

Saxifrag 

Gentian 

Convolvulineae 

Convolvulineae 

Pi ttosporineae 

Gentian 

Santal 

in Olacaceae 

in Olacaceae 

Santal 

Celastr 

Celastr 

Santal 

Santal 

Santal 

Eric 

Santalineae 

Santalineae 

lcacinineae 

Icacinineae 

Santalineae 

Loranthineae 

Santalineae 

Celastr 

Celastrineae 

in Thymelaeaceae 

Thymelae 

Gerani 

Linineae 

Gerani 

Linineae 

Gerani 

Linineae 

in Rutaceae 

Gerani 

Gerani 

Gerani 

Gerani 

Gerani 

Ole 

Polygalineae 

Pol ygalineae 

Polygalineae 

Pol ygalineae 

Polygalineae 

Ole 

Gentian 

in Loganiaceae 

Gentian 

in Apocynaceae 

Solan 

in Convolvulaceae 

Solanineae 

Gentian 

in Saxifragaceae 

Gentian 

Gentian 

Santal 

- in Olacaceae 

in Olacaceae 

Santal 

The 

Santal 

Santal 

Theineae 

Santal 

Pittospor 

Pittospor 

in Thymelaeaceae 

Euphorbi 

Bruniineae 

Bruniineae 

Gerani 

Gerani 

Gerani 

Gerani 

Polygal 

Polygal 

Polygal? 

Polygal 

Polygal 

Ole 

Salvador 

Gentian 

- 

Gentian 

Gentian 

Solan 

Solan 

Gentian 

Corn 

Gentian 

Gentian 

Santal 

- 

in Olacaceae 

Santal 

- 

Corn? 

or Celastr? 

Santal 

Santal 

Santal 

Cunoni 

Geissolomat 

Thymelae or 

in Thymelaeaceae 

Gerani 

Gerani 

Gerani 

in Zygophyllaceae 

Gerani 

Terebinth 

in Caesalpiniaceae 

Terebin th 

Terebinth 

Ligustr 

Celastr 

Rubi 

in Loganiaceae? 

Contort 

Contort 

Tubiflor 

in Convolvulaceae 

Rubi 

Tubiflor? 

Contort 

Contort 

Olac 

in Olacaceae 

Santal 

Olac 

Olac? 

Celastr 

Santal 

Santal 

Santal 

Santal 

Thymelae 

Malv 

Thymelae Thymelae 
NUMBER 58 

TABLE 1 .-continued. 

21 

Faniil) 

I.in 

Erythi-oxyl 

%ygophy11 

Balanit 

Malpighi 

POlygdl 

Krameri 

Trigoni 

Vochysi 

Ole 

Sal~ador 

Logani 

Plocospermat 

Apocyn 

Asclepiad 

Convolvul 

c u scu t 

Rubi 

<:olunielli 

Gentian 

Men! an th 

Olac 

Aptandr 

Octoknem 

Opili 

Medusandr 

Cardiopterid 

Santal 

Loranrh 

Misodendr 

Grubbi 

(;eissoloniat 

(;onyatyl 

Thynielae 

Hutchinson Melchior Stebbins Rouleau Young Benson 

1973 1964 1974 1981 1981 1979 

Malpighi 

Malpighi 

Malpighi 

Malpighi 

hlalpighi 

Polyp1 

Polygal 

POlygdI 

Polygal 

Logan i 

Celastr 

Logani 

Apocp 

Apocyn 

Apocyn 

Solan 

Polemoni 

Rubi 

Person 

Gentian 

(;en tian 

Olac 

Olac 

Olac 

Olac 

Olac 

Celastr 

Santal 

Santal 

Santal 

Gerani 

Geraniineae 

Gerani 

Geraniineae 

Gerani 

Geraniineae 

in Zygophyllac 

Rut 

Rut 

Ros 

Rut 

Rut 

Ole 

Celastr 

Gentian 

Malpighiineae 

Polygalineae 

Leguminosineae 

Malpighiineae 

Malpighiineae 

Celastrineae 

- 

Ge ti t ian 

Gentian 

Tubiflorae 

in Convolvulac 

Gentian 

Tubiflorae 

Gentian 

Convolvulineae 

Solanineae 

Gentian 

Santal 

in Olacaceae 

in Olacaceae 

Santal 

Medusandr 

Celastr 

Santal 

Icacinineae 

Santal 

Santal 

Santal 

Thymelae 

Thynielae 

Thynielae 

Santal 

Thynielae 

in Th\melaeac 

Thymelae 

Gerani 

Gerani 

Sapind? 

- 

Polygal? 

Polygal 

Polygal 

Polygal 

POlygdl 

Gentian? 

Celastr 

Gentian 

- 

Gentian 

in Apocynaceae 

Polemoni? 

Polemoni? 

Rubi 

Scrophulari? 

Gentian 

Polemoni? 

Santal 

- 

- 

Santal 

Santal? 

Celastr? 

Santal 

Santal 

Santal 

Eric? 

Celastr 

- 

Myrt 

Gerani 

Gerani 

Gerani 

Gerani 

Gerani 

Polygal 

Polygal 

Polygal 

Polygal 

Ole 

Celastr 

Gentian 

Gentian 

Gentian 

Gentian 

Polemoni 

Polemoni 

Gentian 

Scrophulari 

Gentian 

Gentian 

Santal 

Santal 

Santal 

Santal 

Santal 

Santal 

Santal 

Santal 

Santal 

Eric 

Celastr 

- 

Thymelae 

Lin 

Lin 

Lin 

- 

Polygal 

Polygal 

Polygal 

Polygal 

Polygal 

Ole 

Ole 

Gentian 

- 

Gentian 

Gentian 

Solan 

in Convolvulac 

Gentian 

Hydrange 

Gentian 

Gentian 

Santal 

- 

- 

- 

Celastr 

The 

Santal 

Santal 

Santal 

Pittospor 

Myrothanin 

- 

Euphorbi 

Gerani 

Gerani 

Cerani 

Gerani 

Polygal 

Polygal 

Polygal 

Polygal 

Gentian 

Sapind 

Gentian 

Oleineae 

Celastrineae 

Gentianineae 

- 

Gentian 

Apocinineae 

Gentian 

Apocynineae 

Polemoni 

Polemoniineae 

in Convolvulaceae 

Rubi 

Gentian 

Gentianineae 

in Gentianaceae 

Santal 

Santal 

Santal 

Santal 

Santal 

Santal 

Myrt 

Myrt 
22 

Family 

Polenioni 

Fouquieri 

Hydrophyll 

Boragin 

Solan 

Nolan 

Scrophulari 

Buddlej 

Globulari 

Lentibulari 

Orobanch 

Acanth 

Bignoni 

Gesneri 

Pedali 

Myopor 

Verben 

Pliryin 

Lanii 

Canipanul 

Goodeni 

Brunoni 

Calycer 

Stylidi 

Aster 

Tremandr 

Elaeagn 

Pittorpor 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 1 .-continued. 

Goldberg 

Polenioni 

Polemoni 

Polenioni 

Polenioni 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scroph u lari 

Scrophulari 

Scrophulari 

Scrophulari 

Lanii 

Lami 

Lanii 

Lanii 

Canipanul 

Campan u I 

Cam pan uI 

Canipanul 

Canipanul 

Aster 

Cronquist 

1981 

Solan 

Viol 

Solan 

Lanii 

Solan 

Solan 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Lami 

in Verbenaceae 

Lami 

Campanul 

Campanul 

Canipanul 

Calvcer 

Cainpanul 

Aster 

Polvgal 

Prow 

Ros 

Takhtajan Thorne 

1983 1983 

Dahlgren Emberger 

1983 1960 

Polenioni 

Tamaric 

Polemoniineae 

Polemoni 

Boraginineae 

Polemoni 

Boraginineae 

Scrophulari 

Solanineae 

in Solanaceae 

Scrophulari 

Scrophulari 

in Scrophulariac 

Scroph u lari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Lanii 

Scrophulariineae 

Scroph ulariineae 

Scrophulariineae 

Scrophulariineae 

Scrophulariineae 

Scrophulariineae 

Scrophulariineae 

Scrophulariineae 

Scroph ulariineae 

in k'erbenaceae 

Lami 

Campanul 

Canipanulineae 

Campanul 

Goodeniineae 

in Goodeniaceae 

Calycer 

Canipanul 

Aster 

Campanulineae 

POlpgdl 

Elaedgn 

Saxifrag 

Pittosporineae 

Solan 

Polemoniineae 

Solan 

Fouquierineae 

Solan 

Boraginineae 

Solan 

Boraginineae 

Solan 

Solanineae 

in Solanaceae 

Bignoni 

Gentian 

in Scrophulariac 

Bignoni 

in Scrophulariac 

Bignoni 

Bignoni 

Bignoni 

Bignoni 

Bignoni 

Lami 

in Verbenaceae 

Lanii 

Canipanul 

Canipanul 

in Goodeniaceae 

Dipsac 

Ros 

Aster 

Pittospor 

Rhamn 

Pittospor 

Saxifragineae 

Pittosporineae 

Pittosporineae 

Solan 

Fouquieri 

Boragin 

Boragin 

Solan 

in Solanaceae 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

in Scrophulariac 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Lami 

in Verbenaceae 

Lami 

Campanul 

Goodeni 

in Goodeniaceae 

Dipsac 

Corn 

Aster 

Pittospor? 

Elaeagn 

Pittospor 

Tubiflor 

Pariet 

Tubiflor 

Tubiflor 

Tubiflor 

Tubiflor 

Tubiflor 

Tubiflor 

Tubiflor 

Tubiflor 

Tubiflor 

Tubiflor 

Tubiflor 

Tubiflor 

Tubiflor 

Tubiflor 

Tubiflor 

Tubiflor 

Tubiflor 

Synanther 

Synanther 

Synanther 

Rubi 

Svnanther 

Synanther 

Terebinth? 

Thvmelae 

Ros 

one or more convergent characters or character 

states in common will differ from one another in 

most respects. I believe convergence is hardly 

ever prevalent enough to mask relationships. 

If a family has more than one state of a character, the state considered primitive for the particular taxon rather than the derived one should 

be considered in attempting to determine the 
NUMBER 58 

Rhariin 

Pittospor 

TABLE 1 .-continued. 

23 

Farnil) 

-~ 

Polenioni 

Fouquieri 

Hydrophyll 

Boragin 

Solan 

Nolan 

Scrophulari 

Buddlej 

Globulari 

1.entibulari 

Orohanch 

Acaiith 

Bignoni 

Gesneri 

Pedali 

M\opol- 

\’erben 

Phr) 111 

L.;I ni i 

(:dllllYdlllll 

Goodeii i 

Brunoni 

Calvcer 

St)lidi 

Aster 

Tremandr 

Elaeagn 

Pittospor 

Melchior 

1964 

Tubiflorae 

Tubiflorae 

Tuhiflorae 

Tuhiflorae 

Tubiflorae 

Tubiflorae 

Tuhiflorae 

Tubiflorae 

Tuhiflorae 

Tubiflorae 

Tubiflorae 

Tubiflorae 

Tubiflorae 

Tuhiflorae 

Tubiflorae 

Tubiflorae 

Tubiflorae 

Tubiflorae 

Tubiflorae 

Campan uI 

Convolvulineae 

Convolvulineae 

Boraginineae 

Boraginineae 

Solanineae 

Solanineae 

Solanineae 

Solanineae 

Solanineae 

Solanineae 

Solanineae 

Solanineae 

Solanineae 

Solanineae 

Solanineae 

M yoporineae 

Verbenineae 

Phryniineae 

Verhenineae 

Canipan u I 

Canipan uI 

Carnpanul 

Campanul 

Canipanul 

Rut 

Thymelae 

Ros 

Pol ygalineae 

Saxifragineae 

Stebbins Rouleau Young Benson 

1974 1981 1981 1979 

Polemoni 

Viol 

Polemoni 

Polernoni? 

Polernoni? 

Polenioni? 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Lami 

Lami 

Lami 

Carnpanul 

Carnpanul 

Campanul 

Dipsac 

Carnpanul 

Aster 

Polygal 

Prote 

Ros 

Polemoni 

Tamaric 

Boragin 

Boragin 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scroph u lari 

Lami 

Lami 

Larni 

Campan u I 

Goodeni 

Goodeni 

Calycer 

Campan u I 

Aster 

Polygal 

Elaeagn 

Saxifrag 

Solan 

Solan 

Boragin 

Boragin 

Solan 

- 

Bignoni 

Gentian 

- 

Bignoni 

in Scrophulariac 

Bignoni 

Bignoni 

Bignoni 

Bignoni 

Bignoni 

Lami 

- 

Lanii 

Campan u I 

Cam pa nu I 

Campanul 

Hydrange 

Aster 

Polygal 

Rhamn 

Pi ttospor 

Dipsac 

Polemoni 

Fouquieri 

Polemoni 

Polemoni 

Polemoni 

Polenioni 

Scrophulari 

in Loganiaceae 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Scrophulari 

Polemoniineae 

Boraginineae 

Boraginineae 

Polemoniineae 

Polemoniineae 

Scrophulari 

Scrophulari 

Scrophulari 

Larni 

Lami 

Lami 

Canipanul 

Campanul 

Canipanul 

Canipan u I 

Canipanul 

Aster 

Elaeagn 

Ros 

family’s ancestor. A descendant has at least one 

more derived character or character state than 

its ancestor. Genera that are intermediate between families are particularly significant in a 

phylogenetic study. They are the knots that tie 

the families together. 

I have constructed a dendrogram suggesting 

relationships of the orders (page 27). Most of my 
24 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 5.-Evolutionary trends in dicotyledons (reversibility: n = not; r = rarely; c = commonly). 

Character 

climate of origin 

plant 

indument 

spines 

tendrils 

latex 

xylem Lessel perforation 

plates 

xylem vessel elements 

and sieve tube elements 

intraxylary and interphloem companion cells 

leaves 

xylary phloem 

leaf margin 

leaf venation 

leaf buds 

stipules 

stomates 

leaves and flowers 

inflorescence 

flowers 

pollinated b) 

pollinators 

in\,olucre 

receptacle 

perianth 

Primitive state Derived state 

Reversibilitv 

warm temperate 

mesoph yte; 

dioecious; 

autotrophic 

absent or sparse; 

simple 

absent 

absent 

absent 

absent; 

scalariform 

long, narrow, angular in 

cross-section; 

end plate slanting; 

pores small 

absent 

absent 

alternate; 

more than 2 ranks; 

thin; 

large 

entire 

open 

small, naked 

absent 

anomocytic 

formed and functional in 

the same growing season 

racemose; 

raceme; 

spike or umbel; 

open cyme 

soli tar);; 

terminal; 

unisexual; 

odorless; 

green, white, or yellow; 

actinomorphic 

wind: 

various pollinators 

Coleoptera; 

short-tongued 

absent 

elongate or convex; 

flat 

absent; 

other 

xerophyte; 

monoecious or bisexual; 

saprophytic or parasitic 

copious; 

stellate or glandular 

present 

present 

present 

present; 

simple 

short, wide, round in 

end plate transverse; 

pores large 

present 

cross-sec tion; 

present 

opposite or whorled; 

distichous; 

thick and sclerophyllous; 

small relative to size of 

plant 

toothed 

closed 

large, perulate 

present 

other 

formed in 1 season and 

functional in the next 

cymose; 

corymb or spike; 

head; 

fascicle 

in inflorescence; 

axillary; 

bisexual; 

scented; 

other colors; 

zygomorphic 

insects, birds, or mamparticular pollinators 

Hymenoptera, Diptera, 

or Lepidoptera; 

long-tongued 

present 

flat; 

concave 

monochlam ydeous; 

mals; 

r 

r 

r-c 

n 

r 

r 

r 

r 

n 

n 

n 

n 

n 

r 

n 

r 

r 

r 

r 

C 

C 

n 

n 

r 

n 

n 

n 

n 

n 

n 

C 

C 

C 

C 

C 

n 

r 

r 

r 

n 

n 

n 

n 

r 
NUMBER 58 25 

TABLE 2.-continued. 

ReversiPrimitive state Derived state bilitv Character 

perianth spur 

perianth parts 

corona 

perianth vasculature 

perianth parts, stamens 

and carpels 

stamens and carpels 

nectaries 

stamens 

staminodes 

anthers 

connective 

microsporangia 

pollen grains 

cells in pollen grains at 

carpels 

ovark 

time of dispersal 

style(s) 

stigma(s) 

monoc hlam ydeous; 

homoiochlamydeous 

absent 

imbricate 

absent 

evident 

indefinite in number; 

dichlamydeous; 

heterochlam ydeous 

present 

valvate 

present 

absent or not evident 

definite in number: 

spirally arranged; 

separate 

pleiomerous; 

2 or more whorls; 

isomerous 

absent 

laminar, not differentiated into anther and 

filament; 

3 trace; 

longer than pistil; 

equal in length; 

inserted on receptacle 

absent 

separate: 

adnate or basifixed; 

oblong; 

dehiscent by slits; 

extrorse or latrorse 

abundant 

4 per anther; 

dorsal or dorsal and ventral 

nionosulcate; 

smooth; 

not sticky; 

free 

2 

open 

number of locules correspond to number of 

carpels: 

superior 

absent; 

separate; 

terminal 

decurrent along the ventral suture of the ovary 

or ventral surface of 

the style; 

whorled: 

connate or adnate 

isomerous; 

1 whorl; 

oligomerous 

present 

differentiated into anther 

and filament: 

1 trace; 

equal to or shorter than 

pistil; 

anisostemonous; 

inserted on perianth 

present 

connate; 

dorsifixed: 

about as long as wide; 

poricidal or valvate: 

introrse 

sparse or absent 

2 or 1 per anther; 

ventral 

colpi 3 or more; 

sculptured; 

sticky; 

united in tetrads, small 

masses, or pollinia 

3 

closed 

unilocular in 2-m carpellate pistil: 

semi-inferior to inferior 

present; 

connate; 

gynobasic 

apical, or style stigmatic 

all around: 

r 

r 

r 

n 

n 

n 

n 

n 

n 

n 

n 

n 

r 

n 

n 

r 

r 

n 

r 

r 

r 

r 

n 

r 

n 

n 

n 

n 

r 

r 

n 

n 

n 

r 
26 

Character 

placentas in 2-m carpellate pistil 

ovules 

fruit 

dehiscence of capsule 

seed 

sarcotesta or aril 

enibrvo 

cotyledons 

radicle 

endosperni 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 2.-continued. 

Revers i - Primitive state Derived state bility 

- 

separate 

axile; 

connate 

parietal; 

n 

r 

axile; apical, basal, or free-cen- n 

not enlarged enlarged and fleshy r 

few; 00, or 1 when carpels C 

crassinucellar; tenuinucellar; n 

bitegmic unitegmic n 

follicle other types n 

ventricidal or septicidal loculicidal n 

medi u m-si zed small or large r 

absent present r 

small; large; r 

straight curved or variously n 

small; large; r 

as wide as radicle wider than radicle r 

long; short; n 

not invested invested n 

copious scanty or absent n 

tral; 

more than 1; 

folded 

study has been of relationships of the families. I 

intend to consider the relationships of the orders 

more fully in a future publication. 

Other families accepted by the above authors, 

are as follows. 

Cronquist, 198 1. Barclayaceae in Nymphaeales; Cecropiaceae in Urticales; Oncothecaceae in Theales; Huaceae in Violales; Anisophylleaceae, Alseuosmiaceae, Chrysobalanaceae, 

Surianaceae, and Rhabdodendraceae in Rosales; 

Dipentodontaceae, Viscaceae, and Eremolepidaceae in Santalales; Mitrostemaceae in Rafflesiales; Hippocrateaceae in Celastrales; Hugoniaceae in Linales; Xanthophyllaceae in Polygalales; Julianiaceae in Sapindales; Retziaceae and 

Saccifoliaceae in Gentianales; Duckeodendraceae 

in Solanales; Medonciaceae in Scrophulariales; 

Pentaphragmataceae and Sphenocleaceae in 

Campanulales; Tepuianthaceae in Celastrales or 

Sapindales. 

Takhtajan, 1983. Glaucidiaceae in Ranunculales; Cecropiaceae in Urticales; Hectorellaceae 

in Caryophyllales; Sauvagesiaceae, Oncothecaceae, and Asteropeiaceae in Theales; Dipentodontaceae in Violales; Koeberliniaceae in Capparales; Huaceae in Malvales; Montiniaceae, Roridulaceae, and Alseuosmiaceae in Saxifragales; 

Chrysobalanaceae in Rosales; Rhabdodendraceae, Nitrariaceae, Kirkiaceae, Ptaeroxylaceae, 

Julianiaceae, and Podoaceae in Rutales; Stylobasiaceae, Emblingiaceae, and Physenaceae in 

Sapindales; Aucubaceae, Melanophyllaceae, Griseliniaceae, Toricelliaceae, and Helwingiaceae in 

Cornales; Sphenostemonaceae, Phellinaceae, and 

Lophopyxidaceae in Celastrales; Viscaceae in 

Santalales; Cynomoriaceae in Balanophorales; 

Morinaceae in Dipsacales; Duckeodendraceae, 

Retziaceae, and Martyniaceae in Scrophulariales. 

Thorne, 1983. Glaucidiaceae in Paeoniales; 

Oncothecaceae, Phellinaceae, and Sphenostemonaceae in Theales; Halophytaceae in Chenopodiales; Ledocarpaceae in Geraniales; Lophopyxidaceae in Celastrales; Eremolepidaceae and 

Viscaceae in Santalales; Cynomoriaceae in Balanophorales; Dipentodontaceae in Violales; Huaceae and Plagiopteraceae in Malvales; Tepuian- 
NUMBER 58 27 

ASTERALES 

fl CAMPANULALES SCROPHULARIALES 

THYMELAEALES ’? POLEMONIALES 

SANTALALES? GENTIANALES 

POLYGALALES 

RHAMNALES 

SAPINDALES 

BALANOPHORAL 

RAFFLESIALES PLANTAGINALES 

POLYGONALES 

CISTALES PRIMULALES 

CARYOPHYLLALES 

STYRACALES PAPAVERALES 

BALANOPALES DILLENIALES 

PROTEALES 

NYMPHAEALES 

PIPERALES 

ARISTOLOCHIALES : 

LAURALES 

=I NEPENTHALES 

SARRACENIALES 

HALORAGALES 

MAGNOLIALES RANUNCULALES 

HAMAMELIDALES 

TROCHODENDRALES 

.ES 

Suggested phylogenetic relationships 

of the orders of dicotyledons. 
28 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

thaceae, Ptaeroxylaceae, and Surianaceae in Rutales; Chrysobalanaceae in Rosales; Roridulaceae 

in Pittosporales; Rhynchocalycaceae and Alzateaceae in Myrtales; Martyniaceae in Bignoniales; 

Pentaphragmataceae in Campanulales; Aucubaceae in Cornales; Helwingiaceae and Torricelliaceae in Araliales. 

Dahlgren, 1983. Kingdoniaceae, and Glaucidiaceae(?) in Ranunculales; Halophytaceae and 

Hectorellaceae in Caryophyllales; Limoniaceae 

in Plumbaginales; Plagiopteraceae and Huaceae 

in Malvales; Cecropiaceae in Urticales; Dipentodontaceae in Violales; Oncothecaceae in Theales; 

Aegicerataceae and Coridaceae in Primulales; 

Corylaceae in Fagales; Brexiaceae in Saxifragales; Lepuropetalaceae in Droserales; Malaceae, 

Amygdalaceae, Surianaceae(?), and Rhabdodendraceae(?) in Rosales; Psiloxylaceae, Heteropyxidaceae, Memecylaceae, Rhynchocalycaceae, and 

Alzateaceae in Myrtales; Chrysobalanaceae in 

Chrysobalanales; Podoaceae, Emblingiaceae, and 

Meliosmaceae in Sapindales; Ptaeroxylaceae and 

Tepuianthaceae in Rutales; Nitrariaceae, Peganaceae, Lepidobotryaceae, Ledocarpaceae, Biebersteiniaceae, and Diracmaceae in Geraniales; 

Lophopyxidaceae in Celastrales; Eremolepidaceae and Viscaceae in Santalales; Cynomoriaceae 

in Balanophorales; Pentaphragmataceae and 

Lobeliaceae in Campanulales; Sclerophylaceae, 

Goetzeaceae, and Cobaeaceae in Solanales; 

Ehretiaceae in Boraginales; Roridulaceae in 

Ericales; Helwingiaceae, Torricelliaceae, Aucubaceae, Aralidiaceae, Phellinaceae, Sphenostemonaceae, Montiniaceae, Alseuosmiaceae, 

Anisophylleaceae, Sambucaceae, Viburnaceae, 

Dulongiaceae(?), and Tribelaceae in Cornales; 

Triplostegiaceae and Morinaceae in Dipsacales; 

Desfontainiaceae and Saccifoliaceae in Gentianales; Selaginaceae, Stilbaceae, Retziaceae, Trapellaceae, and Martyniaceae in Scrophulariales. 

Emberger, 1960. Cynomoriaceae in Santalales; Sphenocleaceae and Dysphaniaceae in Centrospermales; Julianiaceae in Julianiales; Coriariaceae, Cyrillaceae, and Pentaphlacaceae in Terebinthales, with a tendency toward Celastrales; 

Diclidantheraceae in Terebinthales(?); Hippocrateaceae in Celastrales; Humbertiaceae, Selaginaceae, Martyniaceae, and Tetrachondraceae in 

Tubiflorales; Sarcospermaceae in Ebenales; Lepidobotryaceae in Geraniales; Lophopyxidaceae 

in Euphorbiales; Lobeliaceae in Synantherales; 

Chrysobalanaceae and Roridulaceae in Rosales; 

Heteropyxidaceae in Myrtales. 

Hutchinson, 1973. Corylaceae in Fagales; Picrodendraceae in Juglandales; Aquilariaceae in 

Thymelaeales; Dirachmaceae in Tiliales; Huaceae, Ledocarpaceae, and Lepidobotryaceae in 

Malpighiales; Prionotaceae and Vacciniaceae in 

Ericales; Anisophylleaceae in Myrtales; Koeberliniaceae and Hippocrateaceae in Celastrales; Dipentodontaceae in Olacales; Heteropyxidaceae 

in Rhamnales; Aegicerataceae in Myrsinales; Sarcospermataceae in Ebenales; Averrhoaceae in 

Rutales; Podoonaceae and Julianiaceae in Sapindales; Potaliaceae, Antoniaceae, Spigeliaceae, 

and Strychnaceae in Loganiales; Periplocaceae in 

Apocynales; Cobaeaceae and Martyniaceae in 

Bignoniales; Ehretiaceae, Stilbaceae, and Dicrastylidaceae in Verbenales; Helleboraceae in Ranales; Cleomaceae and Oxystylidaceae in Brassicales; Dysphaniaceae in Chenopodiales; Lobeliaceae in Campanulales; Retziaceae in Solanales; 

Salpiglossidaceae in Personales; Selaginaceae in 

Lamiales. 

Melchior, 1964. Dipentodontaceae in Santalales; Dysphaniaceae in Centrospermae; Roridulaceae and Chrysobalanaceae in Rosales; Picrodendraceae in Rutales; Julianiaceae in Julianiales; 

Hippocrateaceae in Celastrales; Cynomoriaceae 

in Myrtales; Sarcospermataceae in Ebenales; Desfontainiaceae in Gentianales; Duckeodendraceae, Henriqueziaceae, and Martyniaceae in 

Tubiflorae; Sphenocleaceae and Pentaphragmataceae in Campanulales. 

Stebbins, 1974. Alseuosmiaceae(?) and Chrysobalanaceae in Rosales; Dipentodontaceae(?) 

and Cynomoriaceae in Santalales; Hippocrateaceae in Celastrales; Julianiaceae in Sapindales; 

Sphenocleaceae in Campanulales. 

Rouleau, 198 1. Sphenostemonaceae, Siparunaceae, Atherospermataceae, and Gyrocarpaceae 

in Laurales; Peperomiaceae in Piperales; Eury- 
NUMBER 58 29 

alaceae and Barclayaceae in Nymphaeales; Kingdoniaceae, Glaucidiaceae, Hydrastidaceae, and 

Leonticaceae in Ranunculales; Pteridophyllaceae 

and Hypecoaceae in Papaverales; Rhodoleiaceae 

and Altingiaceae in Hammelidales; Celtidaceae 

in Urticales; Carpinaceae and Corylaceae in 

Betulales; Tetragoniaceae, Halophytaceae, Hectorellaceae, and Dysphaniaceae in Caryophyllales; Aegialitidaceae and Limoniaceae in Plumbaginales; Lophiraceae, Sladeniaceae, Asteropeiaceae, and Oncothecaceae in Theales; Neumanniaceae in Violales; Tetramelaceae in Begoniales; 

Cleomaceae, Koeberliniaceae, Canotiaceae, and 

Emblingiaceae in Capparales; Populaceae in Salicales; Vacciniaceae in Ericales; Sarcospermataceae and Boerlagellaceae in Ebenales; Coridaceae in Primulales; Huaceae in Malvales; Stylocerataceae, Androstachydaceae, Bischofiaceae, 

Hymenocardiaceae, Peraceae, and Picrodendraceae in Euphorbiales; Tribelaceae, Brexiaceae, 

Dulongiaceae, Montiniaceae, Roridulaceae, Penthoraceae, and Lepuropetalaceae in Saxifragales; 

Chrysobalanaceae in Rosales; Anisophylleaceae, 

Napoleonaeaceae, Foetidaceae, Heteropyxidaceae, Psiloxylaceae, and Memecylaceae in Myrtales; Myriophyllaceae in Hippuridales; Uapacaceae, Pistaciaceae, Blepharocaryaceae, Julianiaceae, Podoaceae, Surianaceae, Stylobasiaceae, 

Rhabdodendraceae, Flindersiaceae, Ptaeroxylaceae, Kirkiaceae, and Aitoniaceae in Rutales; 

Physenaceae and Meliosmaceae in Sapindales; 

Hugoniaceae, Lepidobotryaceae, Nitrariaceae, 

Peganaceae, Averrhoaceae, Hypseocharitaceae, 

Ledocarpaceae, Dirachmaceae, and Biebersteiniaceae in Geraniales; Xanthophyllaceae in Polygalales; Aucubaceae, Curtisiaceae, Griseliniaceae, Melanophyllaceae, Mastixiaceae, Helwingiaceae, and Toricelliaceae in Cornales; 

Hydrocotylaceae in Apiales; Phellinaceae, Stilaginaceae, Lophopyxidaceae, and Hippocrateaceae in Celastrales; Schoepfiaceae, Dipentodontaceae, Viscaceae, and Eremolepidaceae in Santalales; Cynomoriaceae in Balanophorales; Carlemanniaceae, Sambucaceae, Alseuosmiaceae, 

Morinaceae, and Triplostegiaceae in Dipsacales; 

Desfontainiaceae, Spigeliaceae, Strychnaceae, 

Antoniaceae, Potaliaceae, Periplocaceae, Naucleaceae, and Henriqueziaceae in Gentianales; Cobaeaceae and Humbertiaceae in Polemoniales; 

Ehretiaceae and Wellstediaceae in Boraginales; 

Goetzeaceae, Duckeodendraceae, Retziaceae, Ellisiophyllaceae, Trapellaceae, Martyniaceae, 

Mendonciaceae, and Thunbergiaceae in Scrophulariales; Avicenniaceae, Symphoremataceae, 

Dicrastylidaceae, Plagiopteraceae, Stilbaceae, 

and Tetrachondraceae in Lamiales; Pentaphragmataceae, Lobeliaceae, and Sphenocleaceae in 

Campanulales. 

D. Young, 198 1. Glaucidiaceae and Hydrastidaceae in Ranunculales; Phellinaceae, Oncothecaceae, and Sphenostemonaceae in Theales; Hectorellaceae in Caryophyllales; Gisekiaceae and 

Halophytaceae in Chenopodiales; Huaceae and 

Plagiopteraceae in Malvales; Cecropiaceae in 

Urticales; Dipentodontaceae in Violales; Hippuridaceae and Lophopyxidaceae in Celastrales; Eremolepidaceae and Viscaceae in Santalales; Cynomoriaceae in Balanophorales; Ptaeroxylaceae 

in Rutales; Surianaceae in Sapindales; Chrysobalanaceae in Rosales; Penthoraceae and Brexiaceae in Saxifragales; Montiniaceae, Tribelaceae, 

Alseuosmiaceae, and Phyllonomaceae in Hydrangeales; Roridulaceae in Pittosporales; Martyniaceae in Bignoniales; Pentaphragmataceae in 

Campanulales; Sambucaceae in Dipsacales. 

Benson, 1979. Koeberliniaceae in Papaverales; 

Julianiaceae and Hippocrateaceae in Sapindales; 

Martyniaceae in Scrophulariales; Cynomoriaceae 

in Santalales; Dipentodontaceae(?); Huaceae(?). 

Ordinal and Family Descriptions 

T ROCHODENDR A L ES 

Usually trees, sometimes shrubs; xylem vesselless and tracheids long, or vessel perforation 

plates scalariform with many bars; leaves usually 

alternate, crenate, exstipulate or the stipules adnate to the petiole; buds perulate; plants dioecious or the flowers bisexual; flowers minute, 

anemophilous or rarely entomophilous; sepals 0 
30 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

or 4; petals 0; stamens 4-40, in 1, 3 or 4 series, 

rarely adnate to the ovary, the filaments filiform, 

anthers oblong-linear, basifixed, the connective 

often produced apically; pollen 3-colpate, 3(4)- 

colporoidate; pistils 4-18, in 1 whorl, or pistil 1 

and 4-1 l-carpellate, the styles as many as carpels, 

stigmas decurrent ventrally; ovary superior, 4- 

1 1-locular in the compound pistils, the ovules 1 

or several to numerous; fruit follicular or a capsule dehiscing along the ventral sutures, or small 

samaras; seeds small, the embryo usually small; 

endosperm usually copious. 

Mainly temperate China and Japan. 

Hegnauer considers that lack of 

oil-cells and presence of myricetin and ellagic 

acid support placing Cercidiphyllaceae near Hamamelidaceae. The presence of leucodelphinidin 

and a large number of free triterpene alcohols, 

aldehydes and acids in Trochondendron is not a 

very good indication of relation to Magnoliales. 

Trochodendraceae, Cercidiphyllaceae, Tetracentraceae, and Eupteleaceae do not contain alkaloids nor lignans, whereas both of these are 

common in Magnoliales. 

CERCIDIPHYLLACEAE (Figure 1 a).-Trees with 

long and short shoots; xylem vessel perforation 

plates scalariform, with 20-50 very fine bars; 

leaves simple, deciduous, opposite and alternate, 

crenate, the main veins palmately arranged, stipules adnate to the petiole; inflorescence axillary 

fascicles of 1-6 flowers, the plants dioecious; 

sepals 4, small, also interpreted as bracts; petals 

0; stamens 8-20, the filaments long, filiform; 

anthers oblong-linear, basifixed; pollen 3- 

colp(oid)ate; pistils usually 4-6, the style filiform, 

stigma decurrent ventrally, long; ovules numerous, marginal, biseriate, with subchalazal process, 

bitegmic, crassinucellar, anatropous; fruits follicular, the seeds small, numerous, compressed, 

winged; embryo medium-sized to large, spatulate, straight, as long as the endosperm, the 

cotyledons elliptic, flat, moderately thick, as wide 

as, to twice as wide as, the radicle, 0.5 times the 

length of the embryo; endosperm moderate, 

fleshy. 

Distribution: 

Chemistry: 

Composition: 1 genus, 1 species. 

Distribution: Japan, northeastern China to 

Korea. 

TETRACENTRACEAE (FIGURE 1 b-e).-Trees 

with long and short shoots; xylem vesselless, the 

tracheids extremely long, the parenchyma diffuse; leaves simple, deciduous, alternate, crenate, 

the main veins palmately arranged; bud enclosed 

by stipule adnate to the petiole; inflorescence a 

catkin-like spike, the flowers in groups of 4, 

bisexual; sepals 4, small; petals 0; stamens 4, 

hypogynous, the filaments filiform; anthers as 

long as wide, basifixed; pollen 3-colp(oroid)ate; 

pistil 1, carpels 4, united at ovary level, styles 4, 

free, filiform, stigmas decurrent ventrally; ovary 

superior, locules 4, ovules several per locule, 

axile with subchalazal process, bitegmic, crassinucellar, anatropous; fruit a follicetum, each carpel dehiscing ventrally; seeds minute, fusiform, 

the embryo minute, subglobular or ellipsoid, obscurely cleft at the apex, 0.07 the length 

of the endosperm; endosperm copious, oily. 

Composition: 1 genus, 1 species. 

Distribution: Central and southwestern 

China, Nepal, northern Burma. 

EUPTELEACEAE (Figure 2a,b).-Trees or 

shrubs; xylem vessel perforation plates scalariform, with 20-90 bars; leaves simple, deciduous, 

alternate, serrate, exstipulate, the buds perulate; 

inflorescence a fascicle of 6-12 flowers in axils 

of bracts; flowers bisexual; perianth absent; stamens about 10 in 1 whorl, the filaments filiform; 

anther oblong, long, basifixed, the connective 

produced apically; pollen 3-colpate; pistils 6-1 8 

in 1 whorl, not completely closed, the ovules 1- 

3, ventral, bitegmic, crassinucellar, anatropous; 

stigma sessile, ventral; fruitlets small samaras; 

embryo small, straight, 0.5 the length of the 

endosperm; cotyledons as wide as the radicle, 0.6 

the length of the embryo; endosperm copious, 

oily. 

Composition: 1 genus, 2 species. 

Distribution: Central and southwestern 

China, Assam, Japan. 

TROCHODENDRACEAE (Figure 2c,d).-Trees, 

the bark and foliage slightly aromatic (?); xylem 
NUMBER 58 31 

a 

C 

e 

FIGURE 1.-CERCIDIPHYLLACEAE: a, Cercidiphytlum japonicum 6 flowering twig, I.s. of pistil, 6 

flower, 0 flower, 0 flowering twig, twig with dehiscent fruitlets, seed, I.s. of same showing the 

embryo, embryo (after Sargent). TETRACENTRACEAE: b, Tetracentron sinensis twig with 2 

inflorescences at anthesis, twig with inflorescence in bud; c, bud laid open, flower, pistil, carpel; 

d, views of anther, pistil more advanced, I.s. of same; e, I.s. of fruit, fruit, seed with testa 

removed, seed (after Oliver). 
32 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

b 

FIGURE 2.-EUPTELEACEAE: a, Euptelea davidiana twig with ? flowers, parts of a twig with 6 

flowers in clusters of four, twig in fruit; b, stamen, 8 flower, ? flower, 1,s. of one of the pistils, 

fruitlet, 1,s. of a fruitlet, seed, I.s. of same, embryo (after Hemsley). TROCHODENDRACEAE: c, 

Trochodendron aralioides flower, pistil, I.s. of a carpel, views of anther after and before 

dehiscence, twig with inflorescence; d, dehiscent fruits, fruit, I.s. of same showing seeds, section 

of a seed showing minute embryo (after Hooker and Hooker, 1837-1982; Prantl). 
NUMBER 58 33 

vesselless, the tracheids long, the parenchyma 

diffuse; leaves alternate, simple, coriaceous, 

persistent, subverticillate, crenate-serrate, venation pinnate, exstipulate, the buds perulate; inflorescence a terminal racemiform pleiochasial 

cyme; flowers bisexual; perianth absent; stamens 

20-40, 3-4-seriate, adnate to the ovary, the filaments filiform; anthers twice as long as wide, 

adnate; pollen 3(-4)-colporoidate; pistil 1, the 

carpels (4-) 6-1 1, laterally connate in 1 whorl, 

the styles 4-1 1, short, the stigmas decurrent 

ventrally; ovary locules 4- 1 1 ; ovules ventral, 

with subchalazal process, bitegmic, crassinucellar, anatropous, about 18 per locule; a droplet of 

fluid is secreted from the dorsal protuberance of 

each carpel; fruit a capsule, the carpels dehiscing 

ventrally and partly septicidally; seeds minute, 

fusiform, produced at each end; embryo minute, 

undifferentiated, 0.1 the length of the endosperm; endosperm copious, oily. 

Composition: 1 genus, 1 species. 

Distribution: Japan, Taiwan, Korea; at altitudes of 666-3000 m. 

HAMAMELIDALES 

Trees and shrubs; xylem vessel perforation 

plates entirely or mostly scalariform, usually with 

more than 20 bars; leaves usually alternate, 

toothed or entire, stipulate or exstipulate; plants 

dioecious or monoecious, less commonly the 

flowers bisexual; flowers minute or small, anemophilous or entomophilous; sepals 0 or 4-8; 

petals usually 0, sometimes 4-5 (-7); stamens 2- 

14 (-32), the filaments usually short, anthers 

adnate or basifixed, the connective sometimes 

apically produced; pollen 3-colpate, -colporate, 

-colporoidate or polyforate, rarely 4-ruptate or 

6-rugate; pistils usually 1, rarely 5-9, the styles 

as many as carpels, stigma usually decurrent ventrally; ovary 2-3 (4)-locular in the compound 

pistils, the ovules l-many per locule, axile, rarely 

parietal; fruit follicles, loculicidal or septicidal 

capsule, nutlet or drupe; seeds 1-many, the embryo minute to as long as the endosperm; endosperm copious to absent. 

Distribution: Often in eastern Asia, also in 

Africa and North America, some reaching northern South America, in various habitats. 

According to Jay, Hamamelis, Myrothamnus, and Platanus are chemically related 

and related to Saxifragaceae, and Myrothamnus 

leaf pigments are identical to those of Disanthus. 

Platanaceae are highly cyanogenic. 

PLATANACEAE (Figure 3a-c).-Trees, often 

with segments of bark scaling off; xylem vessel 

perforation plates scalariform with fewer than 20 

bars, and simple; leaves simple, deciduous, alternate, dentate, usually palmately rarely pinnately 

veined, usually lobed, the petiole base capping 

the bud; stipules usually large and broad and 

united around the twig; inflorescences globular 

heads, the peduncle pendulous, the plants monoecious; perianth represented by minute scales; 

stamens 3-7, the filaments short; anthers adnate, 

apical connective flattened horizontally; pollen 

usually 3-colpate or -colporate; pistils mostly 5- 

9, not completely closed apically, the style elongated, stigma decurrent ventrally; ovules 1 (-2), 

bitegmic, crassinucellar, orthotropous; fruit an 

obconical nutlet surrounded at the base by long 

hairs; embryo linear, terete, straight, as long as 

the endosperm; cotyledons thin, 0.5 the length 

of the embryo; endosperm scanty, fleshy. 

Chemistry: 

Composition: 1 genus, 7 species. 

Distribution: Eastern and southwestern 

United States, Mexico; part of temperate eastern 

Canada; southeastern Europe to northern Iran; 

Himalayas; Indochina. 

MYROTHAMNACEAE (Figure 3d,e).-Low resinous shrubs; xylem vessel perforation plates 

mostly scalariform with about 45 bars; leaves 

simple, small, opposite, flabellate-cuneate, plicate-veined, the stipules attached to the petiole; 

inflorescence terminal and axillary erect catkinlike spikes, the plants dioecious; perianth 0; stamens 4-8 subtended by a bract, the filament 

short; anthers large, basifixed, the connective 

produced apically; pollen faintly 3-colpoid; pistil 

1, the carpels 3 (-4), the styles 3, recurved, 

stigma decurrent ventrally; ovary superior, 3- 

locular, the ovules numerous, axile; fruits dehisc- 
34 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 58 35 

ing septicidally and ventrally into 3 follicles, the 

seeds numerous, minute; embryo minute, 0.3 the 

length of the endosperm; cotyledons not broadened(!), 0.3 the length of the embryo; endosperm 

copious. 

Composition: 1 genus, 2 species. 

Distribution: Often in arid areas; tropical and 

southern Africa, Madagascar. 

BUXACEAE (Figure 3f,g).-Trees and shrubs, 

rarely subshrubs; xylem vessel perforation plates 

usually exclusively scalariform with 30 or more 

bars; leaves simple, evergreen, mostly entire, opposite or alternate, exstipulate; inflorescences 

mostly axillary dense racemes, spikes, or the flowers solitary, the ? flowers fewer than the 8, the 

plants monoecious or dioecious; sepals usually 4 

or 6, in 2 whorls, basally connate, sometimes 0; 

petals 0; stamens 4 or 6 (to 30 in Styloceras); 

anthers oblong, basifixed, dehiscing by valves or 

longitudinally; pollen usually polyforate; pistil 1, 

the carpels 3 (2-4), styles 3 (2-4), sometimes 

distant, persistent, the stigmas decurrent ventrally; ovary superior, locules 3 (2-4), the ovules 

1-2 per locule, axile, bitegmic, crassinucellar, 

anatropous; fruit a loculicidal capsule sometimes 

dehiscing explosively, or drupe; seeds usually 

black and shiny, sometimes with a caruncle; embryo straight, linear, terete, small to large, 0.2- 

0.9 the length of the endosperm; cotyledons 0.1- 

0.8 the length of the embryo, 4 times wider than 

the radicle; endosperm copious, fleshy. 

FIGURE Q.-PLATANACEAE: a, Platanus sp. 0 inflorescence, 

P. vulgaris C.S. of P inflorescence, Platanus sp. d inflorescence, P. vulgaris C.S. of d inflorescence; b, Platanus sp. 0 

flower, 1,s. of pistil, d flow-er and C.S. anther, P. vulgaris 

infructescence with several fruits removed; c, Platanus sp. 

fruit, 1,s. of same (after Baillon 1866-1895; Le Maout and 

Decaisne, 1873). MYROTHAMNACEAE: d, Myrothamnus moschata diagrams of d and 0 flowers, M.jabellifolius 1,s. of seed 

showing minute embryo, seed; e, M. moschata 6 flower, 0 

flower, I.s. of pistil, branch with 0 inflorescences, branch 

with d inflorescences (after Baillon, Welwitsch). BUXACEAE: 

f, Bums sempemirens flowering twig, inflorescence, 0 flower, 

I.s. of sane, diagram of 0 flower: g, 1.3 flower, diagram of d 

flower, young fruit and remains of 6 flowers, dehiscent fruit, 

seed, 1,s. of same (after Baillon, 1866-1895; Le Maout and 

Ilecaisne, 1873). 

Composition: 5 genera, -80 species. 

Distribution: Disjunct; temperate to tropical; 

Europe to India, China, Japan, and the western 

East Indies; tropical and subtropical southern 

Africa; southeastern United States; southern 

Mexico; West Indies; northern Andes. 

HAMAMELIDACEAE (Figure 4a-e).-Trees or 

shrubs; xylem vessel perforation plates scalariform, with more than 20 bars; leaves simple, 

alternate, rarely opposite, deciduous or evergreen, often with stellate indument, sublobed, 

dentate or entire, the teeth sometimes glandular, 

stipulate (except Rhodoleia); inflorescence an axillary rarely terminal spike or head, rarely raceme; flowers bisexual or the plants polygamous, 

monoecious or dioecious; sepals 4-5 (0, 6, 7), 

sometimes connate, rarely calyptrate, usually k 

adnate to the ovary; petals 4-5 (0, 6, 7), free, 

often ligulate, usually perigynous or epigynous; 

stamens 4-5 (-32), usually perigynous; anthers 

basifixed, dehiscing lengthwise or by valves, the 

connective often apically produced; pollen usually 3-colpate or 3-colporoidate or polyforate 

(occasionally 4-rupate or 6-rugate); glandular 

disk sometimes present between the stamens and 

ovary; pistil 1, the carpels 2 (exceptionally a 

flower may have 1 or 3 carpels), styles free, 

usually elongate, rarely 0 (Mytilaria), filiform, 

the stigma punctate, capitate or foliar, sometimes 

decurrent ventrally; ovary semi-inferior, inferior, rarely superior, bilocular, rarely unilocular, 

the ovules 1-several per locule, axile, rarely parietal, bitegmic, crassinucellar, anatropous; fruit 

usually a woody capsule, loculicidal or septicidal; 

embryo straight, 0.6 to as long as the endosperm; 

cotyledons thin, 2.2-4.0 times as wide as the 

radicle, 0.6-0.8 the length of the embryo; endosperm moderate or scanty, fleshy. 

Composition: 22 genera, -90 species. 

Distribution: Disjunct, chiefly subtropical and 

warm termperate, centered in eastern Asia; eastern North America to Andes of Venezuela and 

Bolivia; Japan, China, northeastern India, Indochina; East Indies; southwestern Asia; eastern 

and southern Africa, Madagascar; northeastern 

Australia. 
36 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

C 

FIGURE 4.-HAMAMELIDACEAE: a, &pidambar StpCtJUU twig with d and ? inflorescences, 1,s. 

of d inflorescence, C.S. of same, anther, C.S. of ? inflorescence; b, part of same enlarged, stigmas 

and staminodes, 1,s. of ovary, dehiscent infructescence, seed, 1,s. of same, embryo. c, Hamamelis 

virginiana flowering and fruiting twigs, floral diagram; d, twig with flowers in winter, seed, I.s. 

of same, embryo; e, 1,s. of flower, pistil, views of stamen, one with an open anther, staminode, 

C.S. of fruit, dehiscent fruit (after Baillon, 1866-1895; Le Maout and Decaisne 1873; Sargent). 
NUMBER 58 37 

a 

FIGURE !j.-DIDYMELACEAE: a, Didymeles madagascarensis twig with d inflorescences, part of ? 

inflorescence, two ? flowers, 1s. of pistil, 6 flower, fruits (after Hutchinson, 1973). DAPHNIPHYLLACEAE: b, Daphniphyllum macropodium d flower, 0 flower, 1,s. of fruit, views of embryo, 

twig with 0 inflorescences (after Rosenthal). 

DAPHNIPHYLLACEAE (Figure 5b).-Trees and 

shrubs, sometimes containing very poisonous alkaloids; xylem vessel perforation plates scalariform with more than 20 bars; leaves alternate, 

often subverticillate, rarely opposite, simple, entire or infrequently lobed or toothed near the 

apex, exstipulate; plants dioecious, rarely polygamodioecious; inflorescences axillary racemes, 

sometimes partly umbelliform, the flowers minute; sepals 0 or 3-8, free or connate, imbricate; 

petals 0; stamens 6-14, free, the filaments short, 

rarely long; anthers introrse or extrorse, oblongish, rarely as long as wide, adnate or basifixed, the connective considerable between the 

sacs, and sometimes apically produced; 0 often 

with 5-10 staminodes; pollen small, 10-27p, 

3(-4) colpor(oid)ate; disk 0; pistil 1, the carpels 

2 (-4), the styles 1-2 (-4), usually short, stigmas 

2 (1-4), divaricate, enlarged, apical or decurrent 

ventrally; ovary superior, unilocular or incompletely 2(3-4)-locular, the ovules (1) 2 per locule, 

axile or parietal, subapical, bitegmic, crassinucellar, anatropous; fruit a drupe, the seeds 1 (-2); 

embryo straight, clyindrical, minute, 0.1 the 

length of the endosperm; cotyledons not broadened, 0.6 the length of the embryo; endosperm 

copious, fleshy and oily. 

Composition: 1 genus, - 10 species. 
38 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Distribution: Japan, China to the Himalayas, 

Indochina and the western East Indies, western 

India, Sri Lanka. 

DIDYMELACEAE (Figure 5a).-Trees; xylem 

vessel perforation plates scalariform with numerous bars; leaves alternate, simple, entire, smooth, 

exstipulate; plants dioecious; cJ inflorescences axillary compound spikes, 0 in racemes, the flowers 

in pairs (1-3); sepals (? bracts) 0-4; petals 0; 

stamens 2, the filaments shorter than the anthers, 

connate(?); anthers oblongish, basifixed, extrorse; pollen 3-colpate, the colpi 2-orate; pistil 

1, unicarpellate, style present or the stigma subsessile, apical, somewhat elongated, papillose, 

grooved so appearing bilobed, persistent in fruit; 

ovary superior, the ovule 1, semi-anatropous, 

adaxial; fruit a large drupe, 3.5 X 1.5 cm., the 

endocarp osseous, seed bitter; cotyledons large, 

plano-convex, sometimes connate in age; endosperm 0. 

Composition: 1 genus, 2 species. 

Distribution: Madagascar. 

MAGNOLIALES 

Trees and shrubs, sometimes climbers; xylem 

vessels rarely absent and the tracheids long, vessel perforation plates usually scalariform, often 

with numerous bars, sometimes simple; leaves 

usually alternate, entire, less commonly toothed, 

usually exstipulate; flowers small to large, usually 

entomophilous, usually bisexual, more rarely the 

plants monoecious or dioecious, the parts usually 

hypogynous, rarely perigynous; perianth homoor heterochlamydeous, rarely absent, the parts 

spirally arranged and indefinite in number or in 

1 or more whorls and fairly frequently 3-merous; 

stamens usually numerous and spirally arranged, 

rarely few and whorled, sometimes laminar; 

staminodes sometimes present; filaments free, 

rarely connate, the anthers adnate or basifixed, 

usually longitudinally dehiscent, rarely valvate; 

pollen usually 1 (-3)-sulcate, less frequently 3- or 

6-colp(oid)ate, 1- or 8-1 2-pored, 2-colpate or 

nonaperturate; pistils many to 1, often spirally 

arranged, more rarely few or 1 and whorled; 

style elongate or the stigma sessile, the stigma 

frequently decurrent ventrally, more rarely apical, the ovules I-many; fruits follicles, berries, 

drupes, achenes, or a fleshy syncarp, rarely a 

capsule; embryo usually minute; endosperm copious, very rarely scanty or absent. 

Most frequently in tropical rain 

forest, less commonly in temperature rain forest 

or drier forests, especially of eastern Asia and 

eastern North America. 

Magnoliales are rich in characteristic aromatic oils such as monoterpenes and sesquiterpenes. Secretory cells and passages occur 

in many of the families. They have aromatic 

aldehydes, mostly phenolic; the Ranunculales 

lack them. Magnoliales, except Schisandraceae 

and Illiciaceae, are rich in alkaloids; aporphines 

are common. They have phenolic esters and 

ethers, but the Ranunulales lack them. Lignans 

are disproportionately common in Magnoliales, 

Ranunculales, Piperales, and Aristolochiales. Calycanthaceae are cyanogenic. 

MAGNOLIACEAE (Figure 6a, b).-Trees or 

shrubs; xylem vessel perforation plates typically 

scalariform with few to 25 bars, more rarely 

simple; leaves simple, alternate, entire, rarely 

lobed, the stipules large, enclosing the buds; flowers usually large, usually solitary, terminal or 

axillary, bisexual (unisexual in Kmeria), the parts 

hypogynous; perianth homoiochlamydeous or 

heterochlamydeous, spirally arranged or in one 

whorl of sepals and 2-4 of petals, and then 

usually trimerous (rarely 5-merous); stamens 

sometimes not differentiated into anther and 

filament, numerous (-30-50), spirally arranged, 

free, staminodes 0; anthers generally adnate and 

oblong, the connective usually apically produced; 

pollen monosulcate; pistils numerous (- 12-60) 

to 1, spirally arranged on an elongated axis, the 

stigma decurrent ventrally; ovules 2 to several, 

bitegmic, crassinucellar, anatropous; fruitlets follicles, samaras, or rarely a fleshy syncarp, or 

woody capsule (Pachylarnax) septicidally or loculicidally dehiscent; seeds usually large, usually 

with an outer stomate sarcotesta and woody inner 

integument; embryo minute, 0.2-0.3 the length 

Distribution: 

Chemistry: 
NUMBER 58 39 

a 

FIGURE ~.-MAGNOLIACEAE: a, Magnolia grandzyora flowering twig, M. yulan I.s. of flower, M. 

grandzflora floral diagram; b, Liriodendron tulipifera two views of anther, fruit, 1,s. of same 

showing minute embryo, Magnolia plan dehiscent fruit, M. purpurea 1,s. of seed showing 

minute embryo (after Baillon, 1866-1895; Martius, 1840-1906). 

of the endosperm; cotyledons hardly differentiated to 1.5 times wider than the radicle; endosperm copious, fleshy. 

Composition: 12 genera, -225 species. 

Distribution: Centered in the Northern Hemisphere; temperate and tropical southeastern 

Asia from the Himalayas to Japan and the East 

Indies; eastern United States through Central 

America to disjunct areas in northern South 

America and southeastern Brazil; fossils recovered in Europe and Greenland. 

WINTERACEAE (Figure 7a-c).-Trees or 

shrubs; xylem vessels absent, the tracheids very 

long; leaves simple, alternate, subverticillate, entire, pellucid-punctate, aromatic, exstipulate; inflorescences subterminal, intercalary or axillary, 

cymose or fasciculate, or the flowers rarely solitary; flowers small, bisexual, or the plants rarely 
40 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 7.-wINTERACEAE: a, Drimgs lanceolata flowering twig, I.s. of flower, S flower, D. 

winteri flower from above and below, pollen grain tetrads; b, petal, essential organs from a bud, 

views of stamen, gynoecium, same from above, I.s. and C.S. of 1 pistil, fruitlets; c, single fruitlet, 

seed, 1,s. of same showing the minute embryo, Pseudowintera axillaris upper and lower views of 

flower, Zygogynurn vieillardi flower with petals removed, floral diagram (after Baillon, 1866- 

1895; Hooker and Hooker, 1837-1982; Martius, 1840-1906). 

polygamous, the parts hypogynous; perianth heterochlamydeous; sepals 2-6 free and valvate or 

connate and calyptrate enclosing the petals in 

bud; petals spirally arranged or in 1-4 whorls of 

3 or more; stamens 12-250+, spirally arranged, 

staminodes rare; anthers adnate or basifixed; pollen uniporate (ulcerate; pore usually on distal 

pole); hypogynous disk rarely present (Tetrathalamus); pistils 1-24, sometimes incompletely 

closed, in 1 whorl, sometimes connate (Zygogynum), disputedly bicarpellate in Takhtajania, 

style short, stigma(s) decurrent ventrally or apical; ovules 1-85 per pistil, 2-seriate, bitegmic, 

crassinucellar, anatropous; fruits berries or a 

fleshy syncarp; embryo minute, not differentiated, 0.04 the length of the endosperm; endosperm copious. 

Composition: 

Distribution: 

7 genera, -1 20 species. 

Wet tropical montane to cool 

temperate rain forest in disjunct areas in Mexico, 
NUMBER 58 41 

a 

C 

FIGURE ~.--ANNONACEAE: a, Guatteria pohliana flower from above and below, pistil, pistils and 

stamens on the convex receptacle, views of stamen, seed, I.s. of same enlarged: b, Rollinza 

cuspidata pistils on receptacle, pistil, I.s. of same; c, Guatteria villosissima fruitlets, Anaxagorea 

phaeocarpa dehiscent fruitlets and seed, Annona squamosa fruit (after Martius, 1840-1 906). 

Central America, northern, eastern, and southern South America, the East Indies, eastern Australia, New Zealand, New Caledonia, Madagascar. 

ANNONACEAE (Figure 8a-c).-Trees, shrubs, 

or climbers; xylem vessel perforation plates simple; leaves simple, alternate, entire, almost always 

distichous, exstipulate; inflorescence axillary, 

leaf-opposed or cauliflorous, sympodial, or the 

flowers solitary; flowers large or small, sometimes 

smelling like carrion, usually bisexual, rarely the 

plants monoecious or dioecious (Stelechocarpus), 

the parts hypogynous (stamens perigynous in Xylopia); perianth heterochlamydeous; sepals usually 3 (2-4), free or connate, rarely calyptrate; 

petals often 6 and trimerous, rarely 12, 8, 4, or 

3, rarely connate; stamens usually numerous (to 

-150), spirally arranged, rarely 3 or 6 and 

whorled, the filaments very short; subpetaloid 

staminodes external to the stamens sometimes 

present (Fusaea); anthers usually adnate, sometimes basifixed, usually extrorse, connective usually apically produced and often broad and truncate; pollen nonaperturate or I-sulcoidate; pistils 

numerous (to -25) to 1, spirally arranged (on an 

elongated torus in Mischogyne) or whorled, very 

rarely open, the styles short or 0 (Tetrameranthus), stigmas decurrent to subapical (united in 

Piptostigma, 3-lobed in Tetrameranthus); ovary superior, rarely sunk in torus (Pseudannona), unilocular, placentas ventral or parietal, the ovules 

usually numerous, rarely 1, 2(3)-tegmic, crassinucellar, anatropous; fruits usually berries, often 

stipitate, sometimes a syncarp, rarely a capsule 
42 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

d 

FIGURE 9.-DEGENERIACEAE: a, Degeneria vitiensis flower, same with petals, stamens and 

staminodes removed, slightly after anthesis, lateral view of pistil, I.s. of same, views of staminode 

and stamens, petal; b, flowering twig, fruiting twig, C.S. of anther greatly enlarged; c, nearly 

mature fruit with part removed to show seeds, seed with endocarpic appendage (after Bailey 

and Smith). AUSTRORAILEYACEAE: d, Austrobaileya maculata views of stamen, a pistil, flower 

from below, flowering twig, fruitlet (after Hutchinson, 1973; Takhtajan, 1969). 

or follicle (Anaxagorea), seeds often arillate tiated or 0.2-0.3 the length of the embryo; en- 

(winged in Richella); embryo minute or small, 

linear, straight, 0.1-0.3 the length of the endosperm; cotyledons not broadened, undifferen- Distribution: Mostly in lowland evergreen 

dosperm copious, ruminate, hard or fleshy. 

Composition: -1 20 genera, -2,000 species. 
NUMBER 58 43 

forest; tropical America; southeast Asia to the 

East Indies and northern Australia; Africa; few 

species in temperate America and Asia. 

DEGENERIACEAE (Figure 9a-c).-Tree; xylem 

vessel perforation plates scalariform with numerous bars; leaves simple, alternate, entire, exstipulate; flowers bisexual, solitary, supra-axillary, 

the parts hypogynous; perianth heterochlamydeous, whorled; sepals 3(-4), persistent; petals 

12-1 8 in 3-5 whorls, fleshy; stamens 20-30, flat, 

in 3 whorls, not differentiated into anther and 

filament, anther sacs immersed, linear, extrorse, 

about 9 staminodes present; pollen 1-sulcate; pistil I(-2) unicarpellate, open, the stigma sessile, 

decurrent along the ventral suture of the ovary, 

the placenta subventral, the ovules numerous, 

bitegmic, crassinucellar, anatropous; fruit large, 

leathery, dehiscent along the ventral suture, the 

seeds large, strongly flattened, with a sarcotesta; 

embryo minute, 0.02 the length of the endosperm; cotyledons 3 (-4), 0.55 the width of the 

radicle, 0.4 the length of the embryo; endosperm 

copious, ruminate, oily. 

Composition: 1 genus, 1 species. 

Distribution: Fiji. 

AUSTROBAILEYACEAE (Figure 9d).-Climbing 

shrubs, xylem perforation plates scalariform, 

sieve tube companion cells sparse or absent; 

leaves simple, opposite, subopposite or alternate, 

leathery, entire, the stipules small; inflorescence 

axillary or terminal, the flowers solitary or in 

cymes of 3 flowers; flowers medium to large, 

bisexual, smelling of dead fish, the parts hypogynous; tepals 19-24, spirally arranged, pale 

green, free, with gradual transition from tepals 

to stamens and staminodes; stamens 7-1 1, laminar, covered with resinous warts, anther sacs 

adnate, introrse, staminodes 9- 16, internal to 

the stamens; pollen 1 -sulcate; pistils shorter than 

the staminodes, 6- 14, incompletely closed, styles 

apically bilobed, stigmas decurrent ventrally; 

ovules 6-8 (4- 14) per pistil, bitegmic, crassinucellar, anatropous; fruitlets large, slightly elongated, stipitate berries; seeds about 6, the embryo 

minute, 0.1 the length of the endosperm, the 

cotyledons 0.4 the length of the embryo; endosperm copious, ruminate, starchy. 

Composition: 1 genus, 2 species. 

Distribution: Rain forest of Queensland, Australia. 

HIMANTANDRACEAE (Figure 1 Oa, b).-Trees; 

xylem vessel perforation plates simple and scalariform; leaves simple, alternate, entire, pellucidpunctate, exstipulate, indument of peltate scales; 

flowers solitary or paired, on short axillary 

branches, bisexual, the parts hypogynous; sepals 

2, connate, calyptrate; petals about 4-7, lanceolate, similar to the stamens; stamens about 40, 

the anther loculi extrorse, separate and adnate 

near the base of the elongated filament, staminodes present; pollen I-sulcate; pistils 7-10 (to 

-25), spirally arranged, free except at the base, 

style elongated, papillose; ovule 1 (-2) per pistil, 

ventral; fruit a globose, red, fleshy syncarp, the 

seeds suborbicular, 3-4 mm in diameter; embryo 

small, endosperm copious, oily. 

Composition: 1 genus, 2 species. 

Distribution: Rain forest of northeastern Australia, New Guinea, and the Moluccas. 

EUPOMATIACEAE (Figure 1 Oc,d).-Shrubs or 

small trees; xylem vessel perforation plates scalariform with 20-100 bars; leaves simple, alternate, entire, exstipulate; flower solitary, terminal, bisexual, the parts perigynous; perianth 

('bracts) calyptrate; stamens numerous (to -50), 

spirally arranged, anthers adnate, the connective 

acuminate; staminodes internal to the stamens 

petaloid and bearing food-bodies eaten by beetles; pollen 2-3-sulculate, sometimes with an annular groove; pistils numerous (- 16), contiguous 

but free, incompletely closed, immersed in an 

urceolate receptacle, the stigmas penicillate, essentially sessile; ovules several per pistil, bitegmic, crassinucellar, anatropous; fruit nutlets 

(achenes?) enclosed in a fleshy receptacle which 

is apically truncate; embryo minute, 0.1 the 

length of the endosperm; endosperm copious, 

ruminate. 

Composition: 1 genus, 2 species. 

Distribution: Temperate and tropical eastern 

Australia, New Guinea. 

Myristicaceae (Figure 1 1 a, b).-Trees; xylem 

vessel perforation plates scalariform, the bars 

few, occasionally to 20, sometimes also simple; 
44 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

b ~ 

t 

a 

I ~ II 
NUMBER 58 45 

leaves simple, persistent, alternate, entire, often 

pellucid-punctate, sometimes distichous, exstipulate; inflorescences axillary or supra-axillary 

panicles, racemes, cymes, fascicles, or heads, the 

plants dioecious; flowers small, the parts hypogynous; sepals 3 (2-5), connate, colored, the calyx 

saucer-shaped to globose, the lobes valvate; petals 

0; stamens 2-45, the filaments usually united 

into a column, sometimes nearly free; anthers 

connate or free, the connective sometimes apically produced; pollen l-sulcate, in some cases 

nonaperturate(?); pistil 1, unicarpellate, the ovule 

1, subbasal, bitegmic, crassinucellar, anatropous, 

rarely suborthotropous, the stigma subsessile, apical; fruit usually a berry, usually splitting along 

both sutures; seed large, often with a laciniate, 

colored aril; embryo minute, 0.2 the length of 

the endosperm; cotyledons divaricate and grading into the radicle, 0.7 the length of the embryo, 

or united; endosperm copious, usually ruminate, 

oily or starchy. 

Composition: 

Distribution: 

- 17 genera, -300 species. 

Tropical lowland rain forest of 

America, Asia, Africa, Madagascar, the East Indies, northeastern Australia. 

CANELLACEAE (Figure 1 lc,d).-Trees; xylem 

vessel perforation plates scalariform with 10-60 

bars; leaves simple, alternate, leathery, entire, 

glandular-punctate, exstipulate; inflorescence a 

terminal or axillary cyme or raceme, or the flowers solitary; flowers bisexual, small, the parts 

hypogynous; sepals 3-5, free, thick, imbricate; 

petals 4-5 (-12) free or connate, or 0, 3-5- 

merous, rarely spirally arranged; stamens 7-40, 

connate in a tube; anthers adnate, extrorse; pollen l-sulcate; pistil l, the carpels 2-6, the style l 

short and thick, stigmas 2-5, apical; ovary uniFIGURE 1 ~.--HIMANTANDRACEAE: a, Galbulimima baccata 

twigs in bud, indunient scales, bud, I.s. of bud, flower, pistil, 

C.S. of same, 1,s. of carpel; b, staminode, stamen, fruit, C.S. of 

same (after Lauterbach, Sprague). EUPOMATIACEAE: c, Eupomatia bennettii floral diagram, flowering twig, E. laurina 

flower, I.s. of same; d, stamen, inner staminode, carpels, I.s. 

of 2 carpels, fruit, I.s. of same, I.s. of seed showing the 

minute embryo, embryo, C.S. of seed (after Baillon, 1866- 

189.3; R. Brown). 

locular, the placentas parietal 2-6, the ovules 2- 

several, bitegmic, crassinucellar, anatropous or 

semi-anatropous; fruit a berry, the seeds shining; 

embryo linear, straight or slightly curved, 0.2- 

0.4 the length of the endosperm; cotyledons not 

broadened(?), 0.4 the length of the embryo; endosperm copious, ruminate, oily and fleshy. 

Composition: 

Distribution: Tropics of northern South 

America, the West Indies; Florida; eastern and 

southern Africa, Madagascar. 

SCHISANDRACEAE (Figure 1 2a-c).-Climbing 

or trailing shrubs; xylem vessel perforation plates 

scalariform with 7- 15 bars, sometimes also simple; leaves simple, alternate, often pellucid-punctate, entire or crenate-serrate, exstipulate; flowers small, axillary, usually solitary, the parts hypogynous, the plants monoecious or dioecious; 

perianth parts 9-1 5, spirally arranged, with gradual transition from sepals to petals; stamens 5 to 

-60, partially on wholly connate into a fleshy 

globose mass, spirally arranged or in 1 or more 

whorls; anthers small, basifixed, extrorse or introrse, as long as wide, with connective separating the thecas but not apically produced, staminodes rare; pollen 3- or 6-colp(oid)ate; pistils 20- 

30 (1 2-1 20), spirally arranged, only partly 

closed, the style short, stigma decurrent ventrally; ovules 2-5, ventral, bitegmic, crassinucellar, anatropous or campylotropous; fruit berries 

or drupaceous; embryo minute, straight, 0.2 the 

length of the endosperm; cotyledons divaricate, 

not broadened, 0.5 the length of the embryo; 

endosperm copious, oily. 

5 genera, - 16 species. 

Composition: 2 genera, 47 species. 

Distribution: Eastern temperate and tropical China to northeastern India, Malaya, the 

western East Indies; southeastern United States. 

ILLICIACEAE (Figure 12d,e).-Shrubs or small 

trees; xylem vessel perforation plates scalariform 

with numerous bars (to 150); leaves simple, alternate or subverticillate, entire, exstipulate; flowers solitary, axillary or supra-axillary, rarely cauliflorous, bisexual, the parts hypogynous; perianth homoiochlamydeous, the parts free, 7-33, 

spirally arranged, the outmost small and brac- 
46 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 11 .-MYRISTICACEAE: a, Myristica surinamensis d inflorescence, bud, 6 flower, d flower 

with sepals removed, ,Vf. fragrans embryo and I.s. of seed; b, 0 flower, 1,s. of same, views of 

pistil, section of part of ovary with ovule, part of twig w,ith 0 inflorescence, I.s. of fruit, seed 

with aril (after Martius, 1840- 1906). CANELLACEAE: c, Cinnamodendron axillare floral diagram, 

twig in bud, bud, 1.s. of same, petal and sepal; d, essential organs, C.S. of same, C. corticosum 

fruit, 1,s. and C.S. of same, seed, I.s. of same showing the minute embryo, embryo (after Martius, 

1840-1 906). 
NUMBER 58 47 

C 

d 

e 

FIGURE I 2.-SCHISANDRACEAE: a, Schisandra coccinea d flowering twig, androecium and d floral 

diagram, 1,s. of 0 flower, 1,s. of pistil, gynoecium, seed and 1,s. of sanie showing the minute 

embryo, fruitlets; b, Kadsura japonica d flower, 1,s. of same, androecium, stamen; c, calyx, 

cluster of fruitlets, a fruitlet, 1,s. of same (after Baillon, 1866-1895; Le Maout and Decaisne, 

1873; Lindley, 1853). ILLICIACEAE: d, Illiciurn micranthurn flowering twigs, 1. anisaturn flower, 

1. rnicranthurn flower, inner petal and 3 stamens, stamen; e, I. anisaturn gynoecium, dehiscent 

fruit, seed, 1,s. of sanie showing the minute embryo (after Martius, 1840-1906; Dunn). 
48 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

teole-like, inner ones becoming larger and 

ligulate or suborbicular, the innermost often 

reduced; stamens usually numerous, 4-50; anthers oblong or as long as wide, introrse, basifixed or adnate, the thecas sometimes separate, 

the connective sometimes glandular, but not or 

only slightly apically produced; pollen 3- 

colp(oid)ate; pistils 5-1 5 (-2 1) in 1 whorl, compressed laterally, narrowed into a style, the 

stigma decurrent ventrally, the ovule ventralsubbasal, bitegmic, crassinucellar, anatropous; 

fruit a follicetum, seeds glossy, the embryo minute, 0.06-0. l the length of the endosperm; 

cotyledons hardly differentiated to 0.5 the length 

of the embryo; endosperm copious, fleshy. 

Composition: 1 genus, -40 species. 

Distribution: Japan, Korea, eastern China to 

Malaya, Borneo, and the Philippines; southeastern United States, eastern Mexico, West Indies. 

AMBORELLACEAE (Figure 1 3c,d).-Shrubs; xylem vesselless; leaves alternate, entire, or repand 

and obscurely toothed, finally distichous, exstipulate; inflorescences axillary d panicle, ? cyme 

or fascicle, the flowers small, unisexual, or sometimes ? flowers with 2-3 stamens, the parts hypogynous to weakly perigynous, perianth homoiochlamydeous, the parts 8, spirally arranged, 

weakly connate basally; stamens about 18, in 

several whorls, broad, the outer ones adnate to 

the perianth at its base; anthers extrorse, the 

thecas dehiscing longitudinally, pollen nonaperturate; pistils 5-6, incompletely closed, in 1 

whorl, the stigmas decurrent ventrally, the ovule 

1 , bitegmic, crassinucellar, orthotropous(?); fruitlets compressed drupelets, stipitate; seed small, 

the embryo minute, not differentiated, 0.1 the 

length of the endosperm; endosperm copious. 

Composition: 1 genus, 1 species. 

Distribution: New Caledonia. 

TRIMENIACEAE (Figure 1 3a,b).-Trees, 

shrubs, or climbers; xylem vessel perforation 

plates scalariform with more than 20 bars; leaves 

simple, alternate or opposite, entire or crenate, 

coriaceous, sometimes glandular punctate, exstipulate; inflorescences axillary or terminal racemes or panicles; flowers bisexual or the plants 

sometimes dioecious, small, numerous, the parts 

hypogynous; sepals 6 or 4, or an indefinite number of bracteoles and sepals; petals 0; stamens 6- 

22 in 1-3 whorls, hardly differentiated into anther and filament, or anthers oblong, basifixed, 

as long as the filament, extrorse, the connective 

apically produced, staminodes sometimes present; pollen 8-12 pored, or with 2 irregular, 

unthickened areas; pistil 1 (-2); stigma sessile, 

capitate, papillose; fruit a berry or drupelet; 

seeds compressed, the embryo small, the endosperm copious, gelatinous. 

Composition: 2 genera, 5 species. 

Distribution: New Guinea, New Caledonia, 

Fiji, and eastern Australia. 

MONIMIACEAE (Figure 14a-c).-Mostly 

shrubs, rarely trees or climbers, with a characteristic aromatic fragrance; xylem vessel perforation 

plates usually predominantly scalariform with 

more than 20 bars (to 100 in Atherosperma), 

sometimes predominately simple, rarely only simple; leaves simple, opposite, rarely alternate, entire or serrate, coriaceous, pellucid-punctate, exstipulate; inflorescences axillary, rarely terminal, 

cyme or raceme, or the flowers rarely solitary; 

flowers bisexual or the plants dioecious or monoecious; perianth parts in 2 or more series or 

acyclic, homoiochlamydeous or the outer sepaloid and the inner petaloid, 4-20 or rarely 0, 

hypogynous to perigynous, the calyx oblique like 

Aristolochia in Glossocalyx; stamens numerous, to 

- 150, or few, rarely 1, in 1-2 series, usually free, 

rarely connate (Tetrasynandra), the filaments 

very short, often flattened, sometimes with a 

gland on each side at the base; anthers adnate or 

basifixed, dehiscing by longitudinal slits or by 

valves from the base upwards, introrse or extrorse, the connective sometimes apically produced, staminodes sometimes present; pollen 

nonaperturate or 2(-3)-sulculate; disk adnate to 

the concave receptacle; pistils numerous, to 

-100, several, or rarely 1, rarely connate or 

adnate to the receptable, the style elongated or 

short, rarely subbasal, the stigma apical; ovule 1, 

apical, basal or marginal, 1 -2-tegmic, crassinucellar, anatropous (orthotropous in Daphnan- 
NUMBER 58 49 

FIGURE 1 Q.-TRIMENIACEAE: a, Trirnenia papuana flowering twig, flower bud, outer, middle 

and inner tepals, flower with tepals removed, stamen, pistil; b, Is. and C.S. of pistil, fruit, T. 

weinmannii$olia 1,s. of bud (d ?), stamen (after Gilg and Schlechter, Perkins and Gilg). AMBORELLACEAE: c, Amborella trichopoda d bud, 6 flower from above and below, I.s. of 6 flower, views 

of stamen; d, part of twig with d inflorescence, infructescence, fruitlet, 1,s. of same showing the 

minute embryo (after Perkins and Gilg). 

dra?); fruitlets drupes or achenes enclosed in or 

borne on a fleshy receptacle; seed may have a 

very fleshy testa (Baldoa), an aril rarely present 

(Siparuna); embryo minute, straight, 0.2-0.5 the 

length of the endosperm; cotyledons erect or 

divaricate, thin, 1-4 times the width of the radicle, 0.3-0.75 the length of the embryo; endosperm copious, oily. 

Composition: -30 genera, -450 species. 

Distribution: Mainly tropics and subtropics of 

the Southern Hemisphere; tropical to south temperate America; Sri Lanka, Malaya, and the East 
50 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 14.-MONIMIACEAE: a, Atherosperma moschata 1,s. of? flower, a pistil, 6 flower, dehiscent 

stamen, ,Mollinedia sancta ? flower, 1.s. of same; b, iMonimia citrina 1,s. of 6 flower, stamen, 

Atherosperma moschata flowering twig, Siparuna apiosyce part of 6 flowering twig; c, d flower, 

views of open, closed and open anthers, I.s. of ? flower, pistil with ovary opened, part of twig 

with 0 flowers and fruit, 1,s. of fruit, S. oligandra 1,s. of drupelet showing the minute embryo, 

embryo enlarged (after Baillon, 1866-1 893; Martius, 1840-1906). 

Indies; eastern Australia, New Zealand; relatively 

poorly represented in tropical rain forest of western Africa and in eastern Africa and Madagascar. 

IDIOSPERMACEAE (Figure 15a,b).-Trees; xylem vessel perforation plates scalariform; leaves 

simple, entire, opposite, coriaceous, exstipulate; 

inflorescence axillary, of 1 (-3) flowers; flowers 

bisexual, the perianth parts 30-40, perigynous, 

spirally arranged, petaloid, the receptacle urceolate; stamens 13-15 the inner ones staminodal, 

the filaments very short; anthers extrorse, laminar; pollen dicolpate; pistil 1 (-2), unicarpellate, 

at the base of the receptacle, the stigma broad, 

subsessile, fleshy, papillose; ovules 1-2, basal; 

fruits 1 -2-seeded nutlets enclosed within the 

somewhat fleshy receptacle; embryo large, the 

cotyledons 3-4, massive, peltate, the radicle 

short, invested or shortly exserted, the plumule 

well developed; endosperm 0. 

Composition: 1 genus, 1 species. 

Distribution: Rain forest of northern Queensland, Australia. 

CALYCANTHACEAE (Figure 15c,d).-Shrubs; 

xylem vessel diameter very small, the perforation 

plates simple; leaves simple, entire, opposite, exstipulate; flowers bisexual, solitary, axillary or 

terminal on axillary shoots, fragrant, the parts 

perigynous, the receptacle urceolate; perianth 

homoiochlamydeous to incompletely heterochlamydeous, the parts numerous, petaloid, spi- 
NUMBER 58 51 

FIGURE 1 .~.--IDIOSPERMACEAE: a, Idiospermum australiense 1,s. of flower, the outer tepals fallen, 

outer and inner tepals and C.S. of same, stamen and C.S. of same; 6, 1.s. of tepal, stamen and 

staniinode, stigma, 1,s. of pistil, embryo showing 4 massive cotyledons (after Blake). CALYCANTHACEAE: c, Caljcanthusf7oridus flowering twig, 1,s. of flower, stamen, I.s. of ovary, C. occidentalis 

flower with perianth removed, I.s. of same; d, Chimonanthus praecox floral diagram, bud, seed, 

C.S. of same showing the convolute embryo (after Baillon, 1866-1895; Le Maout and Decaisne, 

1873). 

rally arranged with gradual transition in size; 

stamens 5-30, the inner ones staminodal (7-17) 

with fleshy apex eaten by beetles, the filaments 

very short; anthers large, adnate, extrorse, the 

connective apically produced; pollen 2 (-3?)-sulculate; pistils about 3-20, at the base of the 

receptacle, the styles long, filiform, the stigmas 

obtuse, apical; ovules 1-2, ventral, bitegmic, sub- 
52 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

crassinucellar, anatropous; fruitlets achenes enclosed in the enlarged, slightly fleshy receptacle; 

embryo large, the cotyledons foliose, convolute, 

much broader than the radicle, 0.7 the length of 

the embryo; endosperm 0 or very scanty. 

Composition: 2 genera, 6 species. 

Distribution: Temperate China; southeastern 

and western United States. 

LAURALES 

Trees and shrubs, rarely twining parasitic 

herbs; xylem vessel perforation plates scalariform 

or simple; leaves usually alternate, entire or 

lobed, simple or rarely digitately compound, exstipulate; flowers small, bisexual or unisexual; 

perianth usually whorled, rarely spirally arranged, frequently 3-merous, usually perigynous, 

more rarely hypogynous or epigynous; stamens 

3-12 (-36) in one or more whorls, the filaments 

with a gland on each side at the base, the anthers 

opening by valves; pollen nonaperturate; pistil 1, 

pseudomonomerous, the stigma usually apical or 

subapical; ovules 1 per ovary or locule; fruit a 

berry, drupe or nutlet, rarely a samara; embryo 

medium-sized to large; endosperm usually absent 

rarely copious. 

Distribution: Centered in southeastern Asia 

and tropical America, in lowland and montane 

rain forest. 

Laurales are rich in characteristic 

aromatic oils, such as monoterpenes and sesquiterpenes. They are cyanogenic. 

GOMORTEGACEAE (Figure 1 Ga,b).-Tree; xylem vessel perforation plates scalariform, usually 

with numerous bars; leaves simple, opposite, entire, coriaceous, exstipulate; inflorescences axillary and terminal racemes or panicles; flowers 

small, bisexual, the parts epigynous; perianth 

homoiochlamydeous with 6- 10 spirally arranged 

parts; stamens 9 (2-ll), the filaments short to 

somewhat elongated, the inner ones with a gland 

on each side, sometimes 3 staminodes present; 

anthers introrse, opening by valves; pollen nonaperturate; pistil 1, the carpels 2-3, the style 1 

with 2-3 stigmatic lobes; ovary 2-3-locular, the 

Chemistry: 

ovules 1 per locule, axile-subapical; fruit a drupe; 

embryo medium-sized, 0.2 the length of the endosperm; cotyledons 3 times the width of the 

radicle, 0.7 the length of the embryo; endosperm 

copious, oily. 

Composition: 1 genus, 1 species. 

Distribution: Central Chile. 

LAURACEAE (Figure 1 Gc,d).-Trees or shrubs, 

very rarely twining parasitic herbs; xylem vessel perforation plates simple, occasionally a few 

sporadic scalariform; leaves simple, alternate, 

rarely opposite or subopposite, evergreen or deciduous, usually entire, rarely lobed, exstipulate; 

inflorescences axillary, rarely terminal, cymes or 

racemes, umbelliform or paniculate, rarely 

heads; flowers small, usually trimerous, rarely 

dimerous, greenish or yellowish, bisexual or the 

plants sometimes polygamous or dioecious; perianth in 2 or 3 trimerous (dimerous) whorls or 

very rarely absent, the parts hypogynous or perigynous, sometimes basally connate; stamens 3- 

12 (-36) in 1-4 whorls, filaments often with a 

gland on each side of the base, staminodes commonly present; anthers basifixed, dehiscing by 

valves from the base upward, very rarely by 

minute pores (Hexapora); pollen nonaperturate; 

annular disk in Synandrodaphne (Ocotea); pistil 1, 

unicarpellate (derived from 3 carpels), the ovary 

superior, very rarely inferior (Hypodaphnis), the 

style elongated or absent, the stigma 1 (2-3) 

FIGURE 16.-GOMORTEGACEAE: a, Gomortega keule twigs 

with inflorescence and infructescence, 1,s. of bud, Is. of 

flower, flower; b, essential organs, stamen and staminodes, 

views of stamen, 1,s. of fruit showing the minute embryo, 1,s. 

of seed (after Baillon, 1866-1 895; Muhoz Pizarro, 1959). 

LAURACEAE:; c, Cryptocarya moschata flower laid open, views 

of stamen, fruit, I.s. of same showing seed, views of Yz the 

embryo; d, h’ectandra gardneri calyx from below, flower, 

pistil, Persea erythropus dehiscent stamen, pistil, floral diagram, Cryptocarya moschata part of flowering twig (after 

Martius). HERNANDIACEAE: e, Hernandia sonora 1,s. of d 

flower, 6 flo\ver without the perianth, 1,s. of 0 flower, 0 

flower without the perianth, fruit;f, H. jamaicensis fruit and 

1,s. of same, H. catalpifolia inflorescence in axil of leaf, 1,s. of 

0 flower, the perianth partly removed, stamen with 1 anther 

sac dehiscent, 0 flower (after Baillon, 1866-1895; Martius, 

1840-1906; Fawcett and Rendle). 
NUMBER 58 53 
54 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

apical, ovule 1 subapical, bitegmic, crassinucellar, 

anatropous; fruit a berry or drupe, sometimes 

partly or entirely enclosed by the enlarged, cupular receptacle; embryo large, straight, the 

cotyledons thick, plano-convex, 9 times the width 

of the radicle, 0.8-0.9 the length of the embryo, 

radicle invested, plumule sometimes evident; endosperm 0. 

Composition: 32 genera, -2250 species. 

Distribution: Centered in southeastern Asia, 

the East Indies, and tropical America, in lowland 

and montane rain forest; less commonly in Africa, the Mediterranean region, Australia, New 

Zealand, and temperate North America. 

HERNANDIACEAE (Figure 1 6e,f).-Trees or 

shrubs, sometimes scandent; xylem vessel perforation plates predominantly simple, occasionally 

scalariform; leaves alternate, entire or lobed, simple or digitately compound, exstipulate; inflorescence axillary cyme, thyrse or corymb; flowers 

small, bisexual, or sometimes the plants monoecious, polygamous, or dioecious, the parts epigynous; perianth parts 3-10 in 1-2 whorls; stamens 3-6 in 1 whorl, sometimes 1-2 whorls of 

staminodes present, the filaments sometimes with 

a gland or two at their base; anthers basifixed, 

dehiscing by valves; pollen nonaperturate; pistil 

1, unicarpellate (derived from 3 carpels), the 

style elongate, stigma enlarged, subapical, the 

ovary inferior, ovule 1, subapical, bitegmic, crassinucellar, anatropous; fruit a nutlet enclosed by 

the enlarged receptacle, or a samara; embryo 

large, the cotyledons large, planoconvex or flat 

and spirally twisted around the radicle; endosperm 0. 

Composition: 4 genera, -65 species. 

Distribution: Most commonly along seacoasts; 

tropical Central America, Venezuela, and Colombia, Africa, southeastern Asia, the East Indies, 

northeastern Australia. 

ARISTOLOCHIALES 

The order is monotypic. 

Chemistry: Aromatic oils and aporphines are 

common. Chemical evidence does not strongly 

favor a relation with Nepenthaceae. 

ARISTOLOCHIACEAE (Figure 1 7a-c).--Shrubs 

and herbs, usually climbing, the tissues containing oil cells with a characteristic spicy fragrance 

similar to that of members of the Magnoliales 

and Laurales; xylem vessel perforation plates 

simple; leaves alternate, simple, mostly entire, 

exstipulate; inflorescence axillary, terminal or 

cauliflorous, the flowers solitary or in racemes; 

flowers bisexual, often fetid, the parts perigynous 

or epigynous; sepals often connate in a petaloid 

symmetrical or asymmetrical tube sometimes 3- 

lobed of the apex; petals usually 0, sometimes 3; 

stamens (4-) 6-40 in 1 or 2 whorls, the filaments 

short, free or more often connate in a tube 

adnate to the style; anthers free or connate and 

adnate to the style; pollen nonaperturate or 1- 

sulcate; pistils 6, or 1 and then the carpels 6 (3- 

5), the styles free or connate, short, the stigmatic 

lobes 6 (3-25); ovary inferior or semi-inferior, 

the locules 6 (3-5) sometimes incomplete, the 

ovules numerous, axile, bitegmic, crassinucellar, 

anatropous; fruit a capsule or follicetum, dehiscing from the base upward or along the ventral 

sutures, or rarely an indehiscent, ribbed nutlet; 

seeds usually numerous; embryo minute, linear, 

0.05-0.2 the length of the endosperm, the cotyledons 0.4 the length of the embryo and as wide 

as the radicle; endosperm copious, fleshy or subhorny. 

Composition: 7 genera, -500 species. 

Distribution: Mainly tropical, but extending 

into temperate regions; America, Eurasia, the 

East Indies, Africa. 

RANWNCULALES 

Usually shrubs and herbs, sometimes climbing; 

xylem vessel perforation plates usually simple, 

occasionally scalariform with few bars; leaves alternate, frequently compound, lobed, dissected 

or dentate, exstipulate; flowers large to small, 

bisexual or unisexual, the parts hypogynous; perianth 3-5-merous in whorls, or indefinite and 

spirally arranged; stamens numerous and spirally 

arranged, less frequently 6 and whorled; anthers 

basifixed, the connective sometimes apically produced; pollen usually 3-colpate or -colporate, 
NUMBER 58 55 

FIGURE 17.-ARISTOLOCHIACEAE: a, Aristolochta acutzfolaa flowering twig, 1,s. of base of perianth 

showing stigmas and androecium, stigmas and androecium enlarged, I.s. of same, C.S. of 

androecium, C.S. of ovary; b, flower, Saruma henryi flower, petal, calyx tube and carpels, 1,s. of 

fruit; c, Aristolochia cymbqera seed, dehiscent fruit (after Martius, 1840-1 906; Oliver). 

more rarely multiporate or nonaperturate; pistils 

usually indefinite in number, often numerous but 

sometimes only 1 and unicarpellate, spirally arranged or whorled, the style elongate or short, 

stigma decurrent ventrally or apical, the ovules 

1 -many; fruits achenes, drupelets, berries, rarely 

follicles or a capsule; embryo usually small and 

linear; endosperm usually copious, rarely scanty 

or absent. 

Distribution: Ranunculaceae cosmopolitan, 

centered in temperate and cold regions, the 

other families are centered in temperate Asia 

except Menispermaceae, which are most abundant in tropical rain forest. 

Chemistry: Aromatic oils are absent. They 

have very few terpenoid substances; Magnoliales 

have many. Aporphines are common. The alkaloids suggest relationship between Ranunculales 

and Magnoliales; several groups of alkaloids occur in both. The Ranunculales, however, have 

some groups that have not been detected in the 

Magnoliales. The alkaloids also suggest relation 

with Aristolochiales, Piperales, Papaverales, and 

Rutales. Morphine occurs in Menispermaceae 

and Lauraceae as well as Papaveraceae. Nandina 

is the only genus outside the Papaveraceae 

known with certainty to have the alkaloid protopine. In other respects it seems to fit well in 

the Berberidaceae. Nandina and Thalictrum have 

p-hydroxy-nandilonitril glucoside. Ranunculales 

and Magnoliales have phenolic esters and ethers, 

the latter order being richer than the former. 

Laticifers occur in the fruit of Decaisnea. 

MENISPERMACEAE (Figure 1 Sd-i).-Usually 
56 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 58 57 

lianas, rarely small trees or herbs; xylem vessel 

perforation plates simple; leaves alternate, usually simple, rarely trifoliolate, sometimes palmately lobed, not glandular, exstipulate; inflorescences axillary or cauliflorous cymes, thyrse, 

raceme, fascicle, or rarely the flowers solitary; 

plants dioecious, rarely monoecious (Albertisia), 

the flowers small, usually trimerous, the parts 

hypogynous; perianth sometimes homoiochlaniydeous; sepals free 6 (1- 12) in 1-4 whorls, or 

sometimes spirally arranged, increasing in size 

from the outside inward; petals free 6 (0-9), 

often in 2 whorls, usually smaller than the sepals, 

minute, rarely connate; stamens small, 6 or 3 (2- 

24) opposite the petals when 6 or 3, the filaments 

sometimes connate; anthers often 4-lobed, small, 

as long as wide, basifixed, sometimes connate, 

the connective not apically produced; pollen 3- 

colporate, occasionally (2-)3-colpate, 4-6- 

rug(or)ate, or nonaperturate; pistils 3 or 6 (1- 

32) in one or more whorls, the style mostly short, 

stigma apical; ovules 2 or 1, ventral, 1-2-tegmic, 

crassinucellar, anatropous, semi-anatropous, 

campylotropous or amphitropous; fruit drupelets, the seed often hippocrepiform; embryo linear or sometimes spatulate, arcuate, rarely spirally contorted (Spirospermum), as long as the 

endosperm; cotyledons semi-terete or flat, 0.4- 

FIGURE 18.-SARGENTODOXACEAE: a, Sargentodoxa CUneata 

part of 6 inflorescence, 6 flower, same with part of calyx 

removed, 1,s. of 6 flower with sepals removed (2 of the glandlike petals, 3 stamens and the pistillodes are visible), views 

of a stamen with glandlike petal at its base: b, 0 flower, I.s. 

of same with perianth removed, 0 flower with perianth 

removed, 1,s. of a pistil, top-view of a seed and same in I.s. 

showing the minute embryo, 1,s. of a fruitlet, embryo, part 

of 0 infloresence: c, part of an infructescence showing the 

fruitlets from 1 gynoecium (after Stapf). MENISPERMACEAE: 

d, Cissampelos fasciculata 0 inflorescence, C. tamoides d inflorescence, C. fasciculata 1,s. of fruit showing large curved 

embryo, seed: e, Sciadotaenia microphyllum diagram of d 

flower, corolla and androeciuni, androecium; J; Hyperbaena 

domingensis 6 bud, flower at anthesis, flower with a perianth 

segment depressed, flower from below: g, corolla and staniens from above, views of stamen, floral diagram; h, 0 calyx 

from below, sepal, floral diagram; i, bud, petal, flower at 

anthesis, calyx and bracteole from below, corolla and pistils, 

pistils and staminodes (after Martius, 1840-1906). 

0.97 the length of the embryo; endosperm moderate, scanty or absent, fleshy. 

Composition: 65 genera, -375 species. 

Distribution: Centered in tropical rain forest, 

few temperate; eastern and southern United 

States to subtropical South America; tropical and 

south temperate Africa, Madagascar; Japan to 

India, the East Indies, northern and eastern Australia. 

SARGENTODOXACEAE (Figure 1 8a-c).-Lianas; 

xylem fibers primitive; leaves alternate mostly 

trifoliolate, sometimes 3-lobed or entire, deciduous, exstipulate; inflorescence axillary racemes; 

plants dioecious, the flowers small, the parts hypogynous; sepals 6, in 2 whorls; petals 6, squamiform, glandular; stamens 6, in 2 whorls opposite 

the petals, the filaments short, free; anthers oblong, the connective shortly apically produced; 

pollen (2-)3-colporoidate; 0 torus elongate; pistils more than 35, spirally arranged, ovule 1; 

fruitlets stipitate berries; embryo minute, 0.3 the 

length of the endosperm, subcylindrical, the 

cotyledons not broadened, 0.5 the length of the 

embryo, the radicle slightly thicker than the 

cotyledons; endosperm copious, fleshy, oily and 

starchy. 

Composition: 1 genus, 1 species. 

Distribution: China and Indochina. 

LARDIZABALACEAE (Figure 19a, b).-Lianas, 

rarely erect shrubs; xylem vessel perforation 

plates simple, occasionally a few scalariform, or 

exclusively scalariform (Decaisnea), with few bars; 

leaves alternate, usually digitately compound, 

rarely 2-3-ternate or pinnately compound, exstipulate; inflorescence axillary racemes or the 

flowers solitary; flowers small to moderate-sized, 

the parts hypogynous; plants polygamous, monoecious or dioecious, the flowers of each sex with 

rudiments of the other sex; perianth homoiochlamydeous; sepals 3 or 6, petaloid; petals 6 or 

0, smaller than the sepals, sometimes 2 whorls of 

3 honey-scales between the perianth and stamens; stamens 6, the filaments short, free or 

connate in a tube or column; anthers basifixed, 

extrorse, free or connate, oblong, the connective 

sometimes apically produced; pollen as a rule 3- 

colp(oroiod)ate; pistils 3 or 6 (-15), in 1-5 
58 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

d 

C 

e 
NUMBER 58 59 

whorls, the stigma sessile, apical; ovules parietal 

or ventral, numerous in 2 or more ranks, or 

solitary, bitegmic, crassinucellar, anatropous, 

campylotropous or orthotropous; fruitlets berries or fleshy follicles; embryo small, linear, 

straight, 0.1-0.3 the length of the endosperm, 

the cotyledons thin, 0.3 the length of the embryo; 

endosperm copious, fleshy. 

Composition: 8 genera, 35 species. 

Distribution: Japan, Korea to the Himalayas, 

to southeastern China, Indochina, Taiwan; central Chile. 

NANDINACEAE (Figure 1 9c-e).-Shrub; xylem 

vessel perforation plates simple; leaves alternate, 

2-3 times pinnately compound, exstipulate; inflorescence a terminal panicle; flowers small, bisexual, the parts hypogynous; sepals numerous, 

spirally arranged, gradually larger from the 

outer to the innermost; petals 6 slightly larger 

than the inner sepals, honey-scales 6; stamens 6 

in 1 whorl opposite the petals; anthers elliptic, 

subsessile, introrse, dehiscing longitudinally, the 

connective slightly apically produced; pollen fossaperturate; pistil 1, unicarpellate, the ovary gradually narrowed into a short style, stigma apical; 

ovule 1, bitegmic, anatropous; fruit a berry; embryo small, linear, 0.3 the length of the endosperm, the cotyledons hardly differentiated; endosperm copious. 

Composition: 1 genus, 1 species. 

Distribution: China and Japan. 

FIGURE 1 9.-LARDIZABALACEAE: a, Lardizabala biternata 

floral diagrams of 13 and P flowers, d flower, pistil, fruit, C.S. 

of ovary, 1,s. of seed showing the minute embryo; b, Akebia 

pinata d flower and I.s. of same, flowering twig, 0 flower 

and 1,s. of same, Holboellia latifolia I.s. of d flower, I.s. of P 

floiver. floral diagram of? flower (after Baillon, 1866-1895; 

Le Xlaout and Decaisne. 1873). NANDINACEAE: c, h’andina 

domestica flower, leaf and infructescence, floral diagram; d, 

infructescence, inflorescence in bud, leaf; e, fruit, flower 

bud, pistil, 1,s. of same, views of stamen and androecium 

(after Steward, 1958; Hutchinson, 1973). BERBERIDACEAE: 

f, Berberis laurina bud, perianth from below, views of stamen 

before and at anthesis, infructescence; g, petal, pistil, I.s. of 

same, 1,s. of seed showing different views of the embryo, 

fruit, 1,s. of same, C.S. of same showing 2 seeds (after Martius 

1840- 1906). 

BERBERIDACEAE (Figure 1 9f,g).-Shrubs and 

herbs; xylem vessel perforation plates simple, 

occasionally scalariform (Berberis) with few bars; 

vascular bundles sometimes scattered in the pith; 

leaves alternate, entire or serrate, pinnately compound, trifoliolate, simple, or pinnately 1 -3-ternate, exstipulate, rarely stipulate (Epimedium); 

inflorescences cymes, racemes, spikes, fascicles, 

panicles or the flowers solitary; flowers bisexual, 

the parts hypogynous; sepals 6 (4), free; petals 6 

(4), free, the perianth trimerous in 4 whorls or 

rarely dimerous, very rarely absent; nectaries on 

the base of 6 petals; stamens 6 (4-15), free, 

opposite the petals; anthers basifixed or adnate, 

dehiscing by valves, the connective not apically 

produced; pollen basically 3-colpate, spiraperturate? in Berberis, 6- 1 2-rugate in Ranzaniu; pistil 

1, unicarpellate (derived from 2-3 carpels!), the 

style short, stigma apical; ovules 1 -several, basal, 

bitegmic, crassinucellar, anatropous; fruit a 

berry, rarely a capsule dehiscing by a transverse 

slit, very rarely a nutlet; pericarp sometimes disappearing before the seeds ripen; seeds sometimes arillate; embryo minute and roundish, linear or spatulate, 0.07 to as long as the endosperm; cotyledons undifferentiated to 0.7 the 

length of the embryo, sometimes thin, 1-3 times 

as broad as the radicle; endosperm copious, 

fleshy, or horny. 

Composition: 

Distribution: Centered in temperate North 

America and Eurasia, extending through Central 

America and along the Andes to the southern tip 

of South America; northern Africa, the highlands of northeast Africa; southeastern Asia to 

Borneo. 

PODOPHYLLACEAE (Figure POu,b).-Herbs; xylem vessel perforation plates simple(?); leaves simple, subopposite, palmately lobed and serrate, 

exstipulate; inflorescence terminal cymes, fascicles, subumbels, racemes, spikes, panicles, or the 

flowers solitary; perianth parts 12 or 18 in 2 or 

3 whorls, heterochlamydeous, honey-scales absent; stamens 12 (1 8), twice as many as the petals, 

the filaments elongated; anthers basifixed, dehiscing longitudinally by slits, the connective very 

-1 3 genera, -500 species. 
60 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 58 61 

FIGURE 2 1 .-PAEONIACEAE: a, Paeonia officinalis I.s. of flower, calyx and pistils, stamen, P. 

moutan I.s. of pistil, dehiscent fruitlets one with 1 seed; b, P. officinalis floral diagram, fruit, 1,s. 

of seed showing the minute embryo (after Baillon, 1866-1 895; Le Maout and Decaisne, 1873). 

shortly apically produced; pollen 3-colpate (brevicolpate); pistil 1, unicarpellate, the ovules numerous, lateral, bitegmic, crassinucellar, anatropous; fruit a berry; embryo small, linear, 0.4 the 

length of the endosperm; endosperm copious, 

fleshy. 

Composition: 2 genera, 9 species. 

Distribution: Himalayas to China and Taiwan; 

eastern North America. 

PAEONIACEAE (Figure 2 la,b).-Herbs or 

shrubs; xylem vessel perforation plates scalariFIGURE ~~.--PODOPHYLLACEAE: a, Podophyllum versipelle 

fruit, leaf and inflorescence, P. peltatum floral diagram. b, P. 

versipelle views of anther, 1,s. of seed showing the minute 

embryo, C.S. of ovary (after Baillon, 1866-1895; Oliver). 

CIRCAEASTERACEAE: c, Circaeaster agrestis monandrous and 

diandrous flowers, plant in fruit; d, 1,s. of pistil, stamen; e, 

embryo, 1,s. of fruit, monocarpellary fruit; f, dicarpellary 

fruit, leaf (after Oliver). RANUNCULACEAE: g, Anemone sellowii 

form with 2-5 bars; sometimes vascular bundles 

present in the pith; leaves alternate, biternate, 

with sublobed segments, exstipulate; flowers almost always solitary, bisexual, large, heterochlamydeous, the parts hypogynous; sepals 5, subfoliaceous; petals usually 5(- 10); stamens numerous 

(45-75), free, spirally arranged, the filaments 

elongated; anthers oblong, basifixed, extrorse, 

dehiscing by longitudinal slits; pollen 3-colpor(oid)ate; glandular disk external to the pistils; 

pistils 2-5, large, the ovary tapering gradually 

into a short style, the stigma decurrent ventrally; 

ovules numerous, ventral, bitegmic, crassinucellar, anatropous; fruitlets follicular, the seeds 

large; embryo small, 0.1-0.2, the length of the 

endosperm, the cotyledons not broadened, 0.25 

the length of the embryo; endosperm copious. 

Composition: 1 genus, 33 species. 

Distribution: Temperate Eurasia, especially 

gynoecium, stigma, 1,s. of ovary; h, Ranunculus apiijiolius 

flower, sepal and petal, views of stamen, pistil, fruit, views 

of fruitlet and 1,s. of same showing seed; i, Clematis dioica 0 

flower, 8 flower, 1,s. of fruitlet showing the minute embryo, 

embryo enlarged, Trollius europaeus 1,s. of pistil, Delphinium 

consolida dehiscent fruit, Eranthis hyemalis fruit, Nigella satiua fruit (after Martius, 1840-1906; Schnizlein). 

China; southern Europe; northwestern United 

States. 

RANUNCULACEAE (Figure POg-i).-Herbs, 

rarely small shrubs or climbers; xylem vessel 

perforation plates simple, rarely also scalariform 

(Hydrastis); vessels frequently in several circles or 
62 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

* irregularly scattered; leaves alternate and also 

basal, or rarely opposite, simple, serrate-crenate, 

sometimes lobed or compound, almost always 

exstipulate; inflorescence a cyme, raceme, panicle, thyrse, or the flowers solitary; flowers bisexual or rarely the plants dioecious or monoecious, 

the parts hypogynous; sepals (3-) 5-8, sometimes 

petaloid, rarely spurred; petals 5 (0-12) often 

with subbasal nectar gland(s), sometimes spurred, 

sometimes honey-scales between the perianth 

and stamens, rarely nectar secreted by the carpels 

(Calthu); stamens usually numerous (5- 1 lo), 

free, rarely the outer staminodal and petaloid; 

anthers usually small, about as long as wide, 

basifixed, the connective rarely apically produced, extrorse or introrse, dehiscing longitudinally by slits; pollen often 3-colpate, sometimes 

multiporate or nonaperturate; pistils usually numerous (to -250) and spirally arranged, more 

rarely few and whorled, rarely only 1, rarely 

partially connate, the style short to elongated, 

the stigma decurrent ventrally or apical; ovules 

numerous to 1, ventral, 1-2-tegmic, crassinucellar or tenuinucellar, anatropous or semi-anatropous; fruitlets achenes, follicles or rarely a berry, 

drupelets or capsule; embryo minute and 

rounded to linear, 0.07-0.3 the length of the 

endosperm; cotyledons undifferentiated to 0.6 

the length of the embryo and 1.5 times the width 

of the radicle; endosperm copious, fleshy. 

Composition: 

Distribution: Cosmopolitan, but centered in 

temperate and cold regions of the Northern and 

Southern Hemispheres, in various habitats but 

mainly moist; few tropical (Clematis spp.). 

CIRCAEASTERACEAE (Figure 2Oc-f).-Small 

annual herb; xylem vessel perforation plates simple; leaves simple, dentate, the venation open 

and dichotomous, exstipulate, arranged in a 

rosette at the apex of the stem; flowers solitary 

in the upper leaf axils, bisexual, the parts hypogynous; sepals 2 (-3), valvate, persistent; petals 0; 

stamens 2 (1-3); anthers small, as long as wide, 

basifixed, the thecas separated by connective, not 

apically produced, dehiscing longitudinally; pollen 3-colpate; pistils 1-3, unicarpellate, the style 

50 genera, - 1900 species. 

very short, stigma slightly elongated, the ovary 

uniovulate, the ovule unitegmic, tenuicellar(?), 

orthotropous; fruit an achene with uncinate prickles, the seeds minute; embryo linear, terete, 

straight, 0.3 the length of the endosperm; cotyledons not broadened, 0.2 the length of the 

embryo, linear-oblong in the seedling; endosperm copious. 

Composition: 1 genus, 1 species. 

Distribution: Temperate northwestern Himalayas to northwestern China. 

HALORAGALES 

Herbs, aquatic or of damp localities; xylem 

vessel perforation plates simple, the vascular tissue sometimes reduced to a weak axile bundle; 

leaves alternate or opposite, simple, often 

toothed, sometimes entire, usually small, stipulate or exstipulate; flowers minute, bisexual or 

unisexual; sepals 0 or 2-4; petals 0 or 2-4; stamens usually 1 or 2, more rarely to 8, in 1 or 2 

whorls, the anthers basifixed; pollen 3-, 5-7- 

colpate, 3-5-rupoidate, 3-5-por(or)ate, 3-6-colpoidate or nonaperturate; pistil 1, the carpels 1- 

2(-4), styles elongate and stigmas decurrent or 

stigmas subsessile; ovary usually inferior, 1-4- 

locular, the ovules 1 per locule; fruit 1-4 nutlets, 

drupelets, or mericarps; embryo minute to linear; endosperm usually scanty, rarely copious. 

Distribution: Often cosmopolitan, most common in temperate regions, but some tropical and 

others in cold regions. 

Gunneru stands apart from Haloragaceae. Hzppuris seems chemically rather out 

of place in Myrtales. It has stachyose not reported 

in Myrtales and lacks ellagic acid so characteristic 

of that order. It has been suggested that the 

Callitrichaceae are related to the Lamiaceae, but 

the latter are rich in terperioids, which are absent 

from the former. 

HALORAGACEAE (Figure 22a,b).-Terrestrial 

or submerged herbs; xylem vessel perforation 

plates simple; leaves simple, small, opposite, alternate or whorled, serrate, the submerged 

leaves pectinately divided, exstipulate; flowers 

Chemistry: 
NUMBER 58 63 

FIGURE 22.-HALORAGACEAE: a, Haloragis erecta flowering shoot, floral diagram, ? flower, 1,s. 

of same, Laurembergia coccinea flower; b, views of stamen, C.S. of anther, inflorescence, I.s. of 

bisexual flower the anthers removed, L. tetrandra fruit (after Le Maout and Decaisne, 1873; 

Martius, 1840-1906). GUNNERACEAE: c, Gunnera petaloidea part of inflorescence and leaf, G. 

chilensis part of inflorescence, G. scabra 6 flowers; d, G. petaloidea bisexual flower, petal; e, 

stamen, G. scabra fruit, 1,s. of same, 1,s. of seed showing the minute embryo, embryo enlarged 

(after Baillon, 1866-1895; Le Maout and Decaisne, 1873). 
64 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

solitary or in fascicles and axillary, or in terminal 

spikes, racemes, or panicles, minute, bisexual or 

the plants monoecious; calyx-tube adnate to the 

ovary; sepals 2-4 or 0; petals 2-4 or 0; stamens 

2-8, in 1 or 2 whorls; anthers basifixed, the 

connective sometimes apically produced; pollen 

5-'7-colpate, 3-5-rupoidate, 3-5-por(or)ate; pistil 1, the carpels 2-4, stigmas 2-4 subsessile, 

papillose or plumose; ovary inferior, 3-4-locular, 

the ovules 1 per locule, axile-subapical, bitegmic, 

crassinucellar, anatropous or semi-anatropous; 

embryo sac of the Polygonum type; fruit 1-4 

nutlets; embryo linear, straight, terete, as long as 

the endosperm, the cotyledons 0.1-0.3 the 

length of the embryo; endosperm moderate or 

scanty, fleshy. 

Composition: 

Distribution: Cosmopolitan, especially Australia. 

GUNNERACEAE (figure 22c-e).-Terrestrial or 

aquatic herbs, sometimes symbiotic with Nostoc; 

stems sometimes polystelic; xylem vessel perforation plates simple; leaves simple, small to very 

large, lobed and serrate, all radical; stipules large, 

axillary; inflorescence scapose, the flowers minute, often in dense clusters, bisexual or the 

plants monoecious or dioecious; sepals 2; petals 

0; stamens 2, the anthers subsessile, basifixed, 

the connective not apically produced; pollen 3- 

colpate; pistil 1, bicarpellate; the styles 2, linear, 

stigmatic along most of their length; ovary inferior, unilocular, the ovule 1, apical, bitegmic, 

crassinucellar, anatropous, embryo sac tetrasporic of the 16 nucleate Peperomia type; fruit a 

drupelet or nutlet; embryo minute, 0.1 the 

length of the endosperm, the cotyledons not 

broadened, 0.4 the length of the embryo; endosperm copious. 

6 genera, - 120 species. 

Composition: 1 genus, -35 species. 

Distribution: Tropical and temperate; Mexico 

to Chile, including Juan Fernandez, reaching 

high in the Andes; Malaysia, Tasmania, New 

Zealand, Solomon Islands, Hawaii; tropical and 

South Africa. 

HIPPURIDACEAE (Figure 23a,b).-Emergent 

aquatic herbs; vascular bundles concentric or an 

axile strand; leaves simple, linear, entire, veticilFIGURE 23.-HIPPURIDACEAE: a, Hippuris vulgaris flowering 

shoot, floral diagram, 0 flower, pistil, stamen: b, I.s. of 

bisexual flower, fruit, 1,s. of same (after Le Maout and 

Decaisne, 1873). CALLITRICHACEAE: c, Callitrich verna fruiting plant, C. dejlexa leaf, C. verna 1,s. of young ? flower, C. 

dejlexa stamen: d, C. verna S flower, 0 flower with prophylls, 

C. dejlexa fruit: e, C.S. of fruit showing embryo and endosperm (after Martius, 1840-1 906). 

late, exstipulate; flowers minute, solitary in the 

axils of the leaves, bisexual or rarely the plants 

monoecious or polygamous; perianth 0 or calyx 

merely a ring on the floral axis; stamen 1, epig- 
NUMBER 58 65 

ynous, the anther large, bilobed, basifixed; pollen 

4-6-colpoidate(-colpate-colporoidate?); pistil 1, 

unicarpellate, the style subulate, stigmatic all 

along one side for its entire length; ovary inferior, the ovule 1, apical, unitegmic, tenuinucellar, anatropous; fruit an achene or drupelet; embryo linear, 0.9 to as long as the endosperm, the 

cotyledons 0.1-0.5 the length of the embryo; 

endosperm scanty. 

Composition: 1 genus, 2 species. 

Distribution: Temperate and arctic North 

America and Eurasia; southern tip of South 

America. 

CALLITRICHACEAE (Figure 23c-e).-Small terrestrial or aquatic herbs, stems slender; vascular 

system a weak axile bundle; leaves simple, opposite, linear, entire, exstipulate; plants monoecious, the flowers minute, solitary or 8 and 0 in 

the same axil; perianth 0; stamen 1, subtended 

by 2 bracteoles, the filament filiform; anther 

small, as long as wide, basifixed; pollen nonaperturate or 3 (-4) colpoidate(?); pistil 1, bicarpellate, the styles 2, filiform, papillose along most 

of their length; ovary superior, 4-lobed, 4-locular, the ovules 1 per locule, axile, unitegmic, 

tenuinucellar, anatropous; fruit a schizocarp of 2 

or 4 mericarps; embryo linear, terete, slightly 

curved, as long as the endosperm, the cotyledons 

0.1-0.3 the length of the embryo; endosperm 

scanty, fleshy. 

Composition: 1 genus, -25 species. 

Distribution: Almost cosmopolitan, centered 

in the temperate zones. 

SARRACENIALES 

The order is monotypic. 

Chemistry: Gibbs considers that it is clear that 

the Sarraceniaceae and Nepenthaceae are chemically alike, but that the Droseraceae differ from 

them in important respects. 

SARRACENIACEAE (Figure 24a-d).-Insectivorous herbs of bogs; rhizome xylem vessel perforation plates scalariform; leaves pitcher-shaped 

or tubular, usually hooded, radical; inflorescence 

scapose, a few-flowered raceme or the flower 

solitary; flowers bisexual, the parts hypogynous; 

sepals 5 (3-6), free, imbricate, persistent, often 

colored; petals 5 (4-6) or 0, free; stamens -80- 

12, free, the filaments considerably shorter than 

the pistil; anthers about as long as wide, dorsifixed, dehiscing longitudinally; pollen 5-9-colporoidate; pistil 1, the carpels 3 or 5 (-6), the 

style 1, 5-rayed toward the apex, or umbrella 

shaped, or shortly 3-lobed apically, the stigmas 

at the tips of the umbrella ribs or apical and 

papillose on the style arms; ovary 3 or 5(-6)- 

locular, the ovules numerous, axile, unitegmic or 

also bitegmic (?) (Darlingtonia), tenuinucellar, 

anatropous; fruit a loculicidal capsule, the seeds 

small, numerous; embryo linear, straight, 0.1- 

0.8 the length of the endosperm; cotyledons not 

broadened, 0.2-0.5 the length of the embryo; 

endosperm copious, fleshy. 

Composition: 3 genera, 17 species. 

Distribution: Eastern North America; California and Oregon; Guayana Highlands. 

NEPENTHALES 

The order is monotypic. 

NEPENTHACEAE (Figure 24e,J).-Herbs and 

shrubs, often climbing; xylem vessel perforation 

plates simple, medullary and cortical bundles 

present in some species; leaves simple, entire, 

very rarely denticulate, some developing into 

insectivorous lidded ascidia, exstipulate; inflorescences terminal or axillary elongate racemes, 

sometimes with cincinnal branches, the plants 

dioecious; flowers small, actinomorphic; sepals 4 

(3-6), usually imbricate in pairs, free or rarely 

basally connate, nectariferous on the inner surface; petals 0; stamens 8-24 (4-6) uniseriate or 

subbiseriate, the filaments connate into a column, the anthers 2-celled, apical, extrorse, about 

as long as wide or oblong, dehiscing longitudinally; pollen in tetrads, the apertures obscure, 

sexine spinuliferous; disk 0; pistil 1, the carpels 

4 (3), the 4 (3) stigmas simple or bifid, sometimes 

capitate, apical, papillose, sessile or subsessile; 

ovary superior, 4( 3)-locular, the ovules numerous, axile-septal, bitegmic, crassinucellar, anatropous; fruit a leathery loculicidal capsule, the 

seeds numerous, minute, usually bicaudate; em- 
66 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

h 

FIGURE ~~.--SARRACENIACEAE: a, Sarracenia purpurea flower and scape, pitcher, Is. of flower, 

pistil and 2 of the stamens, 1,s. of pistil, floral diagram; b, dehiscent fruit, C.S. of same, seed and 

1,s. of same. c, Heliamphora nutans androecium, stamen, pistil, C.S. of ovary, seed, I.s. of seed, 

embryo; d, Darlingtonia calqornica essential organs, C.S. of ovary, seed and I.s. of same, (after 

Le Maout and Decaisne, 1873; Wunschmann). NEPENTHACEAE: e, Nepenthes sp. leaf terminating 

in a pitcher, 1L’. gracilis 0 inflorescence, N. distillatoria d flower, P flower, 1,s. of pistil, h‘epenthes 

sp. dehiscent fruit, valve of same;f, seed, 1,s. of same, d floral diagram, 0 floral diagram (after 

Le ,Maout and Decaisne, 1873). 
NUMBER 58 67 

bryo straight, cylindrical, 0.8-0.9 the length of 

the endosperm; cotyledons linear, 0.4-0.8 the 

length of the embryo; endosperm copious, fleshy 

or starchy. 

Composition: 1 genus, -70 species. 

Distribution: Wet tropical jungle; centered in 

Borneo, extending to southern China, Indochina, northern Australia, New Caledonia; AsSam, Sri Lanka; Madagascar. 

PIPERALES 

Herbs, shrubs, and small trees, the stems often 

jointed and the tissues aromatic; xylem vessel 

perforation plates scalariform with very numerous bars to simple, rarely vessels absent; leaves 

alternate or opposite, simple, entire or toothed, 

stipulate or exstipulate; flowers minute or small, 

bisexual or unisexual; perianth absent or very 

rarely 3 sepals present; stamens 1-10, rarely in 

2 whorls of 3 each, the anthers basifixed; pollen 

1 -sulcate, ulceroid or nonaperturate; pistils 1-4, 

the stigmas decurrent ventrally or a single sessile 

tuft; ovary 1-4-locular, the ovules 1-8 per placenta, axile or parietal; fruit drupelets, rarely 

small follicles or a ventricidal capsule; embryo 

minute; perisperm or endosperm copious. 

Most frequently in the understory of tropical rain forest, often in other wet 

habitats. 

Piperales are rich in characteristic 

aromatic oils such as monoterpenes and sesquiterpenes, probably present in secretory cells. 

‘They have phenolic esters and ethers and a peppery taste. Alkaloids are prominent in Piperaceae 

but not in the other members of the order. 

Myristicin occurs in Macropiper. 

SAURURACEAE (Figure 25a,b).-Perennial 

herbs of wet places, the stems jointed; xylem 

vessel plate perforations not clearly differentiated; tissues containing oil cells that give the 

plant a characteristic spicy fragrance; leaves alternate, simple, entire, the stipules united to the 

petiole; inflorescence terminal, a dense or loose 

spike or raceme, the upper leaves sometimes 

forming a white involucre; flowers small, bisexDistribution: 

Chemistry: 

ual; perianth absent; stamens 3-8, free or adnate 

to the ovary, the filaments longer than the anther; anther slightly longer than wide, basifixed, 

the connective sometimes slightly apically produced; pollen 1 -sulcate, occasionally faintly trichotomosulcate; pistils 3-4 sometimes connate 

basally or more completely, the styles free, the 

stigmas decurrent ventrally; ovary unilocular 

with parietal placentas or 3-4-locular with axile 

placentas, the ovules 1-8 per placenta, bitegmic, 

crassinucellar or tenuinucellar, orthotropous; 

fruit dehiscing into cocci, or a fleshy capsule 

dehiscing ventrally at the apex; embryo minute, 

0.1 the length of the perisperm; cotyledons not 

differentiated; perisperm copious, starchy; endosperm scanty. 

Composition: 5 genera, 7 species. 

Distribution: Temperate and subtropical; 

United States, Mexico; Japan, China, Indochina, 

Philippines. 

PIPERACEAE (Figure 25c).-Herbs, shrubs, or 

small trees, sometimes scandent, often with swollen nodes, the tissues containing oil cells which 

give the plant a characteristic pungent taste and 

odor; vascular bundles f scattered as in the monocots; xylem vessel perforation plates simple, or 

occasionally scalariform with few bars (Piper); 

leaves simple, usually alternate, sometimes opposite or verticillate, entire, often the main veins 

extending from the base of the leaf nearly to the 

apex, or the leaves succulent and the veins inconspicuous; stipules united to the petiole or absent; 

inflorescence leaf-opposed or axillary dense 

spike, or umbellate spikes, rarely a raceme (Ottonia); flowers minute, bisexual or unisexual; perianth absent; stamens 1-10 hypogynous (to perigynous?), the filaments short; anthers basifixed, 

the sacs united or spreading at the base; pollen 

f distinctly 1 -sulcate (nonaperturate in Peperomia); pistil 1 with up to 5 short styles, the stigmas 

decurrent ventrally, or stigma a single sessile tuft; 

ovary unilocular, ovule 1, 1-2-tegmic, crassinucellar, orthotropous; embryo sac with 8-16 nuclei, or up to 100 antipodals; fruit a drupelet; 

embryo minute, 0.04-0.2 the length of the perisperm, the cotyledons 0.4 the length of the 
68 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 25.-sAURURACEAE: a, Saururus cernuus flowering stem, floral diagram, flower, I.s. of 

same, 1,s. of fruit showing the minute embryo, endosperm and perisperm; b, Houttuynia cordata 

infloresence, flower, I.s. of same, Gyrnnotheca chinensis flower, C.S. of ovary (after Baillon, 1866- 

1895; Le Maout and Decaisne, 1873). PIPERACEAE: c, Piper nigrum 3 flowers, floral diagram 

and 1s. of fruit showing the minute embryo, endosperm and perisperm, P. cubeba fruiting twig, 

Peperomia blanda flowering stem, flower and 1,s. of same (after Baillon, 1866-1 895). 

embryo; perisperm copious, starchy; endosperm 

scanty. 

East Indies, Australia, New Zealand; the woody 

species are especially Asiatic, the herbaceous (PeLACTORIDACEAE (Figure 26a,b).-Shrubs with 

swollen nodes; xylem vessel perforation plates 

simple; leaves alternate, simple, distichous, small, 

Composition: 6 genera, - 1,500 species. peromia) American. 

Distribution: Mostly in the understory of rain 

forest in tropical America, the West Indies and 

Africa; eastern Africa; Japan, China, India, the 
NUMBER 58 69 

v e 

FIGURE %.-LACTORIDACEAE: a, Lactorzs fernandezzana flowering twig, base of blade and 

stipule, floral diagram, flower, 1,s. of a carpel, the same cut open dorsally, C.S. of a carpel; b, 

anther, dehiscent fruit, dehiscent segment of same, seed, I.s. of same showing the minute 

embryo (after Hooker, Engler). CHLORANTHACEAE: c, Sarcandra chloranthoides part of flowering 

and fruiting twigs, part of inflorescence enlarged, flower further enlarged, pistil, 1,s. of same, 

1,s. of fruit showing the minute embryo; d, Hedyosmum arborescens d infloresence, stamen, 0 

flower, I.s. of same, I.s. of fruit showing the minute embryo; e, Chloranthus tnconspzcuus 1,s. of 

flower, stamens, C.S. of same, 1,s. of fruit showing the minute embryo, embryo enlarged (after 

Baillon, 1866-1895; Le Maout and Decaisne, 1873; Wight). 

emarginate, glandular-punctate, the stipules petals 0; stamens in 2 whorls of 3 each: anthers 

large, membranous, united to the petiole, and subsessile, extrorse, the connective apically prosurrounding the twig; plant polygamo-monoe- duced: pollen with 1 distal ulceroid aperture; 

cious; inflorescence axillary with 1-5 flowers; pistils 3, the ovary grading into a small style, 

flowers small, the parts hypogynous; sepals 3; stigmatic ventrally near the apex; ovules ventral- 
70 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

basal, 6-8 per pistil, bitegmic, tenuinucellar(?), 

anatropous; fruitlets follicles, 4-6 seeded, embryo minute, 0.1 the length of the endosperm; 

endosperm copious, oily. 

Composition: 1 genus, 1 species. 

Distribution: Juan Fernandez Islands of Chile. 

CHLORANTHACEAE (Figure 2 6c-e). --Shrubs, 

trees, and herbs with aromatic tissues; xylem 

vessel perforation plates scalariform with very 

numerous bars, to 100 or more, or vessels absent; 

leaves opposite or subwhorled, simple, crenate 

or serrate, the stipules small; inflorescence spikes, 

heads, panicles or cymes; flowers minute, bisexual or pseudohermaphrodite or the plant monoecious or dioecious; 6 perianth absent; stamens 

1, or 3 with the middle anther bilocular and the 

laterals unilocular, the connective sometimes apically produced; stamens connate to one another 

and adnate to the ovary, or 50-250 free stamens 

in a Hedyosmum flower (inflorescence ?); pollen 

nonaperturate, 1 -sulcate or polycolpoidate; 0 calyx adnate to the ovary and minutely 3 dentate, 

or absent; pistil 1, the styles 3, very short, or the 

stigma sessile and not divided; ovary unilocular, 

the ovule 1, apical, bitegmic, crassinucellar, orthotropous; fruit a drupelet; embryo minute, 

0.06-0.1 the length of the endosperm, the cotyledons divaricate, 0.4 the length of the embryo; 

endosperm copious, oily. 

Composition: 5 genera, 65 species. 

Distribution: Tropical and subtropical, from 

low elevations to high in mountains; America, 

Japan, China, India, the East Indies, New Zealand. 

NYMPHAEALES 

Aquatic herbs; xylem vessels absent, the tracheids long in Nymphaeaceae; leaves usually alternate, often large, floating or emergent, stipulate 

or exstipulate; flowers minute to very large, usually bisexual; sepals 3-6; petals 3 to -25 or rarely 

absent; stamens 5 to -120, whorled or spirally 

arranged, the filaments sometimes laminar; anthers basifixed or adnate, elongate; pollen 1-4- 

sulcate, more rarely 3-colpate or nonaperturate; 

pistils 1 to -25, the carpels 1-35, style long and 

the stigma decurrent ventrally or the stigmas 

sessile and linear or apical; ovules I-many, usually scattered over the ovary wall; fruit usually a 

berry or capsular syncarp, sometimes a nutlet; 

embryo minute or large; perisperm copious or 

absent. 

Distribution: Almost cosmopolitan, but absent from cold regions. 

Chemistry: Benzylisoquinoline occurs in Nelumbo, Magnoliales, Ranunulales, Fagara and one 

Croton. Aporphine occurs in Nelumbo, Magnoliales, and Ranunculales. 

CERATOPHYLLACEAE (Figure 27a,b).-Plants 

submerged herbs, rootless; xylem vessels absent; 

leaves simple, verticillate, dichotomously divided 

into linear serrulate segments, exstipulate; plants 

monoecious; flowers minute, axillary, solitary; 

sepals (bractlets?) 8-1 5, basally connate in one 

whorl, hypogynous; petals 0; stamens 5-27, spirally arranged; anthers oblong, subsessile, adnate, the connective apically produced; pollen 

nonaperturate; pistil 1, unicarpellate, the style 

fairly long, the stigma decurrent ventrally; ovule 

1, unitegmic, crassinucellar, orthotropous; fruit 

a nutlet; embryo large, the cotyledons oblong, 

the radicle shorter than the cotyledons, the plumule well developed; endosperm 0. 

Composition: 1 genus, -5 species. 

Distribution: Cosmopolitan except in the 

colder parts of the Northern Hemisphere. 

CABOMBACEAE (Figure 27c,d).-Aquatic herbs 

with laticiferous tubes or sacs in the parenchyma; 

xylem vessels absent; floating leaves alternate, 

peltate, entire, simple, the submerged ones opposite and finely dissected, exstipulate; flowers 

axillary, solitary, bisexual, the parts hypogynous; 

sepals 3, petaloid; petals 3; stamens 3-18, cyclic, 

the filaments longer than the anthers; anthers 

extrorse, about twice as long as wide, basifixed, 

the connective not apically produced; pollen 1- 

sulcate; pistils 1-18, the ovary grading into a 

short style, the stigma apical; ovules 1-3, bitegmic, crassinucellar, orthotropous, lamina1 

(distant from the ventral suture); fruit nutlets; 

embryo minute, 0.07 the length of the per- 
NUMBER 58 71 

n 

b 

FIGURE 27.-cERATOPHYLLACEAE: a, Ceratophyllum demersum habit, leaf, 8 flower, stamen, ? 

flower; b, fruit, embryo laid open, I.s. of pistil (after Martius, 1840-1906). CABOMBACEAE: c, 

Cabomba aquatica flowering shoot, floral diagram, C. piauhyensis flower, petal, stamen, pistils: 

d, 0’. aquatica pistil laid open, C. piauhyensis fruitlets, C. aquatica I.s. of seed showing the minute 

embryo, endosperm and perisperm (after Baillon, 1866- 1895; Martius). 

isperm, the cotyledons slightly differentiated; NY MPHAEACEAE (Figure 2 8a-c). -Aquatic 

perisperm copious, starchy; endosperm scanty. herbs with laticiferous tubes or sacs in the 

parenchyma and scattered vascular bundles; xylem vessels absent, tracheids long; leaves simple, 

radical, alternate, large, floating, peltate or cordate, entire or margin sinuate, sometimes stipuComposition: 2 genera, 7 species. 

Distribution: Eastern North America to Argentina; eastern Asia, India, Australia; formerly 

in Europe. 
72 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

\ 

I 

FIGURE 28.-NYMPHAEACEAE: a, Nymphaea ampla flower and part of a leaf, flower from below; 

b, inner, middle and outer stamen, 2 carpels viewed from above; c, Nuphar luteum floral 

diagram, flower, I.s. of same, fruit, I.s. of seed showing the minute embryo, endosperm and 

perisperm (after Martius, 1840-1 906). 

late; flowers solitary, bisexual, large, often fra- scale-like (Nuphar), rarely absent (Ondinea), spigrant, the parts hypogynous or perigynous; se- rally arranged, sometimes nectariferous and 

pals 4-6, free or adnate to the receptacle; petals grading into stamens; stamens (15-) 50-100, 

5 to -25, usually large, sometimes small and spirally arranged, the filaments often flattened; 
NCMBER 58 73 

anthers basifixed, the connective not apically 

produced; pollen f monocotyledonoid, 1 -4-sulcate or zonisulculate, or nonaperturate(?); pistil 

1, the carpels (3) 5-35, sometimes incompletely 

closed, the stigmas sessile, linear; ovary superior 

to inferior, the ovules numerous, parietal (scattered over the ovary walls), bitegmic, crassinucellar, anatropous; fruit a berry or capsular syncarp; embryo minute, 0.1-0.2 the length of the 

perisperm, the cotyledons undifferentiated; perisperm copious, starchy; endosperm scanty. 

Composition: 5 genera, -75 species. 

Distribution: Temperate North America to 

tropical South America; temperate and tropical 

Asia, Africa and Australia, New Guinea. 

NELUMBONACEAE (Figure 29a,b).-Aquatic 

herbs; xylem vessels absent; leaves simple, peltate, entire, floating or emergent, the stipule 

ochreate; flower solitary, the scape arising in the 

axil of a scale leaf, bisexual, the parts hypogynous; perianth entirely petaloid or sepals 4 and 

petals numerous (-20), spirally arranged; stamens numerous (1 00- 120), spirally arranged; 

anthers linear, basifixed, the connective apically 

produced; pollen usually 3-colpate; pistils usually 

numerous (-6-25) embedded separately in an 

obconic receptacle, but not adnate, the style 

short, stigma apical; ovary uniovulate, the ovule 

bitegmic, crassinucellar, anatropous; fruit an achene or nutlet; embryo large, the cotyledons thick, 

plano-convex, the radicle invested, the plumule 

foliaceous; perisperm and endosperm 0. 

Composition: 1 genus, 2 species. 

Distribution: Eastern United States, Mexico, 

West Indies, Guayana to Colombia, Iran to eastern Asia; northern Australia. 

PROTEALES 

The order is monotypic. 

Chemistry: Juglone occurs in Proteaceae. 

PROTEACEAE (Figure 29c-f.-Trees and 

shrubs, rarely herbs; xylem vessel perforation 

plates typically simple, a few species with an 

occasional scalariform plate with few bars; leaves 

alternate, rarely verticillate or opposite, simple, 

entire, lobed, to deeply dissected, exstipulate; 

inflorescences racemes, spikes, umbels or heads, 

rarely the flowers solitary, often subtended by an 

involucre of colored bracts; flowers usually 

bisexual, sometimes the plants dioecious or polygamous; sepals 4, free or basally connate, valvate, petaloid; petals 0; stamens 4, rarely 1 or 3 

of these staminodal, opposite to and inserted 

upon the sepals, rarely hypogynous, the filaments 

rarely connate apically (Symphyonema); anthers 

oblong, dithecal, rarely some monothecal, basifixed, adnate or dorsifixed, sometimes apiculate, 

rarely connate (Stirlingia); pollen 3(2-5)-porate, 

3-colpoidate, rarely 3-colpor(oid)ate(?); disk usually present, consisting of 4 (-2) alternisepalous 

glands or scales around the base of the ovary or 

gynophore, or sometimes connate; pistil 1, the 

carpel 1, style 1, long, filiform, the stigma apical 

or lateral near the apex of the style; gynophore 

sometimes present; ovary superior, unilocular, 

the placenta ventral, the ovules 1 or more, subbasal or subapical, bitegmic, the integuments developing very slowly and not enclosing the nucellus until after fertilization, crassinucellar, orthotropous, semi-anatropous or anatropous; fruit 

a nut, drupe or follicle; seeds often large, sometimes winged; embryo often large, the cotyledons 

sometimes 3-8, often unequal, thin or thick and 

plano-convex, 3-5.8 times wider than the radicle, 0.7-0.9 the length of the embryo and gradually grading into the radicle, the latter sometimes invested; endosperm 0 except in Bellendena. 

Composition: 

Distribution: 

62 genera, - 1,100 species. 

Most in warm, dry habitats, few 

in moist; centered in Australia and South Africa; 

Australia to New Zealand, the East lndies, Indochina, China, India, Japan; southern % of Africa, Madagascar; Mexico to temperate South 

America. 

BALANOPALES 

The order is monotypic. 

BALANOPACEAE (Figure 3Oa,b).-Trees or 

shrubs; terminal bud perulate; xylem vessel perforation plates scalariform, with numerous bars; 

leaves alternate, subopposite or subverticillate, 
74 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE L)g.-NELUMBONACEAE: a, Nelumbo lutea receptacle containing fruits, leaf and flower, 

1,s. of receptacle and pistils, stamen, pistil, 1,s. of same; b, 1,s. of fruit, embryo with 1 cotyledon 

removed to show the plumule, embryo laid open, plumule enlarged showing young leaves, N. 

nucqera floii,er without the perianth, 1,s. of same (after Le Maout and Decaisne, 1873; Baillon, 

1866- 1895). PROTEACEAE: c, Embothrium (Oreocallis) grandiflorum I.s. of bud, dehiscent fruit, 

1,s. of seed, floral diagram; d, Roupala rhombijolia bud, R. brasiliensis flower, calyx laid open to 

show insertion of stamens and hypogynous glands, 1,s. of ovary and hypogynous glands; e, fruit, 

seed, Persooniaferruginea fruit, same with upper half of fleshy part removed, I.s. of the stone 

showing the seed, Franklandiafucijoliu upper part of style with stigma, anther, 1,s. of flower, 

fruit with tuft of hair; f. Bellendena montana flower, Symphyonema montanum essential organs, 

Grevillea sp. 1,s. and C.S. of embryo (after Baillon, 1866-1895; Martius, 1840-1906; Le Maout 

and Decaisne, 1873; Endlicher; Engler; Schnizlein). 
NUMBER 58 75 

simple, entire or obscurely crenate apically, exstipulate; plants dioecious, the flowers small, perianth absent; 8 inflorescence in short, loose spikes 

below the leaves, the flower subtended by 2-6 

scales; stamens 3-6 (1 - 12); anthers subsessile, 

oblong-ellipsoid, the connective often slightly apically produced; pollen 3-4(-5)-colpoidate; 9 

flower solitary, subtended by numerous imbricate bracteoles; pistil 1, the carpels 3 (-2), the 

styles 3 (-2), subulate, each 2-partite nearly to 

the base, the stigmas decurrent; ovary superior, 

3(-2)-locular, sometimes incompletely so, each 

locule with 2 anatropous basal ovules; fruit an 

acorn-like drupe, nearly a nut, subtended by the 

involucre: pyrenes 1-2, each 1-seeded; embryo 

large, straight, spatulate, 0.85 the length of the 

endosperm, the cotyledons thin, plano-convex, 

0.8 the length of the embryo; endosperm scanty, 

fleshy. 

Composition: 1 genus, 10 species. 

Distribution: Northeastern Queensland, New 

Caledonia, Fiji. 

FAGALES 

Trees and shrubs, sometimes aromatic due to 

resinous glands; xylem vessel perforation plates 

scalariform or simple; leaves usually alternate, 

simple or compound, entire, toothed or lobed, 

often exstipulate; plants dioecious or monoecious, the flowers minute, usually anemophilous, 

sometimes entomophilous, usually in catkins, 

rarely heads or a panicle; sepals or calyx lobes 3- 

6 or 0; petals 0; stamens 3-40 (-105), the filament often short, sometimes filiform, inserted 

on a bract or flattened receptacle, the anthers 

usually basifixed, sometimes dorsifixed; pollen 

3-6 (7)-colpate or colporate, 3-16-porate to colporate, 3-pororate; pistil 1, the carpels 1-3 (6), 

with as many styles, the stigmas usually decurrent 

ventrally, more rarely punctiform; ovary superior or inferior, l-3-locular, the ovules 1-2 per 

locule; fruit a nut, drupe or samara, rarely a 

tardily dehiscent capsule; embryo large; endosperm absent or rarely scanty. 

Distribution: Primarily north temperate to 

tropical, more rarely south temperate or tropical, 

in wet to dry habitats. 

Chemical evidence is strongly in 

favor of relationship between Fagaceae and 

Betulaceae, including Corylus. Juglandaceae have 

amides and show the Betulaceae type of metabolism (show citrulin), but amides occur in a variety of families. Myricaceae are related to Betulaceae; both have members that develop bluegreen or green color in the NHs layer of the 

Juglone test. This reaction has rarely been observed elsewhere. Myrica fat is said to have large 

amounts of myristic acid, as does the Myristicaceae. 

SIMMONDSIACEAE (Figure 3 Oc-e). -Divaricate 

branched shrub; xylem vessel perforation plates 

simple; growth of stem in thickness by development of successive concentric rings of xylem and 

phloem, the rings separated by narrow circles of 

parenchyma; leaves opposite, simple, entire, exstipulate; plants dioecious; 8 flowers in capitate 

clusters; sepals 5 (4-6); corolla 0; stamens 10- 

12, inserted on a flat receptacle, the anthers 

subsessile, oblong; pollen 3-colpate; 9 flowers 

solitary, more rarely in a 2-7-flowered pendulous raceme; sepals 4-7, imbricate; corolla 0; 

pistil 1, the carpels 3, the styles 3, free, papillose, 

the stigma decurrent along the entire length of 

the style; ovary superior, 3-locular, the ovules 1 

per locule, subapical, bitegmic, crassinucellar, 

anatropous; fruit an ovoid loculicidal capsule, 

with a thick beech-fruit-like surface; embryo 

large, the cotyledons large, thick, the radicle 

short, invested; endosperm 0. 

Chemistry: 

Composition: 1 genus, 1 species. 

Distribution: Dry habitats; southwestern 

United States and northwestern Mexico. 

LEITNERIACEAE (Figure 30Jg).-Shrub; xylem 

vessel perforation plates simple; secretory canals 

with resinous contents in stem and leaves; leaves 

alternate, simple, entire, exstipulate; plants dioecious; 8 inflorescence axillary, cymose, pendulous catkin-like spikes; 8 flower: perianth absent; 

stamens 3-12, inserted on a bract; anthers subsessile, basifixed, slightly longer than wide; pollen 

3 (-6)-colporate; 9 inflorescence erect, a shorter 
76 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 58 77 

catkin than the 8; perianth of 4 (3-8) minute 

scales; pistil 1, the carpel 1, the style elongated, 

stigma decurrent all along the ventral surface; 

ovary superior, the ovule 1, ventral, integuments 

2 (l?), crassinucellar, amphitropous or subanatropous; fruit a drupe with scanty flesh and a stony 

endocarp; embryo large, straight, spatulate, 0.9 

the length of the endosperm, the cotyledons flat, 

fairly thick, 0.7 the length of the embryo; endosperm scanty, fleshy. 

Composition: 1 genus, 1 species. 

Distribution: Swampy areas; southern Missouri to Texas and Florida. 

MYRICACEAE (Figure 3 1 a,b).-Aromatic trees 

and shrubs; xylem vessel perforation plates exclusively scalariform or sometimes also simple; 

leaves alternate, simple, entire, dentate or pinnately lobed, with sessile, resinous glandular 

hairs, stipulate or exstipulate; plants monoecious, 

dioecious or polygamous; inflorescences axillary, 

simple, more rarely compound, catkin-like 

spikes; c3 flower without a perianth; stamens 4-8 

(2-16) inserted on a bract, the filaments short, 

sometimes a pair connate, the anthers basifixed, 

as long as wide; pollen 3-pororate; ? inflorescence 

erect, very short, the flowers sometimes provided 

with sepal-like bracteoles; pistil 1, carpels 2, the 

styles 2, filiform, the stigmas decurrent along 

ventral surface of styles; ovary superior, unilocular, the ovule basal, unitegmic, crassinucellar, 

orthotropous; fruit a drupelet or nutlet, often 

covered with waxy worts, the endocarp bony; 

embryo straight, the cotyledons thick, plano-convex, 3.5-3.7 times wider than the radicle, 0.7- 

FIGURE 3O.-BALANOPACEAE: a, Balanops vieillardi d catkin, 

d flower, 0 flower, 1,s. of 0 flower, pistil; b, part of fruiting 

twig, 1,s. of fruit (after Baillon, 1866-1895). SIMMONDSIACEAE: c, .‘jimmondsia chinensis d flowering twig, d flower, 

views of stamen, 1,s. of d flower; d, 0 flower, pistil; e, I.s. and 

(1,s. of pistil, dehiscent fruit with aborted ovules (seed not 

present), 0 flowering twig (after Le Maout and Decaisne, 

1873; Nuttall). LEITNERIACEAE:~, Leitneriafloridana d inflorescences, P inflorescences; g, I.s. of 0 inflorescence, 6 flower, 

P flower with bract and involucre, the style cut across, 1,s. of 

9 flower, fruiting twig, Is. in 2 planes and C.S. of fruit, 

embryo (after Sargent; Baillon, 1866-1 895). 

0.9 the length of the embryo, the radicle partly 

invested; endosperm 0. 

Composition: 3 genera, 40 species. 

Distribution: Temperate and subtropical, in 

wet and dry habitats; America, northeastern Siberia to China and India, Indochina, the East 

Indies, New Caledonia, tropical and southern 

Africa; northwestern Europe. 

JUGLANDACEAE (Figure 3 1 c-f).-Aromatic 

resinous trees lacking resin vessels; xylem vessel 

perforation plates usually simple, 2 genera have 

scalariform plates with few bars; sieve tubes 

sometimes with lateral sieve areas, recalling those 

of gymnosperms; leaves alternate or rarely opposite (Alfaroa, Engelhardtia spp.), pinnately 

compound, the leaflets serrate or rarely entire, 

with shortly stalked resin-secreting peltate 

glands, exstipulate; plants monoecious (dioecious 

in Engelhardtia spp.); c3 inflorescences usually catkins (few flowers in Alfaroa), erect or pendulous, 

1-8 in a cluster; 8 flower calyx 3-6-lobed or 

absent, adnate to a bract; stamens 3-40 (-105), 

the filaments short, inserted on the flattened 

receptacle; anthers basifixed, slightly longer than 

wide, the connective sometimes apically produced; pollen 3- 16-porate to colporate; ? inflorescence a spike or the flowers subsolitary, pendulous or erect; ? flower calyx 4-toothed, adnate 

to the ovary; pistil 1, the carpels 2 (3-4 in an 

occasional flower), the style usually with 2 large 

branches, plumose and stigmatic on the ventral 

surface; ovary inferior, 1 (-2) locular, the ovule 

1, basal, unitegmic, crassinucellar, orthotropous; 

fruit a nut or samara; embryo usually large, the 

cotyledons thick, broad, and often much contorted, 0.75-0.9 the length of the embryo and 

investing the radicle, the plumule evident, sometimes the cotyledons thin and folded toward the 

apex (Engel hardtia, Pterocarya); endosperm 0. 

Composition: 7 genera, -50 species. 

Distribution: Primarily north temperate and 

subtropical; United States to temperate South 

America, extending furthest south along the Andes; southeastern Europe to India, China, Indochina, the East Indies, Japan. 

RHOIPTELEACEAE (Figure 32a,b).-Tree; xylem vessel perforation plates exclusively scalari- 
78 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 58 79 

form with 4-1 1 bars; leaves alternate, pinnately 

compound, serrate, glandular beneath and aromatic, stipulate; inflorescences large, lax axillary 

and terminal panicles, the flowers in clusters of 

3 (1 bisexual and 2 sterile 0); sepals 4 (2 interior 

and 2 exterior), free; petals 0; stamens 6, the 

filaments shorter than the anthers; anthers basifixed, oblong, slightly longer than wide, the thecas separate at apex and base; pollen 3-colporate, 

brevicolpate, granorate; disk 0; pistil 1, the carpels 2, the styles 2, short; ovary superior, bilocular, one locule aborting, the other with 1 axile 

ovule; fruit a small samara; embryo straight, the 

cotyledons ellipsoid, thick, 2.7 times wider than 

the radicle, 0.7 the length of the embryo; endosperm 0. 

Composition: 1 genus, 1 species. 

Distribution: Southwestern China, Indochina. 

FAGACEAE (Figure 32c,d).-Trees and shrubs, 

the buds peru late; xylem vessel perforation plates 

predominantly scalariform to simple; leaves alternate, very rarely whorled, lobed, dentate, 

crenate or entire, rarely with sessile resinous 

glands (Castanea japonica), stipulate; plants monoecious, rarely dioecious (Nothofagus) or bisexual; 8 inflorescences usually catkin-like spikes or 

heads, 8 flower calyx 4--6(-7)-lobed; corolla 0; 

stamens 4-40, the filaments free, filiform, the 

anthers basifixed or dorsifixed; pollen 3-6(-7)- 

FIGURE 3 1 .-MYRICACEAE:: a, .Vyrica gale twig with 0 inflorescences, twig with 6 inflorescences, M. californica inner 

and outer vieivs of staminate flower with scale and bractlets, 

0 floiver with scale and bractlets, 1,s. of pistil, 1,s. and C.S. of 

fruit, 'M. asplenifolia 1,s. of 0 florver with scale and bractlets; 

b, ,M. aethiopica twig with 0 and 6 flowers, ,M. gale 1,s. of fruit, 

Comptonia sp. inner and outer views of 0 flower, I.s. of fruit, 

embryo (after Sargent; Raillon, 1866-1895; Engler; Le 

Maout and Decaisne). JUCLANDACEAE: c, Juglans regia 6 

inflorescence, 0 inflorescence, P flower, diagram and I.s. of 

same, 6 flower and 1,s. of same, stamen; d, fruit, same with 

husL renioved, seed, 1,s. of same; e, Carpa mpristicaeformis 

fruiting twig, Platyarqa strobilacea 0 flowers with staminodes, 1,s. of 0 flo\\,er, P strobilus; P. chinensis 0 flower, 6 

flower: f, fruit, 1,s. of same, Pterocarya fraxinifolia fruit, 

Engelhardtia spicata young fruit with 3-lobed bract (after 

Le Maout and Decaisne, 1873). 

colpate or colporate; 0 flowers solitary or in 

dichasia or spikes, 1-7 (or more) in an involucre; 

calyx 4-6-lobed, adnate to the ovary; corolla 0; 

pistil 1, the carpels 3 (6), the styles 3 (6), I' inear 

or filiform, the stigmas decurrent to punctiform; 

ovary inferior, the locules 3 (6), the ovules 2 per 

Iocule, axile, 1 -2-tegmic, crassinucellar, anatropous; fruit a nut, partly to completely enclosed 

in the involucre; embryo large, the cotyledons 

thick and plano-convex, or thin and folded, 0.8- 

0.9 the length of the embryo, the radicle invested; endosperm 0. 

Composition: 8 genera, -900 species. 

Distribution: Widespread in temperate and 

subtropical Northern Hemisphere; mainly in 

mountains in tropical regions; one genus occurs 

in New Guinea, southeastern Australia, New Zealand, and southwestern South America; Mediterranean region of Africa. 

BETULACEAE (Figure 32e-h).-Trees and 

shrubs, the buds perulate; xylem vessel perforation plates scalariform, sometimes with more 

than 20 bars; leaves alternate, simple, serrate, 

sometimes with sessile resinous glands, stipulate; 

plants monoecious; inflorescences pendent catkins or the ? sometimes an erect short dichasial 

spike; 8 flower: Calyx membranous, usually 4 (1 - 

5) partite or absent; stamens (1 -) 2-6, inserted 

on a bract, the filaments very short; anthers 

oblong, basifixed?, the thecas connate or separate; pollen often aspidote, 3(-7)-poroid; ? 

flower: perianth absent or adnate to the ovary; 

pistil 1, the carpels 2, the styles 2, filiform; ovary 

inferior or naked, bilocular, the ovules 2-1 per 

locule, axile, unitegmic, crassinucellar, anatropous; fruit a nutlet or samara; embryo straight, 

as long as the seed; cotyledons moderately to 

very thick and plano-convex, 4-5.3 times wider 

than the radicle, 0.8-0.9 the length of the embryo, the radicle sometimes invested; endosperm 

0. 

Composition: 

Distribution: 

6 genera, species - 150. 

Dry and wet habitats; temperate 

Northern Hemisphere, extending into the arctic 

and subtropical zone; Andes to Bolivia. 
80 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

h 

f 
NUMBER 58 

SALICALES 

The order is monotypic. 

SALICACEAE (Figure 33a,b).-Trees, shrubs, 

and subshrubs; xylem vessel perforation plates 

simple; leaves simple, deciduous, alternate, crenate, stipulate, the buds perulate; inflorescences 

erect or pendulous catkins, the plants dioecious, 

rarely individual specimens monoecious or the 

flowers bisexual; perianth absent; 6 flowers sometimes with a cupular disk, sometimes with 1-2 

small linear glands; stamens 2-3 (-60), the filaments free, rarely connate, filiform, long; anthers slightly longer than wide, basifixed or 

dorsifixed near the base; pollen non-aperturate, 

3-colporoidate or 3-colporate; 0 flowers with distinct glands or a cupule which is sometimes 

fleshy; pistil 1, the carpels 2 (-4), the styles 2 (- 

4), short or long, stigmatic ventrally; ovary 1- 

locular with 2-4 parietal or basal placentas, the 

ovules few -m, 1 (2)-tegmic, crassinucellar, anatropous; fruit a loculicidal 2-4-valved capsule; 

seeds small or minute, enveloped in fine hair; 

embryo straight, the cotyledons 3.3 times as wide 

as the radicle, 0.5-0.8 the length of the embryo; 

endosperm 0. 

Composition: 2 genera, -350 species. 

Distribution: Centered in temperate Northern Hemisphere, extending into the arctic zone; 

commonly in moist habitats and on mountains; 

few in the Southern Hemisphere, absent from 

Australia. 

81 

CASUA RINAL ES 

FIGURE 32.-RHOIPTELEACEAE: a, Rhoiptelea chiliantha hist:xual floi\.er, pistil, anther; b, fruit, C.S. of same, stipule, 

fruiting twig (after Hutchinson). FAGACEAE: c, Quercus robur 

twig with 0 inflorescences, 6 inflorescences, P flower, I.s. of 

same, d flower, fruit, I.s. of same, Fugus sylvatica C.S. of fruit, 

Castanea vulgaris flowering twig; d, 0 flower, 1,s. of same, d 

flower, Is. of same, Nothofagus clifortioides d flower, N. 

gunnii fruits, Castanea vulgaris d and 0 inflorescences (after 

Baillon, 1866-1893; Le Maout and Decaisne, 1873; Hooker 

and Hooker, 1837-1982). BETULACEAE: e, Corylus avellana 

tivig ivith 0 and 6 infloresences, 0 inflorescence, 1,s. of young 

fruit;f, 6 flower, fruits enveloped by accrescent bracteoles; 

g, Betula pumila scale with 3 d flowers, d flower, Alnus 

cordfolia scale with 2 0 flowers, 1,s. of fruit; h, Carpinus 

betulus stamen, bract with fruit, 1,s. of fruit (after Baillon, 

1866-1 895; Le Maout and Decaisne, 1873). 

The order is monotypic. 

Chemistry: Casuarinales have hinokiflavone, a 

biflavonyl reported elsewhere only in gymnosperms, but biflavonyls occur in several other 

unrelated families. Casuarina and gymnosperms 

have estolide waxes. 

CASUARINACEAE (Figure 3 3c-e).-Trees and 

shrubs with longitudinally grooved, jointed 

branchlets; xylem vessel perforation plates simple, occasionally also scalariform, with 2-30 bars; 

leaves scale-like, reduced to a 4- 16-toothed 

sheath; plants monoecious or dioecious; inflorescence terminal, the 6 usually a spike, the Q a 

head; perianth 0 (2 sepals in 6 flower?), the 

flowers bracteolate, in a cup-like involucre; stamen 1, the filament elongate at anthesis; anther 

about 1.5 times longer than wide, basifixed; pollen 3(2-5)-pororate; pistil l, the carpels 2, one 

aborting, the style with 2 filiform branches, 

stigma decurrent; ovary superior, initially 2- 

locular, unilocular at anthesis, ovules 2 (-4), collateral, axile, bitegmic, crassinucellar, orthotropous or k anatropous; bracts and bracteoles 

hardening into a cone; fruit samaras enclosed in 

woody bracteoles; embryo straight, the cotyledons thin, 3.2 times wider than the radicle, 0.8 

the length of the embryo; endosperm 0. 

Composition: 1 genus, -50 species. 

Distribution: Mainly in dry habitats; centered 

in Australia, extending to the East Indies, Malaya, New Caledonia, and Fiji. 

URTICALES 

Trees, shrubs, and herbs, sometimes with 

tough fibers or laticiferous; xylem vessel perforation plates simple, rarely also with a few bars; 

leaves usually alternate, simple, usually serrate, 

sometimes lobed, usually stipulate; flowers minute, usually anemophilous, the plants usually 

monoecious or dioecious, more rarely the flowers 

bisexual; sepals 0-5 (-8), sometimes connate; 

petals 0; stamens 4-1 0 (1- 12), the filaments short 

or elongate, anthers basifixed or dorsifixed, the 

connective sometimes apically produced; pollen 
82 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 33.-sALICACEAE: a, Sdix alba twig with 6 inflorescences, ? inflorescence, d flower, 1,s. 

of same, 1,s. of pistil, 0 flower, I.s. of seed; 6, S. humboldtiana dehiscent fruit, Populus nigra d 

flower, 1,s. of same, P. nivea 0 flower, 1,s. of same (after Baillon, 1866-1895; Martius, 1840- 

1906). CASUARINACEAE: c, Casuarina quadrivalvis d inflorescence, 0 inflorescences; d, d flower 

(bracts) with anther removed, C. nodzjlora diagram of 8 flower, C. angulata 1,s. of ovary; e, C. 

equisetifolia 0 flower with ovary opened to show ovules, Casuarina sp. fruiting strobilus, fruit 

cut to shot7 the seed, seed (after Baillon, 1866-1895; Le Maout and Decaisne, 1873). 
NUMBER 58 83 

3-colporate or -colpate, or 3 (2-1 5)-porate; pistils 

1-3, the carpels 1-3, styles 1-3, the stigmas 

decurrent ventrally or short or fimbriate; ovary 

superior, l(2)-locular, the ovules 1, more rarely 

2; fruit a nutlet or samara, more rarely a drupelet, achene or multiple fruit; embryo large, endosperm usually scanty or absent, rarely copious. 

Distribution: Cosmopolitan, most abundant in 

the tropics, in moist to dry habitats. 

Chemistry: Cardenolides and rubber are 

prominent in Moraceae. The Ehrlich test is + in 

Eucommiaceae, but - in the rest of the order; 

ellagic acid is present in Eucommiaceae but not 

in the rest of the order. Eucommia has polyisoprenes and gutta. It is less happily placed here 

than Moraceae, Ulmaceae, and Urticaceae. 

EUCOMMIACEAE (Figure 34a, b).-Trees with 

laticiferous cells; xylem vessel perforation plates 

simple, some scalariform in primary xylem (Solereder); leaves alternate, simple, serrate, exstipulate; plants dioecious; perianth 0; 8 inflorescence a few-flowered glomerule; flowers with 

10 (4- 12) stamens, the filaments very short, the 

anthers long, linear, basifixed, apically produced; 

pollen 3-colporoidate; ? flowers solitary, in the 

axil of a bract or bract-like leaf; pistil 1, the 

carpels 2, one aborting, the styles 2, stigmas 

decurrent ventrally; ovules 2, collateral, apical, 

unitegmic, crassinucellar or subtenuinucellar, 

anatropous; fruit a samara; embryo straight, as 

long as the endosperm; cotyledons thin, 1.5 times 

wider than the radicle, 0.7 the length of the 

embryo; endosperm copious. 

Composition: 1 genus, 1 species. 

Distribution: Montane forests of temperate 

China. 

BARBEYACEAE (Figure 34c,d).-Trees; xylem 

vessel perforation plates simple, phloem sieve 

plates compound; leaves opposite, simple, entire, 

exstipulate; plants dioecious; inflorescence a 

short cyme; 8 flower calyx of 3-4 nearly free, 

valvate sepals; petals 0; stamens 6-9 (1 0, 12), the 

filaments very short, anthers basifixed, twice as 

long as wide, the connective slightly produced 

apically; pollen 3-colpor(oid)ate; ? flower pistils 

1-3, sometimes slightly connate laterally; sepals 

3-4, accrescent; petals 0; ovary superior, style 

linear, the stigma plumose, decurrent all around, 

the ovule 1, subapical; fruit a nutlet; embryo 

straight, the cotyledons flattened, radicle short; 

endosperm 0. 

Composition: 1 genus, 1 species. 

Distribution: Northeastern Africa, Arabia. 

URTICACEAE (Figure 3 4e). -Herbs, shrubs, 

and soft-wooded trees, the stems often fibrous; 

xylem vessel perforation plates simple; laticiferous canals rarely present; sometimes possessing 

stinging hairs; leaves alternate or opposite, simple, usually serrate, cystoliths often abundant, 

usually stipulate; inflorescence axillary, cymose 

spikes or heads, the flowers minute; plants usually monoecious or dioecious, or rarely the flowers bisexual; calyx 4-5(0,2, 3)-lobed or the sepals 

free, often accrescent; petals 0; stamens 4-5 (1- 

3), the filaments elongate, inflexed in bud; anthers dorsifixed, slightly longer than wide; pollen 

3(2-15)-porate; pistil 1, the carpel 1, stigma often 

fimbriate, often sessile; ovary superior or inferior, the ovule 1, basal, bitegmic, crassinucellar, 

orthotropous or semi-anatropous; fruit an achene or drupelet; embryo straight, 0.9 to as long 

as the endosperm; cotyledons moderately thick, 

2.3-4.0 times wider than the radicle, 0.5-0.7 the 

length of the embryo; endosperm scanty, fleshy 

and oily. 

Composition: 45 genera, -800 species. 

Distribution: Worldwide; more genera and 

species in tropical than temperate regions. 

CANNABACEAE (Figure 34j).-Fibrous herbs, 

sometimes scandent; laticiferous or resiniferous 

canals present; xylem vessel perforation plates 

simple; leaves alternate or opposite, simple, serrate, sometimes palmately lobed, cystoliths present, stipulate; plants dioecious; inflorescence axillary, 8 cymose panicle, ? cymose spike; calyx 5- 

partite (8) or entire (0); petals 0; stamens 5, short; 

anthers erect in bud; pollen 3-porate; pistil 1, the 

carpels 2, styles 2, filiform, stigmas decurrent; 

ovary superior, unilocular, the ovule solitary, 

bitegmic, crassinucellar, anatropous, inserted 

midway in the locule; fruit an achene; embryo 

arcuate or coiled, 1.7-2.3 times longer than the 
84 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 58 85 

endosperm; cotyledons moderately thick, 2-3 

times wider than the radicle, 0.5-0.7 the length 

of the embryo; endosperm scanty or moderate, 

fleshy. 

Composition: 2 genera, 3 species. 

Distribution: Temperate and subtropical 

Northern Hemisphere. 

MORACEAE (Figure 35a-c).-Trees and 

shrubs, rarely herbs, laticiferous; xylem vessel 

perforation plates simple; leaves alternate, rarely 

opposite, simple, serrate or entire, sometimes 

lobed, cystoliths often present, stipulate; plants 

monoecious or dioecious; inflorescences axillary, 

cymose or dense spikes, umbel or head, the flowers minute; sepals 4 (8-0), free or connate, sometimes fleshy in fruit; petals 0; stamens usually 4 

(1-2), the filaments inflexed or straight in bud; 

anthers dorsifixed, as long as wide; pollen 2-4(- 

5)-porate; pistil 1, the carpels 2, one frequently 

aborting, the styles 2 (I), filiform, stigmas short 

or decurrent; ovary superior or inferior, unilocular, the ovule 1, usually apical, rarely basal, 

bitegmic, crassinucellar, anatropous, semianatropous or campylotropous; fruit achenes, nutlets 

or drupelets, often a multiple fruit, the receptacle 

sometimes fleshy; embryo often arcuate; 1.5-1.7 

times longer than the endosperm; cotyledons 

thin or moderately thick, 2-2.7 times wider than 

the radicle, 0.5-0.6 the length of the embryo; 

endosperm moderate or scanty, rarely 0, fleshy. 

Composition: 

Distribution: Mainly pantropical and -sub- 

-55 genera, - 1,400 species. 

tropical, few temperate. 

FIGURE 34.-EUCOMMIACEAE: a, Eucommia ulmoides 6 flowering twig, ? flowering twig, d flower and bracts, anther, ? 

flower and bract, I.s. of ? flower; b, fruits, upper part of 

fruit, I.s. of fruit, C.S. of seed, embryo (after Oliver). BARBEYACLAE: c, Barbeya oleoides d flowering twig, ? flowering 

twig, 0 flower; d, d flower, stamen, pistil, 1.s. of fruit, seed 

and C.S. of same, embryo (after Hutchinson, 1973; Penzig). 

URTICACEAE: e, Boehrneria arguta ? inflorescence, part of 

same enlarged, 0 flower with bracts, ? flower, B. caudata 

part ofd infloresence, 6 bud, d flower, stamen (after Martius, 

1840- 1906). CANNABACEAE: f, Cannabis sativa 0 flowering 

twig, 6 inflorescence, ? flower enclosed in its bract, ? flower, 

1,s. of same, 6 flower, fruit, 1,s. of same (after Baillon, 1866- 

1895). 

ULMACEAE (Figure 35d).-Trees and shrubs, 

not laticiferous; xylem vessel perforation plates 

simple, rarely also scalariform, with 1-6 bars 

(Planera); leaves alternate, rarely opposite, often 

asymmetrical, 2-ranked, serrate or entire, cystoliths present in a few species, stipulate; inflorescence axillary cymose fascicles or the 0 flower 

solitary; flowers bisexual or the plants monoecious; sepals 4-5 (3-8), free or connate; petals 0; 

stamens 4-8 (1 0- 16) erect in bud; filaments hypogynous or adnate to the base of the perianth, 

longer than the anthers, the latter slightly longer 

than wide, the connective rarely apically produced; pollen 2-5(-6)-porate or colpate; pistil 1, 

the carpels 2, the styles 2 (I), the stigma decurrent ventrally, rarely bi- or trifid apically; ovary 

superior, the locules 1 (-2), ovule 1, subapical, 

bitegmic, usually crassinucellar, anatropous or 

semi-anatropous; fruit a samara or subdrupaceous nutlet; embryo straight, arcuate or folded, 

2.0-2.5 times longer than the endosperm; cotyledons thin or moderately thick, 1.5-4.7 times 

wider than the radicle, 0.6-0.8 the length of the 

embryo, the radicle sometimes invested; endosperm moderate to 0. 

Composition: 15 genera, -200 species. 

Distribution: Pantemperate and -tropical but 

generally absent from deserts and the Amazon 

basin. 

FABALES 

Trees, shrubs, and herbs, sometimes lianas; 

xylem vessel perforation plates simple; leaves 

alternate, very rarely opposite, usually compound, rarely with tendrils or reduced to phyllodes, the margin usually entire, sometimes serrate, rarely lobed, stipulate, rarely exstipulate; 

inflorescence usually racemose, a raceme, spike, 

head, or panicle; flowers small to large, bisexual 

or rarely unisexual, the parts hypogynous to perigynous, perianth actinomorphic to zygomorphic; sepals 5 (3-6, 0), free or connate; petals 

5 (-O), free or connate; stamens 1 to -500, often 

10, the filaments free or variously connate, the 

anthers basifixed or dorsifixed; pollen 3 (-6)- 

colpate or -colporate, or 2-4-porate; pistils 1, 
86 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

C i: 

d 

FIGURE Q?i.-bfORACEAE: a, Morus alba d flowering twig, d inflorescence, h'orus sp. ? flower, 

I.s. of fruit, Sorocea sp. diagram of d flower; b, Morus alba d flower, M. nigra multiple fruit; c, 

Dorstenia sp. infloresence, Cecropia adenopus d flower, views of stamen, C. pachystachya ? flower, 

1,s. of fruit, Ficus carica 1.s. of 0 flower, I.s. of multiple fruit (after Baillon, 1866-1895; Le 

Maout and Decaisne 1873; Martius, 1840- 1906). ULMACEAE: d, Ulmus campestris inflorescences, 

flower, I.s. of same, 1s. of fruit showing seed and embryo, Celtis australis flower, I.s. of same, 

I.s. of fruit showing embryo (after Baillon). 

very rarely 2- 15, sessile or stipitate, unicarpellate, the style elongate or short, stigma apical or 

subapical; ovules 1-21 in 2 ventral rows; fruit 

usually a legume, sometimes samaroid, rarely a 

follicle, somewhat baccate, a drupe or loment; 

embryo large, straight or bent; endosperm moderate to absent. 

Distribution: Cosmopolitan, most abundant in 

the tropics and subtropics. 

Chemistry: Fabales have quite a few alkaloids, 

a greater variety than the Rosales. Quinalizidines 

are overwhelmingly commonest in legumes, but 

also occur in Ranunculales, Nymphaeales, Magnoliales, Laurales, Papaveraceae, and other taxa. 

Erythrine alkaloids occur in Erythrina and Cocculus. Fabales and Rosaceae are highly cyanogenic. 

CAESALPINIACEAE (Figure 36a-e).-Trees, 
NUMBER 58 87 

shrubs, rarely lianas or herbs, rarely with 1-2 

tendrils derived from twigs (Bauhinia spp.); xylem vessel perforation plates simple; leaves alternate, very rarely opposite, usually paripinnately, 

sometimes bipinnately compound, rarely simple, 

very rarely a phyllode, the margins almost always 

entire, rarely lobed or serrate, stipulate; inflorescence a raceme, spike, or rarely cyme; flowers 

bisexual or rarely the plants polygamous or dioecious (Gymnocladus), subactinomorphic to zygomorphic, the parts hypogynous to perigynous; 

sepals 5 (6, 0), free or rarely completely connate, 

imbricate or rarely valvate; petals 5-0, imbricate, 

the adaxial one internal; stamens 10 (1 -307, the 

filaments free or variously connate, staminodes 

present or absent; anther basifixed or dorsifixed, 

rarely dimorphic (Maldenhauera), dehiscing longitudinally or by terminal pores; pollen 3-colpate, 

3-colporate, 3-6-colporoidate; disk sometimes 

present; pistil 1, very rarely 2-3 in Swartzia, 

sessile or stipitate, the carpel 1, style elongated, 

stigma apical or subapical, ovules 2-2 1 in 2 rows 

along the ventral suture, bitegmic, crassinucellar, 

anatropous or campylotropous, the funicules 

often long; fruit usually a legume, rarely a follicle, samara or drupe; seeds sometimes arillate, 

generally without areoles, rarely winged (Melanoxylon), the hilum apical or subapical; embryo 0.9 

to as long as the endosperm, straight, or rarely 

folded, the radicle sometimes invested, the cotyledons thin or moderately thick, 2-7 times as 

wide as the radicle, 0.7-0.8 the length of the 

embryo; endosperm moderate, scanty or rarely 

absent, hard and glassy. 

Composition: -200 genera, -2800 species. 

Distribution: Pantropical and -subtropical, 

relatively few pantemperate. 

FABACEAE (Figure 37a-e).--Herbs7 shrubs, 

rarely trees or vines, latex very rarely present 

(Galactia); xylem vessel perforation plates simple; 

leaves alternate or very rarely opposite or 

whorled, usually once compound and then usually imparipinnate, sometimes ending in a tendril, rarely simple, the margins usually entire, 

sometimes serrate, stipulate, rarely exstipulate; 

inflorescence a raceme, panicle, or spike, rarely 

a cyme or the flower solitary; flowers bisexual, 

zygomorphic, the parts hypogynous to slightly 

perigynous; sepals 5 (-4), connate part way; petals 5, very rarely 1 or 0, imbricate, the adaxial 

one exterior and enlarged (standard), 2 lateral 

(wings), and 2 lower interior usually connate by 

their inner margin into a keel; stamens generally 

10, rarely 9 or 5, monadelphous or diadelphous 

usually 9 + 1, rarely (5)+(5) or (4 + 1)+(4 + l), 

rarely free; anthers dorsifixed or basifixed, sometimes dimorphic, mostly dehiscing longitudinally, 

rarely by terminal pores (Fissicalyx); pollen 3(2- 

6)-colporate, 2-4-porate, 3-colporoidate; pistil 1, 

sessile or stipitate, the carpel 1, style elongated 

or short, stigma apical or subapical, ovules about 

2-1 4, ventral, bitegmic, crassinucellar, anatropous, semi-anatropous, campylotropous or amphitropous, the funiculus short; fruit usually a 

legume or sometimes indehiscent and somewhat 

baccate, drupoid, samaroid or a loment, rarely 

with a secondary longitudinal septum; seeds without areoles, the hilum often large and elongated; 

embryo 1-2 times as long as the endosperm, bent 

or rarely straight and the radicle invested, the 

cotyledons moderately to markedly thick, 0.8- 

5.5 as wide as the radicle, 0.6-0.9 the length of 

the embryo; endosperm scanty or 0, sometimes 

moderate, usually hard and glassy, not starchy. 

Composition: -450 genera, - 10,000 species. 

Distribution: Pantropical, -subtropical, and 

-temperate. 

Mimosaceae (Figure 38a-d).-Trees and 

shrubs, sometimes lianas, very rarely herbs; xylem vessel perforation plates simple; leaves alternate, very rarely opposite (Tetrapleura), mostly 

bipinnately compound, rarely once pinnate, paripinnate, the leaflets entire, very rarely ending 

in a tendril (Entada), sometimes reduced to phyllodes, rarely the rachis winged, stipulate or rarely 

exstipulate (some Acacia); inflorescence a head, 

spike or raceme; flowers bisexual or rarely the 

plants monoecious (Neptunia, Archidendron), 

small, actinomorphic, the parts hypogynous or 

rarely perigynous (Parkia); sepals small, 5 (3-6, 

0) partly connate, rarely free, valvate or very 

rarely imbricate (Parkieae); petals 5 (3-6), usu- 
88 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 58 89 

ally connate or very rarely free, valvate; stamens 

4-500, usually monadelphous, sometimes free, 

usually long-exserted, staminodes rarely present 

(Pentaclethra); anthers small, dehiscing longitudinally, often with an apical gland; pollen monads, tetrads, and polyads (to 32 cells), 3-colporate, rarely 3-porate; pistil 1 (-15), sessile or stipitate, the style elongated, stigma apical, the 

ovules several to 1, ventral, bitegmic, crassinucellar, anatropous or semianatropous, the funicle 

often long; fruit a legume or indehiscent with 

sweet pulp, rarely a loment leaving a replum; 

seeds & flattened, with an areole on each side 

(i.e., a line demarcating a central area), sometimes arillate or the testa fleshy, rarely winged, 

the hilum apical or subapical, & punctiform; embryo straight 0.8 to as long as the endosperm, 

the radicle invested; cotyledons thin or moderately to markedly thick, 2.3-5.0 times wider than 

the radicle, 0.7-0.8 the length of the embryo; 

endosperm moderate, scanty or 0, hard and 

glassy. 

Composition: -50 genera, -2400 species. 

Distribution: Pantropical and -subtropical, 

few pantemperate. 

PAPAVERALES 

Herbs and shrubs, rarely lianas or small trees, 

sometimes laticiferous; xylem vessel perforation 

plates simple or very rarely scalariform; leaves 

FIGURE 36.-CAESALPINIACEAE: a, SWUrtZiU WLiCrOCUrpU 

flower, Aldina latifolia bud, I.s. of flower (the filaments 

connate), floral diagram, views of anther, Exostyles uenusta 

1,s. of flower, floral diagram; 6, Cadia uaria flower, I.s. of 

same, floral diagram, seed and I.s. of same, Zollernia ilicifolia 

C.S. and 1,s of fruit, fruit; c, Cassia setosa flower, petals, seed 

and 1,s. of same, C. multtjuga floral diagram, Bauhiniaforficata 1,s. of ovary, stigma, floral diagram; d, Copaifera martii 

flower, C. langsdorfii arillate seed, Apuleia praecox flowering 

twig, floral diagram, flower, 1,s. of same; e, Ceratonia siliqua 

0 inflorescence, young P flower with staminodes, 0 flower 

showing large stigma and disk, 0 flower from below and 

fruit, d flowering twig, d flower, views of anther, disk with 

pistillode in center, Caesalpinia echinata dehiscent fruit (after 

Martius. 1840-1 906; Baillon, 1866-1 895; Johnson). 

alternate, rarely opposite, simple, entire or lobed 

to dissected, sometimes compound with entire 

leaflets, usually exstipulate; flowers actinomorphic or zygomorphic, bisexual or rarely unisexual, the parts hypogynous to perigynous; sepals 2-5 (-9), usually free, rarely connate, sometimes biseriate; petals 2-12 (0, 14) in 1-3 whorls, 

2-3-merous, free or rarely connate; stamens 1 to 

-100, usually free, very rarely connate, the anthers basifixed or dorsifixed; pollen mostly 3 (2- 

9)-colpate or colporate, sometimes rupate, rugate, forate or nonaperturate; glandular disk 

often present, of separate parts or lining an hypanthium; pistil 1, sometimes stipitate, the carpels 2-20, rarely free in Resedaceae, the style 

usually 1, rarely several and the stigmas apical, 

or the stigmas sessile and linear or apical; ovary 

superior, unilocular or secondarily bilocular by 

the placentas meeting or by a secondary septum, 

the ovules usually numerous, rarely only 1, usually parietal; fruit usually a capsule, silique, or 

berry, rarely a nutlet or drupe or transversely 

dehiscent into joints; embryo minute to long, 

linear, arcuate or folded; endosperm copious, 

scanty or absent. 

Distribution: Cosmopolitan, centered in the 

north temperate zone, often in dry habitats, 

more rarely in wet forests. 

Chemistry: Myrosin-cells containing mustardoil glucosides are present except in Papaveraceae 

and Tovariaceae(?). They are present in Bretschneidera. Mustard-oil glucosides also occur in 

Tropaeolaceae, Limnant haceae, Zy gophy llaceae, 

and rarely in Euphorbiaceae. Alkaloids are an 

important feature of Papaveraceae but the other 

families lack most of them; aporphine alkaloids 

are numerous in Papaveraceae. Protoberberines 

are most frequent in Papaveraceae, also in Magnoliales, Laurales, Ranunculales, Nandina, and 

Aristolochia. Phthalene-isoquinoline occurs 

mostly in Papaveraceae, Ranunculales, and Brassica, but also in various other unrelated groups. 

Cistaceae has monoterpines, sesquiterpenes, and 

diterpenes; the HC 1 /Methanol Test is strongly 

+. Bixaceae and Cochlospermaceae have triterpenoid saponins and/or sapogenins. These are 
90 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 58 91 

not characteristic of the Violales. The alkaloids 

in Fabales and Papaverales seem to be different 

in many cases, rarely the same. 

PAPAVERACEAE (Figure 39a-e).-Herbs, 

shrubs, or small trees, often with white, yellow, 

or red sap, the stem bundles in some species of 

Papaver in concentric zones and tending to be 

scattered, recalling Ranunculaceae; xylem vessel 

perforation plates simple, diameter of vessels 

very small, the vessels sometimes tracheid-like 

(Romneya); leaves simple, alternate, those just 

below the flowers rarely opposite or whorled, 

entire, lobed, or much divided, exstipulate; flowers mostly solitary, sometimes in dichasia, racemes, or panicles, essentially actinomorphic, bisexual, the parts hypogynous or rarely perigynous; sepals 2-3, caducous, free or rarely connate and calyptrate; petals 4-12 (0) in 2 (3) 

whorls, free, often 2-merous, sometimes 3-merous, often crumpled in bud, without nectaries; 

stamens usually numerous (-6-85), the filaments 

filiform, anthers basifixed; pollen mostly 3 (2- 

O)-colpate, also rupate, rugate, forate, or nonaperturate; pistil 1, the carpels 2-20, the style 

short or absent, rarely several, the stigmas usually 

linear, sometimes apical; ovary superior, unilocular or several locular by the placentas meeting 

at the center of the ovary, or rarely bilocular by 

a secondary septum; ovules usually numerous, 

parietal, rarely 1 and basal, bitegmic, crassinucellar, anatropous or subcampylotropous; fruit a 

FIGURE ~~.-FARACEAE: a, Pisum sativum flower, essential 

organs, pistil, L’igna luteola androecium laid open, anther, 

views of seed, views of embryo, plumule; b, Dalbergia variabilis calyx, flowers, petals, pistil and calyx, androecium, views 

of anther, fruit: c, Bowdichia virgtlioides calyx laid open 

shotving insertion of the stamens, 1,s. and C.S. of seed, embryo, Pterocarpus violaceus androecium, fruit, Machaerium 

aculeatum androecium: d, Harpalyce brasilianu androecium 

laid open, fruit with part removed to show the seeds, Aeschynomene falcata androecium laid open, fruit, Andira anthelminthica section of fruit showing the seed; e, Isodesmia tomentom fruit with part removed to show the seeds, inflorescence 

in leaf axil, Clitoria stipularis flower, views of anther, 1 valve 

of fruit showing seeds, Centrolobium robustum fruit, Sophora 

tommtosa fruit (after Martius, 1840-1906, Baillon, 1866- 

1895). 

capsule dehiscing by valves or pores, rarely a 

silique, rarely the carpels separate and torulose 

in fruit; seeds often arillate; embryo minute or 

long and linear, straight or arcuate, 0.07-0.9 the 

length of the endosperm; cotyledons hardly differentiated to nearly 0.5 the length of the embryo and as wide as the radicle; endosperm copious, fleshy and oily. 

Composition: 26 genera, -200 species. 

Distribution: Temperate and subtropical 

North America and Eurasia; few in eastern Australia and South Africa. 

FUMARIACEAE (Figure 40a).-Herbs, sometimes scandent, with watery sap; xylem vessel 

perforation plates simple; leaves simple, radical 

or ascending the stem, alternate or rarely subopposite, usually much divided, exstipulate; inflorescence usually a raceme; flowers bisexual, zygomorphic, the parts hypogynous; sepals 2, small, 

caducous; petals 4, biseriate, f connivent, 1 or 2 

outer ones often saccate or spurred, the 2 inner 

narrower and sometimes apically connate, or all 

the petals connate in a tube (Adlumia); stamens 

4 free and opposite the petals, or 6 connate in 2 

bundles, with 1 or 2 nectaries at the base, sometimes extending into the spur; filaments elongated, the anthers small; pollen 2-4-colpate, 3- 

colp(oroid)ate, occasionally 4-rupate, 6- or 12- 

rugate, 6-12-forate; pistil 1, the carpels 2, the 

style 1, slender, stigmas 2, apical; ovary unilocular, placentas 2, the ovules 1 to many, parietal, 

bitegmic, crassinucellar, anatropous or campylotropous; fruit often an elongated capsule, sometimes transversely septate and breaking into 1- 

seeded indehiscent segments, or a bivalved silique, sometimes a nutlet; seeds shiny, usually 

arillate; embryo small, straight, linear, 0.1-0.2 

the length of the endosperm; cotyledons 0.2-0.4 

the length of the embryo, twice as wide as the 

radicle; endosperm copious, fleshy. 

Composition: 16 genera, 450 species. 

Distribution: Temperate North America and 

Eurasia; few in the highlands of eastern Africa 

and in South Africa. 

TOVARIACEAE (Figure 40b).-Herbs or shrubs 

with a strong odor; xylem vessel perforation 
92 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 58 93 

plates simple; leaves trifoliolate, the leaflets entire, stipulate; inflorescence a terminal raceme; 

flowers bisexual, actinomorphic, the parts hypogynous; sepals 8 (7-9), deciduous; petals 8 (7-9); 

stamens 8 (7-9) oppositisepalous, the filaments 

filiform, nectaries absent; anthers twice as long 

as wide, basifixed; pollen (2-)3-colporoidate; pistil 1, the carpels 6-8, stigma 6-8-lobed, subsessile; ovary 6-8-locular, the ovules numerous, parietal, bitegmic, crassinucellar, campylotropous; 

fruit a berry, the seeds minute, shiny; embryo 

linear, arcuate, 0.7 the length of the endosperm, 

the cotyledons 0.3 the length of the embryo; 

endosperm copious. 

Composition: 1 genus, 2 species. 

Distribution: Mexico to Andes of Peru; Jamaica. 

CAPPARACEAE (Figure 4 1 a-e).-Small trees, 

shrubs, or herbs, rarely lianas; xylem vessel perforation plates simple, small tracheid-like members sometimes numerous; leaves alternate or 

rarely opposite, simple or palmately 3-7-foliolate, rarely reduced to scales 1 mm long (Koeberlinia), the stipules represented by thorns or 

glands; inflorescence usually a raceme; flowers 

actinomorphic or zygomorphic, mostly bisexual, 

the plants rarely monoecious or dioecious, the 

parts hypogynous; sepals 4 (-8) in 2 whorls, free 

or partly to wholly connate and calyptrate; petals 

4 (-14, or 0), sometimes connate; disk sometimes 

present between the perianth and stamens, often 

thicker to one side; stamens 100 + to 1, dimorphic in Cladostemon, the filaments filiform, 

rarely widened in the middle (Koeberlinia), the 

anthers oblong, dorsifixed; an androphore someFIGURE 38.-bfIMOSACEAE: a, Mimosa somnians flowering 

twig, fruit entire and dehiscent, M. vellosiana flower, 1,s. of 

same with part of the style and filaments cut off, Acacia 

miersii flower, C.S. of ovary, 1.s. of flower; 6, M. somnians 

views of anther, pistil, I.s. of ovary; c, Affonsea bullata stigma, 

1,s. of ovaries, Stryphnodendron 6arbatimiAo inflorescence, 

seed, flower, pistil, essential organs, petal and calyx, 1 valve 

of the fruit; d, Calliandra longipes flower, androecium laid 

open, C. pamiflora part of fruit with 2 seeds, seed, 1,s. of 

same, embryo, Pentaclethra jlamentosa flower, 1 .s. of same 

with part of the filaments cut off, views of anther (after 

Martius, 1840-1 906). 

times present; pollen 3(2-4)-colporate (-colporoidate); pistil 1, the carpels 2 (-12) in 1 or 2 

whorls, the style filiform or absent, the stigma 

bilobed or capitate, apical; ovary usually stipitate, 

unilocular or sometimes with a secondary septum, the ovules many to few, parietal or very 

rarely axile, bitegmic, crassinucellar, campylotropous; fruit a berry, sometimes torulose, silique, 

apically dehiscing capsule, rarely a drupe or 

transversely locellate (Maerua spp.), or nutlets; 

seeds often reniform; embryo linear, arcuate, as 

long as the endosperm, the cotyledons 0.3-0.7 

the length of the embryo; endosperm scanty, 

fleshy. 

Composition: -42 genera, -900 species. 

Distribution: Mostly in arid regions, mainly 

tropical and subtropical, few temperate; temperate North America to temperate South America; 

Japan to eastern Mediterranean region and south 

to South Africa and Australia. 

PENTADIPLANDRACEAE (Figure 42a-b).- 

Large shrubs and climbers; xylem vessel perforation plates simple(?); leaves alternate, simple, 

entire, exstipulate; inflorescences axillary and 

terminal few-flowered racemes, rarely also umbels; plants polygamous; sepals 5, free, imbricate 

in bud; petals 5, free, cupular and connivent 

basally; stamens 9-1 3, the filaments free, filiform; anthers small, basifixed, the connective 

produced into an apical knob; pollen (2-)3-colpate; disk thick, cupular, external to the stamens; 

pistil 1, the carpels 5-4, the style 1, elongated, 

the stigmas 5-4; ovary superior, stipitate, 5-4- 

locular, the ovules several, in 2-3-series per 

locule, campylotropous, axile; fruit a berry, the 

seeds reniform, embryo curved, cotyledons 

short; endosperm 0 or very scanty. 

Composition: 1 genus, 2 species. 

Distribution: 

RESEDACEAE (Figure 42c-e).-Herbs or rarely 

shrubs with watery sap; xylem vessel perforation 

plates simple; leaves simple, alternate, entire or 

pinnately dissected, the stipules gland-like; inflorescence a raceme or spike; flowers bisexual or 

rarely the plants monoecious, usually zygomorphic, the parts hypogynous; sepals 2-8, 

sometimes basally connate; petals small, 2-8 (0), 

Tropical west Africa, in forest. 
94 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

a 

0 a; 

FIGURE 39.-PAPAVERACEAE: a, Papaver somniferum floral diagram, fruit, I.s. of seed, Roemeria 

sp. Is. of seed; b, Platystemon californicus flower, pistil, fruit, floral diagram, Sanguinaria 

canadensis flowering plant, floral diagram; c, Eschscholria sp. pistil, I.s. of flower with petals cut 

off, fruit; d, Chelidonium majus floral diagram, pistil and 1 stamen, dehiscent fruit, replum with 

some seeds still attached; e, Bocconiafrutescens leaf, part of inflorescence-infructescence, flower 

bud, flower with 1 sepal removed, fruit with 1 valve gone, fruit with both valves gone, 1,s. of 

seed showing the minute embryo (after Rendle, 1938; Baillon, 1866-1895; Le Maout and 

Decaisne, 1873). 
NUMBER 58 95 

0 

b 

FIGURE ~~.-FUMARIACEAE: a, Dicentra canadensis interior petals, rest of flower laid open, 

floral diagram, Corydalis aurea interior petals, rest of flower dissected, floral diagram, replum 

with seeds, seed, Is. of same showing the minute embryo, embryo enlarged (after Gray). 

TAVARIACEAE: b, Tonaria pendula flowering shoot, flower, calyx and pistil, C.S. of ovary, petal, 

views of stamen, fruit and C.S. of same, I.s. of seed, T. dz;ffusa C.S. of fruit (after Hooker and 

Hooker, 1837-1982; Pax). 

often laciniate, rarely connate, sometimes with a 

scale at the base; disk often eccentric; androgynophore short; stamens 3-45, the filaments elongated; anthers as long as wide, dorsifixed?; pollen 

3-colpate (-colporoidate); pistil 1, the carpels 2- 

7 rarely free; ovary unilocular, often open apically, the stigmas 2-7, sessile; ovules numerous 

to 1, bitegmic, crassinucellar or tenuinucellar, 

anatropous or campylotropous, parietal or basal; 

fruit a capsule open apically, berry, or separate 

fruitlets; seeds reniform, carunculate; embryo 

linear, arcuate, as long as the endosperm, the 

cotyledons 0.4-0.8 the length of the embryo and 

as wide as the radicle; endosperm scanty, fleshy, 

or absent. 

Composition: 6 genera, 70 species. 

Distribution: Dry habitats, centered in the 

Mediterranean region; most of Europe to India; 

South Africa; southwestern United States. 

BRASSICACEAE (Figure 43a-c).-Herbs, very 

rarely undershrubs, the sap watery; xylem vessel 

perforation plates simple; leaves alternate or 

rarely subopposite, opposite and alternate in 

Chamira, simple to compound, entire to variously 

cut, exstipulate; inflorescence usually a raceme 

or corymb; flowers bisexual, mostly actino- 
96 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

"i 

[I I 

i 

FIGURE 4 1 .-CAPPARACEAE: a, Capparis jacobinae inflorescence, flower, calyx and disk segments, petal, stamen, pistil, c.s., of ovary; b, C. auicennijolia C.S. and 1,s. of ovary, fruit: c, Cleome 

rosea bud laid open, I.s. of seed, C. siliculijera replum with seeds; d, Koeberlinia spinosa part of 

flowering twig, leaf (minute), flower, 1,s. of same, floral diagram, part of fruiting wig; e, I.s. 

and C.S. of fruit, section of seed, embryo (after Martius, 1840-1906; Sargent). 
NUMBER 58 97 

a 

I' d 

FIGURE ~~.-PENTADIPLANDRACEAE: a, Pentadiplandra brazreana flowering twig, 8 flower, Is. 

of same with pistillode, sepal, petal showing thickened basal portion; b, corolla showing 

connivent basal portions of petals, stamen, C.S. of ovary, fruits (after Hutchinson and Dalziel). 

RESEDACEAE: c, Reseda odorata flowering shoot, floral diagram, front view of flower, rear view, 

flower with petals removed, a posterior petal, a lateral petal, an anterior petal; d, calyx, disk 

and young pistil, R. lutea 1,s. of flower, R. odorata fruit, Is. of seed, Ochradenus sp. fleshy fruit, 

Randonia africana floral diagram; e, Astrocarpus canescens flower with 6 pistils, 1,s. of flower, 

floral diagram, fruit (after Baillon, 1866-1895; Le Maout and Decaisne, 1873). 
98 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

a 

C 

FIGURE 43.-BRASSICACEAE: a, Megacarpaea polyandra flower, same with perianth removed, 

pistil and receptacle, seed, embryo, fruit; b, Warea cuneijolza flower, essential organs, Stanleya 

pinnatzjida flower, embryo, stipitate pistil, Erysimum sp. 1,s. and C.S. of seed to show incumbent 

cotyledons, Lunaria sp. I.s. and C.S. of seed to show accumbent cotyledons; c, Barbarea sp. floral 

diagram, Rorippa barbaraefolia C.S. of fruit, dehiscent fruit, R. bonariensis flower, sepal, petal, 

essential organs, views of anther, fruit, replum with seeds, seed and C.S. of same (after Hooker 

and Hooker, 1837-1982; Gray; Le Maout and Decaisne, 1873; Martius, 1840-1906). 

morphic, the parts hypogynous; sepals 4, free, 

rarely connate (Garnosepalurn), biseriate; petals 4 

(0); stamens 6, tetradynamous, very rarely to 16 

or 4, free or connate in pairs, the anthers dorsifixed; pollen usually (2-)3-colpate, 4-rupate, 6- 

rugate or nonaperturate; glands usually present 

on the receptacle, the nectar secreted into often 

gibbous bases of the inner sepals; pistil 1, sessile 

or rarely stipitate, the carpels 2, very rarely 3 or 

4 in some specimens, the style short, stigma(s) 

apical; ovary divided longitudinally by a secondary septum, rarely undivided or divided transversely, the ovules usually numerous, bitegmic, 

crassinucellar or tenuinucellar, campylotropous 
NUMBER 58 99 

or anatropous, parietal; fruit a silique or silicle, 

bivalved or rarely an indehiscent nutlet, or transversely dehiscent into joints; embryo coiled or 

folded, the cotyledons 2-10 times as wide as the 

radicle, thick or thin; 0.4-0.85 the length of the 

embryo; endosperm 0, very rarely very little 

present . Composition: -375 genera, -3000 species. 

Distribution: Cosmopolitan, centered in the 

north temperate zone, especially the Mediterranean region, central Asia, and western United 

States; rarer in the tropics. 

MORINGACEAE (Figure 44a,b).-Trees exuding a gum resembling tragacanth from bark 

wounds; xylem vessel perforation plates simple; 

leaves alternate, 2-3 times pinnately compound, 

exstipulate or stipules represented by glands; inflorescence an axillary panicle; flowers honeyscented, bisexual, zygomorphic, the parts perigynous, the hypanthium short; sepals 5, petaloid; 

petals 5, differing in size; stamens 5, staminodes 

5, inserted on the margin of a disk which lines 

the hypanthium, the filaments free; anthers 1.5 

times longer than wide, finally unilocular, dorsifixed; pollen (2-)3-colporate; pistil 1, stipitate, 

the carpels 3 (-4), the style elongated, stigma 

apical; ovary unilocular, the ovules numerous, 

parietal, bitegmic, crassinucellar, anatropous; 

fruit a thick-walled, long, beaked, torulose, loculicidal capsule, 3 (-4)-valved; embryo straight, 

the cotyledons broad, thick, plano-convex, 0.9 

the length of the embryo, radicle invested, plumule evident; endosperm 0. 

Composition: 1 genus, 12 species. 

Distribution: Arid habitats; Mediterranean 

region to western India; eastern and southwestern Africa, Madagascar. 

BRETSCHNEIDERACEAE (Figure 44c).-Trees; 

xylem vessel perforation plates scalariform with 

about 20 bars; leaves alternate, compound, imparipinnate, the leaflets entire, exstipulate; inflorescence a terminal raceme; flowers bisexual, 

slightly zygomorphic, the parts perigynous; hypanthium present; sepals 5; petals 5, unequal; 

stamens 8, biseriate, the filaments filiform, anthers dorsifixed; pollen (2-)3-colpate; disk absent; pistil 1, sessile, the carpels 3, style 1, long, 

curved, the stigma apical; ovary 3-locular, the 

ovules 2 per locule, axile; fruit an obovate, thickwalled, 3-valved capsule, the seeds red; embryo 

straight, the cotyledons large, radicle very short; 

endosperm 0. 

Composition: 1 genus, 2 species. 

Distribution: Mountains of western China. 

BATALES 

The order is monotypic. 

Chemistry: Batales have myrosin-cells. They 

have no betalains. 

BATACEAE (Figure 45a,b).-Coastal fleshy subshrubs; xylem vessel perforation plates simple; 

leaves opposite, simple, entire, linear, subterete, 

exstipulate or stipules small; inflorescences axillary and terminal dense spikes or the ? flowers 

sometimes solitary, the plants dioecious or monoecious, the flowers minute; S flower: sepals 

(bracts?) connate, the calyx bilabiate; petals 4, 

free, clawed; stamens 4 (2-5), free, the filaments 

elongated, anthers about twice as long as wide, 

dorsifixed, the connective sometimes apically 

produced; pollen 3-4-colporoidate (-ruporoidate); ? flower: sepals and petals 0; pistil 1, the 

carpels 2, stigma papillose, subsessile, apical, obscurely bilobed; ovary 4-locular, one of the 2 

septa being secondary, the ovules 1 per locule, 

basal, the funicle elongated; fruits subdrupaceous and united into a fleshy mass, or 4 nutlets; 

embryo large, almost straight, oblanceolate; 

cotyledons oblong, plano-convex, slightly flattened, grading gradually into the radicle, twice 

as broad as the radicle, 0.6-0.8 the length of the 

embryo; endosperm 0. 

Composition: 1 genus, 2 species. 

Distribution: Coastal regions; warm temper- 
100 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

a Lii b 

FIGURE 44.-MORINGACEAE: a, Moringa oleijera flowering twig, floral diagram and part of 

androecium, part of fruit cut open; b, M. aptera flower, I.s. of same, seed, 1,s. of same (after 

Martius, 1840- 1906; Le Maout and Decaisne, 1873). BRETSCHNEIDERACEAE: c, Bretschneidera 

sinensis part of twig in bud, views of anther, flower, Is. of same showing insertion of petals and 

stamens, I.s. and C.S. of ovary (after Hemsley). 
NUMBER 58 101 

a 

\ b 

FIGURE 45.-BATACEAE: a, Batis rnaritirna flowering twig of d plant, flowering twig of 0 plant, 

d inflorescence, 0 inflorescence, d flower; 6, I.s. of d bud, d calyx, stamen, I.s. of 0 flower, I.s. 

and C.S. of fruit, seed and embryo (after Martius, 1840-1906; Le Maout and Decaisne, 1873). 

ate to tropical eastern and western America; New 

Guinea, Queensland, Hawaii, Galapagos. 

DILLENIALES 

Trees and shrubs, rarely herbs, sometimes 

climbing; xylem vessel perforation plates mostly 

scalariform with few to many bars, sometimes 

also simple; leaves alternate, rarely opposite, simple, usually toothed, rarely pinnatifid or lobed, 

often with numerous parallel veins extending 

from the midrib, mostly exstipulate; flowers bisexual or more rarely unisexual, the parts hypogynous; sepals 5 (3-17), free, imbricate; petals 5 

(-2), free or shortly connate; stamens numerous, 

to -160, rarely as few as 7-10, free, or basally 

connate or adnate to the base of the petals, the 

filaments usualy filiform, elongate, the anthers 

adnate, basifixed or dorsifixed, dehiscing longitudinally or by apical pores; pollen 3-colpate or 

3-colporate; pistils 20 to rarely 1, the carpels 1- 

16 per pistil, styles usually free, rarely connate, 

elongate, the stigmas apical or slightly decurrent, 

the ovules numerous to 1 per locule, ventral or 

axile; fruits follicles or a berry, the seeds minute 

or moderate-sized and mostly arillate; embryo 

minute, more rarely elongate and linear; endosperm copious. 

Distribution: Pantropical, extending into temperate Australia and temperate eastern Asia. 

Chemistry: Alkaloids have not been reported 

from Dilleniaceae. 

DILLENIACEAE (Figure 46a,b).-Trees, shrubs, 
102 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

a 

-, 

I\ 

b 

FIGURE 46.-DILLENIACEAE: a, Tetracera alnifofolia flower with bracts, T. laszocarpa corolla in 

bud, 1,s. and C.S. of gynoecium, fruit and calyx, single dehiscent fruitlet, 1.s. of fruitlet, seed 

with aril; b, T. alnifolia views of stamen, views of anther, pistil, views of stigma, Doliocarpus 

grandzjlorus part of flowering twig, petal, flower from above and below (after Martius, 1840- 

1906). 

and twiners, rarely herbs; xylem vessel perforation plates typically scalariform with few to 130 

bars, sometimes both simple and scalariform; 

leaves alternate, rarely opposite, simple, entire 

or dentate, rarely pinnatifid or trilobed, usually 

with numerous parallel lateral veins, mostly exstipulate, sometimes the stipules wing-like and 

adnate to the petiole; inflorescences cymose, racemes, panicles, or the flowers solitary; flowers 

bisexual or rarely the plants polygamodioecious, 

the parts hypogynous; sepals 5 (3-17), free, imbricate, or spirally arranged, persistent, often 

accrescent; petals 5 (-2), free, imbricate, yellow 

or white, often crumpled in bud; stamens numerous, to -160, rarely 7-10, free or basally 

connate, development centrifugal, usually persistent, staminodes often present; filaments filiform 

or thickened apically, as long as or longer than 

the anthers; anthers oblong, nearly twice as long 

as wide, or small or didymous, usually adnate or 

basifixed, the connective sometimes apically produced, dehiscing longitudinally or by apical 

pores; pollen 3-colpate and 3-colporate; receptacle sometimes convex; pistils several, 20 to 

rarely 1, sometimes slightly united to a columella, 

the styles free, elongate, and divergent, the stigmas apical; ovule 1 or several, basal, ventral, or 

axile, bitegmic, crassinucellar, anatropous, campylotropous, or amphitropous; fruitlets follicles, 

dehiscent along the ventral and rarely also dorsal 

suture, or baccate; seeds 1 or few, mostly with a 

crested or laciniate aril; embryo minute; linear, 

straight, 0.05-0.1 the length of the endosperm; 

cotyledons not broadened, 0.3-0.5 the length of 
NUMBER 58 103 

d 

FIGURE ~~.-ACTINIDIACEAE: a, Actinzdia polygama flower, stamen, pistil, fruit, C.S. and Is. of 

same; b. A. holotricha sepal, stamen, petal, pistil, another view of stamen; c, A. strzgosa flower, 

floral diagram; d, fruit, seed, I.s. of same (after LavalEe; Finet and Gagnepain; Baillon, 1866- 

1895). SAURAUIACEAE: e, Saurauia bullosa Is. of flower, calyx from below, floral diagram, 

stamens from a bud, stamen at anthesis, part of inflorescence;f, pistil, ovary with 1 carpel 

opened to show ovules, C.S. of ovary, columella, fruit, seed, embryo, I.s. of seed (after Martius, 

1840- 1906). 

the embryo; endosperm copious, fleshy, oily and 

proteinaceous. 

Composition: 14 genera, -400 species. 

Distribution: Pantropical; temperate Australia; centered in Australia, rare in Africa. 

ACTINIDIACEAE (Figure 47a-d).-Shrubs, 

often climbing; xylem vessel perforation plates 

mostly scalariform, with numerous bars, simple 

plates also present; leaves alternate, simple, serrate, exstipulate; inflorescences axillary cymes or 

fascicles, rarely the flowers subsolitary; flowers 

bisexual or the plants polygamous or dioecious, 

the parts hypogynous; sepals 5, free, imbricate; 

petals 5, free; stamens numerous (-50) to 10, the 

filaments filiform, elongated; anthers twice as 

long as wide, dorsifixed, inflexed in bud, dehiscing longitudinally or by pores; pollen usually 3- 

colporate; pistil 1, the carpels 5-16, the styles 

free and divaricate, or united (Clematoclethra), 

the stigmas papillose, apical, slightly decurrent; 

ovary 5- 16-locular, the ovules numerous, axile, 

unitegmic, tenuinucellar, anatropous; fruit a 
104 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

berry, the seeds small; embryo linear, straight, 

0.75-0.8 the length of the endosperm; cotyledons about as broad as the radicle, 0.25-0.5 the 

length of the embryo; endosperm copious, fleshy. 

Composition: 2 genera, 50 species. 

Distribution: Eastern Asia. 

SAURAUIACEAE (Figure 47e,f).-Trees or 

shrubs; xylem vessel perforation plates scalariform, with numerous bars (to 50); leaves alternate, simple, usually serrate, the major lateral 

veins marked and parallel, exstipulate; inflorescence axillary or cauliflorous panicles (cymose?); 

flowers usually bisexual, small to medium-sized, 

the parts hypogynous; sepals 5, imbricate, free; 

petals 5, free or shortly connate, stamens numerous (-150) adnate to the base of the petals, the 

filaments filiform, elongate; anthers oblong, 

about 5 times longer than wide, dorsifixed, dehiscing by apical pores or short slits; pollen usually 3-colporate; pistil 1, the carpels 3-5, the 

styles 3-5, free or united, the stigmas apical; 

ovary 3-5-locular, the ovules numerous, axile, 

unitegmic, tenuinucellar, anatropous; fruit a 

berry, the seeds small; embryo straight or bent, 

0.8 the length of the endosperm; cotyledons as 

wide as the radicle, 0.4 the length of the embryo; 

endosperm copious, mostly mealy. 

Composition: 1 genus, 300 species. 

Distribution: Tropical Asia and America. 

THEALES 

Trees and shrubs, more rarely herbs, sometimes climbing, sometimes with resinous or mucilaginous sap or gland-dotted or resinous tissues; 

xylem vessel perforation plates scalariform, with 

numerous or few bars, or simple, sometimes cortical vascular bundles present; leaves alternate or 

opposite, simple, rarely pinnatifid or compound, 

toothed or entire, stipulate or exstipulate; flowers bisexual or more rarely unisexual, the parts 

hypogynous or more rarely perigynous, very 

rarely epigynous; sepals 5 (2-14), free or connate, usually imbricate, sometimes unequal, 

sometimes accrescent; petals 5 (0, 2-16) in 1 or 

rarely 2 whorls, free or shortly connate, rarely 

calyptrate; stamens usually numerous, 30 to 

-1000, more rarely 3-15, in 1-several whorls, 

free or basally connate, sometimes in bundles, 

sometimes adnate to the base of the petals; staminodes sometimes present; anthers basifixed or 

dorsifixed, rarely adnate, sometimes didymous, 

the connective sometimes apically produced, dehiscent longitudinally, rarely by apical pores; pollen 3(2-6)-colporate, less commonly 2-3-colpate 

or 3(2-4)-porate, rarely 4-rupate or ruporate; 

pistils 1, very rarely only connate basally, or 3, 

the carpels 3-5 (2-25), styles free, connate half 

way or to the apex, the stigmas usually apical, 

more rarely sessile, very rarely decurrent; ovary 

sometimes basally surrounded by a glandular 

disk, 3-5 (2-25)-locular, the ovules I-many per 

locule, axile or rarely parietal or basal, sometimes 

tenuinucellar; fruit a loculicidal or septicidal, 

rarely also ventricidal capsule, often leaving a 

columella, or berry or drupe, rarely a nut or 

schizocarp; embryo usually large, straight or 

curved; endosperm most commonly absent, 

sometimes moderate, least often copious. 

Distribution: Pantropical, especially America 

and Asia, uncommon in temperate regions. 

Chemistry: Secretory cavities of various types 

occur in most of the families. Theobromine occurs in Theaceae, most Sterculiaceae, Aquifoliaceae, and Sapindaceae. Alkaloids are rare. Some 

alkaloids highly characteristic of the “Polycarpicae,” such as protoaporphines, aporphines, berberin, and other quaternary bases, are absent 

from the Theales. The order contains trihydroxylated phenolic substances. These are absent 

FIGURE 48.-THEACEAE: a, Ternstroemia alni$olia fruiting 

twig, floral diagram, 1,s. of seed, embryo, T. candolleana 1,s. 

of flower, calyx from below; b, corolla laid open, views of 

stamen, part of style and stigma, I.s. and C.S. of ovary; c, 

Laplacea semiserrata flo\ver from below, pistil, fruit opened 

to show seeds, seed on columella, 1.s. of seed, views of embryo 

(after Martius, 1840-1 906). PENTAPHYLACACEAE: d, Pentaphylax euryoides stamen and petal, infloresence, 1s. and C.S. 

of ovary, flower without stamens, 1,s. of seed, P. malayana 

corolla and stamens, fruit (after Ridley; Hutchinson, 1973). 

BONNETIACEAE: e, Bonnetia paniculata 1,s. of flower, C.S. and 

1,s. of ovary, one of the 3 stigmas, B. sessilis petal, three of 

the sepals, views of stamen; J B. anceps dehiscent fruit, 

embryo, 1,s. of seed attached to placenta (after Martius, 

1840-1 906). 
NUMBER 58 105 

C 

d 
106 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

from the “Polycarpicae.” Quercetin occurs in 

many. Saponins are often present in the Clusiaceae. The Juglone test is negative and no cyanogenesis occurs. Aucubin (iridoid) glycosides are 

very rare. Theaceae and Symplocaceae show 

strong tendency to accumulate aluminum. 

THEACEAE (Figure 48a-c).-Trees and 

shrubs; xylem vessel perforation plates scalariform, often with many bars (to IOO), vessel members very long; leaves alternate, simple, dentate 

or crenate, often distichous, exstipulate; flowers 

mostly solitary and axillary, rarely a panicle or 

raceme, bisexual or rarely the plants dioecious 

(Eurya), often large, the parts hypogynous, very 

rarely perigynous or epigynous; sepals 5 (4-7), 

free or shortly connate, imbricate; petals 5 (4- 

8), free or shortly connate, sometimes spirally 

arranged; stamens numerous (to -85), rarely 5, 

10, or 15, free or shortly connate, sometimes 

adnate to the base of the petals, sometimes in 5 

bundles, the filaments filiform, usually longer 

than the anthers; anthers about twice as long as 

wide, basifixed or dorsifixed, the connective 

sometimes apically produced, dehiscing longitudinally, very rarely by apical pores; pollen 3- 

colporoidate; pistil 1, the carpels 3-5 (2-lo), 

very rarely only united basally, the styles free 

half way, or connate, the stigmas decurrent or 

apical; ovary usually 3-5-locular, the ovules 

(1 -)2 or several per locule, axile, bitegmic, 

tenuinucellar, anatropous, rarely becoming campylotropous; fruit a loculicidal or septicidal capsule, often leaving a columella, or berry or drupe; 

embryo large, straight or arcuate, 0.9 as long as 

the endosperm; cotyledons terete or thin, sometimes folded or thick and plano-convex, 1-3.5 

times the width of the radicle, 0.5-0.9 the length 

of the embryo, the radicle sometimes invested, 

the plumule evident; endosperm moderate or 0. 

Centered in tropical and subtropical America and Asia; few species in temperate regions; southeastern United States; temperate China; tropical western Africa, Madagascar; northeastern Australia. 

PENTAPHYLACACEAE (Figure 48d).-Shrubs 

Composition: 17 genera, -500 species. 

Distribution: 

and small trees, the buds perulate; xylem vessel 

perforation plates scalariform, usually with about 

15 bars but sometimes more than 25; leaves 

alternate, coriaceous, simple, entire, exstipulate; 

inflorescences axillary and terminal racemes or 

racemiform; flowers bisexual, small; sepals 5, 

free; petals 5, free or shortly connate; stamens 5, 

the filaments thick, broad, connivent, sometimes 

adnate to the petals; anthers basifixed, didymous, 

dehiscing by apical lidded pores; pollen 3-colporoidate (-colporate); disk absent; pistil 1, the carpels 5, style 1, stigmas 5, apical; ovary superior, 

5-locular, the ovules 2 per locule, axile-subapical, 

bitegmic, crassinucellar, anatropous or campylotropous; fruit a loculicidal capsule, with a columella; embryo U-shaped; cotyledons as wide as 

the radicle, 0.5 the length of the embryo; endosperm scanty. 

Composition: 1 genus, 2 species. 

Distribution: Southern China to Malaya and 

Sumatra. 

BONNETIACEAE (Figure 48e,f).-Trees, rarely 

shrubs, sometimes resinous; xylem vessel perforation plates simple, rarely also scalariform, with 

a few bars; leaves alternate, rarely opposite, entire, exstipulate; inflorescence terminal or axillary toward the tip of the branches, panicles, 

racemes or umbelliform; flowers bisexual, the 

parts hypogynous; sepals 5, imbricate, unequal, 

free; petals 5, contorted, free; stamens numerous 

(-1 00), the filaments filiform, free or connate 

basally into 5 bundles; anthers small, about as 

FIGURE 49.-PELLICIERACEAE: a, Pelliciera rhirophorae sepal, 

I.s. of flower, 1,s. and C.S. of ovary, flower with bracts, flower 

with bracts removed, stamen, stanlens and pistil, petal, fruit, 

longisections of fruit showing plumule and cotyledons, C.S. 

of seed (above) (after Hemsley). MEDUSAGYNACEAE: b, .Viedusagyne oppositfolia flowering branch, partially open 

flower, fully open flower, essential organs, essential organs 

\vith sonie stamens removed, pistil and some filaments; c, 

views of dehisced fruit, seed (enlarged), views of anther, C.S. 

and I.s. of pistil, 1,s. of pistil (after Hemsley, Engler and 

Melchior). EUCRYPHIACEAE: d, Eucryphia billardieri flowering twig, calyptriform calyx, pistil, C.S. of ovary, views of 

stamen; e, B. cordfoliu 1,s. of ovary and base of androecium, 

part of dehiscent fruit, seed, 1,s. of same (after Hooker and 

Hooker, 1837- 1982; Focke). 

108 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

long as wide, dorsifixed; pollen 3-colporate; pistil 

1, the carpels 3 or 5, the style 1 or 3- or 5-fid; 

ovary 3-5-locular, the placentas axile, thick, the 

ovules usually numerous; fruit a septicidal capsule, leaving a columella; embryo straight, the 

cotyledons as wide as or narrower than the radicle, 0.2-0.3 the length of the embryo; endosperm present or 0. 

Composition: 3 genera, 22 species. 

Distribution: Mostly in swampy or coastal regions; tropical Asia and America. 

PELLICIERACEAE (Figure 49a).-Buttressed 

mangrove tree; xylem vessel perforation plates 

simple; leaves alternate, simple, involute in bud, 

coriaceous, exstipulate; flowers axillary, solitary, 

large, bisexual, the parts hypogynous; sepals 5, 

imbricate, short, free, at first white, then rose 

like the petals; petals 5, elongated, free; stamens 

5, the filaments free, slightly shorter than the 

large, linear anthers; anthers at least 10 times 

longer than wide, basifixed; pollen 3-colporate; 

pistil 1, the carpels 2, the style 1, stigma bifid, 

apical; ovary conic-cylindric, bilocular, the ovules 

1 per locule, axile; fruit a large, long-acuminate, 

coriaceous nut, the seed 1, large; cotyledons 

broad, thick, plano-convex, radicle short, plumule long; endosperm 0. 

Composition: 1 genus, 1 species. 

Distribution: Pacific shores from Costa Rica 

to Ecuador. 

MEDUSAGYNACEAE (Figure 49b,c).-Shrubs, 

xylem vessel perforation plates simple, cortical 

bundles present in the stem; leaves opposite, 

simple, obscurely crenate, exstipulate; inflorescence a short, lax, deltoid, terminal panicle; flowers bisexual, the parts hypogynous, the stamens 

and styles included; sepals 5, imbricate, basally 

connate; petals 5, free, imbricate and contorted 

in bud; stamens about 100, at 4-5 levels, the 

filaments filiform, free; anthers small, as long as 

wide, basifixed, sometimes the sacs slightly separated at base by connective and not at the same 

height; pollen 3(2-4)-porate; pistil 1, the carpels 

17-25, the styles free, on the shoulders of the 

carpels, the stigmas capitate; ovary 17-25-locular, the ovules 2(3) per locule, axile; fruit a capsule, septicidally dehiscent from the base, leaving 

a columella; seeds small, winged; embryo clavate, 

the cotyledons ovate, nearly equal to the radicle 

in length(?); endosperm very scanty. 

Composition: 1 genus, 1 species. 

Distribution: Seychelles Islands. 

EUCRYPHIACEAE (Figure 49d,e).-Trees and 

shrubs; xylem vessel perforation plates typically 

all scalariform, usually with 20 or fewer bars but 

occassionally to 40; leaves opposite, simple or 

pinnately compound, serrate, stipulate; flowers 

axillary, solitary, bisexual, the parts hypogynous; 

sepals 4 (-5), imbricate, connate apically, deciduous; petals 4 (-5), large, free; stamens numerous 

(-200), free, the filaments filiform; anthers 

small, about as long as wide, dorsifixed; pollen 

2-colpate; pistil 1, the carpels 5-12 (4-18), the 

styles free, filiform, stigmas apical, inconspicuous; ovary 5- 12(4- 18)-locular, the ovules several per locule, axile; fruit a septicidal capsule, 

each carpel also ventrally dehiscent, leaving a 

columella; seeds winged, the embryo large; cotyledons foliaceous, moderately thick, plano-convex, the radicle short; endosperm scanty, fleshy. 

Composition: 1 genus, 5 species. 

Distribution: Temperate Southern Hemisphere; New South Wales, Tasmania, Chile. 

PARACRYPHIACEAE (Figure 50a,b).-Shrubs or 

small trees; xylem vessel perforation plates scalariform with more than 100 bars; leaves simple, 

subverticillate, serrate, exstipulate; inflorescence 

a terminal compound spike; flowers bisexual and 

8; sepals (bracts?) 2-3; petals 0; stamens 8-10 

(-1 6), the filaments hypogynous or subperigynous; anthers oblong, twice as long as wide, basifixed; pollen tricolporate; pistil 1, costate-cylindrical, the apex truncate, the carpels 12-15, the 

stigmas 12- 15, sessile; ovary superior, 12- 15- 

locular, the ovules few per locule, 1-seriate; fruit 

a septicidal capsule, leaving a columella; seeds 

not seen. 

Composition: 1 genus, 2 species. 

Distribution: New Caledonia. 

SYMPLOCACEAE (Figure 5Oc-e).-Trees and 

shrubs; xylem vessel perforation plates scalariform, usually with more than 20 bars; leaves 
NUMBER 58 109 

FIGURE 50.-PARACRYPHIACEAE: a, Paracryphia suaveolens flowering twig, flower, flower with 

part of perianth removed, pistil, C.S. of ovary; b, stamens, single carpel of fruit, dehiscent fruit 

(after Baker). SYMPLOCACEAE: c, Symplocos lanceolata 1,s. of bud, flower, flower laid open, sepal, 

corolla lobe, stigma; d, floral diagram, C.S. of ovary, I.s. of ovary, views of stamen; e, fruits, I.s. 

of same, S. paniculata pistil with epigynous glandular disk, I.s. of fruit (after Martius, 1840- 

1906; Johnson). 

alternate, simple, serrate or entire, exstipulate; numerous (to -loo), in 1-4 whorls, the filaments 

inflorescences axillary or terminal panicles, free or connate basally, sometimes in 5 bundles, 

spikes, racemes or fascicles, or the flowers soh- inserted on the base of the corolla; anthers small, 

tary; flowers bisexual, or rarely the plants poly- as long as wide; pollen 3(2-4)-colporate, the colpi 

gamodioecious, the parts perigynous or epigy- short; pistil 1, the carpels 2-5, the style 1, elonnous; sepals 5, connate, persistent; petals 5 (3- gate, stigmas apical, capitate or 2-5-lobed; ovary 

1 l), connate basally in 1-2 series; stamens 4- semi-inferior, 2-5-locular, the ovules 2-4 per 
110 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

a 

W 

b 

C 

FIGURE 5 1 .-TETRAMERISTACEAE: a, Tetramerista glabra flowering twig, bud, flower, I.s. of 

same, C.S. of ovary; b, stamen, views of anther, I.s. and C.S. of fruit; c, immature pyrene, the 

same opened (after Boerlage and Koorders). 

locule, axile, unitegmic, tenuinucellar, anatropous; fruit a berry or drupe; embryo linear, 

straight or arcuate, 0.7-0.95 the length of the 

endosperm; cotyledons 0.1 the length of the 

embryo; endosperm copious. 

Composition: 1 genus, -400 species. 

Distribution: Centered in tropical and subtropical America and Asia; southern United 

States; Japan; northeastern Australia, southwestern Pacific Islands; absent from Africa and western Eurasia. 

TETRAMERISTACEAE (Figure 5 1 a-c).-Trees 

and shrubs; xylem vessel perforation plates usually simple, sometimes also scalariform with 

about 8 bars; leaves alternate, simple, entire, 

asymmetric, coriaceous, glandular-punctate below, exstipulate; inflorescences axillary umbelliform racemes; flowers bisexual, the parts hypogynous; perianth glandular-punctate; sepals 4, imbricate in 2 series, shortly connate basally; petals 

4; stamens 4, the filaments elongate, flattened 

basally; anther thecas separate and divergent basally, the connective basally produced and glandular; pollen 3-colporate; pistil 1, the carpels 4, 

the style 1, subulate, stigmas 4, minute, apical; 

ovary 4-locular, the ovules 1 per locule, basal; 

fruit a berry, the exocarp coriaceous, mesocarp 

fleshy; seeds 4, oblong; endosperm 0. 

Composition: 1 genus, 3 species. 

Distribution: Malaya and western East Indies. 

CLUSIACEAE (Figure 52a-c).-Trees and 

shrubs, the sap resinous; xylem vessel perforation 
NUMBER 58 111 

FIGURE ~~.-CLUSIACEAE: a, Clusia arrudea d bud, 0 flower, d flower; b, C.S. and 1,s. of ovary, 

androeciuni, pistil and staminodes; c, C. cambessedeseii inflorescence, views of stamen, C. 

planchoniana dehiscent fruit (after Martius, 1840-1 906). QUIINACEAE: d, Quiina glaziovii 

flowering twig, Q. macrostachya d flower, views of stamen, Q. rhytidopus pistil, immature fruit, 

Q. sessilis I.s. of fruit showing the seed; e, Q. obovata fruit, Is. of same showing the seed, 1,s. of 

seed, C.S. of embryo (after Martius, 1840-1906). 
112 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

plates simple to scalariform with numerous bars; 

leaves opposite, simple, entire, exstipulate; inflorescences cymes, sometimes umbellate, or the 

flowers solitary; flowers bisexual, or the plants 

polygamous or dioecious, the parts hypogynous; 

sepals 6-2 (-14), imbricate, free; petals 6-2 

(- 14), free; stamens numerous (to -1 000), rarely 

few (8 in Oedematopus), free or all connate basally, or connate in bundles opposite the petals; 

anthers often oblong, adnate or basifixed; staminodes often present in 0 flowers; pollen 3(2-5)- 

colporate; pistil 1, the carpels 5 or 3 (1- 15), the 

styles free or connate or absent, the stigmas often 

shield-shaped, lobed or radiate, sometimes small; 

ovary l-several locules, the ovules l-many per 

locule, axile or basal, rarely parietal, bitegmic, 

crassinucellar or tenuinucellar, anatropous or 

semianatropous; fruit often a septicidal capsule 

or berry, rarely a drupe (Endodesmia); seeds 

large, often arillate; embryo straight, sometimes 

conferruminate; cotyledons sometimes 4 times 

wider than the radicle, 0.9 the length of the 

embryo, sometimes 0.05 the width of the macropodal radicle and 0.07 the length of the embryo; 

endosperm 0. 

Composition: 35 genera, 400 species. 

Distribution: Pantropical in moist regions, relatively few in Africa. 

QUIINACEAE (Figure 52d,e).-Trees, shrubs, 

and climbers; xylem vessel perforation plates simple, rarely also scalariform, lysigenous mucilaginous cavities often in the pith, sometimes resin 

canals in traumatic tissue, sometimes resin in fruit 

and petals; leaves opposite or whorled, simple, 

sometimes pinnatifid or imparipinnately compound, the margins entire or crenate, lateral 

veins numerous, tertiary veins feather-veined, 

stipulate; inflorescences axillary or terminal racemes or panicles; flowers bisexual or the plants 

polygamodioecious, the parts hypogynous; sepals 

4-5, sometimes unequal in size, free or nearly 

free; petals 4-8, free; stamens 15-200+, the 

filaments filiform, free or basally connate, sometimes inserted on the base of the petals; anther 

thecas small, about as long as wide, didymous; 

pollen 3-colporate; pistils 1 or 3, the carpels 

2-14, the styles 2-14, filiform or short, free, 

stigmas apical; ovary 2- 14-locular, the ovules 2 

per locule, axile near the base of the locule; fruit 

baccate but eventually dehiscent; seeds 1 -few, 

often densely hairy; embryo straight, the cotyledons thick, plano-convex, the radicle very short; 

endosperm 0. 

Composition: 4 genera, 50 species. 

Distribution: Tropical American forests, especially the Amazon region. 

HYPERICACEAE (Figure 53a,b).-Mostly 

shrubs and herbs, sometimes trees, with schizogenous secretory cavities, sometimes resinous 

(Vismia); xylem vessel perforation plates usually 

simple, sometimes also scalariform; leaves opposite or verticillate, very rarely the lower leaves 

alternate (Psorospermum), simple, entire, usually 

gland-dotted, exstipulate; inflorescences mostly 

terminal, cymose-paniculate, or the flowers solitary; flowers bisexual, the parts hypogynous; sepals 4-5, imbricate, free or basally connate; petals 4-5, free, sometimes with a basal nectar scale; 

stamens numerous (- 50-200), rarely less than 

10 (5), the filaments elongate, free or often connate in 3-5 bundles, or basally monadelphous; 

anthers dorsifixed, often small, slightly longer 

than wide; pollen 3-colporate; pistil 1, the carpels 

3-5, the styles mostly free, very rarely connate 

high, stigmas apical; ovary 1- or 3-5-locular, the 

ovules numerous, axile or rarely parietal, bitegmic, tenuinucellar, anatropous; fruit a loculicidal or septicidal capsule, or berry; seeds minute, 

not arillate; cotyledons hemi-cylindric, flat or 

rarely curved, 0.3 the length of the embryo; 

endosperm 0. 

Composition: 8 genera, 350 species. 

Distribution: Mainly pantemperate and 

mountains of the tropics; tropical lowland. 

ELATINACEAE (Figure 53c,d).-Herbs and 

shrublets, sometimes aquatic; xylem vessel perforation plates simple; leaves opposite or verticillate, simple, entire or dentate, stipulate; inflorescences axillary, dichasia or the flowers solitary; 

flowers bisexual, minute, the parts hypogynous; 

sepals 3-5 (2-6), imbricate, free or basally connate; petals 3-5 (2-6) or 0, free; stamens 3-10, 
NUMBER 58 113 

FIGURE 53.-HYPERICACEAE: a, Hypericum brasiliense bud, 1,s. of flower, 2 sepals, petal, stamina1 

phalanx, views of anther, part of leaf showing glands; b, floral diagram, pistil, C.S. of ovary, 

fruit with calyx, seed and C.S. and I.s. of same, Vismiajapurensis pistil, one of the stigmas, seed 

(after Martius, 1840-1906). ELATINACEAE: c, Elatine lindbergii views of stamen, seed, I.s. of 

same, apical view of seed, floral diagram, flower, same from above, young flower; d, Bergia 

arenarioides part of plant in flower, seed, 1,s. of same, floral diagram, flower, petal, stamens, 

pistil, dehiscent fruit, part of fruit and columella (after Martius, 1840-1906). 

in 1 or 2 whorls, free, the filaments sometimes merous per locule, axile, bitegmic, crassinucellar, 

subulate; anthers small, as long as wide; pollen 3- also tenuinucellar (?), anatropous; fruit a septicicolporate; disk absent; pistil 1, the carpels 3-5 dal or septifragal capsule, sometimes leaving a 

(2-6), the styles short, 3-5 (2-6), free, stigmas columella; seeds minute, sometimes longitudiapical; ovary 3-5 (2--6)-locular, the ovules nu- nally and horizontally sculptured; embryo 
114 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

straight or arcuate; cotyledons not wider than 

the radicle, 0.2-0.6 the length of the embryo; 

endosperm 0. 

Composition: 2 genera, 40 species. 

Distribution: Almost cosmopolitan, absent 

from arctic Northern Hemisphere. 

DIPTEROCARPACEAE (Figure 5 4a ,b). -Trees, 

usually resinous and mucilaginous; xylem vessel 

perforation plates simple; cortical vascular bundles are characteristic of the young stem; leaves 

alternate, simple, entire, stipulate; inflorescences 

axillary or terminal panicles, rarely cymose 

(Upuna and some Vatica); flowers bisexual, the 

parts hypogynous or subperigynous, rarely the 

calyx adnate to the ovary; sepals 5, connate part 

way, the lobes imbricate or valvate, often 2-5 

accrescent and wing-like in fruit; petals 5, markedly convolute, free or shortly connate basally, 

often hairy; stamens usually numerous (to -40), 

sometimes 5, 10 or 15, frequently adnate to the 

base of the petals, the filaments short or elongate, 

usually connate, sometimes in bundles; anthers 

dorsifixed or basifixed, the connective usually 

apically produced, dehiscing longitudinally or apically; pollen 3-colpate or colporoidate; pistil 1, 

the carpels 3 (2 only in Stemonoporus), the style 

simple or 3-lobed, a fleshy stylopodium often 

present, the stigma small, apical; ovary 3-locular 

(rarely incompletely so), the ovules 2 per locule, 

axile, (rarely parietal?), bitegmic, crassinucellar, 

anatropous; fruit a nut, mostly I-seeded; embryo 

large, the cotyledons contorted, and crumpled, 

and enclosing the radicle, sometimes plano-convex, much wider than the radicle, 0.7 the length 

of the embryo; endosperm mostly 0. 

Composition: 15 genera, -500 species. 

Distribution: Tropical, centered in rain forest 

of Malaya and the East Indies, extending to 

India; few in tropical Africa. 

HUMIRIACEAE (Figure 54c-e).-Trees and 

shrubs, sometimes resinous; xylem vessel perforation plates exclusively scalariform, with 15-25 

bars, the vessel members extremely long; leaves 

alternate, simple, crenate or entire, stipulate or 

exstipulate; inflorescences axillary or rarely terminal thyrses; flowers bisexual, the parts hypogynous; sepals 5, imbricate, connate part way; 

petals 5, free; stamens 10, 20, 30, or 50 to 180, 

in 1-4 whorls, the filaments subulate or filiform, 

connate basally; anther sacs small, the connective 

markedly produced apically; pollen 3-colporate, 

the colpi short; disk cupular or of 10-20 free 

glandular scales surrounding the base of the 

ovary; pistil 1, the carpels 5-7, the style simple, 

stigmas apical; ovary 5-7 locular, the ovules 1-3 

per locule, axile, bitegmic, crassinucellar, anatropous; fruit a drupe, the endocarp woody, sometimes ribbed, sometimes with numerous resinfilled cavities; seeds usually 1-2; embryo straight 

or slightly curved; cotyledons oblong or ovate, 

often subcordate basally, less than 0.5 to 0.7 the 

length of the embryo, 1.5 times wider than the 

radicle; endosperm copious, fleshy and oily. 

Composition: 8 genera, 50 species. 

Distribution: Tropical America, centered in 

the Amazon basin; two species in tropical west 

Africa. 

ANCISTROCLADACEAE (Figure 54f,g).-Lianas, 

rarely shrubs, the twigs with coiled hooks; xylem 

vessel perforation plates simple; leaves alternate, 

simple, entire, with scattered minute glandular 

pits, stipulate or exstipulate; inflorescences terminal racemes, panicles or cymes; flowers bisexual, the parts perigynous; calyx adnate to the 

base of the ovary, the lobes 5, imbricate, accresFIGURE 54.-DIPTEROCARPACEAE: a, Dipterocarpus bourdilloni bud, pistil and base of calyx, I.s. of ovary, views of 

anther, Dryobalanops aromatica I.s. of floizer, 1,s. of fruit, 

floral diagram; 6, D. cumphora C.S. of seed, embryo unfolded 

(after Brandis; Baillon, 1866- 1895; Lindley). HUMIRIACEAE: 

c, Vantanea paniculata bud, 1,s. of bud, calyx laid open, petal, 

part of androeciuni, veiws of anther, C.S. of anther, part of 

hypogynous cupule, floral diagram; d, Humiria balsamfera 

pistil, stigma from above, 2 views of anther of longer filament, 3 vie\vs of anther of shorter filament, C.S. of anther of 

longer filament, H. jloribunda 1,s. of seed, embryo; e. H. 

balsamifera 1,s. of ovary, H. jloribunda fruit, putamen, 1,s. of 

same, C.S. through middle of same, Sucoglottis cuspidata floral 

diagram (after Martius, 1840- 1906). ANCISTROCLADACEAE: 

f, Ancistrocladus guineensis 1,s. of flower, A. pinangianus flowering t\+,ig, fruit, A. harmandii long and short stamens, pistil; 

g, bud, 2 sepals, 2 petals (after Baillon, 1866-1 895; Ridley; 

Gagnepain). 
NUMBER 58 115 

A 
116 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

cent, unequal and wing-like in fruit, often with 

conspicous dorsal glandular pits; petals 5, connate basally; stamens 10, rarely 5, uniseriate, the 

filaments basally broad and connate, apically filiform; anthers basifixed, slightly longer than 

wide; pollen 3-(4)-colpoidate; pistil 1, the carpels 

3, the style 1, stigmas 3; ovary unilocular, ovule 

1, basal, bitegmic, semianatropous; fruit a woody 

nut, the seed large; embryo 0.5 the length of the 

endosperm; cotyledons divergent, as broad as the 

radicle, 0.4 the length of the embryo; endosperm 

ruminate, much folded and coiled, brain-like, 

starchy. 

Composition: 1 genus, 20 species. 

Distribution: Tropical western Africa, Sri 

Lanka, India to Malaya. 

MARCGRAVIACEAE (Figure 55a,b).-Shrubs, 

usually climbing and epiphytic, rarely tree; xylem 

vessel perforation plates simple or both simple 

and scalariform, with few to many bars, vessel 

members very long; secretory canals sometimes 

present; leaves simple, alternate, entire, exstipulate; inflorescences terminal cymose umbels, or 

spikes; flowers bisexual, the parts hypogynous; 

bracts modified into pitcher-like, saccate, urnshaped or spurred, colored nectaries; sepals 4-5, 

imbricate, free or connate basally; petals 4-5, 

free or connate basally or calyptrate; stamens 

numerous (to -40) to 3, in 1 whorl, free or 

connate basally and sometimes adnate to the base 

of the corolla, the filaments broadened toward 

the base; anthers oblong, 2-5 times longer than 

wide, basi- or dorsifixed; pollen 3-colporate or 

colporoidate; pistil 1, the carpels 2-8, the style 

1, very short or the stigmas sessile, radiate; ovary 

initially unilocular, later 2-8-locular, the ovules 

bitegmic, tenuinucellar, anatropous, numerous, 

in several series, the placentas thick; fruit a berry 

with leathery epicarp, eventually slightly loculicidally dehiscent basally; seeds numerous, small; 

embryo straight or slightly arcuate; cotyledons 

1-2 times the width of the radicle, 0.3-0.7 the 

length of the embryo; endosperm 0. 

Composition: 

Distribution: Tropical America. 

CARYOCARACEAE (Figure 55c-e).-Trees and 

shrubs; xylem vessel perforation plates simple, 

5 genera, - 120 species. 

occasionally with some scalariform plates; leaves 

opposite or alternate, digitately compound, the 

leaflets 3-5, the margins entire, stipulate or exstipulate; inflorescences terminal racemes; flowers bisexual, the perianth hypogynous; sepals 5- 

6, imbricate or open, connate basally; petals 5- 

6, free or calyptrate; stamens subperigynous, numerous (-30-60), in 5-6 series, the filaments 

long, filiform, colored, often warty, free or connate in 1 whorl basally or in 5 bundles, bent in 

bud, sometimes some lacking anthers; anthers 

small, oblong, dorsifixed; pollen 3-colporate or 

4-ruporate; pistil 1, the carpels 4-20, the styles 

4-20, filiform, usually long (short in Anthodiscus), stigmas punctate; ovary 4-20-locular, the 

ovules 1 per locule, axile, bitegmic, anatropous; 

fruit a drupe, the epicarp sometimes coriaceous, 

endocarp woody, muricate, splitting into mericarps; seeds reniform; cotyledons small, hookedinflexed, 0.04-0.1 the length of the embryo, the 

radicle much thickened, curved or twisted, 7-1 5 

times wider than the cotyledons; endosperm 

scanty or 0. 

Composition: 2 genera, 25 species. 

Distribution: Tropical America, centered in 

the Amazon basin. 

OCHNACEAE (Figure 56a-e).-Trees and 

shrubs, rarely herbs; xylem vessel perforation 

plates typically simple, but those of some genera 

with few or many bars; cortical bundles are 

nearly always present in the stem; leaves alternate 

(also opposite in Rhytidanthera), simple or very 

rarely pinnately compound, often with numerous 

pinnate veins extending from the midrib, mostly 

sharply serrate, more rarely entire, stipulate; inflorescences axillary and terminal panicles, racemes or cymes, sometimes umbelliform; flowers 

bisexual, the parts hypogynous; sepals 5 (3-1 0), 

imbricate, free or basally connate, sometimes 1 

or 2 accrescent and wing-like in fruit (Lophireae), 

sometimes colored; petals 5 (4-1 2), free; stamens 

persistent, numerous (to -80) or 5 or 10, in 1-5 

whorls, the filaments filiform, free or sometimes 

connate basally, or united with staminodes into 

a tube (Lavradia); anthers linear, basifixed, dehiscing longitudinally or by a terminal pore; pollen 3-colporate, occasionally 4-rupate; pistil l, 
NUMBER 58 117 

C 

\ 

d 

i 

FIGURE 55.-MARCGRAVIACEAE: a, Souroubea crasszpes pedicel with bud and bract, pedicel with 

flower and bract, Marcgrauia polyantha floral diagram, bud, Norantea delpiniana views of 

stamen, C.S. of ovary; b, N. adamantium pistil, Is. of ovary, seed, I.s. of same (after Martius, 

1840-1906). CARYOCARACEAE: c, Caryocar crenatum flower, petal and stamens, views of stamen, 

1 style, 1,s. of ovary, C.S. of ovary; d, C. glabrum pistil, C.S. of semimature fruit, putamen, 

Anthodzscus obouatus pistil, calyx and one of the stamens, C.S. of ovary; e, 1,s. of fruit, embryos 

(after Martius). 

the carpels 2-5 (-15), the style 1, elongate, sometimes gynobasic, the stigmas apical; ovary deeply 

lobed or entire, 2-5 (l-l5)-locular, sometimes 

surrounded by an annular basal disk, the ovules 

1-many per locule, axile or parietal, bitegmic, 

tenuinucellar, anatropous or campylotropous; 

fruit of several drupaceous fruitlets on an enlarged, red, fleshy torus, or a berry or a septicidal 
118 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 56.-oCHNACEAE: a, Ouratea spectabilis calyx, 1,s. of flower, fruit, 1,s. of same; b, flower, 

C.S. of 2 ovaries, views of stamen, pistil: c, Luxemburgia nobilis pistil and several stamens, C.S. of 

fruit, floral diagram; d, Sauuagesia sprengelii floral diagram, corolla in bud, calyx and internal 

squamae and external processes, sepal, petal, stamen and C.S. of anther, squama and 2 stamens, 

another view of stamen; e, external processes, C.S. of ovary, pistil, dehiscent fruit, valves, C.S. of 

fruit, seed, 1,s. of same (after Martius, 1840-1906). 

capsule; seeds 1 -many; embryo usually straight, 

in Ochna large, the cotyledons thick, plano-convex, 0.9 the length of the embryo, the radicle 

slightly invested, and endosperm 0; in Luxemburgieae the embryo is linear or spatulate, 0.7 

the length of the endosperm, the cotyledons 0.5 

the length of the embryo. 

Composition: -35 genera, -500 species. 

Distribution: Pantropical, centered in South 

America. 
NUMBER 58 119 

STRASBURGERIACEAE (Figure 57a).-Tree 

with cortical vascular bundles and large mucilage 

cells and passages in tissues; xylem vessel perforation plates simple and scalariform; leaves alternate, simple, remotely toothed, coriaceous, stipulate; flowers bisexual, solitary, axillary; sepals 

8-10, free, unequal, spirally arranged; petals 5, 

free, rather fleshy; stamens 10, the filaments 

elongated, free; anthers large, elongated, oblong, dorsifixed; pollen 3(4)-colporate, brevicolpate; disk thick, annular, 10-lobed, surrounding 

the base of the ovary; pistil 1, the carpels 5, style 

1, elongated, the stigma apical, capitate; ovary 

superior, 5-locular, the ovules 2 per locule, axile; 

fruit a large berry, the seeds trigonous; cotyledons thick, subelliptic, the radicle short; endosperm fleshy. 

Composition: 1 genus, 1 species. 

Distribution: New Caledonia. 

DIEGODENDRACEAE (Figure 57b).-Shrub or 

small tree; xylem unknown; leaves alternate, simple, entire, pellucid-punctate, smelling of camphor when crushed, the stipules long, intrapetiolar, coiled around the bud; inflorescence a terminal cymose panicle; flowers bisexual, actinomorphic, the parts hypogynous, fragrance of 

rose; sepals 5 (-6), free, imbricate, unequal; petals 5 (-6), free, large, slightly unequal; stamens 

very numerous (to -434), free, the filaments 

elongate, filiform; anthers small, about as long as 

wide, basifixed; pollen tricolporate; pistil 1, the 

carpels 2 (-4), the style 1, elongate gynobasic, 

the stigma apical, punctiform; ovaries 2 (-4), the 

ovules 2 per carpel, basal; fruit of 1-3 dry coriaceous mericarps, 3 x 2.5 X 2 cm, with very 

numerous verrucosities, and very numerous 

small peltate glands; seeds 2.5 X 2 X 1.2 cm, 

coriaceous, smooth; cotyledons foliaceous, 2.3 X 

2.0 X 0.3 cm, narrowly notched for 5-6 mm 

basally; radicle invested, 0.7 cm long; endosperm 

0. 

Composition: 1 genus, 1 species. 

Distribution: Madagascar. 

SPHAEROSEPALACEAE (Figure 57c,d).-Trees, 

sometimes large, and shrubs, rich in mucilage, 

with extremely tenacious and fibrous bark; xylem 

vessel perforation plates simple; leaves alternate, 

simple, entire, the stipules intrapetiolar and caducous; inflorescences axillary and terminal umbelliform cymes arranged in a panicle, the pedicels stout; flowers bisexual; the parts hypogynous, densely streaked with short lines (? mucilaginous or resinous); sepals 4 in 2 whorls, rarely 

6, free, markedly imbricate and differing in size, 

coriaceous; petals 4 (5-8), in 2 whorls, free, 

unequal, imbricate, about the size of the sepals 

or smaller, sometimes caducous; stamens 25- 

160, in 2-4 whorls, the filaments elongate, free 

or shortly connate, the anthers small, dorsifixed, 

didymous, the sacs separated by a broad glandular connective, dehiscing longitudinally; pollen 3-4 (-6)-colpor(oid)ate, finely echinate; disk 

large or rarely absent (Rhopalocarpus thouarsianus), cupular, below and surrounding the base 

of the ovary; a small gynophore present; pistil 1, 

the carpels 2-4 (-5), the style 1, elongate, rarely 

gynobasic (Dialyceras), slightly geniculate, the 

stigma capitate or very shortly 3-4-lobed; ovary 

2-4 (-5)-locular and lobed or of 4 separate parts 

(Dialyceras), the ovules 2-9 per locule, basal or 

basal-axile, bitegmic, anatropous; fruit a nut or 

dryish berry, some locules aborting, or a schizocarp of horn-shaped parts, the seeds mostly 1 per 

locule, large, initially enclosed in a translucent 

glutinous mass; embryo straight, filling most of 

the seed, the cotyledons separate from one another (right and left), thin, undulate, foliaceous, 

basally cordate, bilobed apically, the lobes lobulate, sometimes laciniate, 0.65-0.8 the length of 

the embryo, 4-9 times wider than the terete 

radicle; endosperm moderate, sometimes ruminate. 

Composition: 2 genera, 14 species. 

Distribution: Madagascar. 

SCYTOPETALACEAE (Figure 58a,b).-Trees 

and shrubs; xylem vessel perforation plates simple or with few bars (to 12); leaves alternate, 

simple, entire or dentate, often distichous, exstipulate; inflorescence terminal panicles or axillary 

racemes, or umbelliform or racemiform cauliflorous fascicles; flowers bisexual; calyx short, flattish or cupular, the margin entire or 3-4- 
120 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

a 

FIGURE 57.-sTRASBURGERIACEAE: a, StrasbUrgerZU calliantha flower, 1.S. Of Same, floral diagram, calyx from below, petal (after Engler). DIEGODENDRACEAE: b, Diegodendron humberti 

flowering branch, stipules and C.S. of a pair of stipules (after Capuron, 1963a). SPHAEROSEPALACEAE: c, Dialyceras parufoliurn 2 carpels opened to show ovules, fruit, mericarp opened showing 

1,s. of seed, Rhopalocarpus excelsus flowering twig, flower; d, pistil, I.s. of ovary, R. thouarianus 

fruit, 1,s. of a fruitlet, embryo, R. alternfolius pistil and disk, stamen, group of stamens, dorsal 

view of anther, 1,s. of ovary, petal (after Capuron, 1963b; Hutchinson, 1973). 
NUMBER 58 121 

b 

e 

FIGURE 58.-sCYTOPELTALACEAE: a, Scytopetalum klazneanum twigs with flowers and young 

fruit, I.s. of bud, views of anther, stigmas, C.S. of ovary; b, fruit, I.s. of same showing the embryo 

and endosperm of the seed (after Engler; Pierre). SARCOLAENACEAE: c, Sarcolaena multzjlora 

flower, 1,s. same, same without the perianth, Schirolaena rosea I.s. of flower with part of corolla 

cut off, pistil with 1 ovary locule opened, 2 fruits one dehiscent; d, Leptolaena multzjlora 

flowering branch, floral diagram, pistil, 1,s. of same, part of disk and androecium, fruit; e, I.s. 

of fruit, seed, C.S. of same (after Baillon, 1866-1 895). 

toothed; petals 3- 16, sometimes basally connate, long as wide, basifixed, dehiscing by a pore or 

valvate; stamens numerous (-80), rarely 10, slit; pollen 3-colpate or 3-colpor(oid)ate; pistil 1, 

sometimes on an annular disk, sometimes shortly the carpels 3-8, the style 1, stigma small, apical; 

connate, the filaments filiform; anthers about as ovary superior, 3-8-locular, the ovules 2-several 
122 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

per locule, axile, subapical, unitegmic, crassinucellar (?), anatropous; fruit a loculicidal woody 

capsule, rarely a l-seeded drupe; seeds sometimes long-haired; embryo 0.8 the length of the 

endosperm; cotyledons broad and thin, 0.5 the 

length of the embryo; endosperm copious, often 

ruminate. 

Composition: 5 genera, 30 species. 

Distribution: Tropical western Africa. 

SARCOLAENACEAE (Figure 58c-e).-Shrubs 

and trees; xylem vessel perforation plates simple; 

leaves alternate, simple, entire, stipules present, 

often large; inflorescences cymes or panicles, or 

the flowers solitary or paired; flowers bisexual, 

fairly large, often subtended by bracts; sepals 3- 

5, free, sometimes unequal, imbricate; petals 5- 

6, free; stamens numerous (to -50), rarely 5-10, 

the filaments filiform, inserted on a disk, free or 

sometimes shortly connate or in 5 bundles; staminodes external to the stamens; anthers oblongish, slightly longer than wide, basifixed or dorsifixed, dehiscing longitudinally; pollen 3- 

(demi)colporate; pistil 1, the carpels 3-5, style 1, 

the stigmas 3-5, apical; ovary 3-5-locular, the 

ovules 2-several per locule, axile; fruit a loculicidal capsule or indehiscent and l-seeded; embryo 

0.5 the length of the endosperm; cotyledons 

foliaceous, plane or undulate, twice as wide as 

the radicle, 0.5 the length of the embryo; endosperm copious, fleshy or corneous. 

Composition: 8 genera, 40 species. 

Distribution: Madagascar. 

EBENALES 

Trees and shrubs; xylem vessel perforation 

plates simple, rarely also a few scalariform with 

few bars; laticifers sometimes present; leaves alternate, rarely opposite, simple, entire, toothed 

in only 1 species, usually exstipulate; flowers 

bisexual or unisexual; sepals 3-8 (1 2), connate; 

petals 3-8, connate; stamens 3-50 (-120?), usually 2-4 times the number of corolla lobes, one 

series sometimes staminodal, hypogynous or adnate to the corolla, the filaments free, rarely 

connate; anthers adnate, basifixed or dorsifixed; 

pollen 3-4 (-6)-colporate; pistil 1, the carpels 2- 

16, styles 1-8, free or connate, the stigmas apical; 

ovary superior, 2-1 6-locular, the ovules 1 (2) per 

locule, axile; fruit a berry, rarely a capsule; embryo straight, usually moderate-sized; endosperm 

copious to absent. 

Distribution: Pantropical, mainly in rain forest, few in temperate and dry regions. 

Chemistry: Kubitzki mentions the occurrence 

of protoaporphine and aporphine in some Ebenales; Gibbs does not mention them. The Ebenaceae are unique in having a + Juglone test and 

naphthaquinones and other naphthalene derivatives. The Sapotaceae differ in having pyrrolizidine alkaloids and d-quercitol. 

EBENACEAE (Figure 59a,b).-Trees and 

shrubs; xylem vessel perforation plates simple; 

hairs occasionally 2-shanked; leaves alternate, 

rarely opposite or whorled, simple, entire, exstipulate; inflorescences axillary or rarely cauliflorous cymes or the flowers solitary; flowers mostly 

unisexual, the plants often dioecious, sometimes 

polygamous; sepals 3-7, connate; petals 3-7, 

connate, usually contorted and imbricate, rarely 

valvate; stamens 3-50 (-1 20?), usually 2-4 times 

the number of corolla lobes, rarely equal in number and alternate with them, hypogynous or inserted on the corolla, the filaments free or connate in pairs; anthers elongated, adnate or basifixed, dehiscing longitudinally, more rarely by 

apical pores, apiculate; pollen 3-colporate; pistil 

1, the carpels 2-1 6, styles 2-8, free or connate 

part way, the stigmas apical; ovary superior, 2- 

16-locular, sometimes incompletely so, the ovules 

1 (2) per locule, axile-subapical, bitegmic, tenuinucellar, anatropous; fruit a berry, rarely tardily 

dehiscent apically by valves; embryo straight, 

0.5-0.6 the length of the endosperm; cotyledons 

2-3 times wider than the radicle, 0.5-0.7 the 

length of the embryo; endosperm copious, very 

hard. 

Composition: 2 genera, 500 species. 

Distribution: Tropical and subtropical, few 

temperate; centered in lowland rain forest of the 

East Indies, extending to India, Japan, and Australia; next most abundant in Africa and then 
NUMBER 58 123 

a 

d 

e 

FIGURE 59.-EBENACEAE: a, Diospyros coccolobaefolia Is. of pistil, stigma, corolla laid open 

showing staminodes, D. sericea flower in axil of leaf, C.S. of ovary, D. guianensis part of corolla 

with stamens, views of stamen; 6, calyx, corolla, fruit, I.s. of same, views of seed, I.s. of same, 

embryo, same with cotyledons laid open, axillary flowers (after Martius, 1840-1906). SAPOTACEAE: c, Manilkara zapota axillary flowers, floral diagram, flower, calyx, corolla, same laid 

open showing insertion of stamens and broad staminodes; d, I.s. of pistil, stigma, C.S. of ovary, 

views of stamen, views of seed showing long hilum, I.s. of seed without testa, embryo laid open, 

hairs from pistil; e, Lucuma caimito 1,s. of bud, C.S. of fruit (after Martius, 1840-1906). 
124 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

America; eastern United States; southwestern 

Asia. 

SAPOTACEAE (Figure 59c-e).-Trees and 

shrubs with laticiferous canals in the pith, cortex, 

and leaves; xylem vessel perforation plates simple, rarely also occasional scalariform plates with 

1-7 bars; hairs commonly 2-shanked; leaves alternate, very rarely subopposite or verticillate, 

simple, entire, very rarely toothed (Chrysophyllum 

imperiale), coriaceous, usually exstipulate; inflorescences axillary or cauliflorous cymose fascicles, or the flower solitary; flowers bisexual, more 

rarely the plants polygamous; sepals 4-8 (12), 

sometimes in 2 series or spirally arranged, shortly 

connate; petals 4-8 in 1 or 2 series, connate, 

imbricate; stamens 4-24, epipetalous, rarely also 

connate in a tube (Aulandra), 1-3 times the 

number of petals, in 1-3 series, if uniseriate then 

opposite the petals, one series often staminodal 

and petaloid; anthers about as long as wide or 

elongated, basifixed or dorsifixed, commonly extrorse; pollen 3-4 (-6)-colporate--slightly ruporate; hypogynous annular disk rarely present; 

pistil 1, the carpels 4-12 (2), style 1, elongated 

or short, the stigma(s) apical; ovary superior 4- 

12 (2)-locular, the ovules 1 per locule, axile, 

unitegmic, the integument thick, tenuinucellar, 

anatropous or suborthotropous; fruit a berry, 

rarely a capsule; embryo straight, as long as the 

endosperm; cotyledons thin, or thick and planoconvex, rarely conferruminate, 4.5-7.0 times 

wider than the radicle, 0.5-0.9 the length of the 

embryo; endosperm moderate or 0, fleshy or 

hard. 

Composition: -50 genera, 800 species. 

Distribution: Pantropical, mainly in lowland 

and lower montane rain forest, few temperate. 

STYRACALES 

Trees and shrubs; xylem vessel perforation 

plates scalariform and simple, with fewer than 20 

bars; laticifers sometimes present; leaves alternate, simple, entire or toothed, the indument 

commonly stellate or lepidote, exstipulate; flowers bisexual, rarely unisexual; sepals 4- 10, connate, the lobes short; petals 4-10, free or basally 

connate; stamens 4-10 (-32), the filaments free 

or connate, hypogynous or inserted on the corolla, the anthers linear, basifixed or adnate; 

pollen 3-4 (2-5)-colporate or -porate; pistil 1, 

the carpels 2-5, style 1, the stigma(s) apical, 

rarely decurrent ventrally; ovary superior to inferior, 1 -5-locular, the ovules 1 -several per locule, axile or apical; fruit nut-like, sometimes 

winged, or a loculicidal capsule or drupe; embryo 

moderate-sized; endosperm usually copious. 

Distribution: Mostly in eastern Asia, the 

United States, tropical South America, and tropical Africa. 

Chemistry: Huber considers Styracaceae and 

Symplocaceae related to his Cornales, which are 

derived from relatives of Theaceae. Alangium 

has isoquinoline alkaloids of the emetine group 

and certain pyridine alkaloid, not known to occur 

in Cornales. The former occur principally in 

Rubiaceae, but also in Icacinaceae; the latter 

occurs in various families. 

STYRACACEAE (Figure GOu,b).-Trees and 

shrubs; xylem vessel perforation plates scalariform with fewer than 20 bars, some simple perforations in Bruinsmiu; leaves alternate, simple, 

usually entire, sometimes toothed, the indument 

mostly stellate or lepidote, exstipulate; inflorescences axillary or terminal, mostly racemes or 

panicles, sometimes cymes, rarely the flowers 

solitary; flowers bisexual, rarely the plants polygamodioecious; sepals 4-5 (-7), connate, the 

lobes valvate or the calyx open in bud, often 

truncate; petals 4-5 (-7), mostly basally connate, 

rarely free?; stamens 4- 10 (- 14), alternate with 

the corolla lobes if equal to them in number, the 

filaments usually connate and often inserted basally on the corolla, rarely hypogynous; anthers 

usually elongated, rarely round, basifixed, sometimes the connective produced apically; pollen 3- 

colpor(oid)ate (a constricti-colpate); pistil 1, the 

carpels 3-5, style 1, the stigma apical, punctiform 

or 3-5-lobed; ovary superior, semi-inferior or 

inferior, 3-5-locular, often unilocular above, 

sometimes at length entirely unilocular, the 

ovules 1 to several per locule, axile, l-2-tegmic, 
NUMBER 58 125 

1 a 

--===-- 

... ., 

Li 

FIGURE 60.-sTYRACACEAE: a, Styrax martii floral diagram, bud, flower (calyx removed), part 

of corolla laid open showing insertion of stamens, views of stamen; 6, Is. of pistil, C.S. of ovary, 

fruit and calyx, 1,s. of fruit showing the embryo in copious endosperm, seed, S. leprosum 

indument scale from leaf, C.S. of ovary (after Martius, 1840-1906). LISSOCARPACEAE: c, 

Lissocarpa benthami axillary inflorescence, corolla laid open, bud, anther, 1.s. of ovary and calyx, 

C.S. of ovary, fruit, apex of fruit showing the persistent calyx lobes; d, seed, I.s. of same showing 

the embryo (after Oliver). ALANGIACEAE: e, Alangium decapetalum 1,s. of flower, A. begoni@lzum 

twig with axillary inflorescences, bud, 1,s. of flower, stamen, pistil, fruit and C.S. of same, A. 

salvzi$olium 1,s. of fruit (after Baillon, 1866-1895; Wangerin). 
126 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

tenuinucellar, anatropous or semi-anatropous; 

fruit nut-like, sometimes winged, or a loculicidal 

capsule, rarely a drupe, the seeds 1 or few, rarely 

numerous (Huodendron, Alniphyllum); embryo 

straight or slightly curved, 0.8 the length of the 

endosperm; cotyledons thin, (1) 2-5 times wider 

than the radicle, (0.2) 0.5-0.8 the length of the 

embryo; endosperm copious, fleshy. 

Composition: 12 genera, 180 species. 

Distribution: Japan to southeastern Asia, 

western East Indies; eastern and western United 

States to the northern % of South America, West 

Indies; one species in the Mediterranean region. 

LISSOCARPACEAE (Figure GOc,d).-Small trees; 

xylem vessel perforation plates simple and scalariform; leaves alternate, simple, entire, exstipulate; inflorescences axillary, cymose; flowers bisexual; sepals 4, connate, imbricate; petals 4, 

connate, the lobes contorted in bud; corona of 8 

teeth at the apex of the corolla-tube; stamens 8, 

in one series, inserted on the corolla-tube, the 

filaments short; anthers linear, basifixed, the connective produced apically; pollen 3-porate; pistil 

1, the carpels 4, style 1, the stigma apical, simple 

or indistinctly 4-lobed; ovary inferior, 4-locular, 

the ovules 2 per locule, axile-subapical; fruit a 

drupe, the seeds 1-2; embryo subterete, straight, 

0.5 the length of the endosperm; endosperm 

copious, horny. 

Composition: 1 genus, 2 species. 

Distribution: Tropical South America. 

ALANGIACEAE (Figure 6Oe).-Trees and 

shrubs, rarely a liana, rarely spiny; xylem vessel 

perforation plates scalariform, typically with 

fewer than 10 bars, occasionally more, some species with simple perforations; laticifers present in 

flowers, fruit and leaves; leaves alternate, simple 

entire or with 1-3 large teeth or with small lobes, 

exstipulate; inflorescences axillary, cymose-paniculate; flowers bisexual, the plants rarely dioecious or polygamous (Alangium grisolleoides); sepals 4- 10, connate, the calyx truncate or shorttoothed; petals 4-10, linear, valvate, free or basally connate; stamens 4-1 0 (-32), alternipetalous 

when equal to the number of petals, free or 

slightly connate basally, the filaments sometimes 

inserted on the base of the petals; anthers linear, 

adnate with connective between the sacs or basifixed; pollen 3-4 (2-5)-colporate, colpoidorate 

or porate; disk epigynous; pistil 1, the carpels 2- 

3, style 1, the stigma punctiform or 2-3-lobed, 

apical or decurrent ventrally; ovary inferior 1 

(-3)-locular, sometimes 1-locular at the apex and 

2-3-locular at the base, the ovules 1 per locule, 

apical, unitegmic, crassinucellar, anatropous; 

fruit a drupe, the seed 1; embryo straight, 0.9 

the length of the endosperm; cotyledons thin, 

foliaceous, 3-6 times wider than the radicle, 0.6- 

0.9 the length of the embryo; endosperm copious 

or moderate, fleshy, ruminate, or friable. 

Composition: 1 genus, 17 species. 

Distribution: Japan, Korea, China to India, 

Malaya, the East Indies and northeastern Australia; tropical Africa, Madagascar. 

VIOLALES 

Trees, shrubs, and herbs, sometimes climbing; 

xylem vessel perforation plates scalariform with 

many or few bars, simple, or both; leaves usually 

alternate, simple, rarely compound, entire, 

toothed, lobed or divided, sometimes with extrafloral nectaries, sometimes distichous, usually 

stipulate; flowers bisexual or unisexual, actinomorphic or rarely zygomorphic, the parts hypogynous, perigynous or rarely epigynous; perianth 

rarely not differentiated into sepals and petals; 

sepals 3-5 (0, 2-1 5), free or connate; petals 3-5 

(0-1 5), rarely 2-4-seriate, very rarely spirally 

arranged, very rarely transitional to stamens, 

free, more rarely connate, sometimes inserted on 

the calyx, sometimes a corona present; stamens 

usually numerous, to -300, or 10-1, in 1-5 

series; staminodes sometimes present; filaments 

free or connate, sometimes in bundles or rarely 

in a column, hypogynous or inserted on the 

calyx, more rarely on the corolla; anthers dorsifixed, basifixed or rarely adnate, very rarely connate, sometimes the connective produced apically; pollen usually 3 (2- 12)-colporate, rarely 3 

(7-9)-colpate, very rarely 2-pantoporate; glandular disk often present; pistil 1, the carpels 3 
NU.MBER 58 

(2-lo), styles 1, 3 (-lo), very rarely repeatedly 

branched, the stigma(s) apical, very rarely decurrent ventrally, rarely sessile; ovary superior, 

rarely semi-inferior or inferior, usually 1 -locular, 

rarely 2- or 4-locular, the ovules usually numerous, rarely only 1, parietal, rarely axile or apical; 

fruit a berry or loculicidal, rarely septicidal capsule, rarely a drupe, samara or nutlet; seeds 

usually numerous, rarely only 1, often arillate, 

rarely winged; embryo straight, often nearly as 

long as the endosperm; endosperm usually copious, rarely scanty or absent. 

Distribution: Mostly pantropical and -subtropical, few temperate, in various habitats. 

Chemistry: The order is not aromatic, only 

very few terpenoids are recorded. Saponins are 

not characteristic of the order. Coumarins are 

absent or nearly absent. The order is not alkaloid-rich. Flacourtiaceae have fatty-acids of the 

chaulmoogric series in their seed-fats. Myrosincells containing isothiocyanate glycosides and 

glucotropaeolin are said to be present in Caricaceae. 

FLACOURTIACEAE (Figure 6 1 a-f.-Trees and 

shrubs, rarely climbing; xylem vessel perforation 

plates usually simple, but some species wholly 

scalariform; leaves alternate, very rarely opposite, simple, often distichous and coriaceous, serrate or entire, sometimes pellucid-dotted or 

lined, stipulate, very rarely exstipulate (Abatia); 

inflorescences axillary, terminal or below the 

leaves, racemes, cymes or rarely heads or sometimes catkin-like, or the flowers solitary, rarely 

epiphyllous or cauliflorous; flowers bisexual or 

the plants polygamous, monoecious or dioecious, 

the parts hypogynous, perigynous or rarely epigynous; perianth sometimes not clearly differentiated into sepals and petals, sometimes accrescent; sepals 6-3 (2-15), free or connate, imbricate or open in bud or nearly valvate; petals 8-3 

(0-1 5), free, sometimes 2-seriate, those of the 

inner series smaller, sometimes 2-3 times as 

many as the sepals, rarely spirally arranged, 

sometimes gradually transitional to stamens, 

sometimes with an opposite basal glandular scale, 

or corona (from staminodes?) extra- or intrastam127 

inal; stamens usually numerous (to 1 OO+), rarely 

only 1, in several or 1 series, the filaments elongate, free or connate in antepetalous bundles or 

all connate into a tube, or rarely united into a 

column; staminodes sometimes present; anthers 

linear to about as long as wide, dorsifixed or 

basifixed, sometimes the connective produced 

apically; pollen 3 (-6)-colporate; annular disk, 

glands or scales present between the corolla and 

stamens or between the stamens and pistil; pistil 

1, the carpels 2-10, the styles 1-10, free or 

connate, rarely repeatedly branched and with 

numerous stigmas (Dendrostigma, Mayna), the 

stigmas sometimes apical; ovary superior, rarely 

semi-inferior or inferior, usually unilocular, 

rarely 2- or 4-locular, rarely imperfectly 2-6- 

locular, the placentas 2- 10, usually parietal, 

rarely axile or apical, the ovules usually numerous, sometimes only 1 per placenta, bitegmic, 

crassinucellar, anatropous, semi-anatropous, or 

orthotropous; fruit usually a berry, sometimes a 

loculicidal capsule, rarely a drupe or samara, 

sometimes densely prickly or spiny; seeds numerous to 1, rarely winged or with sarcotesta, rarely 

covered with long hairs (Calantica), often arillate; 

embryo straight, usually large, 0.9 the length of 

the endosperm, rarely small; cotyledons thin, 6- 

20 times wider than the radicle, rarely terete, 

usually 0.8-0.9 the length of the embryo, rarely 

shorter than the radicle (Streptothamnus); endosperm copious or moderate, rarely scanty, fleshy. 

Composition: 

Distribution: Mostly tropical and subtropical, 

few temperate; tropical America, southern */3 of 

Africa, Arabia, India to southern China, the East 

Indies and northeastern Australia. 

LACISTEMATACEAE (Figure 62a-d).-Shrubs 

and small trees; xylem vessel perforation plates 

scalariform, often with more than 20 bars, vessel 

elements very long (1.6 mm.); leaves simple, alternate, entire or toothed, distichous, stipulate; 

inflorescences often clustered, axillary dense 

spikes or looser racemes; flowers minute, bisexual or the plants polygamous; sepals 4-6 or 0, 

unequal; petals 0; stamen 1, the filament relatively long, the anther small, about as long as 

-90 genera, - 1200 species. 
128 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

\ 
NUMBER 58 129 

wide, the anther sacs separate, sometimes distant, 

basifixed or adnate, dehiscing longitudinally or 

transversely; pollen 3-colporate; disk cupular, the 

stamen and pistil inserted upon it; pistil 1, the 

carpels 3 (2), the style 1, sometimes 0, the stigmas 

3 (2), slightly elongate; ovary superior, the ovules 

1-2 per placenta, parietal, bitegmic, crassinucellar, anatropous; fruit a loculicidal often fleshy 

capsule, the seeds 1 (-3) with aril (?) or easily 

removed outer testa; embryo straight, 0.9 the 

length of the endosperm, the cotyledons foliaceous, 0.4-0.6 the length of the embryo, 5.5- 

6.0 times the width of the terete radicle; endosperm (‘perisperm) copious, fleshy. 

Composition: 2 genera, -25 species. 

Distribution: Tropical America from Mexico 

to the northeastern half of South America. 

PASSIFLORACEAE (Figure 62e).-Shrubs, herbaceous climbers with axillary tendrils, rarely 

trees; xylem vessel perforation plates simple, 

rarely also scalariform(?); leaves alternate, simple, 

entire or lobed, rarely compound, the petiole 

often with stalked glands, stipulate, rarely exstipulate; inflorescences axillary cymes or racemes 

or the flowers solitary, rarely the middle flower 

replaced by a hook; flowers bisexual or the plants 

monoecious or dioecious; sepals 5 (3-8), free or 

shortly connate, often petaloid; petals 5 (0, 3-8), 

free or shortly connate; corona of 1 or more 

series of thread-like filaments or scales or annular; stamens 5 (3-lo), the filaments shortly conFIGURE 6 1 .-FLACOURTIACEAE: a, Banara brasiliensis flower, 

calyx, stamens and sepal, I.s. of flower, views of anther and 

C.S. of ovary, floral diagram; b, B. guianensis fruit, C.S. of 

same, seed, I.s. of same showing minute embryo and embryo 

enlarged, Carpotroche brasiliensis views of stamens, pistil, C.S. 

of ovary and views of embryo, I.s. of seed with large aril; c, 

Abatia tomentosa floral diagram, I.s. of flower showing ovules, 

corona and filaments (anthers removed), C.S. of ovary, sepal 

with coronal filaments, views of anther, dehiscent fruit, valve 

of fruit, 1,s. and C.S. of seed; d, Homaliumpedicellatum flower, 

I.s. of same. H. racoubea fruit, I.s. of same, C.S. of seed, 

embryo, floral diagram; e, Oncoba latqolia floral diagram, d 

flower, I.s. of same, Prockia crucis I.s. of flower, C.S. of ovary; 

f, .4phaerema spicata floral diagram, 2 views of flowers, pistil, 

1,s. of same, C.S. of ovary (after Martius, 1840-1 906). 

nate or in bundles, sometimes inserted on a gynophore; sometimes 5 staminodes present; anthers 

oblong, dorsifixed; pollen 3- 12-colporate; nectar 

is secreted at the base of the androgynophore; 

pistil 1, the carpels 3 (4-5), the styles 3 (4-5), 

free or connate into 1, the stigmas apical, often 

capitate; ovary superior, unilocular, the ovules 

usually numerous, rarely only 3, parietal, rarely 

apical, bitegmic, crassinucellar, anatropous or orthotropous; fruit a berry or loculicidal capsule; 

seeds with pitted testa surrounded by fleshy aril; 

embryo straight, 0.85-0.9 the length of the endosperm; cotyledons thin, 3-5.5 times wider 

than the radicle, 0.6-0.7 the length of the embryo; endosperm moderate, fleshy. 

Composition: 12 genera, -600 species. 

Distribution: Chiefly tropical and subtropical, 

few temperate; centered in America and Africa; 

few species from northeastern India to the East 

Indies, northern Australia, New Zealand, and 

Madagascar. 

MALESHERBIACEAE (Figure 62f).-Subshrubs 

and herbs; xylem vessel perforation plates mostly 

simple, scalariform plates with a few bars also 

present; leaves alternate, simple, entire, toothed 

or pinnatifid, exstipulate; inflorescences axillary 

or terminal racemes, panicles or cymes, or the 

flowers solitary; flowers bisexual; calyx-tube 

long, the lobes 5, short; petals 5, separate, inserted on the calyx-tube; corona denticulate or 

membranous, at the throat of the calyx-tube; 

stamens 5, the filaments filiform, inserted on the 

gynophore immediately below the ovary; anthers 

oblong, dorsifixed; pollen 3-colporate; pistil 1, 

the carpels 3 (4), the styles 3 (4), distant, inserted 

below the apex of the ovary, filiform, free, the 

stigmas apical, sometimes capitate; ovary superior, stipitate, unilocular, the ovules numerous, 

parietal; fruit a membranous capsule, loculicidal 

from the apex to the middle; seeds pitted, exarillate; embryo straight, 0.7-0.8 the length of the 

endosperm; cotyledons twice as wide as the radicle, 0.4-0.5 the length of the embryo; endosperm copious, fleshy, dotted. 

Composition: 1 genus, 25 species. 

Distribution: Dry habitats in the Andes of 
130 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 58 131 

southern Peru, northern Chile and western Argentina. 

TURNERACEAE (Figure 6 3 a ,b). -Shrubs, 

herbs, and trees; xylem vessel perforation plates 

mostly simple, occasionally also scalariform with 

numerous bars; leaves alternate, simple, sometimes lobed, dentate, the teeth sometimes glandular, often with a pair of glands at the base of 

the blade, stipulate or exstipulate; inflorescences 

axillary racemes or cymes or the flowers solitary; 

flowers bisexual; sepals 5, connate, rarely free; 

petals 5, separate, inserted on the calyx-tube, 

sometimes with a scale attached basally on the 

ventral surface, or with a corona; stamens 5, the 

filaments inserted on the calyx-tube; anthers as 

long as wide or elongate, dorsifixed near the 

base; pollen 3-colporate; pistil 1, usually heterostyled, the carpels 3, the styles 3, free, elongate, 

stigmas terminal, fringed; ovary superior, unilocular, the ovules numerous to 3, parietal, bitegmic, crassinucellar, anatropous; fruit a loculicidal 3-valved capsule; seeds pitted, arillate; embryo straight, 0.9 to as long as the endosperm; 

cotyledons thin, 2.5 times wider than the radicle, 

0.6 the length of the embryo; endosperm moderate, fleshy. 

Composition: 

Distribution: Tropical and subtropical America; southern 2/3 of Africa, Madagascar. 

ACHARIACEAE (Figure 63c).-Herbs and 

shrublets, sometimes scandent; xylem vessel perforation plates mostly simple, but also scalariform 

with few bars; leaves alternate, sometimes radical, simple, often palmately lobed, crenate or 

7 genera, - 120 species. 

FIGURE 62.-LACISTEMATACEAE: a, LUCiStemU robustum inflorescence (enlarged), part of twig with inflorescences; b, L. 

polystachyum flower, 6 flower, perianth, views of stamen, 

pistil; c, L. grandijiolium disk, L. myricoides 1.s. of ovary, L. 

intermedium fruit, L. grandijiolium dehiscent fruit, valve of 

fruit; d, L. pubescens aril, seed with aril removed, I.s. of seed, 

embryo (after Martius, 1840- 1906). PASSIFLORACEAE: e, Passijora serrata flower, P. data C.S. of ovary, 1,s. of flower, 

seed, 1,s. of fruit (after Martius, 1840-1906). MALESHERBIACEAE:~, Malesherbia rugosa flowering twig, I.s. of flower, M. 

taltalina flower laid open, dehiscent fruit, M. rugosa seed, 

Is. of same (after Baillon, 1866-1895; Ricardi, 1967). 

serrate, exstipulate; inflorescences axillary racemes, fascicles, or the flowers solitary; plants 

monoecious; sepals 3-5, free or adnate to the 

corolla; petals 3-5, connate into a campanulate 

corolla; stamens 3-5, inserted on the corolla; 

staminodes 3-5, short, fleshy; anthers about as 

long as wide, the sacs separated by connective; 

pollen 3 (2)-colporate; pistil 1, the carpels 3-5, 

the styles 3-5, shortly connate, sometimes bifid 

apically, the stigmas apical; ovary superior, subsessile or stipitate, unilocular, the ovules 2-many 

per placenta, parietal; fruit a 3-5-valved capsule, 

the seeds pitted, arillate; embryo small, straight, 

0.3 the length of the endosperm; cotyledons not 

broadened, 0.3 the length of the embryo; endosperm copious. 

Composition: 3 genera, 3 species. 

Distribution: South Africa. 

CARICACEAE (Figure 63d,e).--Small trees and 

shrubs, rarely herbs, the sap milky; xylem vessel 

perforation plates simple; leaves alternate, simple, toothed, pamately lobed, or sometimes compound and 5- 12-foliolate, exstipulate; inflorescences axillary racemes, panicles, cymes, fascicles, or the flowers solitary; plants mostly dioecious or monoecious, sometimes the flowers bisexual; sepals 5, connate; petals 5, connate, rarely 

free; stamens 10, in 2 series, rarely 5, inserted 

on the corolla, the filaments free or basally connate; anthers longer than wide, adnate, the connective often produced apically; pollen 3-colporate; pistil 1, the carpels 5, the styles 5, free or 

shortly connate, the stigmas apical, often fringed; 

ovary superior, 1- or 5-locular, the ovules numerous or few, parietal, bitegmic, crassinucellar, 

anatropous; fruit a berry, the seeds pitted or 

smooth, arillate; embryo straight, as long as the 

endosperm; cotyledons thin, 4.5 times wider than 

the radicle, 0.6-0.7 the length of the embryo; 

endosperm moderate or copious, fleshy. 

Composition: 4 genera, -50 species. 

Distribution: Tropical America and west Africa; centered in South America. 

VIOLACEAE (Figure 64a,b).--Shrubs and 

herbs, sometimes small trees or scandent; xylem 

vessel perforation plates simple, or scalariform 
132 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 63.-TURNERACEAE: a, Turnera lamiifolia flower, 1.s. of part of a longistylous flower, 

Is. of part of a brevistylous flower, views of anther, C.S. of ovary, dehiscent fruit; b, views of 

seed, T. difusa I.s. of seed, embryo (after Martius, 1840-1895). ACHARIACEAE: c, Acharia 

tragiozdes 6 flower, 1s. of same, ? flower, I.s. of same, dehiscent fruit, seed, I.s. of same (after 

Baillon, 1866- 1895). CARICACEAE: d, Carica papaya 6 inflorescence, ? inflorescence, bisexual 

flower with corolla removed, opened flower, I.s. of young fruit; e, seed with succulent testa 

removed, seed, diagrams of 2 kinds of bisexual flowers, Jacaratia dodecaphylla views of stamen 

(after Martius, 1840- 1906). 

with few to numerous bars, or both; leaves alternate, rarely opposite, simple, entire or toothed, 

rarely lobed, stipulate; inflorescences axillary racemes, spikes, panicles, or the flowers solitary; 

flowers actinomorphic or zygomorphic, bisexual, 

or the plants rarely polygamous or dioecious; 

sepals 5, free or rarely slightly connate; petals 5, 

free, rarely shortly connate, one often gibbous 
NUMBER 58 133 

C 

c1. 

/, lib 

FIGURE 64.-vIOLACEAE: a, viola subdimidiata views of flower, 3 sepals and 3 petals, essential 

organs, Is. of ovary and stamens, 3 views of stamen, 2 stamens with basal glands; b, pistil, C.S. 

of ovary, seed and Is. of same showing the small embryo, floral diagram, Alsodeia macrocarpa 

flower, Is. of same, views of stamen, A. guianensis dehiscent fruit, valve of fruit with a seed 

(after Martius, 1840-1 906). STACHYURACEAE: c, Stachyurus praecox inflorescence, flower, same 

with part of perianth removed, essential organs, petal, 1,s. of pistil, C.S. of ovary; d, views of 

stamen, fruits, seed, Is. of same, flora diagram (after Hooker and Hooker, 1837-1982; Siebold 

and Zuccarini). 

or spurred; stamens 5, mostly hypogynous, the 

filaments very short, free or connate; anthers 

basifixed or adnate, the connective produced 

apically, the 2 abaxial anthers sometimes spurred 

by a gland which secretes nectar into the petal 

spur; pollen 3 (-5)-colporate; pistil 1, the carpels 

3 (2-5), the style 1, stigma(s) usually 1, apical, 

rarely 3-5-fid, rarely subsessile; ovary superior, 

sessile, unilocular, the ovules numerous to 1 per 

placenta, parietal, bitegmic, crassinucellar, anatropous; fruit a loculicidal capsule, rarely opening 

very early and exposing the unripe seeds, rarely 

densely echinate, rarely a berry or nutlet; seeds 

smooth or tomentose, often arillate, sometimes 

winged, rarely tomentose; embryo straight, 0.9 

the length of the endosperm; cotyledons thin, 
134 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

3.4-6.0 times wider than the radicle, 0.3-0.9 the 

length of the embryo; endosperm moderate or 

copious, rarely scanty, fleshy. 

Composition: -20 genera, -900 species. 

Distribution: Variety of habitats; cosmopolitan, but mainly north temperate, many Andine 

and some in the understory of rain forest in the 

tropics. 

STACHYURACEAE (Figure 64c,d).-Shrubs or 

small trees, sometimes climbing; xylem vessel 

perforation plates scalariform, with few to many 

bars; leaves alternate, simple, serrate, stipulate; 

inflorescences axillary racemes or spikes; flowers 

bisexual or the plants polygamous, the parts hypogynous; sepals 4, free; petals 4, free; stamens 

8, in 2 series, the filaments subulate, free; anthers 

small, about as long as wide, dorsifixed; pollen 

3-colporate, occasionally 4-ruporate; pistil 1, the 

carpels 4, the style 1, stigma apical, very shortly 

4-lobed; ovary superior, 4-locular, the ovules numerous, axile, or partly parietal, bitegmic, crassinucellar, anatropous; fruit a berry-like capsule; 

seeds numerous, small, arillate; embryo straight, 

as long as the endosperm; cotyledons elliptic, the 

radicle long or short; endosperm scanty or copious. 

Composition: 1 genus, -6 species. 

Distribution: Himalayas to Japan, Taiwan. 

SCYPHOSTEGIACEAE (Figure 65a,b).-Small 

trees with soft wood; xylem vessel perforation 

plates simple, few scalariform with few bars; 

leaves alternate, simple, pinnately veined, with 

numerous close transverse tertiary veins, distichous, serrate, stipulate; inflorescences terminal 

and axillary racemes or panicles; plants dioecious; perianth parts 6, similar, in 2 series, connate in the 8, very shortly connate in the ?; 

stamens 3, the filaments connate into a column, 

the anthers with common globose apical connective; pollen 3-colpate; 3 large glands at the base 

of the stamina1 column; pistil 1, the carpels 8-1 2 

(- 16), the stigmas sessile, forming a large fleshy 

disk with a small central ostiole; ovary superior, 

unilocular, the ovules numerous, basal, bitegmic, 

crassinucellar, anatropous; fruit fleshy, dehiscing 

apically by 8- 12 (-1 6) valves; seeds numerous, 

the embryo linear, straight; cotyledons as wide 

as the radicle, 0.5 the length of the embryo; 

endosperm scanty. 

Composition: 1 genus, 1 species. 

Distribution: Northwestern Borneo. 

PERIDISCACEAE (Figure 65c,d).-Tree; xylem 

vessel perforation plates all or predominantly 

scalariform; leaves alternate, simple, distichous, 

entire, with 2 large pits at the base of the blade 

on the dorsal surface, stipulate; inflorescences 

clusters of short racemes, axillary, or at nodes of 

fallen leaves; flowers bisexual; sepals 4-5 (-7), 

free; petals 0; stamens -16-20, the filaments 

elongated, basally connate; anthers small, about 

as long as wide, monothecous; pollen 3-colporoidate; disk large, cupular, lobed, surrounding 

the base of the ovary; pistil 1, the carpels 3-4, 

styles 3-4, filiform, short, free; ovary superior, 

unilocular, the ovules 6-8, apical; fruit a drupe, 

the seed 1, large; embryo straight, 0.3 the length 

of the endosperm; cotyledons thin, 1.2 times 

wider than the radicle, 0.5 the length of the 

embryo; endosperm copious, corneous. 

Composition: 2 genera, 2 species. 

Distribution: Amazon Brazil, Venezuela, 

Guayana. 

HOPLESTIGMATACEAE (Figure 66a,b).-Trees; 

xylem vessel perforation plates simple; leaves 

alternate, simple, entire, exstipulate; inflorescences terminal subscorpioid cymes; flowers bisexual; sepals completely connate, the calyx splitting into 2-4 lobes at anthesis; petals 11-14, 

shortly connate, 3-4-seriate; stamens 20-35, 3- 

seriate, the filaments filiform, inserted on the 

base of the corolla; anthers elongated, oblong, 

dorsifixed; pollen 3-colpate; pistil 1, the carpels 

2, styles 2, shortly connate, stigmas apical, reniform; ovary superior, unilocular, the ovules 2 

per placenta, parietal; fruit a drupe, the seeds 4, 

oblong; embryo large, nearly straight, 0.9 the 

length of the endosperm; cotyledons broad, tapering gradually into the radicle; endosperm 

scanty. 

Composition: 1 genus, 2 species. 

Distribution: West tropical Africa. 

LOASACEAE (Figure 66c-h).-Herbs, fre- 
135 

FIGURE 65.-SCYPHOsTECIACEAE: a, Scyphostegia borneensis d inflorescence, d raceme, same 

with lower bract laid open, d flower and bracts, 1,s. of 6 flower, 0 flower, 1,s. of same; b, 0 

flowering shoot, fruit, seed, apex of same, 1.s. of same, embryo (after Hutchinson, 1973). 

PERIDISCACEAE: c, Peridiscus lucidus group of stamens, views of anther, flower, pistil with adnate 

fleshy disk, I.s. of pistil, C.S. of ovary, inflorescence; d, fruit, seed, embryo (enlarged) (after 

Kuhlmann, Oliver). 

quently twining, or shrubs and low trees, mostly 

scabrid, the hairs often hooked, sometimes stinging; xylem vessel perforation plates simple; leaves 

alternate or opposite, simple, entire or divided 

and toothed, exstipulate or rarely stipulate; inflorescences axillary, terminal or oppositifolious 

cymes, often sympodial (monochasial), or heads, 

or the flower solitary; flowers bisexual, actinomorphic; sepals 5 (4-7); petals 5 (4-7), sometimes 

5 petaloid staminodes present, free or connate, 

inserted on the calyx; stamens numerous (to 

-300) rarely 10 in 2 series, or 5 or only 2 fertile, 

in 1-5 series, the filaments long or short, mostly 

free, sometimes basally connate, often in bundles 

opposite the petals, sometimes adnate to the petals; anthers small, about as long as to 4 times 
136 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 58 137 

longer than wide, basifixed or dorsifixed, rarely 

1 -locellate, the connective sometimes produced 

apically; staminodes often present, petaloid or 

connate into nectar-collecting scales; pollen 3- 

colporate; sometimes disk around base of style 

or lining the calyx-tube; pistil 1, the carpels 3-5 

(1-7), style 1, filiform, the stigma 1, punctiform, 

capitate or 3-7 and linear; ovary inferior, less 

frequently semi-inferior, 1 -3-locular, the placentas 3-5 (-7), parietal, rarely apical or axile, the 

ovules numerous to 1 per placenta, unitegmic, 

tenuinucellar, anatropous or semi-anatropous; 

fruit a capsule, dehiscent apically, basally or laterally, loculicidal or septicidal or rarely indehiscent, woody and nut-like; seeds usually minute, 

rarely large, the testa smooth, reticulate, warty 

or punctate; embryo straight or bent, 0.3-1.4 

times the length of the endosperm; cotyledons 

flat or plano-convex, 1-3 times as wide as the 

radicle, 0.3-0.8 the length of the embryo; endosperm copious, moderate or 0, fleshy or corneous. 

Composition: 15 genera, 250 species. 

Distribution: Centered in tropical and subtropical America, especially the Andes, extending slightly into temperate America; one genus 

in southern Arabia and southwestern Africa. 

CUCURBITACEAE (Figure 67a-d).-Herbs, 

climbing or prostrate, usually with tendrils, 

rarely shrubs or small trees, often scabrid, the 

hairs rarely hooked; xylem vessel perforation 

FIGURE 66.-HOPLESTIGMATACEAE: a, Hoplestigma pierreanum inflorescence, bud, I.s. of flower, C.S. of anther, views 

of stamen, stigma and C.S. of ovary; b, views of fruit, C.S. of 

fruit, 1,s. of seed (after Engler). LOASACEAE: c, Mentzelia 

aurea flower, 1,s. of same, AM. albescens floral diagram, Blumenbachia hieronymi hairs, anther, style with disk, floral diagram; d, scale, staminode, fruit, C.S. of same and C.S. of 

endosperm and embryo; e, floral diagrams of Petalonyx crenatus and Sympetaleia rupestris; J Loasa parvzflora petal, scale 

and staminodes, staminode, style and disk, seed, 1,s. of endosperm and embryo, L. argemonoides floral diagram; g, 

Blumenbachia lateritia flowering shoot, Gronovia scandens 

flower and I.s. of same, floral diagram, fruit, Is. of same; h, 

Kissenia spathulata ovary laid open, seed, embryo, C.S. of 

fruit, fruit (after Martius, 1840-1 906; Baillon, 1866-1 895; 

Urban). 

plates simple; sometimes the vascular bundles 

bicollateral, frequently arranged in 2 rings, and 

with some sieve tubes scattered in the cortex; 

leaves alternate, simple, entire or palmately 

lobed, divided or compound (Zanonia), often 

with extrafloral nectaries, exstipulate; inflorescences axillary cymes, racemes, thyrses, or the 

flowers solitary; plants monoecious or dioecious, 

or rarely the flowers bisexual (Schizopepon); sepals 5 (3-6), free or connate; petals 5 (4-6), 

connate or rarely free (Fevillea), rarely fringed 

or with ventral scales; stamens 3 (1-5), the filaments rarely free (Fevillea), usually inserted on 

the corolla-tube and united in pairs, the 5th free, 

sometimes all connate in a single central column; 

anthers free or connate, often unilocular, often 

sinuate, the connective often produced apically, 

staminodes rarely present in the d (Fevillea); pollen 3-poly-colporate, 2-pantoporate, 7-9-colpate; 

sometimes a disk present at the base of the style; 

pistil 1, the carpels 3 (2-5), styles mostly 1, rarely 

3 and free, the stigmas apical, or sometimes 

decurrent ventrally, thick, usually forked, rarely 

multifid (Cucurbitella); ovary inferior, unilocular 

or imperfectly 3-locular, the placentas 3 (2-5), 

parietal but often meeting in the axis, rarely 

subapical-axile, the ovules numerous to 1, bitegmic, crassinucellar, anatropous; fruit usually 

a berry, often with a firm wall, rarely a capsule 

(Zanonia), rarely circumscissile or a samara; seeds 

often flattened, sometimes winged; embryo 

straight, the cotyledons thin or moderately thick, 

3-6 times wider than the radicle, 0.8-0.9 the 

length of the embryo, the radicle sometimes invested; endosperm 0. 

Composition: -100 genera, -700 species. 

Distribution: Pantropical and -subtropical, especially rain forest of South America and drier 

parts of Africa; few species in Australasia and 

temperate regions. 

CISTALES 

Shrubs, herbs, and small trees; xylem vessel 

perforation plates simple; leaves opposite or alternate, simple, entire, serrate, rarely lobed, stip- 
138 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 67.-cUCuRBrTACEAE: a, Fedlea trilobata d inflorescence and tendril, d flower (2 

views), I.s. of same, androecium and views of stamen; b, Luff. operculata androecium, 2 views 

of bilocular stamen, 2 views of unilocular stamen, seed, I.s. of same; c, 0 flower, 1,s. of same, 

stigmas, style and staminodes, C.S. of ovary, fruit; d, Cayaponia joribunda I.s. of d flower, 

Cyclanthera elegans d flower and I.s. of same, Is. of androecium and androecium from above, 0 

flower, 1,s. of same (after Martius, 1840-1906). 

ulate; flowers bisexual, often showy, the parts 

hypogynous; sepals 5, sometimes differing in 

width, free; petals 5 (3, O), free; stamens usually 

numerous, -25-200, rarely 3-6, the filaments 

free or rarely shortly connate, usually filiform, 

the anthers basifixed; pollen 3 (2-4)-colporate; 

glandular disk sometimes present; pistil 1, the 

carpels 2-5 (-lo), style 1, the stigma(s) apical; 

ovary 1 (3-5)-locular, the ovules numerous (-2), 

parietal, less commonly axile; fruit a loculicidal 

capsule; embryo arcuate or folded, rarely 

straight; endosperm usually copious. 

Distribution: Temperate and tropical, most 

common in the Mediterranean region, in various 

habitats. 

CISTACEAE (Figure 6 8 a ,b). -Glandular herbs 
NUMBER 58 139 

and shrubs; xylem vessel perforation plates simple; leaves opposite or rarely alternate, simple, 

entire, exstipulate or with stipules adnate to the 

petiole; inflorescence a cyme or the flower solitary, sometimes cleistogamous; flowers bisexual, 

actinomorphic, the parts hypogynous; sepals 5, 

free, the outer 2 usually smaller than the inner 

3; petals 5 (3, 0), free, usually crumpled in bud, 

sometimes convolute, caducous; stamens numerous, -25-100, or rarely 3-6, the filaments filiform, free, the anthers slightly longer than wide; 

pollen 3 (2-4)-colporate; disk present; pistil 1, 

the carpels 3 or 5 (-lo), the style 1, long, short 

or absent, the stigma(s) apical; ovary unilocular 

or several locular due to placentas meeting in the 

center of the ovary, the ovules 2-numerous, 

parietal, bitegmic, crassinucellar, orthotropous 

or subanatropous; fruit a loculicidal capsule, dehiscing by 10-5 or 3 valves from the apex downward; embryo spatulate, bent, arcuate, coiled, 

folded or rarely straight, the cotyledons 2-6 

times as wide as the radicle, 0.5-0.7 the length 

of the embryo; endosperm moderate, hard and 

transparent, starchy. 

Composition: 8 genera, -200 species. 

Distribution: Dry sunny habitats, mostly 

north temperate zone, centered in the Mediterranean region; temperate and subtropical North 

America; temperate South America; Europe, 

western Asia. 

COCHLOSPERMACEAE (Figure 68c,d).-Trees, 

shrubs, or subshrubs with yellow-red sap; xylem 

vessel perforation plates simple; leaves alternate, 

simple, palmately lobed, serrate, stipulate; inflorescence a raceme or panicle; flowers large, bisexual, the parts hypogynous; sepals 5, free; petals 5, free; stamens numerous (-80), free, the 

anthers linear, basifixed, dehiscing by apical 

pores or short slits; pollen 3-colporate; pistil 1, 

the carpels 3-3, the style 1, stigma(s) apical; ovary 

1- or 3-3-locular, the ovules numerous, parietal 

or axile, bitegniic, crassinucellar, campylo-anatropous, the funicle rather long; fruit a 3-5- 

valved loculicidal capsule, the seeds often pilose, 

usually reniforni; embryo large, arcuate, 0.8 the 

length of the endosperm, the cotyledons broad, 

0.8 the length of the embryo; endosperm copious, oily. 

Composition: 2 genera, -25 species. 

Distribution: Mostly dry habitats; tropical 

America from southwestern United States to 

northern half of South America, West Indies; 

tropical west Africa; India to Indochina, northern and northeastern Australia. 

BIXACEAE (Figure 68e,f).-Shrubs or trees 

with mucous secretory canals; xylem vessel perforation plates simple; leaves alternate, simple, 

entire, basally palmatinerved, stipulate; inflorescence a terminal panicle, often thyrsoid; flowers 

bisexual, showy, the parts hypogynous; sepals 5, 

free, with 2 basal glands on the outside of each, 

5 remaining after the fall of the sepals; petals 5, 

free; stamens numerous, -200, free or shortly 

connate basally, inserted on an annular disk; 

anther sacs hippocrepiform, dehiscing apically by 

short slits; pollen 3-colporate; pistil 1, the carpels 

2, the style elongated, filiform, stigma bilobed, 

apical; ovary unilocular, the ovules numerous, 

parietal, bitegmic, crassinucellar, anatropous, the 

funicle rather long; fruit a loculicidal capsule, the 

seeds covered by a red, fleshy testa; embryo 

arcuate, 0.9 the length of the endosperm, the 

cotyledons flat, thin, 2.8 times wider than the 

radicle, 0.7 the length of the embryo; endosperm 

copious, fleshy. 

Composition: 

Distribution: Tropical America, West Indies, 

1 genus, 1 or 4 species. 

widely cultivated in the tropics. 

ERICALES 

Shrubs, small trees, and herbs, relatively rarely 

chlorophylless; xylem vessel perforation plates 

scalariform with many or few bars, simple, or 

both; leaves mostly alternate, simple, entire or 

toothed, relatively rarely reduced to scales, rarely 

stipulate; flowers bisexual, more rarely unisexual, the parts hypogynous, rarely perigynous or 

epigynous; sepals 5 (2-lo), free or connate, imbricate; petals 5 (2-8, 0), free or connate; stamens usually twice as many as the petals or corolla lobes, sometimes as many, hypogynous or 

rarely inserted on the corolla, the filaments usu- 
140 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 58 141 

ally free, rarely shortly connate; anthers basifixed 

or dorsifixed, sometimes with basal appendages, 

often dehiscent by apical pores; pollen 3 (2-5)- 

colporate or -colpate, sometimes in tetrads; glandular disk sometimes present: pistil 1, the carpels 

5 (2-20), style 1, the stigma(s) apical; ovary 

locules 5 (2-20), very rarely 1, the ovules numerous to 1 per locule, axile, very rarely parietal, 

unitegmic, tenuinucellar; fruit usually a septicidal 

or loculicidal capsule, more rarely a berry or 

drupe, very rarely achene-like or a samaroid 

nutlet, the seeds often minute: embryo usually 

linear, sometimes minute; endosperm copious or 

moderate. 

Distribution: Cosmopolitan, most common in 

temperate and arctic regions of the Northern 

Hemisphere, often in mountains in the tropics. 

Chemistry: Chemistry favors separation of 

Diapensiaceae from Ericales. The chemistry of 

Cyrillaceae is little known, but in line with a 

position in Ericales. 

ERICACEAE (Figure 69a,b).-Shrubs or subherbaceous, rarely trees or vines: xylem vessel 

perforation plates exclusively scalariform, with 

up to 20 or more bars, or simple, or both: leaves 

mostly alternate, rarely opposite or whorled, simple, entire, more rarely crenate or serrate, exstipulate; inflorescences terminal or axillary racemes or panicles or the flowers solitary: flowers 

usually bisexual, rarely the plants dioecious, the 

parts hypogynous, rarely perigynous or epigynous; sepals 5 (3-8), free or connate; petals 5 

FIGURE 68.-cISTACEAE: a, Helianthemum brasiliense flowering shoot, flower from below, corolla in bud, calyx laid 

open, petal, pistil and C.S. of ovary, 1,s. of flower with apices 

of all parts except pistil cut off; b, views of anther, floral 

diagram, fruit and calyx, dehiscent fruit, seed showing long 

funicle, 1,s. of seed in 2 planes (after Martius, 1840-1906). 

COCHLOSPERMACEAE: c, Cochlospermurn insigne flower, floral 

diagram, 1,s. of ovary, C. codinae valve of fruit, fruit; d, 

floa.er from below, corolla in bud, C.S. of ovary, views of 

anther, pistil, seed, seed with hair removed, I.s. and C.S. of 

seed (after Slartius, 1840- 1906). BIXACEAE: e, Bixa orellana 

flowering twig, C.S. of ovary, petal, views of anther, C.S. of 

same, valve of fruit with seeds;f, 1,s. of flower, floral diagram, 

1,s. and C.S. of seed, views of seed and 1,s. of same, embryo 

(after Martius, 1804-1906; Le Maout and Decaisne, 1873). 

(3-8), mostly connate, sometimes free, rarely 

zygomorphic, usually imbricate, more rarely valvate; stamens usually twice as many as the corolla 

lobes, sometimes as many, obdiplostemonous, inserted on a nectariferous disk or rarely adnate to 

the corolla, the filaments free or rarely connate: 

anthers sometimes with basal tails and produced 

as tubes apically, often adnate, dehiscing by apical pores, very rarely by longitudinal slits; pollen 

usually in tetrads, 3 (2-4)-colporate; pistil 1, the 

carpels 5 (2-20), the ovary with as many locules 

(rarely 1 -1ocular); placentas axile, rarely subbasal, 

often protruding into the locule, the ovules numerous to 1 per locule, unitegmic, tenuinucellar, 

anatropous, semi-anatropous or almost campylotropous, the style 1, the stigma apical, entire 

or lobed; fruit a septicidal or loculicidal capsule, 

berry or drupe: seeds small, often minute, the 

embryo usually straight, rarely curved, 0.5 to as 

long as the endosperm; cotyledons 1-2.5 times 

as wide as the radicle, 0.1-0.5 the length of the 

embryo; endosperm copious or moderate, fleshy. 

Composition: 

Distribution: Cosmopolitan; cool to subtropical regions and mountains of the tropics. 

CLETHRACEAE (Figure 69c,d).-Shrubs and 

trees, the indument often stellate; xylem vessel 

perforation plates exclusively scalariform with 

20-50 bars; leaves alternate, simple, serrate, exstipulate; inflorescences terminal racemes or panicles; flowers bisexual, the parts hypogynous; calyx deeply 5-lobed, the lobes imbricate, persistent: petals 5, free, imbricate; stamens 10 (12), 

obdiplostemonous, free; anthers not appendaged, the thecas separate half way, dehiscing by 

apical pores: pollen grains single or rarely in 

tetrads, 3-colporate; disk 0; pistil 1, the carpels 

3, the style 1, 3-fid near the apex, the stigmas 

terminal; ovary 3-locular, the ovules numerous, 

axile, unitegmic, tenuinucellar, anatropous; fruit 

a loculicidal capsule; seeds minute, the embryo 

linear, straight, terete, 0.9 the length of the 

endosperm: cotyledons not broadened, 0.25 the 

length of the embryo; endosperm moderate, 

fleshy. 

-60 genera, -1 500 species. 

Composition: 1 genus, -40 species. 
142 

a 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

C 

FIGURE ~~.-ERICACEAE: a, Ledum groenlandicum flower, seeds, dehiscent fruit, pistil, Pernettya 

myrtilloides flower, same laid open, views of stamen, pistil and calyx, C.S. of ovary; b, Gaylussacia 

rhododendron 1,s. of flower, views of stamen, fruit, 1.s. and C.S. of same, pyrene and I.s. of same 

(above), seed, two I.s. of sanie (after Johnson; Martius, 1840-1906). CLETHRACEAE: c, Clethra 

laevigata flower, C. spicigera flower, pistil, stigma, I.s. of base of flower, floral diagram; d, petal 

and 2 stamens, views of stamen, C.S. of fruit, C. laevigata views of seed, fruit from below (after 

Le Maout and Decaisne, 1873). 

Distribution: Eastern United States to northwestern quarter of South America; Japan, China 

to the East Indies; Madeira. 

PYROLACEAE (Figure 70a).-Herbs or slightly 

woody; xylem vessel perforation plates scalariform with numerous bars; leaves coriaceous, radical or on a short stem and alternate or subverticillate, rarely reduced, toothed or entire, exstipulate; inflorescence a terminal raceme, corymb or umbel or the flowers solitary and scapose, 

bisexual, the parts hypogynous; calyx 5 (4)-partite, persistent; petals 5 (4), free or rarely very 
NUMBER 58 143 

FIGURE 70.-PYROLACEAE: a, Pyrola minor flower, 1,s. of same, stamen, pistil, C.S. of ovary, 

ovule, embryo taken out of the testa, floral diagram (after Le Maout and Decaisne, 1873). 

MONOTROPACEAE: b, Monotropa hypopitys flower, flowering plant, 1,s. of flower, essential organs 

and perianth segment, dehiscent fruit, seed, undivided embryo; c, Pterospora andromeda 1,s. of 

flower, I.s. of seed, Monotropa hypopity floral diagram (after Le Maout and Decaisne, 1873; 

Baillon, 1866-1895). EMPETRACEAE: d, Empetrum nzgrum fruiting twig, flower, I.s. of same, 

pistil, stamen, floral diagram; e, fruit, 1,s. of same, pyrene, I.s. of same (after Le Maout and 

Decaisne, 1873). 
144 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

shortly connate, imbricate, rarely with nectaries; 

stamens 10 (8), obdiplostemonous, free; anthers 

dorsifixed, dehiscing by 2 terminal pores; pollen 

single or in tetrads, 3-colporate; disk present or 

absent; pistil 1, the carpels 5 (4), the style 1, not 

divided apically, the 5 (4) stigmas apical; ovary 

imperfectly 5 (4)-locular (unilocular in upper 

part), the placentas axile, or parietal in the upper 

unilocular part of the ovary, fleshy, the ovules 

very numerous, unitegmic, tenuinucellar, anatropous; fruit a capsule, basally or apically loculicidally dehiscent; seeds minute, the embryo minute, undifferentiated, 0.7 the length of the endosperm; endosperm moderate or copious. 

Composition: 4 genera, -30 species. 

Distribution: Temperate and arctic regions of 

the Northern Hemisphere. 

MONOTROPACEAE (Figure 7Ob,c).-Chlorophylless herbs, with hardly any vessels; leaves 

alternate, reduced to scales; inflorescence a terminal raceme or head, or the flower solitary; 

flowers bisexual, the parts hypogynous; sepals 2- 

6, free, imbricate; petals 3-6 (0), free or connate, 

imbricate; stamens 6-1 2, the filaments free or 

connate basally; anthers rarely spurred (Pterospora), dehiscing by chinks or slits; pollen grains 

single, 3-4 (2, 5)-colporate or colpate; disk present or absent; pistil 1, the carpels 4-6, the style 

1, sometimes slightly lobed apically, the stigmas 

apical; ovary 1 -6-locular, placentation parietal in 

the unilocular ovary, axile in those with more 

than 1 locule, the ovules very numerous, unitegmic, tenuinucellar, anatropous; fruit a loculicidal capsule; seeds minute, the embryo minute, 

0.2-0.3 the length of the endosperm, the cotyledons 0.2 the length of the embryo; endosperm 

copious or moderate. 

Composition: 12 genera, 20 species. 

Distribution: Cool to warm temperate regions 

of the Northern Hemisphere, extending south 

to Colombia in mountains, and to Malaya. 

EMPETRACEAE (Figure 70d,e).-Small ericoid 

shrubs; xylem vessel perforation plates scalariform with up to 10 bars, sometimes also simple; 

leaves crowded, coriaceous, alternate, entire, exstipulate; inflorescences axillary or terminal 

heads or racemes; flowers bisexual or the plants 

dioecious, monoecious or polygamous, the parts 

hypogynous; sepals 3 (2), sometimes subtended 

by 2 pairs of bracts, imbricate, free, somewhat 

petaloid; petals 3 (2) or 0, free; stamens 2-4, the 

filaments filiform, free, long-exserted; anthers 

dorsifixed, dehiscing longitudinally; pollen usually in tetrads, 3-colporate; disk 0; pistil 1, the 

carpels 2-9, the style short, divided apically into 

2-9 segments; ovary 2-9-locular, the ovules 1 

per locule, axile, unitegmic, tenuinucellar, anatropous; fruit a drupe with 2-9 pyrenes; embryo 

linear, straight or slightly curved, terete, 0.7 to 

as long as the endosperm; cotyledons not broadened, 0.1-0.2 the length of the embryo; endosperm copious, fleshy. 

Composition: 3 genera, 10 species. 

Distribution: Heathlands of cool temperate 

regions; disjunct between northern North America and southern South America, and between 

eastern North America and southwestern Europe; southeastern United States; Azores and 

Canary Islands. 

EPACRIDACEAE (Figure 7 1 a$).-Shrubs and 

small trees; xylem vessel perforation plates exclusively scalariform, sometimes with more than 20 

bars, or simple or both; leaves alternate or rarely 

opposite, simple, coriaceous, entire or rarely 

crenate ( Wittsteinia), exstipulate; inflorescences 

mostly racemes; flowers usually bisexual, sometimes the plants monoecious; sepals 5 (4), free, 

persistent; petals 5 (4), connate, rarely free (Ly 

sinema) or calyptrate, imbricate or valvate; stamens 5 (4), rarely only 2 fertile, hypogynous or 

epipetalous, alternate with the corolla lobes; anthers rarely connate, dorsifixed, dehiscing longitudinally by 1 (2) slits; pollen in tetrads or 

single, 3-colporate; disk usually present, surrounding the base of the ovary; pistil 1, the 

carpels 5 (2- lo), the style 1, elongate, the stigma 

capitate, apical; ovary superior, rarely inferior 

(Wittsteinia), 1-1 O-locular, the ovules l-several 

per locule, axile, unitegmic, tenuinucellar, anatropous; fruit a loculicidal capsule or drupe with 

10-1 pyrenes; embryo linear, terete, straight, 0.9 

the length of the endosperm; cotyledons not 
NUMBER 58 145 

FIGURE 7 1 .-EPACRIDACEAE: a, Cyathodes colensoi flowering twig, 2 flowers, flower with corolla 

removed, corolla laid open, pistil, I.s. of same; b, views of anther, fruit, 1,s. and C.S. of same, 

embryo (after Cheeseman). DIAPENSIACEAE: c, Diapensia lapponica flowering plant, flower, 

corolla laid open, pistil, part of style and stigma, C.S. of ovary: d, fruit, C.S. of same, dehiscent 

fruit, 1,s. of seed, floral diagram (after Le Maout and Decaisne, 1873). CYRILLACEAE: e, Cyrilla 

racemzflora part of twig with infructescence and inflorescence, floral diagram, flower, 1,s. of 

same, I.s. and C.S. of fruit (enlarged);f, Clij?onia monophylla pistil and calyx, Is. of flower, I.s. of 

seed (after Baillon, 1866-1899; Le Maout and Decaisne, 1873; Lindley, 1853). 
146 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

broadened, 0.6 the length of the embryo; endosperm copious. 

Composition: 30 genera, -400 species. 

Distribution: Centered in Australia; few species in New Zealand, Indochina and the East 

Indies, one in southern South America; Hawaii. 

DIAPENSIACEAE (Figure 7 1 c,d).-Small subshrubs or subherbaceous; xylem vessel perforation plates simple, rarely also scalariform; leaves 

coriaceous, simple, entire or toothed, alternate 

or opposite, often basal, exstipulate; infloresence 

a terminal raceme or subcapitate, or the flowers 

solitary; flowers bisexual; sepals 5, free or connate, the lobes imbricate, persistent; petals 5, 

connate, sometimes almost free, the lobes imbricate, sometimes fringed; stamens 5, epipetalous 

or hypogynous, alternate with the lobes, free or 

connate, the filaments short; 5 staminodes sometimes present; anthers basifixed, 2( 1)-locular, dehiscing longitudinally or transversely by slits; pollen simple, usually 3-colporate; disk 0; pistil l, 

the carpels 3, the style 1, elongate or short, the 

stigma apical, 3-lobed; ovary superior, 3-locular, 

the ovules few to many per locule, axile, unitegmic or also bitegmic(?), tenuinucellar, semianatropous or campylotropous; fruit a loculicidal 

capsule; seeds minute, the embryo linear, terete, 

straight or slightly curved, 0.5 the length of the 

endosperm; cotyledons not broadened, 0.3-0.4 

the length of the embryo; endosperm copious, 

fleshy. 

Composition: 6 genera, 20 species. 

Distribution: Arctic and temperate Northern 

Hemisphere. 

CYRILLACEAE (Figure 7 1 e,f).-Shrubs and 

small trees, buds perulate; xylem vessel perforation plates exclusively scalariform with 20-50 

(-70) bars; leaves alternate, simple, entire, 

exstipulate or stipules small and scarious; inflorescences racemes, terminal or at the apex of the 

previous year’s wood; flowers bisexual, actinomorphic, the parts hypogynous; sepals 5 (6), free 

or rarely very shortly connate, quincuncial, imbricate, sometimes unequal; petals 5 (6), free or 

shortly connate, imbricate; stamens 5, or 10 in 2 

whorls, the filaments elongate, sometimes broadened, free or adnate to the corolla at the very 

base (Purdiaea), the anthers small, dorsifixed, 

about as long as wide or oblong (Purdiaea), dehiscing longitudinally or by apical pores (Purdiaea); pollen 3 (4)-colporate, simple; disk weak, 

under the ovary; pistil 1, the carpels 2-4 (5), the 

style short, or elongated (Purdiaea), stigma(s) 

apical; ovary 2-4 (5)-locular, the ovules 1-2 (3) 

per locule, axile-subapical, unitegmic, tenuinucellar or weakly crassinucellar, anatropous, the 

funicle rather long; fruit small, indehiscent, the 

pericarp thin and dry, sometimes a samaroid 

nutlet, the mature seed devoid of a seed coat; 

embryo straight, usually cylindrical, 0.6-0.8 the 

length of the endosperm; cotyledons 0.25-0.3 

(to 0.7 in Purdiaea) the length of the embryo, 

twice as wide as the radicle; endosperm copious, 

fleshy-hard. 

Composition: 3 genera, 13 species. 

Distribution: Marshy habitat; southeastern 

United States to the northern half of South 

America and the West Indies. 

LENNOACEAE (Figure 7 2a ,b).-Chlorophylless, colored, parasitic, fleshy herbs; xylem vessel 

perforation plates simple, or xylem with only 

tracheids; leaves reduced to scales; inflorescence 

a terminal thyrse or head; flowers bisexual; sepals 

5-10, free and linear or connate nearly to the 

apex, persistent; petals 5-8, connate, the lobes 

short, imbricate; stamens epipetalous, 5-10, in 1 

or 2 whorls, sometimes 2 staminodial, the filaments short or absent; anthers dehisce longitudinally; pollen simple, 3-colporate; disk 0; pistil 

1, the carpels 6- 15, the style 1, stigmas apical, 

6- 15; ovary 6- 15-locular, superior, the ovules 2 

per locule, axile, unitegmic, tenuinucellar, anatropous; fruit a capsular-drupe, the dehiscence 

circumscissile; seeds small; embryo globose, undifferentiated; endosperm copious. 

Composition: 3 genera, 4 species. 

Distribution: Deserts and coastal sand dunes; 

southwestern United States and northwestern 

Mexico; northern coast of Colombia and Venezuela; Revilla Gigedo Islands. 
NUMBER 58 147 

/ 

C 

FIGURE 72.-LENNOACEAE: a, Lennoa madreporoides flower, 1.s. of same, L. coerulea stamen, 1.s. 

of seed showing embryo; b, Ammobroma sonorae part of inflorescence with buds, flower and 

young fruit, flower laid open, fruit showing the numerous pyrenes, I.s. of pyrene, plant (after 

Baillon, 1866- 1895; Solms-Laubach; Torrey). HYDNORACEAE: c, Hydnora africana flower laid 

open, Prosopanche burmeisteri C.S. of ovary with 3 groups of placental lamellae, I.s. of seed, 

Hydnora sp. 1,s. of fruit, (after Le Maout and Decaisne, 1873; R. Brown). 
148 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

RAFFLESIALES 

Chlorophylless usually colored parasitic herbs; 

xylem vessel perforation plates simple, sometimes the vascular system reduced to tracheids; 

scale leaves present or absent, usually alternate; 

flowers usually unisexual; perianth segments 3- 

10, in 1-2 whorls, usually connate; stamens -50- 

3, in 1-3 series, adnate to the perianth or connate 

in a column, the anthers connate; pollen 2 (3)- 

sulculate, 2-3-pored, 3 (4)-colpate or nonaperturate; pistil 1, the carpels 3-20, style 1, short, 

or the stigma(s) sessile; ovary inferior or semiinferior, rarely superior, rarely partly several 

locular, ovules numerous, the placentas lamellate 

and parietal, or apical, or the ovules covering the 

wall of the ovary; fruit a berry, the seeds numerous, minute, the testa hard; embryo minute, undifferentiated; endosperm copious or scanty; perisperm copious or scanty. 

Distribution: Mainly tropical and subtropical, 

western United States to southern South America; Asia, Africa, and western Australia, in wet 

and dry habitats. 

HYDNORACEAE (Figure 72c).-Chlorophylless, 

colored, parasitic, fleshy herbs; vascular bundles 

of the rhizome-like structures may be in 1 or 

more irregular rings, the vessel perforation 

plates simple; sieve tubes definitely seen only in 

Prosopunche; scale leaves 0; flowers solitary, bisexual, very rarely unisexual, arising from a rhizome-like structure, actinomorphic, fairly large; 

perianth parts 3-4 (5) in one series, connate, 

thick, valvate; stamens 3-4 (5), the anthers large, 

sessile or subsessile, adnate to the perianth tube, 

connate and divided into numerous sinuous locelli, longitudinally dehiscent; pollen grains usually bilateral, 2 (3)-sulculate; pistil 1, the carpels 

3-4 (5), the stigma subsessile, capitate, 3-4 (5)- 

sublobed; ovary inferior, unilocular, the ovules 

numerous, unitegmic, tenuinucellar, orthotropous, on numerous apical placentas or numerous 

lamellate, parietal placentas; fruit a berry, the 

seeds numerous, minute, the testa hard; embryo 

minute, undifferentiated, 0.3 the length of the 

endosperm; endosperm copious; perisperm copious or scanty. 

Composition: 2 genera, 18 species. 

Distribution: Dry habitats; tropical and southern Africa, Madagascar; warm temperate Paraguay and Argentina. 

RAFFLESIACEAE (Figure 73u-h).-Herbs, 

often fleshy, chlorophylless, colored or white (Mitrustemon) parasites of roots and stems of woody 

plants; vascular system very much reduced, 

merely tracheids, or vascular strands with simple 

perforation plates; leaves scales, spirally arranged 

or rarely opposite (Mitrustemon) or whorled; 

flowers usually solitary, terminal, sometimes 

smelling like carrion, the inflorescences rarely 

spikes, racemes or cymes (Cytineae), actinomorphic, minute to very large, rarely bisexual 

(Mitrustemon, Rhizanthes), usually the plants dioecious or monoecious, rarely polygamous; perianth parts 4- 10, in 1 or 2 whorls, usually basally 

connate, rarely free, sometimes petaloid; anthers 

numerous (-50) to 8 in 1-3 series, round or 

elongate, sometimes the connective produced apically, bilocular or multilocellate, dehiscing longitudinally or by 1-2 apical pores, sessile and 

adnate to a central fleshy column, or the filaments connate in a column; pollen viscous, simple 

or in tetrads, 2-3-pored, 3 (4)-colpate or nonaperturate; nectary sometimes present (Cytinus), 

adnate to the base of the style and base of the 

staminal column; pistil 1, the carpels 4-20, style 

1, short, or the stigma sessile, apical, simple, 

discoid and radially marked off (Bdallophyton) or 

lobate, or in a ring around the upper part of the 

FlCURE ?3.-RAFFLESIACEAE: a, Rafflesia patma I.S. of ? 

flower bud, R. arnoldii flower; b, 1,s. of d flower; c, 1,s. and 

C.S. of anther, several anthers in position; d, R. rochussenii 

1,s. of fruit, R. patma seed testa with endosperm ejected 

(enlarged), endosperm and embryo (enlarged): e, Cytinus 

dioicus I.s. of 6 flower, staminal column, C. hypocistis 1,s. of 0 

flower, C. dioicus C.S. of ovary, C. hypocistis flowering plant, 

C. dioicus 0 perianth, C. hypocistis fruit crowned by the 

perianth, seed, 1,s. of same; f, Pilostyles ingae parasite on a 

legume, Apodantes jlacourtiae flowers; g, Pilostyles caulotreti 

Is. of d flower, I.s. of young d flower, P. blanchetii I.s. of 0 

flower; h, Apodanthes caseariae C.S. of ovary, Pilostyles caulotreti I.s. of P flower with annular stigma shown above, P. 

ingae fruit (after R. Brown; Baillon, 1866-1895; Meijer: 

Solms-Laubach; Harvey: Martius, 1840- 1906; Karsten, 

1958). 
NUMBER 58 149 

e 
150 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 74.-BALANOPHORACEAE: a, Balanophora dioica d plant and bud, 6 flower, B. fungosa 

cluster of 0 floiiers surrounding a hair-like structure, I.s. of P flower; 6, Helosis guyanensis 2 

stamens, d flower, 0 flower, fruit, I.s. of same (greatly enlarged) showing the minute embryo 

and copious endosperm (after Baillon, 1866-1 895; Martius, 1840- 1906). CTENOLOPHONACEAE: 

c, Ctenolophon parvijolius flowering twig, bud, I.s. of flower, C. englerianus pistil, C.S. of ovary, 

woody I-seeded indehiscent fruit, seed with aril and C.S. of same (after Hutchinson, 1973; 

U’inkler; Pierre). 

column; ovary inferior or semi-inferior, rarely cellar, anatropous or orthotropous (Cytinus); 

superior (Mitrustemon), unilocular, or several fruit a berry, sometimes nearly dry, the seeds 

locular in the upper part (Cytinus), with many minute, very numerous, the testa hard; embryo 

twisted locelli, the ovules numerous, on as many minute, undifferentiated, surrounded by a unilamellate, parietal placentas as carpels or cover- cellular layer of endosperm. 

ing the wall of the ovary, I-2-tegmic, tenuinu- Composition: 8 genera, -50 species. 
NUMBER 58 151 

Distribution: Mainly tropical and subtropical; 

western United States to southern tip of South 

America; central and southern Africa, Madagascar; Mediterranean region to Iran; Himalayas, 

Burma to Malaya, the East Indies; Japan; western 

Australia. 

BALANOPHORALES 

The order is monotypic. 

BALANOPHORACEAE (Figure 74a,b).-Chlorophylless rhizomatous fleshy herbs parasitic on 

tree roots, sometimes smelling like carrion; xylem vessel perforation plates simple, also scalariform(?); inflorescence a dense spike, head or panicle; flowers rarely bisexual, the plants usually 

monoecious or dioecious; 8 flowers: perianth 3- 

4 (0, 2-8) parts, free or connate, valvate; stamens 

1-2 in achlamydeous flowers, 3-10 in the others 

and opposite the perianth parts, the filaments 

free or basally connate; anthers 1-multilocellate, 

free or connate, dehiscing by pores or slits; pollen 

3(-5)-colpate, 3-4(2-5)-por(or)ate, or 4-polyfor(amin)ate; 9 flowers: perianth absent or 3- 

lobed; pistil 1, the carpels I-3(-5), the styles 1- 

3 (-5), often filiform, free, the stigmas apical, 

capitate, or absent and the stigma sessile and 

discoid; ovary inferior or superior, 1 -3(-5)-locular, the ovules 1 per locule, 0-1-tegmic; fruit a 

nutlet or drupelet, the seed 1; embryo minute, 

undifferentiated, 0.3 the length of the endosperm; endosperm relatively copious. 

Composition: 

Distribution: 

18 genera, - 100 species. 

Pantropical, usually in moist upland forest. 

CELASTRALES 

Trees and shrubs, sometimes climbers; xylem 

vessel perforation plates scalariform usually with 

fewer than 20 bars, simple, or both; leaves alternate or opposite, simple, entire or toothed, exstipulate; flowers bisexual, rarely unisexual, 

small; sepals 5 (3, 4), free or shortly connate; 

petals 5 (3, 4, 0), free, very rarely shortly connate; stamens 5 (2-20), hypogynous or perigynous, the filaments short or filiform, usually free, 

sometimes shortly connate, the anthers dorsifixed or basifixed; pollen 3(2, 7-9)-colporate; 

glandular disk present, often conspicuous; pistil 

1, the carpels 2-5, styles 1, 2 or 5, usually short, 

the stigmas apical or very rarely decurrent; ovary 

superior or semi-inferior, rarely inferior, 2- 

5( 10)-locular, the ovules 1 -2(to - 14) per locule, 

axile; fruit a drupe, nut, loculicidal or septicidal 

capsule, samara, schizocarp or berry; seeds often 

arillate, the embryo usually large, endosperm 

copious to absent. 

Distribution: Nearly cosmopolitan , mainly 

tropical and subtropical, usually lacking in cool 

and cold regions. 

CTENOLOPHONACEAE (Figure 74c).-Trees; 

xylem vessel perforation plates scalariform, usually with less than 25 bars, but more numerous 

in one species; leaves opposite, coriaceous, simple, entire, stipulate; inflorescences terminal and 

axillary cymes; flowers bisexual , the parts hypogynous; sepals 5, shortly connate, imbricate; petals 

5, free, thick, linear-oblong, spoon-shaped basally; stamens 10, the filaments elongated; anthers 

about as long as wide, dorsifixed; pollen 7-9- 

colp(oroid?)ate; disk cupular, external to the stamens; pistil 1 , the carpels 2, styles 2-1, elongated, 

stigmas apical, capitate; ovary 2-locular, the 

ovules 2 per locule, axile-subapical; fruit a nut, 

the seed arillate. 

Composition: 1 genus, 3 species. 

Distribution: Tropical western Africa, western Malaysia. 

IXONANTHACEAE (Figure 75a).-Trees and 

shrubs; xylem vessel perforation plates simple; 

leaves alternate, simple, entire or serrate, stipulate; inflorescences lateral cymes, racemes, panicles or fascicles; flowers bisexual, the parts hypogynous; sepals 5, free or shortly connate; petals 

5, free, sometimes persistent and becoming indurated; stamens 20, 15, 10, or 5, the filaments 

filiform, elongated, shortly connate; anthers 

small, about as long as wide, dehiscing longitudinally; pollen 3-colporate; disk conspicuous, annular or cupular, internal to the stamens; pistil 

1, the carpels 5-2, style 1, elongated or 5 and 

free nearly to the base; ovary 5-2-locular, the 
152 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE ~~.--IXONANTHACEAE: a, Ixonanthes icosandra bud, I. papuana flower, 1.s. of base of 

flower, part of outer surface of disk with 2 adnate filaments, I. cochinchinensis flower, C.S. of 

fruit, dehiscent fruit, seed (after Lecomte; Ridley; Hutchinson, 1973; Oliver). IRVINGIACEAE: 

b, Desbordesia glaucescens flowering twig, apex of young twig (superimposed on leaf) showing 

large stipules, flower, 1,s. of same, stamen, pistil and disk, fruit, I.s. and C.S. of same, C.S. of seed; 

c, Irvingia barteri part of twig with infructescence (after Hutchinson, 1973; Oliver). DICHAPETALACEAE: d, Dichapetalum rugosum flower from side and from above, 1,s. of same, petal, 

pistil and disk, C.S. of fruit, Stephanopodium engleri I.s. of flower; e, Tapura amazonica flower, 

same with calyx removed, corolla laid open showing insertion of stamens, I.s. of flower (corolla 

cut off), floral diagram, fruit, I.s. of same (after Martius, 1840-1 906). 
NUMBER 58 153 

ovules 2- 1 per locule, axile-subapical, bitegmic, 

anatropous; fruit a septicidal capsule, the seeds 

sometimes arillate; embryo often oblique or lateral; endosperm fleshy. 

Composition: 7 genera, -40 species. 

Distribution: Tropical Asia, America, and Africa. 

IRVINGIACEAE (Figure 75b,c).-Trees; xylem 

vessel perforation plates simple; leaves alternate, 

simple, entire, coriaceous, the stipules large; inflorescences axillary and terminal panicles; flowers bisexual, small, the parts hypogynous; sepals 

5, free or shortly connate; petals 5, free; stamens 

10, the filaments free, elongated; anthers about 

as long as wide, sub-basifixed, dehiscing longitudinally; pollen 3-colporate; disk conspicuous, cupular, surrounding the base of the ovary; pistil 

1, the carpels 5, 4, or 2, style 1, short, the stigma 

apical, punctiform; ovary 5, 4, or 2-locular, the 

ovules 1 per locule, axile-subapical, bitegmic, 

anatropous; fruit 1, 5 or 4 drupes or broadly 

winged samara; embryo straight, as long as the 

endosperm; cotyledons broad, flat, 0.3 the length 

of the embryo; endosperm scanty or 0. 

Composition: 3 genera, 10 species. 

Distribution: Tropical Africa, Indochina, Malaya, Borneo, Sumatra. 

DICHAPETALACEAE (Figure 75d,e).-Small 

trees and shrubs, often lianas; xylem vessel perforation plates simple, sometimes also scalariform 

with up to 10 or more bars; leaves alternate, 

simple, entire, often with a few flat glands near 

the base below, distichous, stipulate; inflorescences axillary and terminal or adnate to the 

petiole; flowers bisexual or the plants polygamodioecious, small, actinomorphic or zygomorphic; 

sepals 5 (4), free or shortly connate; petals 5 (4), 

mostly bilobed, free or united with the stamens 

into a tube; stamens 5 (4), some of these sometimes staminodal, the filaments long or short; 

anthers about as long as wide, the connective 

often dorsally thickened, dehiscing longitudinally; pollen (2) 3-colporate; disk cupular or 

lobed, or of separate glands opposite the petals; 

pistil 1, the carpels 2-3 (4), style 1 or 2-3-fid and 

basally connate, the stigmas 2-3, apical; ovary 

superior to inferior, 2-3-locular, the ovules 2 per 

locule, carunculate, axile-subapical; fruit a 

drupe, sometimes the epicarp splitting, mostly 1- 

seeded; embryo large, straight, the cotyledons 

thick, plano-convex, 12 times broader than the 

radicle, 0.97 the length of the embryo; endosperm 0. 

Composition: 4 genera, -200 species. 

Distribution: Pantropical, few subtropical; 

centered in Africa. 

CELASTRACEAE (Figure 76a-c).-Trees, 

shrubs, and climbers, some genera with secretory 

sacs or laticiferous canals; xylem vessel perforation plates mostly simple, rarely exclusively scalariform with fewer than 10 bars except in Perrottetia, occasionally both simple and scalariform; 

leaves alternate or opposite, rarely 0, simple, 

entire or toothed, stipulate or exstipulate; inflorescences axillary, rarely epiphyllous(?), mostly 

cymose or fasciculate, rarely racemose or solitary; flowers small, often bisexual, rarely the 

plants polygamodioecious; sepals (3) 4-5, free or 

connate; petals 4-5 (3, 0), free, rarely basally 

connate (Microtropis), imbricate or rarely valvate; 

stamens 4-5 (2, 3), alternipetalous, the filaments 

often short, rarely shortly connate (Hippocratea), 

hypogynous or perigynous, sometimes inserted 

on the disk; anthers about as long as wide, longitudinally dehiscent, rarely transversely (Hippocratea); pollen usually 3-colporate; disk usually 

present, cupular or flat, surrounding, or beneath 

the ovary; pistil 1, the carpels 2-5, styles 1 or 5, 

short, the stigmas 2-5, apical; ovary superior, 

rarely semi-inferior (Mortonia), 1 -5-locular, the 

ovules mostly 2 per locule, sometimes 1 to 

- 14 per locule, axile, rarely apical, bitegmic, 

crassinucellar or tenuinucellar, anatropous; fruit 

a loculicidal, rarely septicidal capsule, schizocarp, 

samara, drupe or berry; seeds often with a colored aril, the embryo rather large, rarely minute 

(Perrottia), straight, or the radicle very slightly 

curved, to 0.9 the length of the endosperm; 

cotyledons flat, thin or moderately thick, 1.7- 

4.0 times wider than the radicle, 0.6-0.9 the 

length of the embryo; endosperm moderate, copious or 0, fleshy. 
154 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

f 
NUMBER 58 155 

Composition: -55 genera, -850 species. 

Distribution: Cosmopolitan except arctic and 

cool temperate regions of the Northern Hemisphere; mainly tropical and subtropical. 

GOUPIACEAE (Figure 76d,e).-Trees and 

shrubs; xylem vessel perforation plates scalariform with 3-10 bars, the vessel members very 

long; leaves alternate, simple, entire, (dentate in 

seedlings), stipulate; inflorescences axillary , pedunculate, subumbellate racemes; flowers bisexual; sepals 5, connate, the lobes imbricate; petals 

5, free, valvate, with long, narrow apices; stamens 

5, the filaments very short, inserted on the disk; 

anthers small, the connective produced apically, 

the locules separate; pollen 3-colp(oroid)ate; disk 

cupular, partly enclosing the ovary; pistil 1, the 

carpels 5, styles 5, short, free, the stigmas apical; 

ovary superior or semi-inferior, 5-locular, the 

ovules several per locule, axile-basal; fruit a 

drupe, the seeds several; embryo straight, 0.8 the 

length of the endosperm; cotyledons oblong, not 

broadened?, 0.5 the length of the embryo; endosperm copious, fleshy. 

Composition: 1 genus, species 3. 

Distribution: Guayana, northern Brazil. 

SIPHONODONTACEAE (Figure 7 Gf-h).-Trees 

and lianas; xylem vessel perforation plates simple; 

leaves alternate, toothed or subentire, stipulate; 

inflorescences axillary, cymose-umbelliform, 1- 

4-flowered; flowers bisexual; sepals 5, free or 

basally connate, imbricate; petals 5, free, imbri- 

~xRE 76.-cELASTRACEAE: a, Maytenusfloribunda flower, 

1,s. of same (petals removed), M. communis bud, views of 

stamen, fruiting twig; b, dehiscent fruit, seed enclosed in 

aril, views of seed, I.s. of same, Celastrus scandens 1,s. of 0 

flower with staminodes and disk, floral diagram, fruit with 

calyx; c, Cuema kappleriana floral diagram, bud, I.s. of pistil, 

petal, flower (after Martius, 1840-1906; Johnson). GOUPIACEAE: d, Goupia glabra axillary inflorescence, floral diagram, bud, flower, l.~. of same, cupular disk surrounding 

essential organs and 2 stamens inserted on inner face of disk, 

petal; e, pistil, I.s. and C.S. of ovary, fruits, seed, I.s. of same 

(after Baillon, 1866-1 895; Oliver; Hutchinson, 1973). SrPHONODONTACEAE: 5 Siphonodon celastrineus axillary inflorescence, 1,s. of flower (sepals and petals cut off), view from 

above of anthers appressed to disk and view of disk from 

above showing the 5 stigmas and ostiole, sepal, petal on leaf; 

g, 1,s. of bud, I.s. of fruit, Is. of seed; h, fruit, C.S. of same 

(after Hutchinson, 1973; Lecomte). 

cate; stamens 5, oppositisepalous, the filaments 

short, inserted external to the disk, free or connate; sometimes also 5 staminodes present; anthers small, basifixed; pollen 3-porate; disk present; pistil 1, the carpels 5, styles 5, short, adnate 

to the ventral wall of the ovary, the stigmas apical 

and decurrent; ovary superior, 1 0-locular, 2-6 

ovules per locule; fruit a drupe with 10 or 20 

pyrenes; cotyledons large, flat, foliaceous, 

broadly ovate or orbicular, subcordate basally, 

the radicle short; endosperm copious or moderate, rather bony or horny. 

Composition: 1 genus, 5 species. 

Distribution: Southeastern Asia, Malaysia, 

northeastern Australia. 

R HA MNA L ES 

Trees and shrubs, often climbing, rarely herbs; 

xylem vessel perforation plates simple or scalariform with few or many bars, or both; leaves 

usually alternate, simple or compound, toothed 

or entire, rarely lobed, stipulate or exstipulate; 

flowers small, bisexual or unisexual, the parts 

hypogynous or perigynous; sepals 4-5 (-9), connate, small; petals 4-5 (-9, 0), free, more rarely 

connate; stamens 4-5 (-22), often opposite the 

petals; filaments free or rarely connate or inserted on the calyx-tube or petals, the anthers 

often dorsifixed; pollen 3(4-6)-colporate, -colpate, -porate or nonaperturate; glandular disk 

surrounding the base of the ovary or absent; 

pistil 1, the carpels 2-4 (-22), style 1, usually 

short, the stigmas apical or sometimes sessile; 

ovary superior to inferior, 1 -4(-22)-locular, the 

ovules 1-2 (3) per locule, axile, subapical or 

subbasal; fruit a berry or drupe, rarely a schizocarp, nut, samara or septicidal capsule; embryo 

minute to large; endosperm copious to absent. 

Distribution: Cosmopolitan, most common in 

tropical and subtropical regions, in various habitats. 

Gibbs does not feel there is any 

strong argument from chemistry for close relation of Rhamnales and Celastrales. Rhamnaceae 

are noted for their quinones, which have not 

been reported elsewhere in the order. 

Chemistry: 
156 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

VITACEAE (Figure 77a).-Mostly lianas, often 

with tendrils and swollen nodes; xylem vessel 

perforation plates simple; leaves alternate, rarely 

also opposite, simple or compound, toothed, very 

rarely entire, sometimes lobed, stipulate; inflorescences leaf-opposed, cymose in spikes, racemes, panicles; flowers minute, bisexual or the 

plants dioecious or polygamomonoecious, the 

parts hypogynous; sepals 4-5 connate nearly to 

the apex, small; petals 4-5, free or apically connate, valvate; stamens 4-5, opposite the petals, 

the filaments free, anthers free, dorsifixed; pollen as a rule 3-colporate, sometimes with 4 or 6 

furrows; disk usually annular or cupular, surrounding the base of the ovary; pistil 1, the 

carpels 2, style 1, usually short, the stigmas apical; 

ovary 2-locular, the ovules 2 per locule, axile, 

bitegmic, crassinucellar, anatropous; fruit a 

berry, the seed(s) with hard testa; embryo minute, straight, 0.2-0.4 the length of the endosperm; cotyledons 1.5-2.0 times wider than the 

radicle, 0.4-0.5 the length of the embryo; endosperm copious, fleshy, sometimes ruminate. 

Composition: 12 genera, -700 species. 

Distribution: Pantropical and -subtropical, 

much rarer in temperate zones. 

LEEACEAE (Figure 7’ib).-Trees, shrubs, and 

herbs,, without tendrils; xylem vessel perforation 

plates simple; leaves alternate, very rarely opposite, compound, rarely simple, usually toothed, 

exstipulate; inflorescences terminal, very rarely 

axillary, usually corymbose or cymose, the flowers bisexual; sepals 5-4, connate; petals 5-4, 

basally connate, valvate; stamens 5-4, opposite 

the petals, the filaments connate, inserted on the 

corolla, the anthers extrorse; pollen 3-colporate; 

pistil 1, the carpels 3-8, style 1, stigma apical; 

ovary 3-8-locular, the ovules 1 per locule, axile; 

fruit a berry; embryo small, thin, the cotyledons 

very small, radicle long, straight or slightly 

curved; endosperm fleshy, ruminate. 

Composition: 1 genus, 70 species. 

Distribution: Palaeotropics, mostly southern 

and eastern Asia, in moist forest. 

RHAMNACEAE (Figure 77c,d).-Trees and 

shrubs, sometimes thorny, often climbing, very 

rarely herbs, tendrils rarely present; xylem vessel 

perforation plates simple, also occasionally scalariform with few bars; leaves usually alternate, 

sometimes opposite, rarely absent, simple, 

toothed or entire, mostly stipulate; inflorescences 

axillary, cymose or racemose, usually corymbiform; flowers small, bisexual or the plants rarely 

polygamodioecious; sepals 5 (4), connate, valvate; 

petals 5 (4) or 0, small, free; stamens 5 (4), the 

filaments free or inserted on the calyx-tube, opposite the petals; anthers about twice as long as 

wide; pollen usually 3-colporate; disk mostly present, perigynous, sometimes lining the calyx-tube; 

pistil 1, the carpels 3 (2-5), style 1, sometimes 

shortly lobed, the stigmas apical; ovary superior 

to inferior 3(1-5)-locular, the ovules 1 (2) per 

locule, axile-subbasal, bitegmic, crassinucellar, 

anatropous; fruit a drupe, berry, schizocarp, nut 

or rarely samara or septicidal capsule; embryo 

large, straight, rarely bent, as long as the endosperm; cotyledons thin or moderately thick, 3-6 

times wider than the radicle, 0.9 the length of 

the embryo; endosperm mostly moderate, sometimes scanty or 0, fleshy, sometimes ruminate. 

Composition: 58 genera, -900 species. 

Distribution: Cosmopolitan except part of the 

arctic region; most common in tropical and subtropical regions. 

ERYTHROPALACEAE (Figure 77e).-Scandent 

shrubs or lianas, sometimes with tendrils; xylem 

vessel perforation plates simple; leaves alternate, 

simple, entire, exstipulate; inflorescences axillary, cymose, lax; flowers bisexual, the parts usually perigynous; sepals 5, basally connate; petals 

5, free, valvate; stamens 5, opposite the petals, 

the filaments short, inserted on the base of the 

petals; anthers small, about as long as wide; pollen 3-colpate; disk large, partly enclosing the 

ovary; pistil 1, the carpels 3, style 1, short, the 

stigmas 3, apical; ovary semi-inferior, initially 3- 

locular, becoming unilocular, the ovules 3-2, 

apical, unitegmic, tenuinucellar, anatropous; 

fruit a drupe, the seed 1; embryo minute, 0.2 the 

length of the endosperm; cotyledons not broadened, about 0.6 the length of the embryo; endosperm copious, fleshy. 
NUMBER 58 157 

FIGURE ~~.-VITACEAE: a, cissus scabra part of inflorescence, floral diagram, flower, same 

with petals fallen, I.s. and C.S. of ovary, Vitis vinijera seed, I.s. of same showing the minute 

embryo and copious endosperm, C.S. of seed, Cissus salutaris C.S. of immature fruit (after 

Martius, 1840-1906). LEEACEAE: b, Leea sambucina flower, I.s. of same, L. coccinea flower, same 

with stamens removed from the urceolus, I.s. of flower, pistil and hypogynous disk, stamen 

(after Baillon, 1866-1895; Hooker and Hooker, 1837-1982). RHAMNACEAE: c,Gouania latifolia 

flower, I.s. of same, petal and stamen, stamen, part of inflorescence; d, Rhamnidium elaeocarpum 

infloresences, flower, 1,s. of same, I.s. and C.S. of fruit, I.s. of embryo (after Martius, 1840- 

1906). ERYTHROPALACEAE: e, Erythropalum scandens 1.s. of flower, fruiting twig (after Baillon). 
158 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Composition: 1 genus, 2 species. 

Distribution: Eastern Himalayas to the East 

Indies. 

AQUIFOLIACEAE (Figure 78a,b).-Trees and 

shrubs; xylem vessel perforation plates exclusively scalariform, mostly with more than 20 bars; 

leaves alternate, very rarely opposite, simple, 

usually toothed, stipulate or exstipulate; inflorescences axillary cymes or fascicles, rarely racemes or the flowers solitary; plants mostly dioecious, sometimes polygamodioecious, some of the 

flowers bisexual, the parts hypogynous; sepals 4 

(5-9), connate, imbricate, minute; petals 4 (5-9 

or 0), free or shortly connate, imbricate or valvate (Phelline); stamens 4 (5-22), the filaments 

free or adnate to the base of the corolla; anthers 

about as long as wide; pollen 3-colporoidate; disk 

absent; pistil 1, the carpels 4 (3-22), stigma usually sessile, rarely a short style present; ovary 

4(3-22)-locular, the ovules 1 (2) per locule, axileapical, unitegmic, crassinucellar or tenuinucellar, 

anatropous; fruit a drupe with 4 (3-22) pyrenes; 

embryo minute, 0.1 the length of the endosperm, 

barely differentiated; endosperm copious, fleshy. 

Composition: 3 genera, -400 species. 

Distribution: Nearly cosmopolitan; most numerous in Asia and South America. 

ICACINACEAE (Figure 78c-f).-Trees, shrubs, 

and lianas, rarely herbs; xylem vessel perforation 

plates exclusively scalariform with 20-50 bars, 

or simple or both; interxylary phloem present in 

2 genera; leaves mostly alternate, rarely opposite, 

simple, entire, rarely lobed or toothed, exstipulate; inflorescences axillary or leaf-opposed, usually cymose or thyrsiform, less frequently racemes, spikes or panicles; flowers usually bisexual, rarely the plants dioecious; sepals 4-5 (0), 

connate, small; petals 4-5 (0), free or rarely 

connate, valvate or imbricate; stamens 4-5, alternipetalous, the filaments free or the anthers 

sessile high on the corolla; pollen 4(5)-brevicolpate, 3-porate, 3(-6)-pororate, 3-7-brevicolporate, 3-colporoidate, or nonaperturate; disk 

rarely present; pistil 1, the carpels 3 (2-5), style 

1, usually short, stigma(s) apical, sometimes capitate; ovary superior, usually unilocular, rarely 

2-5-locular, the ovules usually 2 per locule, subapical, unitegmic, crassinucellar or tenuinucellar, 

anatropous; fruit usually a drupe, rarely a samara, the seed 1 ; embryo usually small, straight or 

arcuate, 0.2 the length of the endosperm; cotyledons sometimes foliaceous or folded, rarely 

thick, 2-9 times wider than the radicle, 0.8 the 

length of the embryo; endosperm mostly present, 

rarely 0, sometimes copious. 

Composition: -45 genera, -400 species. 

Distribution: Rain forest and dry habitats; 

pantropical, few subtropical. 

CAR YOPHYLLALES 

Herbs, shrubs, rarely small trees, rarely climbing, often succulent, sometimes spiny; xylem vessel perforation plates simple, very rarely also 

reticulate; anomalous secondary thickening of 

the stem fairly frequent; leaves alternate or opposite, simple, entire, rarely toothed, sometimes 

rudimentary; flowers bisexual, rarely unisexual; 

perianth sometimes not differentiated into calyx 

and corolla, rarely spirally arranged and the parts 

numerous, free or rarely shortly connate; sepals 

2-5 (0-25), free or shortly connate, often colored; petals 0, 4, 5 (2 to -40) (staminodal?), in 

1-6 whorls, rarely transitional to stamens, free 

or connate, sometimes inserted on the calyx tube; 

stamens 5-30 (1 to -700), free or basally connate, rarely spirally arranged or in bundles, insertion hypogynous, perigynous or epigynous or 

on the base of the petals or on a glandular disk, 

the filaments often filiform, rarely very short, 

anthers dorsifixed or basifixed, sometimes pink; 

pollen 3-polycolpate, -rugate or -forate, 4-rupate; 

glandular disk often present; pistils 1 (-1 2), the 

carpels 1-5 (-20), styles 1-5 (-20, 0), the stigmas 

usually decurrent ventrally, sometimes apical; 

ovary superior to inferior, 1-5 (-2O)-locular, the 

ovules 1 -many per locule, axile, basal or parietal, 

usually campylotropous; fruit a berry, capsule, 

nutlet, drupe, achene, or schizocarp; embryo 

usually arcuate, peripheral to perisperm; perisperm, rarely endosperm, copious to absent. 

Distribution: Cosmopolitan, most common in 

dry and disturbed habitats. 
NUMBER 58 159 

r? 

b u 

f 

Q 

FIGURE 78.-AQUIFOLIACEAE: a, Ikx paraguayensis C.S. of bud, bud, flower, corolla and stamens, 

calyx, stamen, inflorescence, fruit in axil of leaf; b, fruit, C.S. of same showing the pyrenes, I. 

loranthoides Is. of pistil, views of stamen, pyrene, seed (after Martius, 1840- 1906). ICACINACEAE: 

c, Ernmoturn nitens flower, I.s. of same, floral diagram, views of petal, petal and 2 stamens; d, 

views of stamen, C.S. of anther, 1.s. of pistil, C.S. of ovary, infructescence; e, Poraqueiba sericea 

bud, petals, calyx, 2 petals and stamen, I.s. of bud, views of petal, views of stamen, pistil and 

calyx, 1,s. of pistil;f, floral diagram, fruit (after Martius, 1840-1906). 
160 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Chemistry: Betalains occur only in this order. 

According to Gibbs, except for lack of betalains, 

the Caryophyllaceae and Molluginaceae seem to 

fit reasonably well in the order. Triterpenoids, 

chiefly as saponins, occur in Phytolaccaceae, Molluginaceae, Caryophyllaceae, Chenopodiaceae, 

Aniaranthaceae, and Cactaceae but not Polygonaceae, but triterpenoids are very widespread in 

plants. Betacyanins are probably absent from 

Polygonaceae. According to Gibbs, Cactaceae 

and Aizoaceae are often, and probably rightly, 

associated. Caryophyllales, except Caryophyllaceae, lack ellagic acid. Fouquieria has ellagic acid, 

which is lacking in the Tubiflorae; otherwise it 

fits well in that group. Ocotitlol, a triterpenoid 

occurs in Fouquieria and Neolloydia texensis, a 

cactus. Gibbs believes the Bataceae and Polygonaceae are separate entities from this order and 

that chemically the Plantaginaceae fits very well 

in or near the Tubiflorae. It would be less happily 

placed as a derivative of the Primulales. 

CACTACEAE (Figure 79a).-Spiny succulent 

herbs, shrubs, trees and scramblers; xylem vessel 

perforation plates simple, very rarely also reticulate; anomalous secondary thickening of the 

stem occurs in at least 2 genera; leaves usually 

rudimentary, sometimes well developed, simple, 

alternate, entire, exstipulate; flowers usually solitary, rarely in panicles, bisexual, rarely unisexual, the perianth usually not differentiated into 

calyx and corolla, spirally arranged, the tepals 

usually many, sometimes few, free or rarely 

basally connate, the floral tube containing nectar; 

stamens spirally arranged, numerous (-30-700), 

sometimes in groups, insertion perigynous or 

epigynous, sometimes adnate to the base of the 

petals, the filaments filiform; anthers small, 

about as long as wide, dorsifixed or basifixed; 

pollen 3-colpate, 6-polyrugate or forate; pistil 1, 

the carpels 2 to -10, the style 1, stigmas 2 to 

-1 0, slightly elongated and together broader 

than the apex of the style; ovary usually inferior 

and the receptacular tube often extending above 

the ovary, or rarely superior (Pereskia), unilocular, the ovules numerous to few, parietal or rarely 

basal (Pereskia spp.), bitegmic, crassinucellar, 

anatropous to f campylotropous or circinotropous, the funicle often long; fruit a berry, rarely 

dry, rarely with longitudinal or circumscissile 

dehiscence, the seeds usually numerous; embryo 

often peripheral and arcuate, rarely straight; cotyledons 0.07-0.7 the length of the embryo, large 

and convolute in Pereskia, reduced in some genera and then the radicle expanded; perisperm 

generally floury-starchy, moderate, scanty or 0. 

Composition: 

Distribution: Chiefly dry regions of warm 

temperate to tropical America; few high in the 

Andes or epiphytic in tropical forest; one species 

may be native in tropical west Africa, Madagascar, and Sri Lanka. 

AIZOACEAE (Figure 79b,c).-Herbs and low 

shrubs, often succulent; xylem vessel perforation 

plates simple; anomalous secondary thickening 

of the stem occurs; leaves alternate or opposite, 

simple, usually entire, rarely toothed, usually exstipulate; inflorescences axillary or terminal pleiochasia, dichasia or the flowers solitary; flowers 

bisexual, rarely unisexual (Glischrothamnus is 

dioecious); calyx lobes 5-8; petals -30-40 (staminodial?) or 0, in 1-6 whorls, sometimes transitional to stamens, sometimes connate, inserted 

on the calyx-tube; stamens numerous (to -120), 

4-5, 8 or 10, rarely 1, free or basally connate, 

sometimes in bundles, perigynous; anthers small, 

about as long as wide; pollen 3-colporoidate, 

occasionally 4-rupate or 4-6-rugate; disk often 

around the ovary; pistil 1, the carpels 2-5 (-20), 

sometimes weakly connate, the styles 1-5 (-20) 

or 0, stigmas apical or decurrent ventrally or 

sessile; ovary superior to inferior 2-5( 1-20)- 

locular, the ovules (1-) several or many, axile, 

basal, or apical, bitegmic, crassinucellar, campylotropous or anacampylotropous, the funicle 

often long; fruit usually a capsule, mostly loculicidal, more rarely a septicidal schizocarp or circumscissile, sometimes a nut or drupe, rarely a 

berry (Carpobrotus); embryo periferal, arcuate, 

as long as the perisperm to more than twice as 

long; cotyledons as wide as the radicle or slightly 

- 100 genera, -2,000 species. 
NUMBER 58 161 

FIGURE ~~.--CACTACEAE: a, Opuntia monacantha flower, fruit, Cereus macrogonus Is. of flower 

(apex of style removed), anther, stigma, seed and I.s. of same (after Martius, 1840-1906). 

AIZOACEAE: b, Sesuvium portulacastrum flowering shoot, flower, lateral view of same, part of 

calyx laid open, views of stamen, pistil; c, floral diagram, dehiscent fruit, part of same, seed, I.s. 

of same. (after Martius, 1840-1 906). PORTULACACEAE: d, Portulaca elatior flower, lateral view 

of same, sepals (2), sepal, petal, 1.s. of pistil, views of stamen, floral diagram; e, fruit, dehiscent 

fruit, seed, I.s. of same (after Martius, 1840-1906). 
162 

wider, 0.4-0.5 the length of the embryo; perisperm moderate or copious, hard and semi-transparent or soft and whitish. 

Composition: 

Distribution: 

130 genera, - 1200 species. 

Centered in the drier parts of 

South Africa, extending into eastern and western 

Africa; southern and western Australia, New 

Zealand; southwestern United States to northwestern Mexico; western middle South America; 

western, southern, and eastern Asia. 

PORTULACACEAE (Figure 79d,e).-Herbs and 

subshrubs, often succulent; xylem vessel perforation plates simple; intraxylary phloem present 

in some genera; leaves alternate or opposite, 

sometimes rosulate, simple, entire, the stipules 

sometimes scarious, rarely 0 (Claytonia); inflorescences cymes, often dichasia to cincinni, panicles or heads or the flower solitary; flowers bisexual or very rarely the plants monoecious (Portulacaria) sepals (bracteoles?) 2 (5-8), free or 

basally connate; petals (tepals) 4-5 (2-16), free 

or basally connate; stamens 4-5 (1 to -50), opposite the petals, the filaments filiform, free or 

rarely connate in bundles, occasionaly adnate to 

the corolla; anthers small, often pink, about as 

long as wide, wall development conforms to the 

Monocot type; pollen 3-colpate, 6-polyrugate, 

polyforate; nectaries receptacular; pistil 1, the 

carpels 3 (2-8), the styles united most of the way, 

with 3 (2-8) apical linear branches, the stigmas 

decurrent ventrally, ovary superior, semi-inferior or inferior, septate when young, becoming 

unilocular, the ovules rn to 1, axile-basal, bitegmic, crassinucellar, anatropous, anacampylotropous, or amphitropous, the funicle sometimes 

long; fruit a capsule, dehiscence by apical valves 

or sometimes circumscissile, or a schizocarp, 

rarely a nut; seeds reniform; embryo peripheral, 

arcuate, more than twice as long as the perisperm; cotyledons about as wide as the radicle, 

0.3-0.5 the length of the embryo; perisperm 

moderate or copious, rarely scanty, generally 

hard and semitransparent, sometimes soft and 

whitish. 

Composition: 19 genera, -540 species. 

Distribution: Cosmopolitan except arctic reSMITHSONIAN CONTRIBUTIONS TO BOTANY 

gion of the Northern Hemisphere, the Amazon 

basin, the Sahara, and part of the Australian 

deserts; centered in South Africa, western North 

America, and the Andes. 

THELIGONACEAE (Figure 80a,b).-Succulent 

terrestrial herbs; xylem forms a continuous cylinder; leaves simple, entire, lower ones opposite, 

upper alternate; stipules attached to the petiole; 

plants monoecious, the flowers axillary in 1-3- 

flowered sessile cymes, anemophilous; calyx 

closed in bud, 2-5-partite at anthesis; petals 0; 

stamens 7-12 (2-30), the filaments filiform, free; 

anthers linear, dorsifixed; pollen 3-7-poratebrevicolpate; pistil 1, carpel 1, the style 1, filiform, stigma apical, punctiform; ovary superior 

(inferior according to some who see adnation to 

perianth), unilocular, uniovulate, the ovule basal, 

unitegmic, tenuinucellar, subanatropous or campylotropous; fruit a nutlet; seed hippocrepiform, 

the embryo axile, arcuate, about as long as the 

endosperm; cotyledons flat, thin, 2.3-5.5 times 

wider than the radicle, 0.5-0.6 the length of the 

embryo; endosperm copious, fleshy. 

Composition: 1 genus, 3 species. 

Distribution: Mediterranean region, southern 

Japan, western China, Canary Islands. 

DIDIEREACEAE (Figure 80c,d).-Spiny trees 

and shrubs; xylem vessel perforation plates simple; leaves alternate, simple, entire, exstipulate; 

plants usually dioecious, rarely gynodioecious; 

inflorescences terminal dichasia or fascicles; sepals 2 (bracteoles?), petaloid; petals 4, in 2 series, 

free; stamens 8- 10, the filaments filiform, 

shortly connate basally, inserted outside an annular disk; anthers oblong, didymous, dorsifixed; 

pollen 4-7-colpate; pistil 1, the carpels 3 (2-4), 

the style 1, stigma often expanded and 3-4- 

lobed; ovary superior, the locules 3 but only one 

fertile, ovule 1, basal, bitegmic, crassinucellar, 

campylotropous or hemicampylotropous; fruit a 

nutlet; seed arillate; embryo folded, the cotyledons about as long as the radicle; endosperm 

very scanty or 0. 

Composition: 4 genera, 11 species. 

Distribution: Semi-desert regions of Madagascar. 
NUMBER 58 163 

FIGURE ~~.--THELIGONACEAE: a, Theligonum cynocrambe flowering shoot, S flower bud, S 

flower, young 0 flower with bracteoles, mature 0 flower; b, fruit, I.s. and C.S. of same, embryo 

(after Le Maout and Decaisne, 1873). DIDIEREACEAE: c, Didierea mirabilis habit, 0 flower, I.s. of 

same, is. of pistil, fruit; d, 8 flowers, stamen (after Drake). 

GYROSTEMONACEAE (Figure 8 la-c).-Trees 

and shrubs; xylem vessel perforation plates simple; stem without anomalous secondary thickening; leaves alternate, simple, entire, succulent, 

stipules rudimentary or absent; plants mostly 

dioecious, sometimes monoecious; inflorescences 

axillary racemes or spikes, or the flowers solitary; 

calyx 2-5-lobed or truncate; petals 0; stamens 6 

to -100 in 1 or more series; anthers oblong, 

subsessile; pollen 3-colpor(oid?)ate, receptacle 

broad; pistil 1, the carpels sometimes loosely 

connate, mostly to 19, rarely as few as 2 or 1, 

the styles 1-19, stigmas apical, linear; ovary superior 19- l-locular, the ovules campylotropous, 

1 per locule, axile; fruit dry, a capsule, or eventually falling apart, the parts dehiscing ventrally, 

dorsally or both, or rarely indehiscent; seeds 

hippocrepiform, arillate; embryo peripheral, 

longer than the perisperm (endosperm ?); cotyledons not broadened, 0.4 the length of the 

embryo; perisperm (endosperm ?) copious or 

scanty, fleshy. 
164 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Composition: 

Distribution: Most of Australia. 

PHYTOLACCACEAE (Figure 8 1 d,e).-Herbs, 

shrubs, rarely trees or scandent; xylem vessel 

perforation plates simple; anomalous secondary 

thickening of the stem occurs in some species; 

leaves alternate, simple, entire, exstipulate; inflorescences terminal, leaf-opposed or axillary 

spikes or racemes, rarely panicles; flowers bisexual or rarely the plants dioecious, the parts hypogynous; sepals 4-10, free, green or colored; 

petals 0; stamens 5-30, in 1 or 2 series, the 

filaments filiform, free or basally connate, sometimes inserted on a fleshy disk; anthers small, 

about as long as wide, dorsifixed; pollen 3-colpate, 4-rupate; pistils 4-12, sometimes k connate, the styles 4-12, free or basally connate, 

stigmas ventrally decurrent; ovary 4- 1 2-locular, 

the ovules 1 per locule, bitegmic, crassinucellar, 

amphi-, anacampylo-, or campylotropous, basalventral; fruit a berry or drupe; seeds & reniform, 

the embryo peripheral, arcuate; cotyledons 1 - 

1.5 times the width of the radicle, 0.4 the length 

of the embryo; perisperm moderate or copious, 

soft or granular and whitish. 

5 genera, 1 6 species. 

Composition: 12 genera, 100 species. 

Distribution: Centered in tropical and subtropical America, extending to temperate America; southern 2/3 of Africa; eastern Mediterranean 

region; Japan to India and the East Indies, northeastern Australia. 

BARBEUIACEAE (Figure 8 If).-Shrubby tree, 

sarmentose; xylem vessel perforation plates simple; stem with anomalous secondary thickening; 

leaves alternate, simple, entire, exstipulate; inflorescence short axillary racemes; flowers bisexual, 

the parts hypogynous; sepals 5, free; petals 0; 

stamens numerous (-50-60), in several series, 

the filaments inserted on an annular disk; anthers 

linear, sagittate, basifixed; pollen 3-colporoidate, 

occasionally 4-ruporoidate or 6-rugoroidate; pistil 1, the carpels 2, the styles 2, linear, very 

shortly connate, the stigmas decurrent ventrally; 

ovary bilocular, the ovules 1 per locule, basal, 

campylotropous; fruit a woody capsule, the seeds 

reniform, arillate; embryo arcuate, axile, about 

as long as the endosperm; cotyledons not broadened, 0.4 the length of the embryo; endosperm 

(perisperm?) copious, mealy. 

Composition: 1 genus, 1 species. 

Distribution: Madagascar. 

ACHATOCARPACEAE (Figure 8 lg).-Trees and 

shrubs, sometimes thorny; xylem vessel perforation plates simple; stem without anomalous secondary thickening; leaves alternate, simple, entire, exstipulate; inflorescences axillary or cauliflorous racemes, short panicles, or fascicles; 

plants dioecious, the flower parts hypogynous; 

sepals 4-5, imbricate, free or very shortly connate, persistent; petals 0; stamens 10-20, the 

filaments filiform, shortly connate; anthers oblong, basifixed; pollen 6-polyforate or 3-colpate; 

pistil 1, the carpels 2, the styles 2, short, free, the 

stigmas decurrent ventrally; ovary unilocular, 

ovule 1, basal, campylotropous; fruit a berry, the 

seed exarillate; embryo peripheral, arcuate, 

about 2.5 times longer than the perisperm; 

cotyledons not broadened, 0.5-0.6 the length of 

the embryo; perisperm copious. 

Composition: 2 genera, 10 species. 

Distribution: Texas and Mexico to Paraguay 

and Argentina. 

PETIVERIACEAE (Figure 82a).-Shrubs, lianas, 

and herbs; xylem vessel perforation plates simple; 

stem with anomalous secondary thickening; 

leaves alternate, simple, entire or rarely crenate, 

the stipules small or 0; inflorescences axillary and 

FIGURE 81 .-GYROSTEMONACEAE: a, Gyrostemon australasicus 6 flowering branch, 0 flowering and fruiting branch; b, 

d flower, seed, G. ramulosus d flower, 1,s. of same; c, G. 

australasicus fruit, G. ramulosus 0 flower, 1.s. of same, dehiscent fruit, Codonocarpus pyramidalis 1,s. of fruitlet (after 

W'alter; Baillon, 1866-1 895). PHYTOLACCACEAE: d, Phytolacca thyrszflora d flower from above, flower not fully open, 

1,s. of flower, views of anther, pistil; e, fruit with perianth, 

seed, 1,s. and C.S. of same (after Martius, 1840-1906). BARBEUIACEAE: J, Barbeuia madagascariensis part of flowering 

twig, bud with 2 tepals removed, flower, pistil with ovary 

opened to show ovule and 1 stamen, I.s. of ovary (after 

Baillon, 1866-1 895). ACHATOCARPACEAE: g, Achatocarpus 

bicornutus flowering twig, young fruit, A. balansae interior 

tepal, external tepal, A. praecox 1,s of fruit (after Schinz and 

Autran). 
NUMBER 58 165 
166 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE ~~.-PETIVERIACEAE: a, Petiveria hexaglochin 2 views of flower, tepal, stamens, views of 

pistil, 1,s. of same, fruit with and without perigone, C.S. of fruit showing embryo (after Martius, 

1840- 1906). AGDESTIDACEAE: b, Agdestis clernatidea flower, views of stamen, I.s. of ovary, part 

of perianth and base of stamens, ripe fruit with persistent perianth (after Hemsley). NYCTAGINACEAE: c, Pisonia tornentosa 6 flower, same laid open, views of stamen, 0 flower, I.s. of same, 

fruit enclosed in perigone, embryo (after Martius, 1840- 1906). STEGNOSPERMATACEAE: d, 

Stegaosperna halim$oliolium flowering twig, flower, views of stamen, 1,s. of ovary, dehiscent fruit, 

seed enclosed in aril and without aril, I.s. of seed (after Fawcett and Rendle; Walter). ILLECEBRACEAE: e, Paronychia brasilianu flower, part of perianth laid open, fruit, Is. of ovary, I.s. of 

seed (after Martius, 1840-1906). 
NUMBER 58 167 

terminal racemes, panicles or spikes; flowers bisexual or rarely the plants dioecious, the parts 

hypogynous; sepals 4-5, sometimes shortly connate; petals 0; stamens 4-25, the filaments free 

or shortly connate; anthers oblong, dorsifixed or 

basifixed; pollen 3-colpate or 6-polyrugate or 

polyforate; disk present or absent; pistil 1, the 

carpels 1-5, the styles 1-5, stigmas decurrent; 

ovary unilocular, the ovule 1, basal, bitegmic, 

crassinucellar, campylotropous; fruit a berry, 

samara, or spiny nutlet; embryo peripheral, arcuate or straight, the cotyledons thin, foliaceous, 

convolute about the upper part of the radicle; 

perisperm moderate, hard and semitransparent. 

Composition: 6 genera, 10 species. 

Distribution: Warm temperate to tropical 

America. 

AGDESTIDACEAE (Figure 82b).-Twining 

herb, liana, or subshrub; xylem vessel perforation plates simple; stem with anomalous secondary thickening; leaves alternate, simple, entire, 

exstipulate; inflorescences axillary loose panicles 

of cymules, the flowers bisexual; sepals 4-5, very 

shortly connate or free, accrescent; petals 0; stamens 15-20, in more than 1 series, the filaments 

filiform, inserted on a disk; anthers dorsifixed 

near the base; pollen 3-colpate; pistil 1, the carpels 3-4, the styles shortly connate, 3-4-fid and 

linear apically, the stigmas decurrent ventrally; 

ovary semi-inferior, 3-4-locular7 the ovules 1 per 

locule, axile, subbasal, campylotropous; fruit a 

unilocular 1-seeded nutlet, winged by the calyx 

lobes; seed exarillate, the embryo peripheral, 

arcuate, the cotyledons broadly oblong, the 

radicle short; perisperm scanty, mealy. 

Composition: 1 genus, 1 species. 

Distribution: 

NYCTAGINACEAE (Figure 82c).-Shrubs, trees, 

herbs, and lianas, sometimes thorny; xylem vessel 

perforation plates simple, occasionally also reticulate: stem with anomalous secondary thickening; interxylary phloem in some genera; leaves 

alternate or opposite, simple, entire, exstipulate; 

inflorescences various, cymose, very rarely a raceme (Cyphomeris): flowers bisexual or unisexual, 

rarely the plants dioecious (Cephalotomandra), 

Mexico to Brazil, West Indies. 

the parts hypogynous; bracts sometimes colored; 

calyx tubular, often petaloid, the lobes 5 (3-7); 

petals 0; stamens 5 (1-30), sometimes in 2 whorls, 

the filaments filiform, free or shortly connate, 

rarely adnate to the calyx-tube; anthers dorsifixed near the base; pollen 3(4)-colpate, 6-polyrugate or polyforate; pistil 1, the carpel 1 (rarely 

a flower with 2 pistils ventrally connected), the 

style filiform, stigma apical, rarely the stigma 

sessile (Reichenbachia) or decurrent ventrally 

(Ramisia); ovary unilocular, ovule 1, basal, 1-2- 

tegmic, crassinucellar, anacampylo- or campylotropous or subanatropous; fruit an achene or 

nutlet, enclosed by the base of the calyx; embryo 

peripheral, arcuate or rarely straight, twice as 

long as the perisperm: cotyledons often unequal, 

rarely reduced to 1, thin, rarely longitudinally 

convolute, 3-9 times wider than the radicle, 0.5 

the length of the embryo; perisperm copious, 

moderate, or rarely scanty, crystalline-granular, 

rarely whitish and mealy or whitish and hard. 

Composition: 30 genera, -290 species. 

Distribution: Mainly pantropical and subtropical, particularly America; temperate North 

America to temperate South America, Africa, 

southern Asia, Australia, New Zealand. 

STEGNOSPERMATACEAE (Figure 8 2d). -Shrub, 

sometimes scandent; xylem vessel perforation 

plates simple; stem without anomalous secondary 

thickening; leaves alternate, simple, entire, 

fleshy, exstipulate: inflorescence a terminal raceme or thyrse or axillary cymes; flowers bisexual, the parts hypogynous; sepals 5, free, persistent; petals 5, free; stamens 10, the filaments 

filiform, shortly connate; anthers about as long 

as wide, dorsifixed; pollen 3-colpate; pistil 1, the 

carpels 3-5, the styles short, linear, shortly connate, the stigmas decurrent ventrally: ovary unilocular with a central column, the ovules 3-5, 

basal, amphitropous; fruit a capsule, the valves 

separating to the base; seeds with a large red aril; 

embryo peripheral, arcuate, 1.3 times longer 

than the perisperm; cotyledons not broadened 

(?), 0.7 the length of the embryo; perisperm 

copious, mealy. 

Composition: 1 genus, 3 species. 
168 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Distribution: Dry or coastal regions; Mexico, 

Central America, West Indies. 

CARYOPHYLLACEAE (Figure 8 3a,b).-Herbs, 

subshrubs, rarely shrubs, the stems often swollen 

at the nodes; xylem vessel perforation plates simple; stem and roots of some genera with anomalous secondary thickening; leaves opposite, simple, entire, stipules absent or present and then 

often scarious; inflorescences usually terminal, 

paniculate, racemiform or capitate dichasial 

cymes or cincinni, or the flowers solitary; flowers 

bisexual or rarely the plants dioecious (Sanctambrosia, Melandrium dioicum), the parts hypogynous, rarely slightly perigynous; sepals 5 (4-25), 

rarely spirally arranged, free or connate; petals 

5 (4-1 2) or 0, free; stamens 10 (3-1 3), usually in 

2 series, mostly obdiplostemonous, the filaments 

filiform, free; anthers about as long as wide, 

dorsifixed; pollen 3-colpate, polyforate; glands at 

the base of the stamens or disk present; pistil 1, 

the carpels 2-5 (6), the styles 2-5 (6), filiform, 

free or shortly connate, rarely mostly connate, 

the stigmas decurrent ventrally; ovary unilocular 

or imperfectly 2-5-locular, the ovules mostly numerous to only 1, bitegmic, crassinucellar, usually campylotropous, or semi-anatropous, the placenta free-central, axile or basal; fruit a capsule, 

dehiscent by apical teeth or by valves, rarely 

circumscissile (Drypis spinosa), or rarely a scarcely 

fleshy berry (Cucubalus); seeds usually numerous 

(l(2) in Drypis spinosa); embryo peripheral, arcuate to spiral (Drypis), or straight (Dianthus), 1 - 

3 times as long as the perisperm; cotyledons 1-2 

times as broad as the radicle, 0.5 the length of 

the embryo; perisperm moderate, copious or absent, usually firm or hard and semitransparent, 

rarely soft and white. 

Composition: 55 genera, 1550 species. 

Distribution: Cosmopolitan, mainly temperate Northern Hemisphere, centered in the Mediterranean region, sparingly arctic, antarctic, and 

in the mountains of the tropics. 

MOLLUGINACEAE (Figure 83c).-Herbs and 

subshrubs, rarely shrubs; stems and roots with 

anomalous secondary thickening; leaves opposite, alternate or subverticillate, simple, entire, 

slightly or not succulent, the stipules scarious, 

small or 0; inflorescences pleiochasial or dichasial 

cymes, or in glomerules to cincinni, or the flowers solitary; flowers bisexual; sepals 5, mostly 

free, rarely shortly connate; petals typically 0, 

sometimes 4-5, free or rarely connate (Corbichonia); stamens 5- 10 (many in Corbichonia), the 

filaments filiform, free or shortly connate, sometimes in bundles, insertion hypogynous or slightly 

perigynous; anthers slightly longer than wide, 

dorsifixed; pollen mainly tricolpate; disk absent 

or annular; pistil 1 (except Gisekia), the carpels 

2-5, the styles 2-5, or sometimes the stigmas 

sessile; ovary superior, 2-5-locular, the ovules 1 

per locule and basal, or numerous and axile, 

bitegmic, crassinucellar, campylotropous to almost anatropous (Macarthuria and Mollugo ?); 

fruit a loculicidal or circumscissile capsule, rarely 

a nutlet or schizocarp; seeds sometimes arillate; 

embryo peripheral, arcuate, about twice as long 

as the perisperm; cotyledons not broadened, 0.4 

the length of the embryo; perisperm moderate 

to copious, hard and semitransparent. 

Composition: 14 genera, 95 species. 

Distribution: Mostly tropical and subtropical, 

in dry habitats, few temperate; centered in Africa. 

ILLECEBRACEAE (Figure 82e).-Herbs, rarely 

subshrubs, xylem vessel perforation plates simple; leaves mostly opposite, rarely some alternate, 

simple, mostly entire, the stipules scarious, sometimes connate, rarely absent; bracts often scarious; flowers bisexual or rarely unisexual, solitary 

or in cymes(?); sepals 4-5, free or shortly connate; petals 0 or 4-5; stamens 3-5, rarely fewer 

or more, mostly perigynous, the filaments free 

or shortly connate; anthers about as long as wide, 

dorsifixed, the sacs separate; pollen 3-porate, 3- 

colpate, 6-polyforate; pistil 1, the carpels (1) 2- 

3, the styles 1-3, free, the stigmas terminal(?); 

ovary superior, unilocular, ovule 1 (2-4), basal, 

bitegmic, crassinucellar, semi-anatropous; fruit a 

utricle or nutlet, 1-seeded; embryo peripheral, 

straight to nearly annular; cotyledons 1-1.5 

times the width of the radicle, 0.4-0.6 the length 

of the embryo; perisperm copious to scanty, usually hard and transparent. 

Composition: 15 genera, 200 species. 
NUMBER 58 169 

ca 

1 

d 

FIGURE 83.-CARYOPHYLLACEAE: a, Cerastiurn humifwurn flower, dehiscent fruit with calyx, 

Polycarpon depressurn I.s. of flower, floral diagram, seed, Is. of same; b, Drymaria cordata 

essential organs, views of stamen, dehiscent fruit, seed, I.s. of same (after Martius, 1840-1906). 

MOLLUGINACEAE: c, Mollugo uerticillata flower, floral diagram, fruit, dehiscent fruit, 2 views of 

seed, C.S. and I.s. of same (after Martius, 1840-1906). BASELLACEAE: d, Basella rubra 1,s. of 

flower, fruit, Is. of same, Anredera baselloides flower, Is. of same, flowering shoot (after Baillon, 

1866-1 895). 

Distribution: Cosmopolitan, most common in 

drier habitats. 

BASELLACEAE (Figure 83d).-Herbaceous twiners or procumbent; xylem vessel perforation 

plates simple; stem without anomalous secondary 

thickening, but with bicollateral bundles in old 

stems(?); leaves alternate, simple, entire, often 

fleshy, exstipulate; inflorescences axillary or supra-axillary spikes, racemes or panicles; flowers 

bisexual, rarely unisexual; sepals 5, nearly free 

to connate in a tube, often colored; petals 0; 

stamens 5, the filaments short, connate basally 
170 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 84.-AMARANTHACEAE: a, Amaranthus hypochondriacus d floMer, views of stamen, 9 

flower, dehiscent fruit, seed, I.s. of same; b, Indobanalia brasiliana infloresence, essential organs, 

I.s. of fruit, views of anther, Alternanthera philoxeroides stamina1 tube laid open and pistil (after 

Martius, 1840-1906). CHENOPODIACEAE: c, Chenopodium retusum flower, I.s. of same, Is. of 

ovary, views of anther, C. spathulaturn I.s. and C.S. of seed (after Martius, 1840-1906). 

or inserted on the base of the sepals and opposite 

them; anthers about as long as wide, dorsifixed, 

rarely basifixed, reversed in bud, usually dehiscing by apical pores or pore-like slits; pollen 6- 

polyrugate or forate or transitional between 

them; annular disk sometimes present; pistil 1, 

the carpels 3, the style 1, often 3-lobed, rarely 

unlobed, the stigmas apical or decurrent; ovary 

superior, unilocular, ovule 1, basal, bitegmic, 

crassinucellar, anatropous, anacampylo- or 

campylotropous; fruit a berry or drupe; embryo 

peripheral, arcuate to subannular, or spirally 

twisted; cotyledons plano-convex or subfoliaceous, 0.5-0.7 the length of the embryo; perisperm copious or scanty. 

Composition: 4 genera, -25 species. 

Distribution: Tropical and subtropical, mostly 

America; Africa, Madagascar, southern India, 

New Guinea. 

AMARANTHACEAE (Figure 84a,b).-Herbs, 

rarely shrubs or smaIl trees; xylem vessel perforation plates simple; stem and roots with anomalous secondary thickening; leaves alternate or 

opposite, simple, entire, exstipulate; inflorescences terminal and axillary spikes, heads, racemes or cymes, glomerules, dichasia or cincinni; 

flowers usually bisexual, rarely the plants polygamous or dioecious, the parts hypogynous (stamens perigynous in Guilleminea); bracts and bracteoles often scarious; sepals (0-1) 3-5, free or 

shortly connate, scarious; petals 0; stamens 5 (1- 

4), often alternate with staminodial membranes, 
NUMBER 58 171 

the filaments free or connate, sometimes inserted 

on the calyx or disk; anthers small, dorsifixed; 

pollen 6-polyforate; disk present or absent; pistil 

1, the carpels 2-3, the styles 1-3, the stigmas 

decurrent or apical; ovary unilocular, the ovules 

1 -few, basal, bitegmic, crassinucellar, campyloor anacampylotropous or circinotropous; fruit a 

pyxis, nutlet or berry; embryo peripheral, subannular; cotyledons 1-1.5 times the width of the 

radicle, 0.5-0.7 the length of the embryo; 

perisperm copious, hard and translucent to granular and whitish. 

Composition: 65 genera, -850 species. 

Distribution: Cosmopolitan except for arctic 

Northern Hemisphere; many in tropical Africa 

and America. 

CHENOPODIACEAE (Figure 84c).-Herbs and 

shrubs, rarely small trees or climbers; xylem vessel perforation plates simple or very rarely with 

some oblique scalariform perforations (Axyris); 

stem and roots with anomalous secondary thickening; interxylary phloem typical; leaves mostly 

alternate, rarely opposite, simple, sometimes 

fleshy or reduced to scales, entire or toothed, 

exstipulate; inflorescences axillary and terminal 

heads, spikes, thyrses, dichasia or cincinni, or 

flowers rarely solitary; flowers minute, bisexual 

or the plants monoecious or dioecious; sepals 3- 

5 (0-4), free or connate, sepaloid or membranous, often accrescent; petals 0; stamens 3-5 (1, 

2) and opposite the calyx lobes, hypogynous or 

inserted on a disk or on the calyx, the filaments 

usually free, sometimes shortly connate; staminodes rare; anthers dorsifixed, incurved in bud, 

rarely with a large apical connective appendage; 

pollen (oligo-) polyforate; disk present or absent; 

pistil 1, the carpels 2 (3-5), the styles 2 (1-5), 

short, the stigmas decurrent ventrally or all over; 

ovary superior, rarely semi-inferior (Beta), unilocular, ovule 1, basal, bitegmic, crassinucellar, 

campylotropous or amphitropous; fruit a nutlet 

or achene, rarely a pyxis or berry; embryo peripheral, arcuate, annular or coiled, as long as to 

more than twice as long as the perisperm, sometimes conduplicate and the radicle incumbent; 

cotyledons 1-2 times the width of the radicle, 

0.5 the length of the embryo; perisperm copious 

to absent, usually firm to flinty and glass-like to 

whitish. 

Composition: 

Distribution: 

102 genera, - 1500 species. 

Principally in desert, saline and 

disturbed habitats; cosmopolitan. 

PRIMULALES 

Herbs, shrubs, and small trees, rarely climbing; 

xylem vessel perforation plates simple, a few 

species of Myrsinaceae also with scalariform 

plates; stem sometimes with anomalous secondary thickening, sometimes resinous; leaves opposite or alternate, simple, entire or toothed, 

exstipulate; flowers bisexual, rarely unisexual, 

sometimes heterostyled; sepals 4-5 (3, 6-9), free 

or connate; petals 4-5 (0, 3, 6-9), usually connate; stamens 4-5 (to -75 in Tamaricaceae), 

free, shortly connate or epipetalous opposite the 

corolla lobes when in one series, rarely 5 (4) 

staminodes present; anthers dorsifixed, rarely 

basifixed; pollen usually 3(2-5)-colporate, less 

often 3- 1 O-colpate, 4-rupate or 6-rugate; glandular disk sometimes present; pistil 1, the carpels 

5 (2-9), styles 1 (5), the stigma(s) apical, more 

rarely decurrent ventrally or sessile; ovary superior, rarely semi-inferior, unilocular, the ovules 

1 -many, basal or the placenta free-central, more 

rarely parietal; fruit a capsule, berry or drupe, 

rarely a nutlet; embryo straight, often cylindrical; 

endosperm copious to absent. 

Distribution: Cosmopolitan. 

Chemistry: Triterpenoid saponins are prevalent in the order. Alkaloids are virtually absent. 

Gibbs feels a close association between Plumbaginaceae and Primulales is justified. Plumbaginaceae and Droseraceae have plumbagin, a 

naphthaquinone. Plumbaginaceae lack betalains. 

Primulaceae lack betacyanins. 

PRIMULACEAE (Figure 85a,b).-Herbs, rarely 

subshrubs; xylem vessel perforation plates simple; stem sometimes with anomalous structure 

(concentric rings of bundles); tissues often with 
172 

a 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

1 

FIGURE 85.-PRIMULACEAE: a, Primula officinalis 1.s. of flower, floral diagram, dehiscent fruit, 

Anagallis tenella bud, corolla (1 segment removed) and staminal tube and apex of style, staminal 

tube laid open to show the pistil, dehiscent fruit; b, A. minimus corolla laid open to show the 

insertion of the stamens opposite the petals, A. latifoliolia placenta and 1,s. of same showing 

insertion of seeds, Pelletiera Vera seed, 1,s. of same, embryo (after Martius, 1840-1906; Le 

Maout and Decaisne, 1873). PLUMBACINACEAE: c, Lzmonzum brasiliense flower, corolla laid open 

to show insertion of stamens opposite the petals, calyx in fruit, Plumbago europaea flowering 

shoot, P. scandens floral diagram, P. europaea flower; d, I.s. of flower, essential organs, fruit, 

seed, 1,s. of same, Armeria sp. pistil (after Martius, 1840-1906; Le Maout and Decaisne, 1873). 

secretory cells and spaces; leaves basal or cauline, 

usually opposite or verticillate, sometimes alternate, simple, often glandular-punctate, entire or 

toothed and lobed or divided (Hottonia), exstipulate; inflorescences terminal umbels, racemes, 

panicles, or the flowers solitary; flowers bisexual, 

often heterostyled, rarely slightly zygomorphic; 

sepals 5 (4-9), connate; petals 5 (4-9), (0 in 
NUMBER 58 173 

Glaux), connate, sometimes only shortly so or 

separate (Pelletiera), rarely unguiculate; stamens 

5 (4-9), the filaments long or short, free or 

basally connate, or epipetalous and opposite the 

corolla lobes, rarely alternating with staminodes 

(scales); anthers dorsifixed; pollen 3(2-4)-colporate or colporoidate, or 3-10-colpate; pistil 1, the 

carpels 5 (4-9), the style 1, stigma apical, capitate, simple; ovary superior, rarely semi-inferior, 

unilocular, the ovules few to numerous, basal or 

free-central, 2( 1)-tegmic, tenuinucellar, anatropous, semi-anatropous, or campylotropous; fruit 

a capsule, dehiscent usually by apical teeth, or by 

valves, rarely circumscissile, rarely subindehiscent, the seeds angular, sometimes minute; embryo linear, straight, terete, 0.4-0.9 the length 

of the endosperm; cotyledons 1-1.7 times the 

width of the radicle, 0.2-0.5 the length of the 

embryo; endosperm copious, hard or firm, semitransparent. 

Composition: -25 genera, -900 species. 

Distribution: Cosmopolitan, centered in temperate Northern Hemisphere; many alpine, a few 

on shores. 

PLUMBAGINACEAE (Figure 85c,d).-Herbs, 

subshrubs, or climbers; xylem vessel perforation 

plates simple; stem often with cortical and medullary vascular bundles and occasionally anomalous secondary thickening; leaves alternate, 

sometimes rosulate, simple, entire, exstipulate; 

inflorescences terminal or axillary spikes, racemes, panicles, subumbels, sometimes involucrate heads, or dichasia; flowers bisexual, sometimes heterostyled (Limonium); bracts often 

sheathing and scarious; sepals 5, connate most of 

the way, the limb sometimes scarious; petals 5, 

connate, sometimes shortly so, contorted and 

imbricate in bud; stamens 5, elongated, free or 

basally connate, or epipetalous, opposite the corolla lobes; anthers oblong, dorsifixed; pollen 3- 

5-colpate, 4-rupate, 6-rugate; disk absent; pistil 

1, the carpels 5, the styles 5, filiform, or 1 and 

shortly 5-lobed apically, the stigmas decurrent 

ventrally, sometimes for the entire length of the 

style, or capitate; ovary superior, unilocular, 

ovule 1, basal, bitegmic, crassinucellar, circinotropous or anatropous, the funicle long; fruit a 

nutlet or capsule at length circumscissile, rarely 

valvate from the base; embryo straight, 0.9 to as 

long as the endosperm; cotyledons thin or moderately thick, 2-3 times the width of the radicle, 

0.4-0.9 (Aegialitis 0.2) the length of the embryo; 

endosperm scanty, moderate or 0, firm crystalline-granular, floury, or fleshy. 

Composition: 

Distribution: 

- 15 genera, -700 species. 

Most commonly in dry or saline 

habitats; cosmopolitan, centered in the Mediterranean region, western and central Asia. 

TAMARICACEAE (Figure 86a-c).-Shrubs, 

trees, and herbs; xylem vessel perforation plates 

simple; leaves alternate, simple, scale-like or 

small, entire, exstipulate; inflorescences spikes, 

racemes, or the flowers solitary; flowers bisexual, 

rarely unisexual, the plants dioecious in Tamarix 

dioica, the parts hypogynous; sepals 4-5 (6), free 

or connate; petals 4-5 (6), free, sometimes with 

2 ligular processes; stamens 4-10 (to -75), 

mostly obdiplostemonous, the filaments filiform, 

free or basally connate, inserted on a disk; anthers small, slightly longer than wide, versatile; 

pollen 3(2-4)-colpate, simple or in tetrads; pistil 

1, the carpels 3 (2-5), the styles 3 (2-5), sometimes nearly absent, free or basally connate, the 

stigmas apical or decurrent ventrally; ovary unilocular, the ovules 2-many, parietal or basal, 

bitegmic, crassi- or f tenuinucellar, anatropous; 

fruit a loculicidal capsule; seeds usually with long 

hairs, sometimes winged; embryo straight, 0.7- 

0.8 the length of the endosperm, the cotyledons 

0.4-0.5 the length of the embryo; endosperm 

moderate or 0. 

Composition: 

Distribution: Chiefly temperate and subtropical, commonly steppe, desert, and shore plants, 

centered in the Mediterranean region and extending to eastern Asia; southern and southwestern Africa; Norway. 

FRANKENIACEAE (Figure 86d,e).-Herbs and 

small shrubs of saline habitats, the stems sometimes jointed; xylem vessel perforation plates 

simple; leaves opposite, simple, entire, exstipulate; inflorescences terminal and axillary di4 genera, - 100 species. 
174 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 86.-TAMARICACEAE: a, Tamarix articdata flowering twig, flower, 1,s. of same, essential 

organs, pistil, C.S. of ovary, dehiscent fruit, T. senegalensis seed; b, Reaumuria hypericoides 

flowering twig, 1,s. of flower, 1,s. of seed, Hololachne songarica flower, petal with ligule and 

stamens; c, Myricaria germanica flower, essential organs, floral diagram (after Marloth, 1925; 

Baillon, 1866- 1895; Niedenzu). FRANKENIACEAE: d, Frankenia pulverulenta flowering shoot, 

floral diagram, flower, I.s. of same, petal, essential organs; e, pistil, C.S. of ovary, dehiscent fruit, 

valve of fruit with seeds, seed, C.S. of same, embryo (after Le Maout and Decaisne, 1873). 
NUMBER 58 175 

chasia, cincinni, sometimes scorpioid, or the flowers solitary; flowers bisexual or rarely the plants 

polygamo-monoecious, the parts hypogynous; sepals 4-6 (7), connate, the lobes short, induplicate-valvate; petals 4-6 (7), free or rarely connate 

(Anthobryum), with a ventral ligular process; stamens 4-6 (3-24), in 2 series, the filaments elongate, free or shortly connate; anthers small, 

about as long as wide, didymous, dorsifixed; pollen 3(2-4)-colpate or 6-rugate; pistil l, the carpels 3 (2-4), the style 1, filiform, stigmas 3 (2- 

4), linear; ovary unilocular, the ovules l-numerous per placenta, parietal, bitegmic, thinly crassinucellar, anatropous, the funicle sometimes 

long; fruit a loculicidal capsule; embryo straight, 

axile-peripheral, 1.1 times the length of the endosperm; cotyledons 1-1.8 times the width of 

the radicle, 0.3-0.6 the length of the embryo; 

endosperm copious or moderate, starchy. 

Composition: 4 genera, -70 species. 

Distribution: Mainly saline habitats, centered 

in the Mediterranean region and southwestern 

Asia; southwestern United States to the western 

half of South America; northern and southern 

Africa; western and southeastern Australia; New 

Zealand. 

MYRSINACEAE (Figure 87a,b).-Small trees 

and shrubs, rarely subherbs, rarely climbing, 

with resin canals, streaks, or dots in most of the 

tissues, including the flowers; xylem vessel perforation plates usually only simple, some scalariform plates in a few species; leaves alternate, 

sometimes subopposite or subverticillate, simple, 

entire or toothed, usually coriaceous, exstipulate; 

inflorescences axillary or terminal racemes, panicles, umbels, fascicles or cymes; flowers rather 

small, actinomorphic, bisexual or rarely the 

plants dioecious; sepals 5-4 (3-6, very rarely 9), 

free or basally connate; petals 5-4 (3-6, very 

rarely 7), connate or rarely free (Embelia), the 

corolla rotate or salverform, the tube short; stamens 5-4 (3-6), opposite the corolla lobes, the 

filaments short, adnate to the base of the corolla, 

rarely nearly free, sometimes shortly connate 

(Aegiceras), often nectariferous; anthers as long 

as wide or longer, rarely connate (Amblyanthus), 

rarely transversely septate into locelli (Aegiceras, 

Ardisia sp.), dorsifixed or rarely basifixed, dehiscing longitudinally or by apical pores; pollen 

usually 3(2-5)-colpor(oid)ate, sometimes 4-rupate; pistil 1, the carpels 5-4 (3-6), the style 1, 

long, short or rarely absent, the stigma punctiform, capitate or 5-4(3-6)-lobed; ovary superior 

or rarely semi-inferior (Maesa), the ovules many 

to few, often embedded in the basal or freecentral placenta, the integuments 2, rarely 1 

(Aegiceras), tenuinucellar or rarely crassinucellar 

(Aegiceras), anatropous or amphitropous; fruit a 

berry or a drupe, or rarely longitudinally dehiscent by 1 slit (Aegiceras), the seeds 1 or several, 

small; embryo cylindrical, straight or arcuate, 

often transversely placed in the seed, 0.6-1 .O the 

length of the endosperm, the cotyledons 0.2-0.3 

the length of the embryo, or rarely elongated 

and connate into a tube (Aegiceras), about as wide 

as the radicle; endosperm copious or 0 (Aegiceras), fleshy and oily or horny and semitransparent. 

Composition: 35 genera, 1000 species. 

Distribution: Warm temperate to tropical 

America and Africa; India to Japan and southeastern Asia, the East Indies, Australia, and New 

Zealand; one genus is a mangrove. 

THEOPHRASTACEAE (Figure 87c,d).-Trees 

and shrubs without resin canals, streaks, or dots; 

xylem vessel perforation plates simple; leaves 

alternate, often subverticillate at the ends of the 

twigs or rarely opposite (Jacquinia sp.), simple, 

entire or toothed, exstipulate; inflorescences terminal or rarely axillary or cauliflorous, racemes, 

corymbs, fascicles, panicles, or rarely the flowers 

solitary, bracteolate; flowers bisexual or the 

plants dioecious, fairly large or small, actinomorphic; sepals 5 (4), free or shortly connate; 

petals 5 (4), connate, the tube usually short; 

stamens 5 (4), inserted on the base of the corolla 

opposite the lobes, the filaments free or basally 

connate (Clavija); staminodes 5 (4), sometimes 

fleshy-glandular or petaloid; anthers longer than 

wide, basifixed or slightly dorsifixed, the connective usually produced apically, dehiscing longitudinally; pollen 3-colporate, the colpi narrow; 
176 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 87.-MYRSINACEAE: a, Rapanea umbellata inflorescence, R. leuconeura flower, calyx, 

corolla laid open to show insertion of stamens opposite the petals, I.s. of fruit, seed; b, Aegiceras 

corniculatum bud, I.s. of flower, I.s. of anther, dehiscent fruit, seed (after Martius, 1840-1906; 

Baillon, 1866-1 895; Mez). THEOPHRASTACEAE: c, Clauzj'a spinosa bud, flower, corolla laid open 

to show insertion of stamens opposite the petals, and scales, views of stamen, pistil, 1,s. of same; 

d, Jacguinia aurantiaca 1,s. of flower, sepal, J. armillaris C.S. of fruit, views of seed, I.s. of seed 

(after Martius, 1840-1906). 

pistil 1, the carpels 5, the style 1, or rarely nearly 

absent, the stigma discoid or rarely conical or 

irregularly lobed; ovary superior, unilocular, the 

ovules numerous or rarely few, bitegmic, tenuinucellar, anatropous, semi-anatropous, or subcampylotropous, the placenta basal or free-central, immersed in mucilage; fruit a berry or rarely 

a drupe or rarely almost dry, the seeds many to 

1, often fairly large; embryo straight, 0.8 the 

length of the endosperm, the cotyledons 0.2-0.5 

the length of the embryo, twice as wide as the 

radicle; endosperm copious, horny. 

Composition: 

Distribution: Tropical to warm temperate 

America from Florida and the West Indies to 

Paraguay. 

5 genera, 1 10 species. 
NUMBER 58 177 

FIGURE ~~.-POLYGONACEAE: a, Polygonum meisnerianum calyx, flower, P. gilesii inflorescence, 

fruit, embryo, Coccoloba peltigera I.s. of flower; b, Oxyria digyna pistil, Atraphaxis spinosa essential 

organs, fruit enclosed by the accrescent calyx, I.s. of fruit, Rumex acetosella I.s. of fruit, Emax 

spinosa 1,s. and C.S. of fruit (after Hemsley; Martius 1840-1906; Dammer). PLANTAGINACEAE: 

c, Plantago brasiliensis inflorescence, P. major 1,s. of flower, P. brasiliensis floral diagram, flower 

and sepal, corolla and stamens, corolla laid open, stamen, corolla and ovary, P. major fruit, 

seed and 1,s. of same (after Martius, 1840-1906; Baillon, 1866-1895). 

POL YGONALES 

The order is monotypic. 

POLYCONACEAE (Figure 88a,b).-Herbs, 

shrubs, small trees, and climbers, the stems 

mostly jointed and with swollen nodes; xylem 

vessel perforation plates simple; anomalous structure as medullary vascular bundles, intraxylar or 

interfascicular phloem in a few species; leaves 

alternate, rarely opposite or whorled, simple, 

mostly entire, mostly with a membranous stipule 

(ochrea) sheathing the stem at the base of the 

petiole; inflorescences terminal and axillary 

cymes disposed in thyrses, racemes, spikes or 

heads; fowers bisexual or the plants polygamous, 

monoecious or dioecious, the parts hypogynous; 

sepals 3-6, free or connate, in 2 series when 4 

or 6, often colored, often accrescent and membranous, rarely fleshy in fruit; petals 0; stamens 

usually 6-9, rarely less (1-5) or more (12-50), 

mostly in 2 series, the filaments filiform, free or 

shortly connate, sometimes adnate to the perianth; anthers about as long as wide, dorsifixed, 

wall development of the Monocot type; pollen 

3-4-colporate, 4-ruporate, 6-rugorate, polyrugate, polyforate, 8-colpate, 3-colporoidate; disk 
178 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

annular, sometimes lobed, or central or glands 

at the base of the stamens; pistil 1, the carpels 3 

(2-4), the styles 3 (2-4), usually free, stigmas 

enlarged, apical, or sometimes decurrent along 

the ventral surface of the style; ovary unilocular, 

the ovule basal, orthotropous, 2( 1)-tegmic, crassinucellar, the funicle sometimes long; fruit a 

nutlet; embryo peripheral or axile, straight, arcuate, bent or folded, rarely convolute, 0.5-1.6 

times the length of the endosperm; cotyledons 

moderately thick, or thin, 1-3 times the width of 

the radicle, 0.3-0.8 the length of the embryo; 

endosperm copious or moderate, hard and semitransparent, or occasionally crystalline-granular, 

farinaceous, rarely soft to firm and whitish. 

Composition: -40 genera, -800 species. 

Distribution: Dry or wet, often disturbed, 

habitats; cosmopolitan, centered in temperate 

Northern Hemisphere, few tropical or subtropical. 

PLANTAGINALES 

The order is monotypic. 

PLANTAGINACEAE (Figure 88c).-Herbs and 

subshrubs; xylem vessel perforation plates simple; stem sometimes with cambial rows of cells in 

the cortex and/or pith, rarely forming medullary 

bundles; leaves radical, alternate or rarely opposite, simple, usually entire, exstipulate; inflorescences scapose spikes or heads or rarely the flowers solitary; flowers usually bisexual, sometimes 

also 0 or the plants monoecious or gynodioecious; 

sepals 4, connate; petals 4 (3), scarious, connate, 

the lobes imbricate; stamens 4 (1-3), the filaments filiform, hypogynous or inserted on the 

corolla; anthers about as long as wide, dorsifixed; 

pollen with (3) 4-1 4 foramina, occasionally pori, 

sometimes slightly colpoid or rugoid; pistil 1, the 

carpels usually 2, the style 1, filiform; stigma 

decurrent all around the style for 0.3 its length 

or punctiform; ovary superior, 2( 1 -4)-locular, 

the ovules 1-40 per locule, unitegmic, tenuinucellar, semi-anatropous to anatropous, axile or 

basal; fruit a pyxis or nutlet; seeds usually peltately attached to the placentas; embryo straight 

and axile or arcuate and peripheral (Bougueria 

and at least Plantago lanceolata), 0.9 the length 

of the endosperm; cotyledons 1-2 times the 

width of the radicle, 0.4-0.6 the length of the 

embryo; endosperm moderate or copious, firm, 

transparent and waxy in appearance. 

Composition: 3 genera, -260 species. 

Distribution: Various, often disturbed, habitats, few aquatic; pantemperate, especially in the 

Northern Hemisphere; in mountains in tropical 

regions. 

E UPHORBIALES 

Shrubs, trees, and herbs, sometimes laticiferous; xylem vessel perforation plates scalariform, 

simple or both; leaves alternate, rarely opposite, 

simple, entire or toothed, rarely lobed or palmately compound, extrafloral nectaries common, mostly stipulate; plants monoecious or dioecious; flowers anemophilous or entomophilous, 

the parts hypogynous, very rarely perigynous; 

sepals 5 (0, 2-6), free or connate; petals usually 

0, sometimes 5 (4-12), free or rarely connate; 

stamens 5 or 10 (1 to -300), the filaments free 

or connate in a column; anthers adnate, basifixed, more rarely dorsifixed, rarely connate; 

pollen 3 (2- 1 0)-colpate, colporate, rugorate, forate or nonaperturate; glandular disk often present; pistil 1, the carpels 3 (1-20), styles usually 

free and bifid, the stigmas apical or decurrent 

ventrally; ovary superior 3 (1-20)-locular, the 

ovules 1-2 per locule, axile; fruit septicidal and 

ventricidal, forming cocci and leaving a columella, rarely a drupe or nut; seeds often carunFIGURE 89.-EUPHORBIACEAE: a, RiCinUS communis 1.S. of d 

flower, stamen fascicle, views of anther, I.s. of 0 flower, C.S. 

of ovary, fruit, dehiscent coccus, seed, I.s. of same; 6, Mabea 

jstuligera 6 flower, 1,s. of same, 1,s. of 0 flower, 0 flower: c, 

Heuea discolor d flower, 1,s. of same, staminal column; d, 1.s. 

of 0 flower, pistil: e, Dalechampia stzpulacea views of anther, 

d flower and 1,s. of androecium, 0 floiver, stigma and C.S. of 

ovary, I.s. of 0 flower, seed, I.s. of same, dehiscent coccus 

(below), inflorescence;f, Hura crepitans stigmas, C.S. of ovary, 

6 inflorescence, staminal column and C.S. of same, 1,s. and 

C.S. of seed (after Martius, 1840-1 906). 
NUMBER 58 179 
180 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE ~~.-EUPHORBIACEAE: a, Acalypha amblyodonta stamen before dehiscence and 1,s. of 6 

flower, 3 0 flowers subtended by a bract,jatropha curcas 0 flower, I.s. of same, pistil, staminodes 

and glands; b, Croton comosus 6 flower, I.s. of same, P flower, 1,s. of same, C. migrans views of 

stamen; c, Actmostemon concolor 3 6 flowers, 0 flower, Euphorbia coecorum 1,s. of inflorescence, 

views of 6 flower; d, Hieronymia alchorneoides 1,s. of pistil, fruit, section of putamen, 6 flower 

and I.s. of same (after Martius, 1840-1906). 

culate, the embryo varied; endosperm copious or loids, e.g., aporphine, pyridine, indole, quinomoderate, rarely scanty or absent. line, and tropine. Mustard oil glycosides occur in 

Distribution: Cosmopolitan, predominantly Putranjiva (= Drypetes). 

tropical to warm temperate. EUPHORBIACEAE (Figure 89a-f, Figure 

Chemistry: Euphorbiales produce many alka- 9Oa-d).-Shrubs, trees, and herbs, sometimes 
NUMBER 58 181 

Iaticiferous; xylem vessel perforation plates scalariform, simple or both; leaves alternate or 

rarely opposite or whorled, simple or sometimes 

palmately compound, the margin entire or serrate, mostly stipulate; extra-floral nectaries are 

common; inflorescences axillary spike, raceme, 

thyrse, cymes, or solitary, occasionally cauliflorous; the plants mostly monoecious, sometimes 

dioecious; flowers anemophilous or entomophilous, the parts hypogynous, very rarely perigynous (Bridelia); sepals 5 or 0 (2-6); petals usually 

absent, sometimes 5 (8-12 in Garcia), free or 

rarely connate; stamens usually 5 or 10 (1 -300?), 

the filaments free or connate in a column, sometimes branched with several hundred anthers 

(Ricinus); anthers adnate, basifixed or more 

rarely dorsifixed, rarely connate, sometimes the 

thecas are separate; pollen 3(2-1O)-colpate, colporate, rugorate, forate, foraminoidate, or nonaperturate; disk often present in both sexes, annular or of separate glands; pistil 1, the carpels 

usually 3 (1-20); the styles free, usually bifid, 

sometimes united, the stigmas apical or decurrent ventrally; ovary 3( 1 -2O)-locular, the ovules 

1-2 per locule, axile, 2( 1)-tegmic, crassinucellar, 

anatropous, rarely orthotropous; fruit usually 

septicidal, dehiscent into 2-valved cocci open 

ventrally, and leaving a columella, sometimes a 

drupe; seeds often carunculate, rarely lobed 

(Picrodendron); embryo spatulate or rarely linear 

and terete, straight or rarely arcuate or folded, 

0.7-1 .O the length of the endosperm; cotyledons 

very thin to moderately thick, 1-5.3 (-7.0) times 

the width of the radicle, 0.5-0.9 the length of 

the embryo; endosperm copious or moderate, 

rarely scanty or 0, fleshy. 

Composition: -300 genera, -6000 species. 

Distribution: Cosmopolitan except for the 

arctic and cool temperate regions of the Northern Hemisphere; predominantly tropical to 

warm temperate. 

AEXTOXICACEAE (Figure 9 1 a).-Trees; xylem 

vessel perforation plates scalariform with many 

bars; leaves alternate or subopposite, simple, entire, the underside covered with scales, exstipulate; plants dioecious; inflorescences axillary racemes; flower parts hypogynous; sepals 5 (4-6), 

free, imbricate; petals 5 (4-6); stamens 5, free, 

alternating with large reniform disk-glands, the 

filaments thick; anthers about as long as wide; 

pollen 3-colporate; pistil 1, the carpels 2, the 

style 1, shortly bifid apically; ovary unilocular, 

the ovules 2, subapical, bitegmic, crassinucellar, 

anatropous; fruit a drupe or nut; embryo 0.5 the 

length of the endosperm; cotyledons ovate or 

orbicular, cordate, 4 times wider than the radicle, 

0.6 the length of the embryo; endosperm copious, ruminate, fleshy. 

Composition: 1 genus, 1 species. 

Distribution: Chile. 

PANDACEAE (Figure 9 1 b,c).-Trees; xylem 

vessel perforation plates simple and scalariform, 

with 6-10 (to 20 if Galearia is included) bars; 

leaves alternate, simple, distichous, toothed, stipulate; inflorescences elongated, cauliflorous in 

clusters of racemes (?), sometimes several flowers 

per inflorescence node (d), the plants dioecious; 

flowers actinomorphic; sepals 5, connate nearly 

to the apex; petals 5, free, imbricate to subvalvate; stamens free, 10 in 2 series of 2 lengths (5- 

15 if Galearia and Microdesmis are included), the 

filaments shorter than the petals, the anthers 

bilocular, about as long as wide, dehiscing longitudinally; pollen 3-colporate; disk small, intrastamina]; pistil 1, the carpels (2-) 3-4, the styles 

(stigmas) (2-) 3-4, 1 mm long, 1 mm wide; ovary 

superior, (2-) 3-4-locular, the ovules 1 per locule, bitegmic, orthotropous in Panda, obturator 

0, axile-apical; fruit a large drupe, the endocarp 

hard, thick, sculptured, containing 3 one-seeded 

chambers, dehiscing by valves at germination, 

the embryo 0.9 the length of the endosperm, the 

cotyledons flattened, thin, curved, truncate apically, cordate basally, 0.9 the length of the embryo, 7 times wider than the radicle, the latter 

slightly invested; endosperm copious, oily. 

Composition: 4 genera, 28 species. 

Distribution: Tropical western Africa, southeastern Asia, Malaysia. 

MA L VALES 

Trees, shrubs, and herbs, often with mucilage 

cells or canals and stellate hairs; xylem vessel 
182 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

a 

FIGURE 91 .-AEXTOXICACEAE: a, Aextoxicon punctatum calyx (bract ?) of 6 flower bursting 

irregularly, S bud with 3 outer sepals removed and 3 inner attached, bud from which the calyx 

has been removed showing the unexpanded petals, open flower from which the calyx has fallen, 

petal, stamen, androecium, 1 stamen and 5 nectaries surrounding the pistillode, pistil opened 

to show the ovules, fruit, embryo (after Baillon, 1866-1895; Hooker and Hooker, 1837-1982). 

PANDACEAE: b, Panda oleosa part of S inflorescence, bud, an outer and an inner stamen, 6 

flower, part of 0 inflorescence, calyx and pistil of 0 flower, I.s. and C.S. of ovary; c, fruit with 

part of the exocarp removed, putamen from beneath and leaf, C.S. of putamen showing the 3 

seeds, I.s. of seed, radial I.s. of seed (after Engler). 

perforation plates simple, rarely also a few scalariform; bast fibers often tough; leaves alternate 

or more rarely opposite, simple, rarely digitately 

compound, entire or toothed, stipulate; flowers 

mostly bisexual, the parts hypogynous; sepals 5 

(3, 4), usually connate and valvate, an epicalyx 

sometimes present; petals 5 (0, 3, 4), free or 

rarely connate; stamens usually numerous, to 

-80, rarely 4, 5, or 10, often connate in a tube 

closely surrounding the pistil and adnate to the 

base of the petals, rarely free or in a column or 

bundles; staminodes sometimes present; anthers 

rarely in bundles, dorsifixed, 2( 1)-locular, longitudinally dehiscent, rarely poricidal; pollen 

3(-8)-colporate, 3-colpate, 3-9-porate or forate, 

4-ruporate or 6-rugorate; glandular disk sometimes surrounding the base of the ovary; pistils 

1-5 (-65), the carpels 5 (1 to -20), styles 1 to as 

many, rarely twice as many, as carpels, free or 

connate, the stigmas apical or decurrent ven- 
NUMBER 58 183 

trally, rarely sessile; ovary superior 5( 1 to -20)- 

locular, the ovules l-many per locule, axile, very 

rarely parietal; fruit a loculicidal, septicidal, and 

sometimes ventricidal capsule, schizocarp, drupe, 

berry, samara or nut; seeds sometimes winged, 

covered with long hair or arillate; embryo large, 

straight, curved, or folded; endosperm copious 

to absent. 

Distribution: Cosmopolitan, especially pantropical, relatively few in temperate regions and 

absent from the arctic, in various habitats. 

Chemistry: Cyclopropenyl fatty acids are 

known from Tiliaceae, Malvaceae, Bombacaceae, and Sterculiaceae. They have not been 

reported outside the order. Gibbs says chemically 

Scytopetalaceae may not belong in the order. 

STERCULIACEAE (Figure 92a-e).-Trees and 

shrubs, rarely herbs or lianas; indument predominantly stellate, but other types also common; 

plants with mucilaginous receptacles; xylem vessel perforation plates simple; bast fibers tough; 

leaves alternate, rarely opposite (Lasiopetalum 

spp.), simple or digitately compound, entire, serrate or crenate, stipulate; inflorescences axillary 

and leaf-opposed, rarely terminal, sometimes 

cauliflorous, cymes and panicles, rarely the flowers solitary (Octolobus); flowers bisexual or the 

plants monoecious, the parts hypogynous (stamens slightly perigynous in some Thomasia); sepals 5 (3, 4), connate (free in Lysiosepalum), valvate; petals 5 or 0, free or adnate basally to the 

stamina1 tube, contorted; stamens 5 or 10 (-4- 

45), connate into a tube in bisexual flowers and 

column in 8 flowers, rarely free, often alternate 

with showy staminodes; anthers sometimes in 

bundles, bilocular, dorsifixed, longitudinally dehiscent, rarely poricidal; pollen 3-9-porate or 

forate, 3(-8)-colporate, or 3-colpate; pistils 1-5 

(-65 in Octolobus), carpels 5 (1-12), the styles 

free to united most of the way; ovary 5( 1-1 2)- 

locular, the ovules 2 or more per locule, axile, 

bitegmic, crassinucellar, anatropous; fruit a follicular capsule, sometimes spiny, schizocarp or 

samara, rarely baccate; seeds sometimes winged 

or covered with long hairs; embryo straight, 

curved or folded, 0.9 the length of the endosperm; cotyledons thin or thick, to 4.4 times 

wider than the radicle, 0.5-0.8 the length of the 

embryo, the radicle sometimes invested; endosperm copious, moderate or 0, fleshy. 

Composition: -65 genera, -800 species. 

Distribution: Pantropical, extending into subtropical regions, very few temperate. 

ELAEOCARPACEAE (Figure 92f;g).-Trees and 

shrubs with mucilage cells but cavities and canals 

absent; xylem vessel perforation plates simple, 

sometimes also a few scalariform; leaves alternate 

or opposite, simple, toothed or entire, stipulate; 

inflorescences racemes, panicles or dichasia; 

flowers bisexual or rarely unisexual, the parts 

hypogynous, rarely subperigynous; sepals 4-5, 

free or connate, valvate or imbricate; petals 4-5 

or 0, rarely connate, often apically fimbriate, 

valvate or imbricate, not contorted; stamens numerous (-25-80), free, the filaments elongate; 

anthers dorsifixed, bilocular, usually dehiscing 

by 2 pores or short slits, rarely by a transverse 

valve or at the base; pollen 3-colporate; disk 

usually surrounding the base of the ovary; pistil 

1, the carpels 2-5 (-7), the style 1 (0), punctate 

or shortly lobed, the stigmas apical, rarely sessile; 

ovary 2-5( 1 -7)-locular, the ovules 2-several per 

locule, axile, rarely subapical, bitegmic, crassinucellar, anatropous; fruit a septicidal or loculicidal 

capsule, sometimes densely echinate, or drupe or 

berry; embryo straight, 0.75-0.9 the length of 

the endosperm; cotyledons thin, 4 times wider 

than the radicle, 0.5-0.9 the length of the embryo; endosperm copious. 

Composition: 12 genera, -350 species. 

Distribution: Mainly tropical and subtropical; 

subtropical China to the East Indies, eastern Australia, New Zealand; Japan; West Indies; southern South America; absent from Africa. 

TILIACEAE (Figure 93a,b).-Trees and shrubs, 

rarely herbs, often with stellate indument, but 

other types also present; plants with mucilage 

cells, cavities or more rarely canals; xylem vessel 

perforation plates simple; bast fibers tough; 

leaves alternate, rarely opposite, simple, usually 

serrate, stipulate; inflorescences axillary, rarely 

leaf-opposed, cymose; flowers bisexual or rarely 
184 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

A 

f 

I 

FIGURE gL).-STERCULIACEAE: a, Sterculia striata I.s. of d flower, staminal column, 1,s. of 0 

flower, pistil and stamens, views of anther, C.S. of ovary, Brachychiton populneum dehiscent fruit 

(reduced); b, Theobroma cacao bud, I.s. of flower, petal, flower with perianth removed, staminalstaminodal tube laid open, views of anther and part of fruit opened to show the seeds; c, 

Melochia simplex flower, petal, staminal tube from brevistylous flower, pistil, essential organs 

from longistylous flower, fruit, coccus, views of seed, C.S. of same, M. graminijolia I.s. of seed; 

d, Hua paruifolia staminode (petal ?), part of flowering twig, bud, flower; e, essential organs, 

views of stamen, stamens and I.s. of pistil (after Martius, 1840-1906; Johnson; Lindley, 1853; 

Engler and Krause). ELAEOCARPACEAE:~, Sloanea garckeana flower, C.S. and I.s. of ovary; g, S. 

eichleri pistil, floral diagram, 1,s. of seed, S. garckeana dehisced fruit, S. alnijolia views of stamen, 

fruit, same opened to show seed (after Martius, 1840-1 906). 
NUMBER 58 185 

FIGURE 93.-TILIACEAE: a, Christians africana pistil and staminodes, stamen, 1.s. of ovary, 

dehiscent fruit, Triumfetta semitriloba I.s. of seed, T. rhomboidea floral diagram, fruit; b, Corchorus 

acutangulus bud, C. hirtus flower, C. acutangulus sepal, petal, C. hirtus views of anther, C. 

acutangulus pistil, C.S. of ovary (after Martius, 1840-1906). 

the plants monoecious or dioecious (Heliocarpus), 

the parts usually hypogynous; sepals 5 (3-4), free 

or basally connate, valvate, an epicalyx sometimes present; nectariferous tissue at base of sepals or petals; petals 5 (3, 4, 0), free, contorted; 

stamens usually numerous (-5-85), free or 

shortly connate, or in 5-10 bundles, the filaments elongate; anthers bilocular, about as long 

as wide, dorsifixed, dehiscing by longitudinal slits 

or rarely by apical pores; pollen usually 3(-4)- 

colporate, more rarely 4-ruporate, 6-rugorate, 

or oligoforate; pistil 1, the carpels 2-10, rarely 

nearly apocarpous (Glabraria, Christiuna), the 

style usually 1, apex divided or not, the stigma(s) 

capitate or lobed, rarely sessile; ovary superior, 

rarely inferior (Neotessmannia, Dicraspidia), 2- 

1 O-locular, the septa rarely incomplete in the 

upper part of the ovary, the ovules 2 (1 -several) 

per locule, axile, very rarely parietal, bitegmic, 

crassinucellar, anatropous or semi-anatropous; 

fruit a loculicidal, sometimes echinate capsule, 

schizocarp, nut or samara, rarely subbaccate; embryo straight, arcuate or folded, 0.9-1.7 times 

the length of the endosperm; cotyledons thin, 8 

times wider than the radicle, 0.5 the length of 

the embryo; endosperm usually copious, rarely 

0, fleshy. 

Composition: 50 genera, 450 species. 

Distribution: Pantropical, especially South 

America and southeastern Asia; few temperate. 

MALVACEAE (Figure 94a,b).-Herbs and 

shrubs, rarely trees, the indument usually stellate; plants with mucilage cells; xylem vessel perforation plates simple; bast fibers tough; leaves 

alternate, simple, serrate or crenate, sometimes 

palmately lobed, stipulate; inflorescences cymose, in cincinni, or the flowers solitary; flowers 

bisexual, or the plants rarely polygamous or dioecious (Napaea), the parts hypogynous; sepals 

5 (-3), connate, valvate, an epicalyx often present; petals 5, contorted or imbricate, usually 

asymmetrical, free from one another but usually 

adnate to the base of the stamina1 column; stamens numerous (to -60), very rarely 5 or 10, 

the filaments connate in a tube closely surrounding the pistil; anthers unilocular, dorsifixed, dehiscing longitudinally; pollen large, spiny, 3-4- 

colporate, polyrugorate or (oligo-) polyforate; 
186 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

a 

b 

d 

FIGURE 94.-MALvACEAE: a, Malua paroijora flower, petal, pistil, androecium, views of 

mericarp, seed; b, Abutilon megapotamicum views of anther, Pavoniaflauispina staminal tube and 

style branches, Hibiscus peterianus staminal tube and style branches, H. spathulatus flower, 2 

views of fruit valke and dehiscent fruit subtended by calyx and epicalyx (after Martius, 1840- 

1906). BOMBACACEAE: c, Bombax gracilipes floral diagram, flower and bud, views of petal, I.s. of 

ovary (enlarged); d, pistil, B. wittrockianum staminal tube and style, views of anther (after 

Martius, 1840-1 906). 

pistil 1, the carpels 5 (1 to -20), the styles usually 

free for a considerable distance, sometimes connate nearly to the apex, sometimes twice as many 

as the ovary locules, the stigmas apical or decurrent on the ventral surface of the style (Malva, 

KitaibeZia); ovary 5( 1 to -20)-locular, the ovules 

1-many per locule, axile, bitegmic, crassinucellar, anatropous or campylotropous; fruit a schiz- 
NUMBER 58 187 

ocarp or loculicidal capsule, rarely a berry (Malvaviscus); embryo folded or rarely arcuate (Malachra), 1.6-1.8 times longer than the endosperm; cotyledons 4-5 times wider than the radicle, 0.5-0.8 the length of the embryo; endosperm scanty or 0, fleshy or hard, sometimes oily. 

Composition: 

Distribution: Cosmopolitan, but absent from 

arctic Northern Hemisphere; centered in tropical South America. 

BOMBACACEAE (Figure 94c,d).-Trees, the indument stellate or lepidote; plants with mucilage 

cells and cavities; xylem vessel perforation plates 

simple; bast fibers tough; leaves alternate, simple 

or digitately compound, the margins entire or 

rarely toothed (Chorisia), stipulate; inflorescences axillary or terminal cymes, or the flowers 

solitary; flowers bisexual, the parts hypogynous, 

very rarely perigynous (Coelostegia); sepals 5, connate, valvate (slightly imbricate in Ochroma), an 

epicalyx sometimes present; petals 5 (0 in Cullenia), convolute, asymmetric; stamens often numerous, 5 to -100, free or connate into a tube 

closely surrounding the pistil, staminodes sometimes present; anthers 1-2 or more locular, dorsifixed, dehiscing longitudinally; pollen usually 

smooth, 3-porate, 3-colporate, 4-colpate; pistil 1, 

the carpels 2-5, the style 1, the apex lobed or 

capitate; ovary 2-5-locular, the ovules 2-many 

per locule, axile, bitegmic, crassinucellar, anatropous; fruit a loculicidal capsule or indehiscent 

and the seeds with a large aril, or a samara; 

embryo contorted or plicate, 1.7 times longer 

than the endosperm; cotyledons flat, thick, 0.5 

the length of the embryo; endosperm scanty, 

fleshy. 

-80 genera, -1 300 species. 

Composition: -25 genera, -200 species. 

Distribution: Pantropical, especially rain forest of South America, but also in dry habitats. 

R OSALES 

Shrubs, herbs, and trees, xylem vessel perforation plates simple, rarely also scalariform; 

leaves alternate, rarely opposite, simple or compound, usually toothed, sometimes dissected, 

lobed or entire, mostly stipulate; flowers bisexual, rarely unisexual, the parts hypogynous, perigynous or epigynous, an hypanthium sometimes 

present; sepals 5 (3-lo), free or basally connate, 

an epicalyx sometimes present; petals 5 (3-10, 

0), free or sometimes inserted on the calyx; stamens usually numerous in 1-4 or more series, to 

-400, rarely 1-5; staminodes very rarely present; filaments free, very rarely connate, very 

rarely inserted on the base of the petals, the 

anthers dorsifixed; pollen 3(2-4)-colporate, 

rarely 3(4)-pororate or unique; glandular tissue 

usually lines the receptacle between the stamens 

and the pistils; pistils numerous, to -1 50, to 1, 

sometimes spirally arranged, the carpels 5 (2-1 5) 

in the compound pistils, styles free or rarely 

connate, the stigmas apical, very rarely decurrent 

ventrally; ovary superior to inferior, 5(2-15)- 

locular, the ovules 2 (1-24) per locule, axile, very 

rarely apical; fruit follicles, achenes, drupe(s), 

pome, rarely a capsule, the seeds very rarely 

arillate; embryo large, spatulate or linear, 

straight, bent, arcuate or folded; endosperm 

mostly scanty or absent, rarely moderate. 

Distribution: Cosmopolitan, centered in temperate and subtropical Northern Hemisphere, 

mostly in open habitats. 

In contrast to Fabales, alkaloids 

and coumarins are remarkably rare in Rosaceae. 

Rosales and Theales, at least the primitive members, contain trihydroxylated phenolic substances such as ellagic acid, gallic acid, myricetin, 

and leucodelphenidin. These are absent from the 

“Polycarpicae.” Some alkaloids highly characteristic of “Polycarpicae,” e.g., protoaporphines, 

aporphines, berberin, and other quaternary 

bases, are absent from the Rosales and Theales. 

Ellagic acid occurs in Rosoideae and Myrtales 

and elsewhere. Rosaceae and Myrtaceae sometimes have the same alcohols; p-coumaroyl-quinic 

acid occurs in Malus and Eucalyptus. Nitrogen 

fixing plants include some Casuarinaceae, Myricaceae, Betulaceae, Rosaceae, Coriariaceae, 

Rhamnaceae, and Elaeagnaceae. The Crossosomataceae seem to be characterized by possession 

of syringin, which also occurs in Magnoliaceae. 

The Rosaceae and Fabales are highly cyanogenic 

groups. 

Chemistry: 
188 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

e 

I1 

FIGURE ~~.-ROSACEAE: a, Rubus imperialis 1,s. of flower, flower from below, Rosa pimpinellifolia 

I.s. of flower, floral diagram; b, Prunus sphaerocarpa I.s. of flower, floral diagram, views of 

stamen, fruit, C.S. and 1,s. of same, Rosa canina fruit, I.s. of same; c, Pyrus communis I.s. of flower, 

floral diagram, P. malus 1,s. of fruit; d, Quillaja saponaria flower, 1,s. of same; e, Kageneckia 

oblonga 1,s. of seed, dehiscent fruit;f, Couepia macrophylla flower with petals fallen, I.s. of same, 

Licania utilis 1,s. of flower (after Martius, 1840-1906; Baillon, 1866-1895). 

ROSACEAE (Figure 95a-f ).-Herbs, shrubs, 

and trees; xylem vessel perforation plates typically simple, but with sporadic multiperforate 

plates in some species, rarely also scalariform 

(Quillaja, Neillia, Polylepis); leaves pinnately compound or simple, alternate or very rarely opposite, mostly serrate, rarely entire, stipulate or 

rarely exstipulate; inflorescence terminal, race- 
NUMBER 58 189 

mose or cymose, of various types; flowers bisexual or the plants rarely dioecious, the parts hypogynous, perigynous or epigynous, an hypanthium often present; sepals 5 (3-lo), imbricate, 

free or basally connate, an epicalyx sometimes 

present; petals 5 (3-10) or 0, free, imbricate; 

stamens often in 1-4 series of 5 or 10 each, 

sometimes more numerous (to -400), rarely only 

1-4; filaments free or rarely connate (Chrysobalanus); anthers small, about as long as wide, dorsifixed; pollen usually 3-colporate; glandular tissue lines the receptacle between the pistils and 

stamens, sometimes as a disk; pistils numerous 

(to - 150) to 1, sometimes spirally arranged, the 

carpels 5 (2-15) in the compound pistils; styles 

free or rarely connate, the stigmas apical, rarely 

decurrent ventrally (Neviusia), not enlarged; 

ovary 5(2- 15)-locular when compound, the 

ovules 2 (1 -24) per locule, axile, superposed, 1- 

2-tegmic, crassinucellar, anatropous or semianatropous, the funicle short or none; fruit follicles, achenes, drupe(s), pome, rarely a capsule; 

embryo spatulate, large, straight or rarely slightly 

bent, linear or folded, sometimes the radicle 

invested; embryo as long as to 1.3 times the 

length of the endosperm; cotyledons moderately 

to markedly thick, rarely convolute, 1-6 times 

the width of the radicle, 0.6-0.9 the length of 

the embryo; endosperm mostly scanty or 0, 

rarely moderate, fleshy. 

Composition: 

Distribution: Cosmopolitan; centered in temperate and subtropical Northern Hemisphere; 

Chrysobalanoideae are pantropical in lowlands, 

especially of America. 

CORYNOCARPACEAE (Figure 96a,b).-Trees 

and shrubs; resin canals absent; xylem vessel perforation plates simple; leaves alternate, simple, 

entire, stipule intrapetiolar; inflorescences terminal and axillary panicles and racemes; flowers 

bisexual; sepals 5, free; petals 5, shortly adnate 

to the base of the sepals; stamens 5, the filaments 

elongated, inserted on the base of the petals and 

opposite them, alternating with as many petaloid 

staminodes; anthers about as long as wide; pollen 

grains bilateral, 2-colporate (?); 5 large disk 

glands inserted on the staminodes; pistil 1, the 

- 120 genera, -2500 species. 

carpels 2, one aborting, the styles 1 (2), stigma 

capitate; ovary superior, unilocular, the ovule 1, 

apical, bitegmic, the integuments short, crassinucellar, anatropous; fruit a drupe; embryo 

large, straight, the cotyledons thick, plano-convex, 0.9 the length of the embryo; endosperm 0. 

Composition: 1 genus, 5 species. 

Distribution: Northeastern Australia, northern New Zealand, New Guinea, New Hebrides, 

New Caledonia. 

CROSSOSOMATACEAE (Figure 96c).--Shrubs; 

xylem vessel perforation plates simple; leaves 

alternate, simple, entire, glaucous, exstipulate; 

flowers solitary, terminal, deliciously fragrant, 

bisexual, the parts perigynous; sepals 5 at the 

edge of an hypanthium; petals 5, free; stamens 

15-50, in 3-4 series, the filaments filiform, free; 

anthers slightly more than twice as long as wide, 

the filament inserted basally between dorsal and 

ventral halves, which are separate basally; pollen 

(2-)3-colporate; disk adnate to the hypanthium; 

pistils 3-5 (6), the style short, stigma apical, capitate; ovules several, biseriate, ventral, bitegmic, 

crassinucellar, campylotropous or amphitropous; 

fruits follicles; seeds with a multifid aril; embryo 

arcuate, the cotyledons thin, twice as wide as the 

radicle, 0.6 the length of the embryo; endosperm 

moderate, about twice the volume of the embryo, 

fleshy. 

Composition: 1 genus, 4 species. 

Distribution: Dry habitats; southwestern 

United States and adjacent Mexico. 

NEURADACEAE (Figure 96d,e).-Annual herbs, 

often woolly, the hairs not stellate; xylem vessel 

perforation plates simple; leaves alternate, simple, variously lobed or pinnatifid, exstipulate or 

the stipules minute; flowers solitary, axillary, bisexual, the parts perigynous; calyx tube broad 

and flat, the lobes 5, k valvate, sometimes accrescent, an epicalyx sometimes present; petals 

5, separate, inserted on the calyx, in drying turning from yellow to blue-black like some Malvaceae (e.g., Althaeafic$olia); stamens 10, in 2 series, the filaments subulate, free; anthers small, 

bilocular, oblong, dorsifixed; pollen a unique 

type, 3-armed demicolpi at each pole, and a pore 

associated with each arm, sometimes 3-colpate, 
190 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

f 

FIGURE 96.-CORYNOCARPACEAE: a, Corynocarpus laevigata flower, 1.s. of same, flower laid 

open, views of stamen, part of inflorescence and smaller part of same enlarged; b, petal with 

stamen, views of staminode, I.s. of ovary, fruit, seed surrounded by endocarp, embryo (after 

Cheeseman; Hooker and Hooker, 1837-1 982). CROSSOSOMATACEAE: c, Crossosoma californicum 

flowering twig, lateral view of flower, seed enveloped in aril and divested of aril, views of 

stamen, pistil, calyx tube and base of stamens, dehiscent fruit with persistent calyx and stamens 

(after Hemsley). NEURADACEAE: d, Neurada procumbens flower, 1,s. of same, C.S. of ovary, views 

of stamen; e, fruit, 1.s. of same showing the persistent styles, embryo (after Baillon, 1866-1895; 

Hooker and Hooker, 1837-1906). CORIARIACEAE:~, Coriaria japonica 6 flower, 0 flower, pistil, 

1,s. of ovary showing ovules, fruitlet, embryo; g, C. myrtfolia fruit, floral diagram, flower with 

calyx removed, C.S. of fruitlet, seed, 1,s. of same (after Hooker, 1837-1982; Le Maout and 

Decaisne, 1873). 
NUMBER 58 191 

syncolpate; disk sometimes present; pistil 1, the 

carpels 3-10 in one series, connate at the ovary 

level, adnate to the calyx tube; styles 3-10, short, 

sometimes spinescent, persistent, the stigmas capitate; ovary 3-lO-locular, the ovules 1-2 per 

locule, axile, bitegmic, crassinucellar, anatropous; fruit capsular, the sometimes spinose 

locules dehiscing ventrally; embryo bent, the 

cotyledons flattened, auriculate, rather thin, 2- 

4 times wider than the radicle, 0.5 the length of 

the embryo; endosperm 0. 

Composition: 3 genera, 10 species. 

Distribution: Deserts; eastern Mediterranean 

region to India; southern Africa. 

CORIARIACEAE (Figure 96f,g).-Shrubs and 

small trees, rarely herbs, the buds sometimes 

perulate; xylem vessel perforation plates simple: 

leaves opposite, rarely also alternate, or verticillate, simple, entire, major veins extending from 

the base to the apex, the stipules very caducous; 

inflorescences axillary or terminal racemes, or 

the flower rarely solitary; flowers bisexual or the 

plants monoecious or dioecious, the parts hypogynous; sepals 5 (6), free, imbricate, persistent; 

petals 5 (6) usually smaller than the sepals, usually 

with a ventral keel; stamens 10 (12) in 2 series, 

the filaments filiform, free or those opposite the 

petals adnate to the keel; anthers large, oblongish, about twice as long as wide, sometimes 

apiculate; pollen 3-(2-4)-colporate or 3(4)-pororate; pistils 5- 10 (- 12), the styles elongate, 

papillose, and stigmatic all around for much of 

their length: ovule 1, ventral and apical, bitegmic, crassinucellar, anatropous; fruitlets achenes, usually enclosed by the accrescent petals 

that become fleshy; embryo straight, the cotyledons thick, plano-convex, 3 times wider than the 

radicle, 0.8 the length of the embryo, the transition between them and the radicle is gradual; 

endosperm scanty or 0. 

Composition: 1 genus, -8 species. 

Distribution: Disjunct; Mexico to Peru, Chile; 

western Mediterranean region, Himalayas to 

China; Japan to the Philippines; New Guinea to 

New Zealand. 

M YR TALES 

Trees and shrubs, sometimes climbing, more 

rarely herbs; xylem vessel perforation plates simple, rarely also scalariform with a few bars, very 

rarely exclusively scalariform with 1 5-60 bars; 

intraxylary phloem usually present, rarely also 

interxylary, and vascular tissue in the cortex or 

pith; tissues sometimes with secretory cavities 

containing oily substances; leaves opposite, more 

rarely alternate, simple, usually entire, rarely 

toothed, lobed or dissected, rarely stipulate; flowers bisexual, rarely unisexual, the parts perigynous or epigynous; sepals 4-5 (0, 2-1 4, 16), free 

or connate, imbricate or valvate; petals 4-5 

(0, 2-14), free or rarely connate, usually inserted 

on the calyx tube; stamens usually numerous, 1 

to -400, sometimes 4-10, in l-several series; 

staminodes rarely present: filaments free or connate, rarely in bundles, usually inserted on the 

calyx tube, more rarely external to a disk or 

epigynous, very rarely adnate to the petals or 

style; anthers dorsifixed, basifixed or adnate, 

sometimes didymous, dehiscence longitudinal or 

by pores, the connective sometimes produced; 

pollen 3(2-4)-colporate or -colpate, rarely 3-porate, sometimes also with 3-5 pseudocolpi; glandular disk sometimes lining the calyx tube or 

epigynous, often intrastaminal, more rarely absent; pistil 1, the carpels 2-5 (10) -22), styles 1 

(3-5), usually elongate, the stigma(s) usually apical, capitate, punctiform or lobed, rarely sessile 

and radiate; ovary usually inferior or semi-inferior, more rarely superior, 2-5( 1-22)-locular, 

the ovules usually numerous, rarely as few as 1- 

4 per locule, the placentas axile, rarely apical, 

free-central, parietal or basal; fruit a loculicidal, 

more rarely septicidal or circumscissile capsule, 

berry, drupe, nut, sometimes 3-4-winged, the 

seeds many, few, or 1; embryo large, straight, 

arcuate, folded or convolute, the cotyledons terete, flattened or reduced; endosperm absent or 

scanty, rarely copious. 

Distribution: Cosmopolitan, mainly tropical 

and subtropical, but also common in temperate 

regions, in a variety of habitats. 
192 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Chemistry: Myrtaceae have many terpenoids, 

largely in the essential oils. Alkaloids are almost 

unknown in the family, and cyanogenesis is rare, 

perhaps very rare. Derivatives of ellagic acid are 

widespread in Myrtales. Ellagic acid occurs in 

Rosoideae, Myrtales, and elsewhere. Rosaceae 

and Myrtaceae sometimes have the same alcohols. Myrtaceae have phenolic esters and ethers 

like Magnoliales. By and large the similarities 

indicate a close relation between Lythraceae and 

Myrtaceae, but in contrast to the latter, the former lack leucoanthocyanins. Lythraceae have dmannitol, a character shared only by Punicaceae. 

Heteropyxis stands out from Myrtaceae. 

MYRTACEAE (Figure 97a,b).-Trees and 

shrubs, rarely subscandent or subherbaceous; xylem vessel perforation plates usually simple, 

Myrceugenia and Myrtus communis exclusively scalariform with 15-25 bars, also scalariform in 

young Eugenia stem; tissue with secretory cavities 

usually containing oily substances; vascular bundles bicollateral, intraxylary phloem universally 

present; leaves simple, entire, very rarely crenulate, opposite, more rarely alternate, glandularpunctate, coriaceous, exstipulate; inflorescences 

panicles, racemes or dichasial cymes, the flowers 

rarely solitary; flowers bisexual or rarely the 

plants polygamous, monoecious or dioecious, the 

parts perigynous or epigynous; sepals 4-5 (0, 3), 

free or basally connate, imbricate, valvate, or 

sometimes calyptrate; petals 4-5 (0, 6), often 

circular, free or calyptrate; stamens inserted at 

the margin of a disk, usually numerous (to -400), 

rarely only 4-8 and either opposite or alternate 

with the petals when in one series, the filaments 

filiform, free or connate basally or in bundles 

opposite the petals; staminodes rarely present; 

anthers usually small, about as long as wide to 5 

times longer than wide, the connective often with 

an apical gland, very rarely apiculate (Acrandra), 

dorsifixed or rarely basifixed (Calothamnus), dehiscing by longitudinal slits, rarely by apical 

pores; pollen 3(2-4)-colporate; disk lining the 

calyx-tube or epigynous; pistil 1, the carpels 2-3 

(1 (?)-16), the style 1, elongate, stigma apical; 

ovary usually inferior, sometimes semi-inferior 

or superior, the locules 1-3 (-16), the ovules 

mostly numerous, rarely few or 1 per locule, 

axile or rarely parietal, 2( 1 )-tegmic, crassinucellar, usually anatropous, some campylotropous or 

semi-anatropous; fruit a loculicidal capsule, 

berry, drupe or nut, the seeds few, very rarely 

many; embryo straight, arcuate or folded, the 

radicle sometimes invested; cotyledons linear, 

foliaceous and flat or variously folded or thick 

and plano-convex, 1-1.8 times the width of the 

radicle, 0.2-0.8 the length of the embryo; endosperm 0 or scant. 

Composition: 100 genera, 3000 species. 

Distribution: Mostly tropical and subtropical, 

but also numerous in temperate Australia; centered in America and Australia; temperate to 

tropical Asia, East Indies; New Zealand; southern 

2/3 of Africa, Madagascar; Mediterranean region. 

LECYTHIDACEAE (Figure 97c-e).-Trees and 

shrubs; xylem vessel perforation plates simple, 

rarely occasionally scalariform in 2 genera; tissue 

without secretory cavities; intraxylary phloem 

absent, but vascular bundles present in the cortex; leaves simple, alternate, entire, not glanddotted but sometimes with large glands on the 

margins, exstipulate or very rarely stipulate; inflorescences racemes or corymbs or the flowers 

solitary, sometimes cauliflorous; flowers bisexual, 

the parts perigynous or epigynous; calyx 4-6 (2- 

3)-lobed, the lobes valvate or slightly imbricate; 

petals 4-6 (8), free or rarely basally connate; 

stamens very numerous (-60-300) in several 

series, either monadelphous or diadelphous, 

often forming a hood over the pistil, part often 

FIGURE 97.-bfYRTACEAE: a, Eugenia aurata flowering twig, 

bud, flower, young fruit, E. brasiliensis 1,s. of flower; b, 

Marlierea estrellensis Is. of bud, Eugenia brasiliensis flower 

with petals and stamens removed, views of stamen, style, C.S. 

of ovary, floral diagram, E. retusus petal (after Martius, 

1 840- 1906). LECYTHIDACEAE: c, Cariniana estrellensis 

flower, bud, petal, views of stamina1 tube, same laid open; 

d, views of stamen and pistil, C.S. and 1,s. of ovary, Gustavia 

augusta floral diagram, 1,s. of ovary, stigma (enlarged); e, 

Couratari rufescens dehiscent fruit, operculum, Couroupita 

surinamensis C.S. of ovary, seed, same enlarged and partly 

opened to sho\v embryo, embryo, same laid open (after 

Martius, 1840-1 906). 

194 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

staminodial; anthers small, about as long as wide, 

or linear, basifixed or dorsifixed sometimes 

didymous, dehiscing by slits or pores; pollen 3- 

colpate, 3-colporoidate, 3-colporate, never syncolpate; disk intrastaminal, and also associated 

with the stamens and calyx; pistil 1, the carpels 

2-6 (7), style 1, rarely 3-4-lobed, short, the 

stigma(s) apical; ovary inferior or semi-inferior, 

2-6(7)-locular, the ovules l-many per locule, 

axile, bitegmic, tenuinucellar, anatropous, the 

funicle often conspicuous; fruit a woody capsule, 

apically dehiscent, or a berry; seeds large; embryo folded or undifferentiated, the cotyledons 

foliaceous or conferruminate, 4-5.5 times wider 

than the radicle, 0.5-0.9 the length of the embryo, the radicle sometimes invested and sometimes much thickened; endosperm 0. 

Composition: 15 genera, 325 species. 

Distribution: Tropical; centered in rain forest 

of South America; Central America to the northern % of South America. 

BARRINGTONIACEAE (Figure 98a).-Trees, 

rarely shrubs; xylem vessel perforation plates 

simple, rarely with an occasional scalariform 

plate; vascular bundles in the cortex; leaves simple, alternate, entire or toothed, exstipulate; inflorescence axillary or terminal raceme or corymb; flowers bisexual, the parts epigynous; calyx 

4 (5) (2-3)-lobed, imbricate; petals 4(5) or 0 

(Foetida), free; stamens numerous (- 100-300), 

the filaments filiform, long-exserted, basally connate, rarely free; anthers small, about as long as 

wide, basifixed; pollen 3-colpate (syncolpate), 3- 

colporoidate; disk intrastaminal; pistil 1, the carpels 2-4, the style 1, filiform, often very long, 

stigma apical; ovary inferior, rarely semi-inferior, 

2-4-locular, the ovules 1 -several per locule, axile; fruit a fibrous berry, drupe or dry, 4-winged 

and indehiscent; embryo entire or divided, 0.9 

the length of the endosperm; cotyledons 1.5 

times wider than the radicle, 0.5 the length of 

the embryo; endosperm copious. 

Composition: 5 genera, 50 species. 

Distribution: Palaeotropics; tropical and 

southern Africa, Madagascar, Hainan, Taiwan, 

Ryu-Kyu, northern India, Indochina, northeastern Australia. 

ASTERANTHACEAE (Figure 98b,c).-Trees and 

shrubs; xylem vessel perforation plates simple, 

rarely an occasional scalariform plate in Asteranthos; vascular bundles in the cortex; leaves simple, 

alternate, entire, exstipulate; flowers axillary or 

cauliflorous, solitary or rarely in a corymb, bisexual; calyx lobes 3 or 5 (1 0- 12 in Asteranthos), 

imbricate or valvate; petals 0; stamens usually 

numerous (-10-200), in 4 or more whorls, the 

outermost whorl or all but the inner staminodial 

and connate into a plicate corona; filaments 

sometimes filiform, free or basally connate; anthers basifixed, 2- l-locular; pollen 3-colpate, 3- 

colporate (not syncolpate); disk intrastaminal; pistil 1, the carpels 3 or 5 (5-8 in Asteranthos, 5- 

20(?) in Napoleonaea); ovary 3 or 5 

(-2O?)-locular, semi-inferior or inferior, the 

ovules several per locule, axile; style elongate, 

filiform, and the stigma small, or short, flattened 

apically, and the stigmas 5; fruit a large berry or 

loculicidal capsule (Asteranthos); embryo large, 

the cotyledons reniform, plano-convex, the 

radicle very small, endosperm 00). 

Composition: 3 genera, 20 species. 

Distribution: Northern Brazil; tropical west 

Africa. 

DIALYPETALANTHACEAE (Figure 98d,e).- 

Tree with oil glands in all parts; xylem vessel 

perforation plates simple(?); leaves simple, opposite, entire, stipulate; inflorescence a terminal 

thyrse; flowers bisexual, sweet-scented (like Jasminum), the parts epigynous; calyx 4-lobed, the 

lobes imbricate, rounded; petals 4, in 2 whorls, 

free; stamens 18 (16-25), in 2 whorls, the filaments short, connate about half way, free from 

the corolla; anthers elongate, the thecas separate 

basally, adnate to the connective; pollen 3-colporate; disk fimbriate, surrounding the base of 

the ovary; pistil 1, the carpels 2, the style 1, 

stigma apical, very shortly bifid; ovary bilocular, 

the ovules numerous, axile; fruit a loculicidal and 

septicidal capsule, partly extending above the 

persistent calyx; seeds numerous, fusiform; embryo straight, terete, the cotyledons short, radicle 

small, thick; endosperm scanty, oily. 

Composition: 1 genus, 1 species. 

Distribution: Tropical eastern Brazil. 
NUMBER 58 195 

b 

FIGURE 98.-BARRINGTONIACEAE: a, Barringtonza racemosa 1.s. of flower, ovary opened to show 

the ovules (after Baillon, 1866- 1895). ASTERANTHACEAE: b, Asteranthos brasiliensis bud in axil 

of leaf, C.S. of ovary; c, flowers, pistil and calyx, stigma from above, I.s. of flower, stamen, views 

of anther (after Martius, 1840- 1906). DIALYPETALANTHACEAE: d, Dialypetalanthus fuscescens 

flowering twigs, base of leaves showing stipules, views of stamen and C.S. of same (above), pistil; 

e, petal, fruit, dehiscent fruit, seeds (much enlarged) (after Kuhlmann). 

SONNERATIACEAE (Figure 99a,b).-Trees; xylem vessel perforation plates simple; intraxylary 

phloem present; leaves opposite, simple, entire, 

coriaceous, exstipulate; inflorescences axillary or 

terminal, the flowers solitary or in 3-flowered 

cymes or corymbs; flowers bisexual or unisexual, 

the parts perigynous; hypanthium campanulate, 

thick and coriaceous, the calyx lobes 4-8, valvate, 
Q 

NVINOSHLINS SNOILnSIKLN03 0.L ANVLOS 
NUMBER 58 197 

persistent; petals 4-8 or 0, free; stamens usually 

numerous 12 to - 175, in several series, inserted 

with the calyx lobes and petals at the apex of the 

hypanthium, the filaments filiform, free; anthers 

small, reniform, dorsifixed; pollen 3-porate or 3- 

colporate; pistil 1, the carpels 4-20, the style 1, 

elongate, the stigma capitate; ovary 4-20-locular. 

free or adnate basally to the hypanthium, the 

ovules numerous, axile, bitegmic, crassinucellar, 

anatropous; fruit a loculicidal capsule or berry, 

the seeds numerous; embryo arcuate, the cotyledons short, foliaceous; endosperm 0. 

Composition: 2 genera, 7 species. 

Distribution: Tropical; in a mangrove or in 

monsoon forest; east Africa, Madagascar; India 

to Indochina, the East Indies, other western Pacific islands, northern Australia. 

PUNICACEAE (Figure 99c,d).-Trees and 

shrubs; xylem vessel perforation plates simple; 

intraxylary phloem present; leaves mostly opposite, simple, entire, not glandular, exstipulate; 

inflorescences terminal and axillary, the flowers 

solitary or in a fascicle, bisexual; hypanthium 

extending above the ovary, thick, the calyx lobes 

5-8, valvate, persistent; petals 5-8, free, crumpled in bud; stamens numerous (-75-125), in 

several series, inserted with the calyx lobes and 

petals at the apex of the hypanthium, the filaments filiform, free; anthers small, about as long 

as wide, dorsifixed; pollen 3(4)-colporate; pistil 

1, the carpels 9, the style 1, elongate, stigma 

capitate; ovary inferior or semi-inferior, the 

locules 9 in 2 superposed series, the placentas 

thick, axile in one series, parietal in the other, 

the ovules numerous, bitegmic, crassinucellar, 

anatropous, with a long funicle; fruit a berry, the 

seeds numerous, outer testa fleshy; cotyledons 

convolute, much broader than the radicle, 0.8 

the length of the embryo, the radicle partly invested; endosperm 0. 

Composition: 1 genus, 2 species. 

Distribution: Southeastern Europe to the Himalayas; Socotra. 

RHIZOPHORACEAE (Figure 1 OOa-f).-Trees 

and shrubs; xylem vessel perforation plates scalariform with 1-62 bars, simple, or both; intraxylary phloem absent; leaves usually opposite, 

sometimes alternate or whorled, simple, usually 

entire, sometimes toothed, often coriaceous, 

sometimes glandular-punctate, the stipules interpetiolar, more rarely absent; inflorescences axillary cymes, racemes, or spikes, rarely the flowers 

solitary; flowers bisexual or rarely the plants 

polygamo-dioecious (Sterigmapetalum) or polygamo-monoecious (Anisophyllea), actinomorphic; 

sepals 3-8 (-1 4), valvate, often fleshy, persistent, 

sometimes the calyx-tube extending above the 

ovary; petals free, perigynous, 3-8 (-14) or 0 

(Pellacalyx), often notched, laciniate or markedly 

hairy on the ventral surface, convolute or inflexed in bud; stamens 8 (-40), mostly in one 

whorl, the filaments short or elongate, free or 

rarely connate basally (Kandelia), inserted on the 

outer edge of a disk, rarely the epipetalous ones 

adnate to the petals (Anisophyllea spp.) or rarely 

adnate to the calyx-tube (Pellacalyx); numerous 

staminodes rarely present; anthers elongate or 

about as long as wide; 4(2 to w)-locular; pollen 

3(4)-colporoidate-~olporate (sometimes zonate); 

disk perigynous or epigynous, lobed, rarely absent; pistil 1, the carpels 2-5 (6), the style 1 (3- 

5), the stigma(s) apical, rarely sessile and 5-6- 

radiate (Sterigmapetalum); ovary 2-5( 1 - 12)-locular, sometimes incompletely so, mostly inferior, 

rarely semi-inferior or superior (Cassipourea), the 

ovules bitegmic, crassinucellar, anatropous or 

semi-anatropous, 2 (or more) per locule, axile, 

sometimes subapical, or the placenta free-central 

in the unilocular ovaries; fruit usually indehiscent, a berry or drupe, or rarely dry and rather 

hard-shelled (Anisophyllea), rarely a septicidal 

capsule, sometimes 3-4-winged; seeds 1 or 1 per 

locule, sometimes arillate, rarely winged, often 

viviparous; embryo small or large, straight, 

curved or folded; cotyledons 2 (3-4), terete or 

flat, sometimes connate, usually small, the radicle 

usually long; endosperm copious, fleshy, or 0. 

Composition: 16 genera, 120 species. 

Distribution: Pantropical; sometimes a mangrove. 

LYTHRACEAE (Figure 10 1 a-e).-Trees, 

shrubs, and herbs; xylem vessel perforation 
198 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

C 

I 

FIGURE 1 ~~.--RHIZOPHORACEAE: a, Rhizophora mangle floral diagram, bud, flower, views of 

petal, 1,s. of fruit with germinated seed; b, views of anther, I.s. of base of flower, fruit and calyx, 

part of embryo; c, Cassipourea guianensis part of a flowering twig, flower without petals, 1,s. of 

same, views of anther, C.S. of ovary, young fruit with calyx and stamens, views of seed; d, petal, 

views of same (one with stamen); e, Anisophyllea guudichaudiana part of floivering twig and leaf, 

glomerules of d and 1 P flower, 6 flower and same with perianth removed, views of stamen;f, 

0 flower, same with perianth lobes removed, I.s. of 0 flower, fruit (after Martius, 1840-1906; 

Oliver). 
NUMBER 58 199 

C 

n 

FIGURE 101 .-LYTHRACEAE: a, Heimia salicijolia flower from above, views of anther, lateral 

view of flower without petals and anthers, part of calyx laid open to show insertion of filaments, 

pistil; b, two C.S. of ovary, dehiscent fruit seen from above and laterally; c, Lythrum hyssopifolia 

1,s. of seed, views of embryo; d, L. maritimum two I.s. of ovary, dehiscent fruit, Lafoensia 

vandelliana flower, 1,s. of base of same, Cuphea longzjlora calyx laid open to show insertion of 

stamens and petals; e, Heteropyxis canescens flowering twig, part of leaf margin, flower, petal, 

flower laid open, the petals removed, Is. of ovary, dehiscent fruit enclosed in calyx (after 

Martius, 1840-1906; Oliver). 
200 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

plates simple; intraxylary phloem present, interxylary phloem rarely recorded; leaves simple, 

entire, opposite or verticillate, rarely alternate, 

stipules minute or absent; inflorescences panicles, 

racemes, dichasia or the flowers solitary; flowers 

bisexual, the parts perigynous, the pistil and stamens usually trimorphic; calyx-tube often long, 

the lobes short, 4 or 6 (8 or IS), valvate, often 

with appendages between the lobes; petals 4, 6 

(8 or 0), free, crumpled in bud, inserted at the 

apex of the calyx-tube; stamens 4-1 2 (1 to -200), 

in 1-2 series, the filaments filiform, often inserted very low on the calyx-tube, below the 

petals; anthers small, about as long as wide, dorsifixed, rarely basifixed (Crenea); pollen 3-colporate, sometimes also with 3 or 6 pseudocolpi, or 

3-porate; hypogynous glands sometimes present; 

pistil 1, the carpels 2-6, the style 1, stigma apical, 

usually discoid or capitate, rarely bifid (Adenaria); ovary superior, often surrounded by an 

annular disk, (1-) 2-6-locular, sometimes incompletely so at the apex of the ovary, the ovules 

numerous, rarely to 2 per locule, bitegmic, crassinucellar, anatropous, axile, or rarely on 1 parietal placenta, or basal, or free-central; fruit a 

capsule, dehiscence circumscissile, valvate, or irregular, or rarely indehiscent (PepZis); seeds usually numerous, small; embryo straight, the cotyledons 1.5 times wider than the radicle, 0.45- 

0.6 the length of the embryo, sometimes investing the radicle; endosperm 0 or rarely scanty. 

Composition: -25 genera, -500 species. 

Distribution: Often in wet habitats; mainly 

tropical, but also temperate; temperate North 

America to temperate South America; temperate 

to tropical Eurasia, the East Indies, Australia, 

and New Zealand; most of Africa, Madagascar. 

CRYPTERONIACEAE (Figure 102 a, b).-Trees; 

xylem vessel perforation plates simple; intraxylary phloem present; leaves opposite, simple, entire, coriaceous, exstipulate; inflorescences axillary racemes or panicles; flowers bisexual or the 

plants polygamo-dioecious; calyx tubular, the 

lobes 4-5, valvate; petals 0; stamens 4-5, alternate with the calyx lobes, the filaments filiform, 

inserted at the apex of the calyx-tube; anthers 

small, as long as wide, the thecas separate, adnate 

to the connective; pollen 2-colporate or colpate(?); disk 0; pistil 1, the carpels 2, the style 1, 

elongate, stigma apical, bilobed; ovary superior 

2-locular, the ovules numerous, axile; fruit a 

loculicidal capsule, the valves connected by the 

persistent base of the style; seeds numerous, minute; embryo terete, straight; cotyledons as 

broad as the radicle, 0.5-0.6 the length of the 

embryo; endosperm 0. 

Composition: 2 genera, 5 species. 

Distribution: Assam, southeast Asia to the 

East Indies and Philippines. 

OLINIACEAE (Figure 1 OPc,d).-Trees and 

shrubs; xylem vessel perforation plates simple; 

intraxylary phloem present; leaves opposite, simple, entire, the stipules minute; inflorescences 

terminal cymes; flowers bisexual; calyx tubular, 

the lobes 4-5, deciduous; petals 5 (4), free, inserted at the apex of the calyx-tube, alternate 

with 5 colored scales; stamens 4-5, the filaments 

very short, inserted near the apex of the calyxtube; anthers small, about as long as wide, didymous, the connective apically produced; pollen 

3-colporate, sometimes also with 3 pseudocolpi; 

disk 0; pistil 1, the carpels 3-5, the style 1, 

elongate, stigma apical, clavate or capitate; ovary 

inferior, 3-5-locular, the ovules 2-3 per locule, 

axile, bitegmic, crassinucellar, semi-anatropous; 

fruit a drupe, the flesh thin or rarely dry, the 

pyrenes 1-5; seeds minute, the embryo convolute, one cotyledon longer than the other, the 

radicle short; endosperm 0. 

Composition: 1 genus, 10 species. 

Distribution: Tropical eastern and southern 

Africa; St. Helena. 

MELASTOMATACEAE (Figure 103a-g).- 

Shrubs, herbs, and trees, rarely climbing or epiphytic, xylem vessel perforation plates simple; 

intraxylary phloem universal, sometimes phloem 

interxylary and in the pith; leaves opposite or 

verticillate, simple, entire or rarely toothed, 

mostly with 3-9 longitudinal arcuate, transversely anastomosed veins, penninerved in some 

Memecyleae, oil glands only in Microlicia, exstipulate; inflorescences terminal, axillary or at 
NUMBER 58 20 1 

a 

b W 

FIGURE 102.-CRYPTERONIACEAE: a, Crypteronia Eeptostachya 6 flower, Is. of same, C. jauanica 

bisexual flower, 1,s. of same; b, C. paniculata flowering twig, flower laid open, views of anther, 

pistil, C.S. of ovary, fruit (after Baillon, 1866-1895; Hutchinson, 1973). OLINIACEAE: c, Olinia 

usambarensis flowering twig, flower, I.s. of upper part of same, views of stamen, pistil, 1,s. and 

C.S. of ovary; d, 0. capensis infructescence, putamen, 0. cymosa flower, C.S. of ovary, Is. of 

flower, floral diagram with 3 bracts (after Gilg; Marloth, 1925; and Link, Klotzsch and Otto). 

nodes of fallen leaves, dichasial cymes, corymbiform to paniculate, or the flowers solitary; flowers bisexual or the plants rarely dioecious; calyx 

tubular or campanulate, often colored, the lobes 

4-5 (3-lo), imbricate, rarely valvate or calyptrate; petals 4-5 (3-lo), free, inserted at the apex 

of the calyx-tube, usually convolute; stamens 8 

or 10 (3-96) in 2 whorls, rarely one whorl staminodial or absent, the filaments elongate, inserted 

at the apex of the calyx-tube; anthers usually 

elongate, frequently curved, often inflexed in 

bud, basifixed, dehiscence by pore(s), rarely by 

short slits or completely longitudinal, the connective often appendaged at or near the base of 

the anther; pollen 3-colporoid plus 3 pseudocolpi; collar sometimes present around the base 

of the style, nectar rarely present; pistil 1, the 

carpels 4-5 (2-22), the style 1, elongate, stigma 
202 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

C 

FIGURE 103.-MELASTOMATACEAE: a, Lavoisiera pectinata flower, 1,s. of same, C.S. of ovary, L. 

serrulata C.S. of ovary, L. pectinata 3 views of anther of smaller stamen, 3 views of anther of 

larger stamen; b, Rhynchanthera laxa flower, 1,s. of same, ovary, C.S. of same; c, Miconia 

ligustroides fruit, 3 views of seed; d, Marcetia gardneri fruit and part of calyx, dehiscent fruit; e, 

Clidemiafrancavillana flower, 1,s. of ovary and calyx, stamen, C.S. of ovary;f, Mouriri regeliana 

bud, flower, I.s. of ovary and calyx; g, views of stamen, C.S. of ovary (2 locules), M. doriana C.S. 

of ovary (1 locule) (after Martius, 1840-1 906). 
NUMBER 58 203 

apical, often punctiform, sometimes capitate, 

rarely with as many very short lobes as carpels; 

ovary superior, semi-inferior or inferior, 4-5 

(1 -22)-locular, the ovules numerous, rarely few 

(2-3 per locule in some Memecyleae), axile, 

rarely basal, or basal-parietal (Kibessia), or 6-1 2 

ovules on a free-central placenta in a unilocular 

ovary (some Memecyleae), bitegmic, crassinucellar, anatropous, sometimes becoming campylotropous; fruit a loculicidal capsule or berry; seeds 

numerous (few in Memecyleae), often minute, 

the embryo usually arcuate, sometimes straight; 

cotyledons 0.2-0.4 the length of the embryo, or 

in Memecyleae large, plano-convex, often conferruminate, or foliose and convolute, the radicle 

short; endosperm 0. 

Composition: 200 genera, -4500 species. 

Distribution: Usually in moist habitats; mainly 

tropical and subtropical; centered in South 

America; temperate United States to warm temperate South America; Japan, southern Asia, the 

East Indies, other Pacific islands, northern Australia; central ?/< of Africa. 

ONAGRACEAE (Figure 104 a,b).-Herbs, rarely 

shrubs and trees; xylem vessel perforation plates 

simple, rarely also a few multiperforate plates; 

intraxylary phloem present, interxylary in a few 

genera; leaves opposite or alternate, rarely 

whorled, simple, entire, rarely pinnatifid or the 

margin sinuate or toothed, rarely stipulate; inflorescences axillary, the flowers often solitary, or 

in spikes, racemes, rarely panicles; flowers bisexual or rarely the plants polygamous; calyx 

tube often produced beyond the ovary, often 

colored, the lobes 4 (2-6), valvate; petals 4 (0, 

2-6), free, usually convolute, or imbricate; stamens 4-8 (1 - 12), usually obdiplostemonous, 

diplostemonous or haplostemonous, rarely 1 

staminodial (Lopezia), the filaments elongate, inserted at the apex of the calyx tube or epigynous, 

rarely adnate to the style; anthers oblong, dorsifixed, dehiscing longitudinally; pollen 3(2-4)-colpate, -colporate, or -perorate; glandular disk 

often surrounding base of style; pistil 1, the carpels 4 (2-6), the style 1, stigmas 4 (1-6), capitate, 

notched or radiating branches; ovary 4 (2-6)- 

Iocular, sometimes incompletely so, inferior, 

rarely semi-inferior, the ovules 1-many per 

locule, axile, bitegmic, crassinucellar, anatropous; fruit a loculicidal capsule, sometimes apically dehiscent, rarely a berry or nutlet; seeds 

mostly numerous, but may be only 1; embryo 

straight or nearly so; cotyledons 0.7-0.9 the 

length of the embryo, grading gradually into the 

radicle; endosperm 0. 

Composition: 20 genera, -650 species. 

Distribution: Open dry to wet habitats; cosmopolitan except in the larger deserts; mainly 

temperate and subtropical, centered in western 

United States and Mexico. 

TRAPACEAE (Figure 104c,d).-Floating herbs; 

xylem vessel perforation plates simple; leaves 

pinnatisect and opposite when submerged, rhomboid, dentate and with an inflated petiole and in 

a rosette, and some alternate, when floating, 

stipulate or exstipulate; flowers axillary, solitary, 

bisexual, the parts perigynous; sepals 4, basally 

connate, valvate, persistent; petals 4, free, inserted at the base of the disk; stamens 4, the 

filaments subulate, inserted at the same level as 

the petals; anthers small, dorsifixed, longitudinally dehiscent; pollen 3-colpate; disk cupular, 

surrounding the base of the ovary; pistil 1, the 

carpels 2, the style 1, elongate, stigma apical, 

capitate; ovary semi-inferior, bilocular, the 

ovules 1 per locule, axile-subapical, bitegmic, 

crassinucellar, anatropous; fruit turbinate, a drupaceous nut, often with 2-4 spines, I-seeded; 

seed large, the cotyledons very unequal, one 

thick and much larger than the other, 4-6 times 

wider than the radicle, 0.8 the length of the 

embryo; endosperm 0. 

Composition: 1 genus, -3 species. 

Distribution: Temperate to tropical Eurasia 

and Africa. 

COMBRETACEAE (Figure 105 a,b).-Trees and 

shrubs, often climbing; xylem vessel perforation 

plates simple; intraxylary and interxylary phloem 

are common; leaves alternate, opposite or rarely 

verticillate, simple, entire, exstipulate; inflorescences spikes, racemes or panicles; flowers usually bisexual; calyx-tube extending above the 
204 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 104.-oNAGRACEAE: a, Ludwzgia anastornosans bud, flower, views of stamen, style with 

disk, flower with petals and stamens removed, C.S. of ovary, 1,s. of pistil and calyx, fruit, 1,s. of 

same; b, Oenothera mollissirnu flower, 1,s. of upper part of same, 1,s. and C.S. of ovary, dehiscent 

fruit, embryo, 1,s. of seed and 3 views of same (after Martius, 1840-1906). TRAPACEAE: c, 

Trapa natans 1,s. of flower, same without the corolla, floral diagram, views of stamen, style and 

stigma; d, petal, fruit, seed and embryo (after Baillon, 1866-1895; Le Maout and Decaisne, 

1873). 
NUMBER 58 205 

\ 

/' 

C 

FIGURE 10 

d 

, 

-COMBRETACEAE: a, Combretum leprosum bud, flower, I.s. of same, petal and views 

of anther, fruit, C.S. of same; b, LaguncuEaria racemosa bud, flower without the petals, I.s. of 

same, part of calyx laid open to show insertion of the stamens and petal, style and epigynous 

disk, 1,s. of fruit showing seed, C.S. of fruit and floral diagram, embryo, I.s. of same (after 

Martius, 1840-1906). PENAEACEAE: c, Penaea myrtfolia flowering twig, flower, 1s. of same, 

pistil, floral diagram, dehiscent fruit, opened seed, embryo; d, P. mucronata views of stamen, 

Endonema retzioides flower, 1,s. of ovary, views of stamen (after Baillon, 1866-1895; Gilg). 

c 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

ovary, the lobes 4-5 (6-8), valvate, rarely accrescent, wing-like (Getonia); petals 4-5 (6-8) or 0, 

small, inserted on the calyx-tube external to the 

disk; stamens 4-1 0 (-20), mostly diplostemonous, the filaments very long, filiform, inserted 

on the calyx-tube or external to the disk; anthers 

small, didymous, about as long as wide, dorsifixed, dehiscing by longitudinal slits; pollen 3- 

colporate, sometimes also with 3 pseudocolpi; 

disk epigynous; pistil 1, carpel 1 (reduced from 

2-3), the style 1, elongate, stigma(s) apical, capitate or punctiform, 4-lobed in Laguncularia; 

ovary inferior, rarely semi-inferior, unilocular, 

the ovules 2 (-6), apical, bitegmic, crassinucellar, 

anatropous; fruit often samaroid, winged at the 

angles, sometimes drupaceous or nut-like, rarely 

dehiscent at apex, 1-seeded; embryo straight, the 

cotyledons foliaceous, coiled or thick, plano-convex, folded longitudinally, or crumpled, 0.9 the 

length of the embryo; endosperm 0. 

Composition: 20 genera, -550 species. 

Distribution: Often in savanna, rarely a mangrove; pantropical, especially Africa, few subtropical. 

PENAE.KEAE (Figure 105c,d).-Small shrubs 

of ericoid habit; xylem vessel perforation plates 

simple; intraxylary phloem present; leaves opposite, decussate, simple, entire, coriaceous, the 

stipules minute or 0; flowers axillary, solitary or 

paired, often crowded, bisexual, actinomorphic, 

the bracts often colored; sepals 4, connate, the 

calyx tubular, persistent, colored, the lobes 4, 

valvate in bud; petals 0; stamens 4, the filaments 

very short, inserted near (at) the apex of the 

calyx-tube; anthers about as long as wide, basifixed, dehiscing longitudinally; pollen 3(-5)-colporate and with 3-5 pseudocolpi; disk 0; pistil 1, 

the carpels 4, the style 1, elongate, stigma capitate or 4-lobed; ovary superior, 4-locular, the 

ovules 2-4 per locule, axile or basal, bitegmic, 

crassinucellar, anatropous, the embryo sac 16- 

nucleate (Penaea type); fruit a loculicidal capsule; 

seeds often 1 per locule; embryo thick, the cotyledons 0.3 the width of the radicle, 0.1-0.4 the 

length of the embryo; endosperm 0. 

Composition: 5 genera, 25 species. 

Distribution: Cape Province, South Africa. 

S A XIFR A GALES 

Herbs or shrubs, more rarely small trees, 

sometimes succulent; xylem vessel perforation 

plates scalariform with few or many bars, simple, 

or both; leaves alternate or opposite, simple or 

compound, entire, toothed, rarely lobed, very 

rarely ascidia, stipulate or exstipulate; flowers 

bisexual, rarely unisexual, the parts hypogynous, 

perigynous or epigynous; sepals 4-5 (3-30), free 

or connate, rarely colored; petals 4-5 (3-30, 0), 

free or rarely connate; stamens usually in 1-2 

whorls of 4-5 each, more rarely numerous or 

fewer (2, 15, 30, 40, 50 to -200), usually free, 

rarely basally connate or adnate to the petals, 

sometimes inserted on the calyx tube; rarely a 

whorl of staminodes present; anthers usually dorsifixed, less commonly basifixed, rarely adnate, 

sometimes didymous, rarely produced apically; 

pollen usually 3(- 1 1)-colporate, 3-colporoidate, 

rarely 3-colpate or 2- 4- 8- or 9-pored; glandular 

disk sometimes present; pistils 1, rarely 2-6, very 

rarely to 30, the carpels 2-5(-7), styles 1-5(-7), 

often 2, usually free, the stigmas apical, rarely 

sessile or decurrent ventrally; ovary superior to 

inferior, 1 -5(-7)-locular, the ovules usually numerous, rarely few to 1 per locule, axile, parietal, 

ventral, rarely apical or basal; fruit a variously 

dehiscent capsule, follicles, more rarely a berry, 

drupe, nutlet or samara; seeds usually numerous, 

more rarely few, minute; embryo often linear, 

minute, rarely large and broad; endosperm 

copious, less frequently moderate, scanty or absent. 

Distribution: Cosmopolitan, in various habitats. 

Chemistry: Gibbs mentioned that Sedoheptulose, a heptose, occurs in Crassulaceae, Saxifragaceae and Penthorum, emphasizing the close relation of Crassulaceae and Saxifragaceae. But it 

also occurs in various other families. 

SAXIFRAGACEAE (Figure 1 OGa).-Herbs, 

rarely fleshy; xylem vessel perforation plates simple; leaves simple or compound, alternate or 

radical, rarely opposite, exstipulate; inflorescences of various kinds, both racemose and cymose, the flowers rarely solitary; flowers usually 
NUMBER 58 207 

FIGURE 106.-sAXIFRAGACEAE: a, Saxzjraga crassqoh 1.s. of flower, floral diagram, s. tridactylites dehiscent fruit, seed and I.s. of same, S. granulata I.s. of flower, S. irrigua I.s. of flower, 

Rodgersia podophylla floral diagram, Astilbe thunbergii fruit with residual floral parts, seed 

(enlarged) (after Baillon, 1866- 1899; Engler). CRASSULACEAE: b, Sedum acre flowering 

branchlet, petal and 2 stamens, pistils and hypogynous scales, flower, floral diagram, 1,s. of 

flower, fruit; c, Crassula magnolii seed, I.s. of same, Kalanchoe brasiliensis flower, corolla laid 

open, views of stamen, pistils and hypogynous scales, I.s. of pistil, dehiscent fruitlet, floral 

diagram: d, Penthorum sedoides flowering branchlet, flower, dehiscent fruit (after Le Maout and 

Decaisne 1873; Martius, 1840-1906; Baillon, 1866-1895: Britton and Brown). 

bisexual, the plants rarely dioecious (Astilbe), the 

parts hypogynous, perigynous, or epigynous; sepals 5 (4-7), free; petals 5 (4-7) or 0, free or 

rarely connate; stamens 5- 10 (3 in Tolmiea), free, 

the filaments filiform; anthers small, about as 

long as wide; pollen usually 3-colporoidate; disk 

present; pistil(s) 1 (-5), carpels 2 (-5), the styles 

usually free, stigmas apical; ovary 1 -2(-5)-locu- 
208 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

lar, the ovules numerous, axile, ventral when the 

carpels are separate, bitegmic or rarely unitegmic, crassinucellar, anatropous; fruit a septicidal? capsule, or rarely follicular; seeds numerous, minute; embryo linear, terete, straight, 0.3- 

0.7 the length of the endosperm; cotyledons 1- 

2 times as wide as the radicle, 0.1-0.3 the length 

of the embryo; endosperm copious, fleshy. 

Composition: 30 genera, 600 species. 

Distribution: Many alpine and arctic, in dry 

and moist habitats; almost cosmopolitan; centered in the north temperate zone, few in the 

Southern Hemisphere. 

CRASSULACEAE (Figure 1 06b-d).-Herbs or 

shrubs, usually succulent; xylem vessel perforation plates simple (scalariform in Penthorum); 

leaves fleshy, alternate, rarely opposite or 

whorled, sometimes in a rosette, simple, rarely 

compound, entire or crenate, exstipulate; inflorescences usually cymose, cincinni; flowers bisexual or rarely the plants dioecious, the parts 

hypogynous to perigynous; sepals 4-5 (3-30), 

free or connate; petals 4-5 (3-30, 0 in Penthorum), free or connate; stamens 4-10 (-60), usually twice as many as the petals, sometimes as 

many, the filaments filiform, free or rarely basally connate, often epipetalous when the petals 

are connate; anthers slightly longer than wide; 

pollen usually 3-colporate; intrastaminal nectariferous scales usually present; pistils 4-5 (3-30), 

usually as many as petals, sometimes connate 

basally (Penthorum, Diamorpha), the styles short 

or elongate, the stigmas apical; ovary superior, 

the ovules numerous, rarely few, ventral, bitegmic, crassinucellar, anatropous; fruits follicles, rarely a capsule (Penthorum, Diamorpha); 

seeds minute; embryo straight, 0.8 the length of 

the endosperm, the cotyledons plano-convex, 3 

times wider than the radicle, 0.5-0.9 the length 

of the embryo; endosperm scanty or rarely 0, 

fleshy. 

Composition: 

Distribution: 

-35 genera, -1 500 species. 

Mostly in warm dry regions, but 

also in cool moist ones, and at least one species 

aquatic; cosmopolitan; centered in South Africa. 

PARNASSIACEAE (Figure 107a,b).-Herbs; xylem vessel perforation plates simple; leaves 

mostly or all radical, alternate, entire, exstipulate; flowers solitary, bisexual, the parts hypogynous or perigynous; sepals 5, shortly connate 

basally; petals 5, free, marcescent, entire or fimbriate; stamens 5, alternating with 5 well-developing often multifid staminodes with nectar-secreting base; anthers longer than wide, dorsifixed, the apex acute; pollen 3-colporate; pistil 1, 

the carpels 3-4, the style 1, very short, stigmas 

3-4, apical; ovary superior or semi-inferior, unilocular, the ovules numerous, parietal, bitegmic, 

tenuinucellar, anatropous; fruit a loculicidal capsule, the seeds minute; embryo linear, straight, 

terete, as long as the endosperm; cotyledons 1 .O1.3 times the width of the radicle, 0.1-0.3 the 

length of the embryo; endosperm scanty or 0. 

Composition: 1 genus, 50 species. 

Distribution: Chiefly in bogs; temperate 

Northern Hemisphere. 

EREMOSYNACEAE (Figure 1 07c,d).-Annual 

herb; leaves radical and cauline, entire to pinnately lobed, alternate, exstipulate; inflorescence 

terminal dichotomous cymules; flowers bisexual, 

the parts hypogynous or slightly perigynous; sepals 3, nearly free, valvate; petals 5, free; stamens 

5, free, the filaments subulate; anthers dorsifixed, small, about as long as wide; pollen 3- 

colporate (-colporoidate); pistil 1, the carpels 2, 

the styles 2, filiform, divergent, stigmas apical; 

ovary bilocular, the ovules 1 per locule, axile 

near the base of the locule; fruit a capsule, subdidymous, loculicidally dehiscent; embryo apparently oblong; endosperm moderate to copious. 

Composition: 1 genus, 1 species. 

Distribution: Southwestern Australia. 

FRANCOACEAE (Figure 107e).-Perennial 

herbs; xylem vessel perforation plates simple; 

leaves alternate, subbasal, pinnatifid or roundedcordate, exstipulate; inflorescences terminal racemes or panicles; flowers bisexual, the parts 

hypogynous; sepals 4 (5), nearly free, persistent, 

valvate; petals 4 (5), 2 of which are sometimes 

smaller than the others or absent; stamens 8, 

obdiplostemonous or 4 (5) and alternating with 

glands (staminodes?), the filaments free; anthers 
NUMBER 58 209 

11 b 

FIGURE 107.-pARNASSIACEAE: a, Parnassia palustris flowering plant, flower, pistil and calyx, 

glanduliferous scale, seed, 1,s. of same without the testa, dehiscent fruit; b, floral diagram, P. 

faberi flower, pistil, C.S. of ovary, views of stamen, staminodes (after Baillon, 1866-1895; Le 

Maout and Decaisne, 1873; Oliver). EREMOSYNACEAE: c, Eremosyne pectinata plant in flower, 

bud, flower; d, I.s. ofovary, views ofstamen (after Hutchinson, 1973). FRANCOACEAE: e, Francoa 

sonchijloolia flower, I.s. of same, floral diagram, dehiscent fruit, F. appendiculata flower with 

perianth removed showing stamens and staminodes, F. ramosa seed and 1,s. of same, F. 

appendiculata plant in flower (after Baillon, 1866-1 895; Engler). 
210 

oblong, subcordate or didymous; pollen 3-colporate; disk present; pistil 1, the carpels 4 (2), 

the stigmas 4, sessile, commissural; ovary 4(2)- 

locular, the ovules numerous per locule, axile, 

bitegmic, crassinucellar, anatropous; fruit an 

elongate capsule, dehiscence apically septicidal 

and slightly loculicidal; seeds very small, striolate, 

the embryo minute, 0.2-0.3 the length of the 

endosperm; cotyledons not broadened(?), 0.5 the 

length of the embryo; endosperm copious, fleshy. 

Composition: 2 genera, 2 species. 

Distribution: Temperate Chile. 

DAVIDSONIACEAE (Figure 108a,b).-Small 

tree, the indument irritant; xylem vessel perforation plates simple and scalariform; leaves very 

large, alternate, im paripinnatel y compound, the 

leaflets dentate, part of the rachis alate, stipules 

large; inflorescence axillary or supra-axillary simple or sometimes compound spike; flowers bisexual, the parts hypogynous; sepals 4-5, connate 

half way; petals 0; stamens 10 and nectariferous 

scales 10, the filaments If: tumid below, inserted 

in a disk; anthers oblong, dorsifixed; pollen 3- 

colporoidate; pistil 1, the carpels 2, styles 2, free, 

stigmas minute, terminal; ovary bilocular, the 

ovules several per locule, axile; fruit a large 

drupe with 2 leathery pyrenes; seeds 2, large, the 

embryo large, straight, ovate; cotyledons planoconvex, the radicle very short, endosperm 0. 

Composition: 1 genus, 1 species. 

Distribution: Tropical and temperate eastern 

Australia. 

HYDRANGEACEAE (Figure 108c,d).-Herbs, 

shrubs, or small trees, rarely climbers; xylem 

vessel perforation plates typically scalariform 

with numerous bars, rarely simple; leaves alternate or opposite, simple, dentate, exstipulate; 

inflorescences cymes, corymbs or heads; flowers 

bisexual, sometimes the outer ones sterile and 

the calyx lobes large and petaloid, sometimes the 

plants polygamo-dioecious; sepals 5 (4-1 0) connate or more rarely free; petals 5 (4-lo), free or 

rarely calyptrate (Pileostegia); stamens 30, 15, 10, 

or 8, the filaments filiform; anthers basi- or dorsifixed, linear to short and didymous; pollen 3- 

colporate; disk 0; pistil 1, the carpels 2-6, the 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

styles 2-6, free or connate basally or nearly to 

the apex, stigmas apical; ovary semi-inferior to 

inferior, 2-6-locular, sometimes incompletely so, 

the ovules numerous, unitegmic, tenuinucellar, 

anatropous; placentas axile or intrusive parietal; 

fruit a capsule dehiscing loculicidally or apically 

between the styles, or rarely a berry; seeds minute, the embryo linear, straight, 0.4-0.9 the 

length of the endosperm; cotyledons not broadened, 0.2-0.3 the length of the embryo; endosperm moderate, fleshy. 

Composition: 

Distribution: Mainly north temperate and 

subtropical; United States, Mexico; Andes to 

southern Chile; Japan, China, the Himalayas, few 

in southeastern Asia, Java, and Sumatra. 

PHILADELPHACEAE (Figure 108e,f).--Shrubs 

and small trees, the indument mostly stellate; 

xylem vessel perforation plates scalariform, simple or both; leaves opposite or verticillate, rarely 

alternate, simple, serrate or entire, exstipulate; 

inflorescences terminal racemes, cymes or heads, 

rarely the flowers solitary; flowers bisexual; sepals 5-4, connate; petals 7-4, free; stamens usually numerous 200-4, the filaments sometimes 

flattened and toothed toward the apex, free or 

connate basally; anthers small, about as long as 

wide; pollen 3-colpor(oid)ate, longicolpate; pistil 

1, the carpels 3-7, styles 3-7, free or rarely 

united into 1; ovary 7-l-locular, superior to inferior, the ovules numerous, rarely 1 per locule 

(Whipplea), axile, rarely parietal or apical, unitegmic, tenuinucellar, anatropous; fruit a septicidal and loculicidal capsule, rarely a berry; seeds 

minute; embryo linear or subspatulate, straight, 

0.9 the length of the endosperm; cotyledons 

oblong, somewhat flattened, 1-2 times the width 

of the radicle, 0.2-0.5 the length of the embryo; 

endosperm moderate, fleshy. 

10 genera, 1 15 species. 

Composition: 7 genera, 135 species. 

Distribution: Western North America to Central America; eastern Asia to the Himalayas and 

Caucasus, Philippines; southern Europe. 

PTEROSTEMONACEAE (Figure 1 OSb).-Shrubs; 

leaves alternate, simple, serrate, stipulate; inflorescences terminal, few-flowered corymb-like 
NUMBER 58 21 I 

FIGURE 1 ~~.-DAVIDSONIACEAE: a, Davidsonia pruriens inflorescence, part of young inflorescence enlarged, flower, same with stamens removed, pistil, views of anther; b, 1,s. and C.S. of 

fruit, ‘2 views of seed, embryo, cotyledon (after von Mueller). HYDRANGEACEAE: c, Hydrangea 

virens inflorescence, flower, H. hortensis fruit, seed, 1,s. of same; d, H. arborescens bud, floral 

diagram, C.S. of ovary, Is. of seed (after Lindley, 1853; Le Maout and Decaisne, 1873). 

PHILADELPHACEAE: e, Philadelphus coronarius flowering twig, flower, 1,s. of same, floral diagram, 

dehiscent fruit, seed, 1,s. of same; f, Deutria scabra flower, floral diagram, I.s. of flower, 

Decumaria barbara bud, 1,s. of flower, C.S. of ovary (after Baillon, 1866-1895; Le Maout and 

Decaisne, 1873). 
212 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

cymes; flowers bisexual; calyx lobes 5, valvate, 

persistent; petals 5, free, persistent; stamens obdiplostemonous, 5 functional and 5 similar but 

lacking anthers, the filaments flattened, toothed 

near the apex; anthers about twice as long as 

wide, dorsifixed, the connective produced apically; pollen 3-colporate; disk absent; pistil 1, the 

carpels 5, style 1, stigmas 5, apical; ovary inferior, 

5-locular, the ovules 6-4 per locule, axile; fruit 

a capsule septicidally dehiscent apically; seeds 1 - 

few, the embryo long; cotyledons ellipsoid; endosperm moderate, fleshy. 

Composition: 1 genus, 2 species. 

Distribution: Mexico. 

ITEACEAE (Figure 1 OSa).-Shrubs and small 

trees; xylem vessel perforation plates scalariform 

with up to 40-70 bars; leaves alternate, simple, 

dentate or spinose, stipulate; inflorescences terminal or axillary, racemiform or cymes; flowers 

bisexual or the plants polygamous, the parts perigynous; sepals 3, free, valvate or open; petals 

5, free, valvate; stamens 5, the filaments subulate; 

anthers dorsifixed; pollen 2-porate, 2(3)- 

por(oroid)ate; disk annular; pistil 1, the carpels 

2, style 1, elongated, stigmas apical; ovary semiinferior, bilocular, the ovules few to many, axile; 

fruit a septicidal capsule; seeds minute, oblong 

or scobiform; embryo straight, terete, linear, 0.9 

the length of the endosperm; cotyledons not 

broadened, 0.2 the length of the embryo; endosperm moderate, fleshy. 

Composition: 2 genera, 17 species. 

Distribution: Tropical and subtropical eastern 

Asia from Japan to the Himalayas; western Malaysia; temperate eastern North America. 

BAUERACEAE (Figure 109c).--Shrubs of wet 

ground; xylem vessel perforation plates simple, 

very rarely scalariform with less than 10 bars; 

leaves opposite, compound, trifoliate, the leaflets 

serrate, exstipulate; flowers solitary, axillary, bisexual, the parts perigynous; sepals 6-8 (4-1 0), 

free or very shortly connate; petals 6-8 (4-lo), 

free; stamens 4-10 (to -40), in 1-2 series, the 

filaments free, filiform; anthers small, about as 

long as wide, didymous, dehiscing by short apical 

slits; pollen 2-colpate(?), 4 pores; disk thin; pistil 

1, the carpels 2, styles 2, free, filiform, recurved, 

stigmas apical; ovary superior or semi-inferior, 

bilocular, the ovules 2-several per locule, axile, 

bitegmic, crassinucellar, anatropous; fruit a subdidymous, compressed, truncate capsule; dehiscence apical and loculicidal; seeds 1 -several; cotyledons not broadened, 0.3 the length of the 

embryo; endosperm copious, fleshy. 

Composition: 1 genus, 3 species. 

Distribution: Wet habitats; temperate eastern 

Australia. 

BRUNIACEAE (Figure 1 OSd).--Shrubs, often 

ericoid; xylem vessel perforation plates scalariform, with numerous bars; leaves small, simple, 

entire, alternate, usually imbricate, mostly exstipulate; inflorescence a terminal spike, raceme, 

head, or the flowers solitary; flowers bisexual; 

sepals 4-5, connate or free; petals 4-5, usually 

free, rarely connate basally; stamens 4-5, the 

filaments filiform, free or adnate to the petals 

basally; anthers about as long as wide or linear, 

dorsifixed, dehiscing longitudinally, the connective sometimes produced apically; pollen 3-, 6- 

8, 10, 1 l-colporate; disk rarely present; pistil 1, 

the carpels 2 (3); ovary semi-inferior or inferior, 

very rarely superior, 2( 1 -3)-locular, the ovules 

1-4 (-10) per locule, axile near the top of the 

locule, unitegmic, ? crassinucellar, anatropous; 

fruit a nutlet or dehiscing into cocci open ventrally; seeds minute, the embryo straight, 0.1 the 

length of the endosperm; cotyledons not broadened, 0.5 the length of the embryo; endosperm 

copious, fleshy. 

Composition: 12 genera, 75 species. 

Distribution: South Africa, especially Cape 

Province. 

VAHLIACEAE (Figure 1 1 Oa).-Annual or biennial herbs with a taproot; xylem vessel perforation plates simple; leaves opposite, simple, entire, 

exstipulate; inflorescences axillary, sympodial, 

cymose, the flowers paired; flowers bisexual, the 

parts epigynous; sepals 5, valvate; petals 5, free, 

short; stamens 5, the filaments subulate or linear, 

sometimes with a basal scale; anthers oblong, 

slightly longer than wide, dorsifixed; pollen tricolporate; disk epigynous; pistil 1, the carpels 2 
NUMBER 58 213 

b 

FIGURE 1 ~~.--ITEACEAE: a, Ztea ilicijolza half of inflorescence, flower, petal, views of stamen, 

fruit, I. virginica flower, 1,s. of same (after Oliver; Baillon, 1866-1 895). PTEROSTEMONACEAE: 

b, Pterostemon mexicanus flower, Is. of same (petals removed), 1,s. of ovary (after Engler). 

BAUERACEAE: c, Bauera rubioides flowering twig, flower, Is. of same, B. capitata flower, 1,s. of 

sariie (after Baillon, 1866-1895). BRUNIACEAE: d, Brunia pinifololia flower, I.s. of same, floral 

diagram, Is. of fruit, B. nodzjZora C.S. of fruit, 1,s. of seed showing the minute embryo, flowering 

twig (after Le Maout and Decaisne, 1873). 

(3), the styles 2 (3), divaricate, the stigmas apical; 

ovary unilocular, the placentas 2 (3), suspended 

from the apex of the ovary, the ovules numerous, 

bitegmic, tenuinucellar, anatropous; fruit a capsule dehiscent apically between the styles; seeds 

numerous minute, oblong; embryo 0.75 the India. 

length of the seed, the cotyledons 0.3 the length 

of the embryo, endosperm copious. 

Composition: 1 genus, 5 species. 

Distribution: Tropical and subtropical Africa, 

Madagascar; southwestern Asia to northwestern 
214 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

a 

FIGURE 1 1 ~.-VAHLIACEAE: a, Vahlia capensis flower, Is. of same, V. viscosa views of stamen, 

1,s. of pistil, V. oldenlandioides fruit, seed, flowering shoot (after Engler). DONATIACEAE: b, 

Donatia rnagellanica plants, flower, same with petals removed, I.s. of ovary (after Hooker and 

Hooker, 1837-1 982). TETRACARPAEACEAE: c, Tetracarpaea tasmannzca flowering and fruiting 

branch, flower, views of anther, essential organs, fruitlets with persistent filaments and calyx, 

dehiscent fruitlet, C.S. of same (after Hooker and Hooker, 1837-1982). 

DONATIACEAE (Figure 1 1 Ob).-Herbs; xylem 

vessel perforation plates scalariform, tracheids 

present; leaves linear, coriaceous, very densely 

spirally arranged, exstipulate; flowers solitary, 

terminal, bisexual, the parts epigynous: calyx 

lobes 5-7: petals 5-10, free, f fleshy: stamens 

2-3, the filament free; anthers small, about as 

long as wide: pollen 3(4)-colporate: disk epigynous; pistil 1, the carpels 2-3, the styles 2-3, 

stigmas apical; ovary 2-3 locular, the ovules numerous, axile; fruit a turbinate capsule, dehiscence circumscissile; seeds few, with membranous testa; embryo minute: endosperm copious, 

fleshy. 

Composition: 1 genus, 2 species. 

Distribution: Tasmania, New Zealand, cool 

TETRACARPAEACEAE (Figure 1 1 Oc).-Shrubs; 

southern South America. 
NUMBER 58 

xylem vessel perforation plates simple; leaves 

alternate, simple, doubly serrate, coriaceous, exstipulate; inflorescence a raceme; flowers bisexual, the parts hypogynous; sepals 4, nearly separate; petals 4, free, orbicular, erose, clawed, 

caducous; stamens 8, obdiplostemonous, the filaments filiform, anthers basifixed or adnate; pollen 3-colpate(?); pistils 4, the stigmas subsessile, 

apical, the ovules numerous, ventral; fruit fusiform follicles, the seeds numerous; embryo minute, almost globose, at the base of copious fleshy 

endosperm. 

Composition: 1 genus, 1 species. 

Distribution: Tasmania. 

ESCALLONIACEAE (Figure 1 1 la,b).-Trees and 

shrubs; xylem vessel perforation plates usually 

scalariform with less than 20 to 125 bars; leaves 

simple, alternate, rarely opposite or subverticillate, mostly with gland-tipped teeth, exstipulate; 

inflorescences mostly racemes, sometimes cymes; 

flowers bisexual or rarely the plants dioecious or 

polygamous, the parts perigynous or epigynous; 

sepals 4-5, mostly basally connate rarely free, 

imbricate or valvate, often persistent; petals 4- 

5, free or rarely shortly connate, imbricate or 

valvate; stamens 4-5 (-9), free, the filaments 

filiform; sometimes 4-5 staminodes present; anthers dehisce longitudinally; pollen 3-colporate; 

disk-lobes alternate with the stamens; pistil 1, the 

carpels 4-5 (2-6), the style 1 (2-6), stigma apical; 

ovary 1 -5(6)-Iocular, the ovules numerous, axile 

in multilocular ovaries, parietal in those unilocular, unitegmic (Escallonia) or bitegmic (Brexia), 

crassinucellar or tenuinucellar, anatropous; fruit 

a septicidal capsule, more rarely a berry; embryo 

linear, straight, 0.4 the length of the endosperm; 

cotyledons not broadened, 0.15 the length of the 

embryo; endosperm copious. 

Composition: 7 genera, 150 species. 

Distribution: Tropical and south temperate; 

mostly South America, especially Andes, and 

Australia, Malaysia, Burma; south and east Africa, Madagascar. 

GROSSULARIACEAE (Figure 1 1 Ic).-Shrubs, 

often spiny; xylem vessel perforation plates 

nearly always scalariform with less than 20 bars, 

rarely both scalariform and simple; leaves simple, 

215 

alternate, lobed and dentate, exstipulate or stipules adnate to the petiole; inflorescences racemes 

or the flowers subsolitary; flowers bisexual 

or the plants dioecious; calyx lobes 5 (4), often 

petaloid, persistent; petals 5 (4), inserted on the 

calyx-tube, small, often smaller than the calyx 

lobes, rarely 0; stamens 5 (4), inserted on the 

calyx-tube; anthers small, about as long as wide, 

dorsifixed; pollen 8-9-pored; pistil 1, the carpels 

2, the styles 2, free to completely connate, stigmas apical; ovary inferior, unilocular, the ovules 

numerous or few, parietal, bitegmic, crassinucellar, anatropous; fruit a berry, the seeds usually 

numerous; embryo linear, straight, 0.2 the 

length of the endosperm; cotyledons not broadened, 0.2 the length of the embryo; endosperm 

copious, fleshy. 

Composition: 2 genera, 150 species. 

Distribution: North temperate zone, cool regions of mountains in Central America and the 

Andes to southern South America; Eurasia, 

northwestern Africa. 

BRUNELLIACEAE (Figure 1 12a-c).-Trees; xylem vessel perforation plates usually scalariform, 

rarely also simple; leaves opposite or verticillate, 

pinnately compound, trifoliolate, or simple, usually dentate, stipulate; inflorescences terminal 

and axillary panicles; plants usually dioecious 

rarely polygamous or the flowers bisexual; sepals 

4-5 (-7), shortly connate, valvate in bud; petals 

0; disk adnate to the ovary and calyx; stamens 

8- 10 (- 1 S), the filaments filiform, free; anthers 

small, apiculate, twice as long as wide, dorsifixed; 

pollen 3-colporate; pistils 4-5 (2-3), the styles 

subulate, stigmas decurrent along ventral surface 

of style; ovary superior, rarely semi-inferior, the 

ovules paired, collateral, axile; fruits follicles, 

often with irritant hairs, separating into exocarp 

and cartilaginous endocarp; embryo straight, 0.8 

the length of the endosperm; cotyledons thin, 

3.3 times wider than the radicle, 0.8 the length 

of the embryo; endosperm copious, fleshy. 

Composition: 1 genus, 45 species. 

Distribution: Tropical America from Mexico 

to Peru and Bolivia in the Andes; West Indies. 

CUNONIACEAE (Figure 1 13a,b).-Shrubs and 

trees; xylem vessel perforation plates scalariform 
216 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 1 11 .-ESCALLONIACEAE: a, Escallonia monteuidensis I.s. of flower, 1.s. and C.S. of ovary, 

views of anther; b, E. chlorophylla flower with parts removed, petal, fruit, I.s. of same showing 

the placentas, valve of fruit, seed, I.s. of same, inflorescence (after Johnson; Martius, 1840- 

1906). GROSSULARIACEAE: c, Ribes speciosum flower, R. rubrum floral diagram, 1,s. of flower. R. 

nigrum views of stamen, R. rubrum infructescence, seed, 1,s. of same showing the minute embryo 

(after Baillon, 1866-1895; Engler). 

with usually fewer than 20 bars, sometimes both 

scalariform and simple, rarely only simple; leaves 

pinnately or trifoliately compound, rarely simple, 

serrate, mostly opposite, rarely verticillate, very 

rarely alternate if Gumillea is included, coriaceous, the stipules often large; inflorescences 

panicles to heads, or rarely the flowers solitary; 

flowers small, mostly bisexual, sometimes the 

plants dioecious, the parts hypogynous or rarely 

epigynous (Ceratopetalum); sepals 4-5 (3-6), free 

or basally connate, sometimes accrescent; petals 

4-5 (3-6) or 0, free; stamens 8-10 (4 to -SO), 

the filaments free; anthers small, about as long 

as wide, dorsifixed; pollen 2-3-colporate or 3- 

colpate; disk mostly present, often annular, sometimes epigynous; pistils 2 (3-5) or more com- 
NUMBER 58 217 

a 

FIGURE 1 12.-BRUNELLIACEAE: a, Brunellia sibundoya floral diagram, B. pinnata floral diagram 

of d flower, B. colombiana d flower (2 sepals removed), same with sepals and stamens removed 

to show the disk and pistillodes, B. pallida 0 flower; b, B. oualijolia pistil, apex of style enlarged 

to show the decurrent ventral stigma, Brunellia sp. ovary laid open to show the collateral ovules, 

C.S. and 1,s. of ovary, B. oualijolia 1,s. of P flower, B. sibundoya 1,s. of d flower; c, B. standleyana 

part of infructescence, dehiscent fruitlets, B. mexicana 1,s. of fruitlet, B. costaricensis dehiscent 

fruitlet showing the naturally exposed seed, dehiscent endocarp, B. tomentosa I.s. of seed, lateral 

view of embryo (after Cuatrecasas, 1970). 

monly 1 and the carpels 2 (3-5), the styles 2 (3- 

5), free, often filiform, the stigmas apical; ovary 

superior, semi-inferior or inferior, 2(3-5)-locular 

when compound, the ovules numerous or few, 

rarely 1 per locule, axile or apical, bitegmic, 

crassinucellar, anatropous or semi-anatropous; 

fruit usually a capsule, dehiscing ventrally, rarely 

follicles, drupe, nut, or samara; seeds glabrous 

or long-pilose, sometimes winged; embryo 0.5- 

0.7 the length of the endosperm; cotyledons 1.7- 

4.0 times the width of the radicle, 0.3-0.9 the 

length of the embryo; endosperm copious, fleshy. 

Composition: 26 genera, 250 species. 

Distribution: Mostly Southern Hemisphere, 

especially Australia and Oceania; Malaysia; a few 

species in South Africa, Madagascar, and southern South America to Mexico and the West 

Indies. 
218 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

b 

FIGURE 1 1 3.-cUNONIACEAE: a, \l’einmannza guianensis flower, pistil, w. humilis seed, 1,s. of 

same, Lamanonia tomentosa floral diagram, flower; b, I.s. of pistil and 2 stamens, C.S. of ovary, 

C.S. of fruit, valves of fruit, 1,s. of fruit valve, inflorescence (after Martius, 1840-1906). 

GREYIACEAE: c, Greyia sutherlandi flower, calyx, petal, flower with perianth removed, nectary 

laid open and C.S. of ovary, pistil, stamen, dehiscent fruit, Is. of seed and embryo (after Hooker 

and Hooker, 1837-1982; Harvey). 

GREYIACEAE (Figure 1 ISc).-Shrubs and small 

trees; xylem vessel perforation plates simple; 

leaves alternate, simple, shallowly lobed, toothed, 

with resinous glands, the petiole adnate to the 

twig for about 0.6-2.5 cm., exstipulate; inflorescences terminal racemes; flowers bisexual, actinomorphic or slightly zygomorphic; sepals 5, 

free or very shortly connate, imbricate; petals 5, 

free, perigynous, imbricate; stamens 10, obdiplostemonous, free, the filaments elongate, filiform; anthers about as long as wide, didymous, 

basifixed; pollen 3-colporate; disk cupular, external to the stamens, coronoid with 10 staminodial 

appendages, connected to the ovary by 5 interstamina1 ridges, secreting abundant nectar; pistil 

1, the carpels 5, the style 1, elongate, filiform, 

stigma punctiform or shortly 5-lobed; ovary superior, unilocular, the ovules numerous on 5 

parietal placentas sometimes meeting in the axis, 

bitegmic, crassinucellar, anatropous; fruit a chartaceous capsule dehiscing along the ventral sutures and also septicidal; seeds numerous, minute, the embryo straight, very small; endosperm 

copious, fleshy. 
NUMBER 58 219 

b 

FIGURE 1 14.-CEPHALOTACEAE: a, Cephalotus follicularis inflorescence (scapose), floral diagram, 

flower, I.s. of same; b, flower with stamens and calyx removed, I.s. of pistil, dehiscent fruitlet, 

seed, 1,s. of same, pitcher (after Le Maout and Decaisne, 1873). DIONCOPHYLLACEAE: c, 

Dioncophyllum tholoni flower with petals removed, petal, I.s. of pistil, C.S. of ovary, Triphyophyllum 

peltatum essential organs, part of a gland-leaf, very young fruit showing young seeds developing 

on thickened funicles; d, Habropetalum dawei leaves and part of a flowering twig, bud and 1,s. 

of flower, stamen, petal and anther, pistil and C.S. of ovary, Triphyophyllum paltaturn mature 

seed and parts of leaves (after Hutchinson and Dalziel, Gilg). 

Composition: 1 genus, 3 species. 

Distribution: Southeastern South Africa. 

CEPHALOTACEAE (Figure 1 14a,b).--Insectivorous herb of swamps; xylem vessel perforation 

plates scalariform; leaves alternate, in a rosette, 

some flat and entire, others developed into ascidia, exstipulate; inflorescence a terminal raceme of cymules; flowers bisexual, the parts pe- 
220 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

rigynous; sepals 6, free, valvate, accrescent, colored; petals 0; stamens 12, of 2 lengths, the 

filaments filiform; anthers small, about as long as 

wide, produced apically, the connective glandular; pollen usually 3-colporate; intrastaminal disk 

present; pistils 6, the styles about as long as the 

ovary, stigma apical; ovule 1 (2), basal, bitegmic, 

crassinucellar, anatropous; fruit follicles; embryo 

straight, 0.5 the length of the endosperm; cotyledons plano-convex, twice as wide as the radicle, 

0.6-0.8 the length of the embryo; endosperm 

copious, fleshy, oily. 

Composition: 1 genus, 1 species. 

Distribution: Southwestern Australia. 

D ROSERA L ES 

Herbs, less commonly shrublets, sometimes 

climbing; xylem vessel perforation plates simple, 

rarely scalariform with numerous bars; leaves 

alternate, rarely whorled, entire, crenate or pinnatifid, sometimes thalloid, often with stalked 

insectivorous glands, sometimes circinate in bud, 

exstipulate or stipulate; flowers bisexual, less 

commonly unisexual; sepals 5 (0, 2-4), free or 

basally connate; petals 5 or 0 (4), free, rarely 

shortly connate, hypogynous or very rarely perigynous; stamens 1-10 (-30) in l or more 

whorls, the filaments free or connate, hypogynous or less commonly adnate to the base of the 

petals; anthers basifixed or dorsifixed, sometimes 

didymous; pollen sometimes in dyads or tetrads, 

3-4-colpate or colporoidate, oligo- 12-forate, occasionally 4-rupate, 6-rugate or nonaperturate; 

pistil 1, the carpels 2, 3, or 5, styles 1, 2, 3, or 5, 

free or partially connate, the stigmas apical or 

decurrent, sometimes branched; ovary superior, 

unilocular, rarely 2- 3- or 5-locular, the ovules 

usually numerous, rarely few to 1 per locule, 

parietal, basal, axile or rarely the placentas freecentral; fruit a loculicidal or septicidal capsule, 

the seeds usually minute, rarely large, the embryo minute to large; endosperm copious or absent. 

Distribution: Almost cosmopolitan in wet to 

aquatic habitats. 

Chemistry: Naphthaquinones are present in 

all 4 genera of Droseraceae. Droseraceae may 

show cyanogenesis. Ellagic acid and quercitin are 

present in some. 

DIONCOPHYLLACEAE (Figure 1 14c,d).- 

Lianas; xylem vessel perforation plates simple; 

leaves alternate, entire or crenate, the midrib 

often excurrent with 2 recurved hooks, sometimes with numerous conspicous mucilaginous 

glands, vernation sometimes circinate, exstipulate; inflorescence a lax supra-axillary cyme; flowers bisexual, the parts hypogynous; sepals 5, free 

or shortly connate; petals 5, free; stamens 10 

(-30), the filaments short or long, the anthers 

basifixed; pollen 3(4)-colp(oroid)ate; pistil 1, the 

carpels 2 or 5, the styles 2 or 5, filiform, free or 

shortly connate, stigmas apical, sometimes plumose; ovary unilocular, the ovules numerous, 

parietal, bitegmic, crassinucellar, anatropous; 

fruit a loculicidal capsule dehiscing before the 

seeds are ripe; seeds large, discoid, winged; embryo large, spread out, 0.7 the width of the 

endosperm; endosperm copious. 

Composition: 3 genera, 3 species. 

Distribution: Rain forest of tropical west Africa. 

DROSERACEAE (Figure 1 15a,b).-Herbs, rarely 

subshrubs (Drosophyllum); xylem vessel perforation plates simple, successive bundles of xylem 

and phloem without regular arrangement; leaves 

often in rosettes, alternate, rarely whorled, usually circinate in bud and covered with sticky 

stipitate tentacles or with marginal bristles, stipulate or exstipulate, insectivorous; inflorescences 

usually circinate cymes (cincinni), rarely racemes 

or the flowers solitary; flowers bisexual; sepals 5 

(4), shortly connate; petals 5 (4), free, hypogynous, very rarely perigynous; stamens 4-20 in 1 

or more whorls, hypogynous, the filaments filiform, free or rarely shortly connate, the anthers 

small, about as long as wide, the thecas sometimes 

divergent basally and separated by connective; 

pollen in tetrads except Drosophyllum, 6-polyforate; disk 0; pistil 1, the carpels 3 or 5, the styles 

3 or 5 (1 in Dionaea), filiform, mostly free, the 
NUMBER 58 22 1 

v 11 

FIGURE 115.-DROSERACEAE: a, Drosera sessdfoh2 leaf, D. zntermedia bud, I.s. of bud, sepal, 

petal, viehs of stamen, pistil, branched style, C.S. of ovary; b, floral diagram, D. sessilfolia C.S. 

of ovary, single valve of fruit showing the numerous minute seeds, D. intermedia dehiscent 

fruit, seed, I.s. of same showing the minute embryo (after Martius, 1840-1906). BYBLIDACEAE: 

c, Byblisgzgantea part of leaf and views of flower, glandular hair, calyx, essential organs, stamen, 

pistil, C.S. of ovary, dehiscent fruit and part of calyx, seed, 1,s. of same; d, Roridula dentata I.s. 

of flower, essential organs, floral diagram, dehiscent fruit and perianth, seed, I.s. of same (after 

Hooker and Hooker, 1837-1982; Baillon, 1866-1895). 

stigmas apical, sometimes branched; ovary superior, unilocular, rarely 3- or 5-locular, the ovules 

(3-) numerous, parietal or basal, rarely axile or 

free-central, bitegmic, crassi- or tenuinucellar, 

anatropous; fruit a loculicidal capsule or indehiscent (Aldrovanda), the seeds spindle-shaped, mi- Distribution: Almost cosmopolitan, in wet 

nute; embryo straight, subglobular, 0.1-0.3 the 

length of the endosperm; cotyledons barely differentiated, 0.25 the length of the embryo; endosperm copious, crystalline-granular. 

Composition: 4 genera, 100 species. 
222 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

soil, one genus aquatic; centered in Australia and 

New Zealand; much of Eurasia; tropical and temperate southern Africa; North America; northeastern half and southwestern tip of South America. 

BYBLIDACEAE (Figure 1 15c,d).-Herbs and 

shrublets; xylem vessel perforation plates simple 

(Byblis) or scalariform with numerous bars (Roridula); leaves alternate, linear or lanceolate, entire or pinnatifid, circinnate in vernation, with 

stalked, capitate, viscous glands, exstipulate; inflorescences few-flowered racemes, or more commonly the flowers solitary, axillary on long or 

short pedicels; flowers bisexual, actinomorphic; 

sepals 5, free or basally connate, imbricate; petals 

5, free or sometimes very shortly connate, imbricate or contorted; stamens 5, alternipetalous, 

hypogynous or very shortly adnate to the base of 

the petals, the filaments short; anthers basifixed, 

with a swollen rounded base, sometimes didymous, connivent, dehiscent by apical pores or 

short slits; pollen 3-4-colpate or colporoidate, 

occasionally 4-rupate or 6-rugate; pistil 1, the 

carpels 2-3, the style 1, elongate or short, the 

stigma capitate or very shortly 2-3-lobed; ovary 

superior, 2-3-locular, the ovules 1 -several per 

locule, axile, sometimes subapical, bitegmic 

(unitegmic ? in Byblis), tenuinucellar, anatropous; 

fruit a loculicidal capsule; embryo minute, globular and undifferentiated, 0.2 the length of the 

endosperm (Byblis) or straight, 0.4-0.6 the 

length of the endosperm, the cotyledons not 

broadened, 0.2-0.4 the length of the embryo; 

endosperm copious, fleshy. 

Composition: 2 genera, 4 species. 

Distribution: Northern and western Australia, New Guinea; South Africa. 

PODOSTEMACEAE (Figure 1 16a,b).-Submerged fresh-water f thalloid herbs; laticiferous 

secretory canals and cavities in some; vascular 

system in the form of a network; leaves alternate, 

usually simple, linear to lamellate, often basally 

sheathing; inflorescences terminal, cymose, in 

fascicles or dichasia, or the flowers solitary; flowers minute, bisexual, fragrant, often enclosed in 

a spathe, the parts hypogynous; sepals 2, 3, or 5, 

sometimes petaloid, free or basally connate; petals 0; stamens 1-25, in 1 or 2 series, the filaments 

free or connate, the anthers dorsifixed or basifixed, small; pollen grains single or in dyads or 

tetrads, 3-colpate, 3-colporoidate, oligo- 1 2-forate or nonaperturate; pistil 1, the carpels 2 (3), 

the styles 2 (1-3), free or partially connate, filiform or short, often strongly papillate, apex 

punctiform, or capitate when one, the stigmas 

apical or decurrent; ovary superior, sometimes 

stipitate, 2( 1 -3)-locular, the placentas axile, 

thick, rarely free-central, the ovules numerous, 

rarely few (2 or 4 in Farmeria), bitegmic, tenuinucellar, anatropous; fruit a septicidal or septifragile capsule or rarely indehiscent (Farmeria 

sp.); seeds minute, the embryo straight, the cotyledons thick, sometimes unequal, the radicle 

short; endosperm 0. 

Composition: 

Distribution: 

45 genera, - 150 species. 

Attached to rocky substrate in 

rushing water, mainly tropical and subtropical, 

especially America; temperate North America to 

tropical South America; central Africa, Madagascar; India to Indochina and the East Indies, 

northern Australia. 

HYDROSTACHYACEAE (Figure 1 1 Gc,d).-Submerged fresh-water herbs; vascular bundles in an 

isolated ring, accompanied by medullary and cortical strands in the inflorescence axis; leaves radical, elongate, simple or pinnatisect, dilated and 

ligulate basally; inflorescence a dense scapose 

spike; plants dioecious, rarely monoecious, the 

flowers sessile in the axil of bracts; perianth 0; 

stamen 1, the filament short, the anther sacs 

separate; pollen in tetrads, probably nonaperturate; pistil 1, the carpels 2, the styles 2, long, 

filiform, free or basally connate, divergent, the 

apex punctiform; ovary superior, unilocular, the 

placentas 2, parietal, the ovules numerous, unitegmic, tenuinucellar, anatropous; fruit a septicidal capsule, the seeds numerous, minute; embryo linear, the cotyledons not broadened(?), 

0.8-0.9 the length of the embryo; endosperm 0. 

Centered in Madagascar, a few 

Composition: 1 genus, -20 species. 

Distribution: 

species in tropical and southern Africa. 
NUMBER 58 223 

FIGURE 1 16.-PODOSTEMACEAE: a, Tulasneantha monodelpha floral diagram, flower, Rhyncholacis linearis part of plant in flower, immature fruit, floral diagram; b, Lophogyne helicandra 

immature fruit, dehiscent fruit, views of anther, Mniopsis weddelliana flower, fruit (after 

Martius). HYDROSTACHYACEAE: c, Hydrostachys verruculosa d flower, 0 flower, I.s. of ovary, 

dehiscent fruit, seed, 1,s. of same; d, H. distichophylla stamen, C.S. of anther, part of plant in 

flower (after Baillon, 1866-1 895; Delessert). 
224 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

f 

FIGURE 1 17.-BEGONIACEAE: a, Begonia saxifraga inflorescence (without the primary peduncle), 

B. luxurians C.S. of fruit, fruit, B. neglecta d flower, 0 flower, Begonia (Barya) sp. androecium; b, 

B. luxurians 2 views of stamen, B. hispzda stamen; c, Begonia sp. diagram of d flower, B. 

(Eupetalum) sp. androecium; d, B. populnea 6 flower, views of stamen, views of style and stigma, 

Begonia sp. 0 floral diagram, seed, I.s. of same (after Martius, 1840-1906; Le Maout and 

Decaisne, 1873). DATISCACEAE: e, Datisca cannabina d flowering shoot, 6 flower lateral view 

and from below, part of 0 inflorescence, 0 flower, I.s. of same, C.S. of ovary;f; seed, 1,s. of same, 

dehiscent fruit (after Baillon, 1866-1895; Le Maout and Decaisne, 1873). 

BEGONIALES simple and scalariform with many bars; leaves 

alternate, very rarely opposite, usually simple, 

Herbs, often succulent, rarely subshrubs, sometimes compound, usually toothed, stipulate 

or exstipulate; flowers unisexual, rarely bisexual; 

sepals 2-6 (-1 0), usually corolline, usually free, 

rarely climbing, the stems sometimes jointed with 

swollen nodes; xylem vessel perforation plates 
NUMBER 58 225 

rarely connate; petals 2-5 or 0; stamens mostly 

numerous, to -100, rarely as few as 4, the filaments free or connate; anthers adnate or basifixed, often didymous and the connective produced apically, dehiscent longitudinally, rarely 

by pores; pollen 3-colporate; pistil 1, the carpels 

3 (2-5), styles 3 (2-5), free or basally connate, 

usually bifid apically, stigmatic for a considerable 

distance; ovary inferior, rarely semi-inferior, 

3 (1-5)-locular, the ovules numerous, axile on 

lamina, rarely parietal; fruit a capsule dehiscing 

apically between the styles or loculicidal, rarely a 

berry, the seeds numerous, minute; embryo 

straight, terete, endosperm scanty or absent. 

Distribution: Pantropical and -subtropical, except Australia, few in temperate regions, in moist 

or dry habitats. 

BEGONIACEAE (Figure 1 17a-d).-Herbs, 

rarely subshrubs, rarely climbing, mostly succulent, the stems sometimes jointed with swollen 

nodes; xylem vessel perforation plates simple and 

scalariform with many bars, both types sometimes in a single vessel; leaves alternate, very 

rarely opposite (verticillate at the apex of the 

stem in Begonia verticillata), simple or rarely palmately or pinnately compound, mostly asymmetrical, sometimes distichous, sometimes basal, usually toothed, stipulate; plants monoecious; inflorescences often axillary dichasia, tending toward 

cincinni or sometimes the flower solitary; 8 

flower: sepals 2 or 4 (5), corolline, usually free, 

rarely connate; petals 5-2 or 0, usually free, 

rarely connate; stamens mostly numerous (to 

-loo), rarely as few as 4, the filaments free or 

connate; anthers adnate or basifixed, the thecas 

often separate, the connective mostly produced 

apically, dehiscing longitudinally, rarely by 

pores; pollen 3-colporate; 0 flower: sepals 5-2; 

petals 5-2 or tepals 5-2 (6-8); pistil 1, the carpels 

3 (2-5), the styles 3 (2-5), free or connate basally, 

the apices often twisted and bifid, papillose and 

stigmatic for a considerable distance; ovary usually completely inferior, rarely semi-inferior (Hillebrandia), 3( 1-5)-locular, the ovules very numerous, axile or rarely parietal (Hillebrandia), 

bitegmic, crassinucellar or tenuinucellar, anatropous; fruit a capsule dehiscing between styles, 

or loculicidal, rarely a berry; seeds minute, very 

numerous, sculptured; embryo straight, the cotyledons not broadened, 0.2-0.5 the length of the 

embryo; endosperm scanty or 0. 

Composition: 

Distribution: Pantropical and -subtropical, except Australia; most frequent in South America. 

DATISCACEAE (Figure 1 17e,f).-Herbs; xylem 

vessel perforation plates simple; leaves alternate, 

pinnately compound, serrate, exstipulate; inflorescences axillary and terminal fascicles forming 

a large panicle, the plants usually dioecious, 

rarely monoecious or the flowers bisexual; calyx 

lobes in 8 4-6 (-lo), in 0 3 or 6; petals 0; stamens 

4-25, the filaments very short, free; anthers 

large, oblong, basifixed; pollen 3-colporate; pistil 

1, the carpels 3-5, the styles 3-5, free, filiform, 

each bifid, papillose and stigmatic for most of 

their length; ovary inferior, unilocular, the 

ovules very numerous, parietal, bitegmic, crassinucellar, anatropous; fruit a capsule dehiscing 

apically between the styles; seeds minute, numerous; embryo terete, cotyledons 0.5 the length of 

the embryo; endosperm 0. 

3 genera, - 1400 species. 

Composition: 1 genus, 2 species. 

Distribution: Dry habitats; southwestern 

United States, Mexico; western Mediterranean 

region to central Asia and the Himalayas. 

APIALES 

Herbs, shrubs, and trees, rarely climbing, the 

tissues with secretory canals containing oily, 

resinous, mucilaginous or gummy material; xylem vessel perforation plates usually simple, but 

scalariform plates predominant in some genera; 

leaves alternate, very rarely opposite or whorled, 

mostly compound, rarely simple, the leaflets 

toothed, rarely entire, stipulate or exstipulate; 

inflorescences often umbellate; flowers small, 

bisexual rarely unisexual; sepals 5 (3-lo), connate and usually reduced; petals 5 (3-1 0, 0), free, 

rarely connate; stamens 5 (3 to -loo), free, the 

anthers basifixed or dorsifixed; pollen 3(2-4)- 

colporate, 3-colpate or 6-rugate; glandular disk 
226 

epigynous; pistil 1, the carpels 2 or 5 (1 to - 160), 

styles 2 or 5 (0 to -160), free, less commonly 

connate, the stigmas apical; ovary inferior, very 

rarely semi-inferior or superior, 2 or 5 (1 to 

-160)-locular, the ovules 1 per locule, axileapical, unitegmic; fruit a schizocarp, drupe or 

berry; embryo usually minute; endosperm copiDistribution: Cosmopolitan in various habitats. 

Chemistry: The Araliaceae and Apiaceae are 

very closely related. Apiaceae have lots of coumarins and monoterpenes (also other terpenes). 

Saponins are common only in Araliaceae and 

Apiaceae, but triterpenoid saponins and/or sapogenins are reported from these and Alangiaceae 

and Cornaceae. Ellagic acid occurs in Cornaceae 

but not Apiaceae. Kaempferol and caffeic acid 

occur in all four families. Acetylenic compounds 

occur mainly in Asteraceae and Apiaceae, but 

also in Araliaceae, Pittosporaceae, Lauraceae, 

Myoporaceae, and legumes. Aucubin (iridoid)- 

type glycosides occur in Cornaceae and Garryaceae but not Araliaceae and Apiaceae. Alkaloids 

of 9 groups are known from the order but no 

group occurs in more than one family. Unlike 

Cornales, Griselinia does not have aucubin (iridoid) glycosides. Griselinia has triterpinoid saponins and/or sapogenins. Chemical evidence 

seems to indicate a relationship of Pittosporaceae 

to Apiales. It has many saponin-containing members. 

ARALIACEAE (Figure 1 18a-c).-Trees and 

shrubs, less commonly herbs or lianas, the tissues 

with secretory canals containing oily, resinous, 

or gummy material; xylem vessel perforation 

plates usually simple, but scalariform plates predominant in some genera; leaves alternate, very 

rarely opposite, very rarely whorled (Panax), 

mostly compound, sometimes simple, the leaflets 

generally toothed, rarely entire, the indument 

often stellate, stipulate or rarely exstipulate; inflorescences terminal and axillary, often compound, racemes, umbels, heads, spikes or panicles, rarely ending in cymes; flowers bisexual or 

the plants polygamous or dioecious; sepals 5 (3- 

lo), connate, the calyx truncate or toothed; petous. 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

als 5 (3-lo), valvate or slightly imbricate, free or 

rarely connate and calyptrate; stamens 5 (3 to 

- IOO), alternipetalous, free; anthers small, about 

as long as wide, dorsifixed; pollen 3(2-4)-colporate, 3-colpate or 6-rugate; disk epigynous, sometimes lobed; pistil 1, the carpels 5 (1 to -160), 

styles 5 (0 to -160), long or short, free or connate, the stigmas apical; ovary inferior, rarely 

semi-inferior, very rarely superior (Tetraplasandra gymnocarpa), 5( 1 to - 160)-locular, the ovules 

sometimes with an obturator, 1 per locule, axile, 

unitegmic, crassinucellar or tenuinucellar, anatropous; fruit usually a drupe or berry, rarely a 

schizocarp (Myodocarpus), sometimes with oily 

vesicles, rarely concrescent into a head (Meryta); 

embryo often minute and undifferentiated, 

sometimes spatulate, 0.06-0.7 the length of the 

endosperm; cotyledons to 3 times the width of 

the radicle, 0.3-0.4 the length of the embryo; 

endosperm copious, fleshy, sometimes ruminate. 

Composition: 60 genera, 700 species. 

Distribution: Tropical and subtropical, relatively few temperate; centered in southeast Asia 

and tropical America. 

APIACEAE (Figure 1 18d).-Herbs, rarely 

shrubs or small trees, very rarely twining or 

scandent, the tissues with secretory canals containing resinous, oily, or mucilaginous material; 

xylem vessel perforation plates usually simple, 

but occasionally scalariform with few bars; leaves 

alternate, rarely opposite (Bowlesia, Apiastrum 

spp.), basally sheathing, mostly compound, rarely 

simple, the leaflets mostly toothed, rarely entire, 

very rarely reduced to a petiole, exstipulate or 

rarely stipulate; inflorescences axillary, simple or 

compound, cymose umbels or rarely heads or 

umbels racemose-paniculate, or the flowers solitary (Xanthosia, Azorella); flowers bisexual or 

rarely the plants monoecious or polygamous, 

rarely dioecious (Arctopus); calyx consisting of 5 

minute teeth or the latter obsolete, or petaloid; 

petals 5 (0), free, valvate or slightly imbricate; 

stamens 5, alternipetalous, free; anthers small, 

about as long as wide, basifixed or dorsifixed; 

pollen 3(2)-colporate; disk epigynous; pistil 1, the 

carpels 2, sometimes 1 aborts (Echinophora), an 

occasional flower is trimerous, the styles 2, free, 
NUMBER 58 227 

A 

b 

C 

n 4 

FIGURE 1 18.-ARALIACEAE: a, Gilibertia resinosa part of infructescence, flower, views of anther, 

I.s. of flower, C.S. of ovary, flower with petals and 1 stamen removed, fruit and C.S. of same; b, 

Griselinia ruscqoh d inflorescence, d bud, d flower, views of anther, 0 flower, Is. of same; c, 

I.s. and C.S. of fruit, embryo, 0 axillary inflorescences (after Martius, 1840-1906). APIACEAE: 

d, Hydrocotyle quingueloba bud, I.s. of bud, views of stamen and fruit, Eryngium pristis flower, 

same from above with anthers and petals removed and C.S. of fruit, Apium ammi fruit and C.S. 

of mericarp (after Martius, 1840-1906). 

stigmas apical; ovary inferior, bilocular, rarely 

unilocular (Lugoecia, Echinophora), the ovules 1 

per locule, sometimes with an obturator, axileapical, unitegmic, pseudocrassinucellar or tenuinucellar, anatropous; fruit a schizocarp with 

oily vesicles, sometimes winged, the seeds 2, 

rarely 1, very rarely the mericarps not separating 

(Thecocarpus); embryo usually minute, sometimes 
228 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

undifferentiated, 0.03-0.6 the length of the endosperm; cotyledons usually thin, sometimes 

moderately thick, 1-3 times the width of the 

radicle, 0.2-0.6 the length of the embryo; endosperm copious, usually firm, fleshy. 

Composition: -300 genera, -2800 species. 

Distribution: Cosmopolitan except part of the 

arctic region and some deserts; chiefly temperate 

Northern Hemisphere; Apioideae are bipolar 

but centered in Eurasia; Saniculoideae are bipolar but better represented in the Southern Hemisphere than the Apioideae; Hydrocotyloideae 

are predominantly Southern Hemisphere. 

CORNALES 

Trees and shrubs, very rarely herbs; xylem 

vessel perforation plates scalariform, usually with 

many bars, occasionally also simple; leaves opposite or alternate, simple, usually entire, sometimes toothed, exstipulate; flowers small, bisexual, less commonly unisexual; sepals 4-5 (0), free 

or connate; petals 4-5 (-8, 0), free; stamens 4-5 

(1 - 12) in 1-2 series; staminodes rarely present; 

filaments usually short, free, the anthers basifixed or dorsifixed; pollen 3-colporate; glandular 

disk usually present; pistil 1, the carpels 2 

(1-lo), style 1 (2), rarely lobed, the stigma usually apical, rarely decurrent ventrally; ovary inferior, 2(1-lO)-locular, the ovules 1 (2) per locule, axile-apical, unitegmic; fruit a drupe or 

berry, rarely subsamaroid; embryo straight, minute to as long as the endosperm; endosperm 

copious or moderate, rarely scanty. 

Distribution: Mainly temperate Northern 

Hemisphere, a few subtropical. 

Chemistry: The HCl/Methanol test is positive 

in Cornaceae, negative in Apiaceae. All Cornaceae have tannin. Garryaceae lack tannin. Tannins are widely distributed in plants, mainly in 

woody ones. The capacity to synthesize tannin is 

a primitive character. Garryaceae and Cornaceae 

have aucubin (iridoid)-type glycosides unknown 

elsewhere in the order. In most respects the 

order agrees well with Apiales. Results with leucoanthocyanins are different from Apiaceae. Triterpinoid saponins and/or sapogenins are reported to occur in Cornus. 

DAVIDIACEAE (Figure 1 19d).-Trees; buds 

perulate; xylem vessel perforation plates scalariform with very numerous bars; leaves alternate, 

toothed, exstipulate; inflorescences in axil of uppermost leaf on short spurs, dense globose heads 

subtended by 2 large white bracts, the plants 

andromonoecious or polygamous; flowers small, 

8 numerous, perianth 0; stamens 5-6 (1 -1 2), the 

filaments long, anthers basifixed; pollen 3-colpoidorate; 9 or bisexual flower solitary, lateralsubterminal, calyx 0 or of minute teeth according 

to some authors; petals 0; staminodes or stamens 

with short filaments inserted around the base of 

the style; disk 0; pistil 1, the carpels 6-10, style 

1 with 6- 10 linear lobes stigmatic ventrally; 

ovary 6- 1 0-locular, the ovules 1 per locule, axileapical, unitegmic, crassinucellar(?), anatropous; 

fruit a drupe with a stony sulcate endocarp and 

hard epicarp; seeds usually 1-5, embryo straight, 

as long as the endosperm, the cotyledons flat, 0.7 

the length of the embryo and 2.7 times the width 

of the radicle; endosperm moderate, fleshy. 

Composition: 1 genus, 1 species. 

Distribution: Southwestern China. 

NYSSACEAE (Figure 1 19a-c).-Trees and 

shrubs; xylem vessel perforation plates scalariform with many bars, usually more than 20; 

leaves alternate, simple, entire or few-toothed, 

exstipulate; inflorescences terminal or axillary, 

umbels, racemes, heads, or rarely the flowers 

solitary; flowers small, bisexual or the plants polygamodioecious; sepals 5; petals 5 (4-8), free, 

imbricate, or 0; stamens 8-10, often in 2 series, 

the filaments short; anthers small, dorsifixed, 

dehiscing longitudinally; pollen 3-colporate; disk 

pulvinate, epigynous in 9, central in d; pistil 1, 

the carpels 1-2, style 1 (Nyssa), or bifid (Camptotheca), the stigma(s) decurrent; ovary inferior, 

1-2-locular, the ovules 1 per locule, apical, unitegmic, crassinucellar(?), anatropous; fruit a 

drupe or subsamaroid, the seed 1; embryo 

straight, as long as the endosperm, the cotyledons 

moderately thin, 4.7 times wider than the radicle, 
NUMBER 58 229 

FIGURE 1 ~~.-NYSSACEAE: a, Nyssa sylvatica twig with 6 inflorescences, twig with 0 inflorescences, A'. aquatica d flower, 0 flower, 1,s. and C.S. of fruit, putamen; b, N. bzyora d flower, I.s. 

of bisexual flower, iV. javanica d flower, infructescence, Camptotheca acuminata bud, flower (the 

petals fallen), stamen before and after dehiscence, 1,s. of ovary; c, infructescence (after Wangerin; Sargent; Baillon, 1866- 1895). DAVIDIACEAE: d, Davidia inuolucrata inflorescence (most 

of the staniens fallen), C.S. of ovary, C.S. of fruit (after Oliver, Hemsley). GARRYACEAE: e, Garrya 

elliptica part of d inflorescence, part of 0 inflorescence, d flower, views of anther, 1,s. of ovary, 

G.fadyenii 1.5. of ovary;f; G. laurzjolia 1,s. of fruit, G. elliptica C.S. of seed, embryo (after Baillon, 

1866- 1895; Le Maout and Decaisne, 1873). 
230 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

0.7 the length of the embryo?; endosperm scanty 

or copious. 

Composition: 2 genera, 9 species. 

Distribution: Temperate and subtropical; 

eastern North America; eastern Asia from temperate China and Japan to western Malaysia and 

the Himalayas. 

GARRYACEAE (Figure 1 19ef).-Trees and 

shrubs; xylem vessel perforation plates scalariform with few bars; leaves opposite, simple, entire, exstipulate; plants dioecious; inflorescences 

catkin-like racemes or spikes; 8 flower: sepals 4, 

valvate, free; petals 0; stamens 4, alternisepalous, 

free, the filaments short; anthers elongated, basifixed, the connective produced apically; pollen 

3-colporate, the colpi rather short, narrow; 9 

flowers: sepals 0 or 2-4, minute; petals 0; disk 0 

or very imperfect; pistil 1, the carpels 2 (3), styles 

2, the stigmas decurrent; ovary inferior, unilocular, the ovules 2, apical, with an obturator, 

unitegmic, the integument very massive, crassinucellar, anatropous, the funicle rather long; 

fruit 1 -2-seeded, the testa thick, fleshy; embryo 

minute, straight, 0.3 the length of the endosperm, the cotyledons 1.1 times wider than the 

radicle, 0.5 the length of the embryo; endosperm 

copious, firm to hard. 

Composition: 1 genus, 15 species. 

Distribution: Dry habitats; western United 

States to Panama; West Indies. 

CORNACEAE (Figure 120a).-Trees, shrubs, or 

rarely herbs; xylem vessel perforation plates 

mostly scalariform with many bars, occasionally 

also simple; leaves opposite, rarely alternate, simple, mostly entire, rarely toothed, exstipulate; 

inflorescences terminal or axillary, corymbose, 

racemes, heads, umbels, rarely panicles, sometimes dichasia or pleiochasia; flowers small, bisexual or rarely the plants monoecious, polygamodioecious or dioecious; calyx minute, 4-5-lobed; 

petals 4-5, free, valvate; stamens 4-5, alternipetalous, the filaments rather short, free; anthers 

small, about as long as wide, dehiscing longitudinally, basifixed or dorsifixed; pollen usually 3- 

colporate; disk epigynous, surrounding the base 

of the style, central in 8 flowers; pistil 1, the 

carpels 2 (-4), style 1, the stigma short, apical, 

subcapitate; ovary inferior 2( 1 -4)-locular, the 

ovules 1 per locule, apical, unitegmic, usually 

crassinucellar, sometimes tenuinucellar, anatropous; fruit a drupe or berry, the seeds 4-1; 

embryo straight, 0.3-0.9 the length of the endosperm; cotyledons thin, 2-3.3 times wider 

than the radicle, 0.4-0.7 the length of the embryo; endosperm copious or moderate, fleshy. 

Composition: 12 genera, 100 species. 

Distribution: Mainly temperate Northern 

Hemisphere, some species in the tropics and subtropics. 

D IPSA CA L ES 

Herbs, shrubs, and small trees, rarely climbing; 

xylem vessel perforation plates simple, less commonly scalariform, rarely with more than 20 

bars; leaves opposite, simple or sometimes compound, sometimes lobed or divided, entire or 

toothed, exstipulate, very rarely stipulate; flowers bisexual, rarely unisexual, often zygomorphic, sometimes involucrate; sepals 5-4 

(1-3), connate, the calyx minute, sometimes pappus-like or plumose in fruit; petals 5-4 (3, 6), 

connate, the corolla sometimes gibbous or 

spurred; stamens 5-4 (1 -6), equal or sometimes 

didynamous, inserted on the corolla, the filaments free or rarely connate in pairs, anthers 

dorsifixed or basifixed; pollen 3(4)-colporate or 

-colporoidate, 3(2-6)-colpate, rarely 4-rupate; 

pistil 1, the carpels 2-3 (-8), style 1 (-several), 

the stigma(s) apical; ovary inferior, the locules 1- 

3 (-8), ovules l(-several) per locule, unitegmic, 

tenuinucellar, rarely crassinucellar; fruit a berry, 

drupe, achene, nutlet, rarely capsule; embryo 

straight, minute to nearly as long as the endosperm; endosperm copious to absent. 

Distribution: Nearly cosmopolitan, centered 

in temperate Northern Hemisphere, extending 

in the mountains of the tropics to temperate 

Southern Hemisphere. 

Chemistry: Gibbs considers that chemistry 

supports a close relation among the families of 

this order. In serodiagnostic tests Adoxa is said to 
NUMBER 58 23 1 

FIGURE 120.-cORNACEAE: a, Cornus mas floral diagram, I.s. of flower, C.S. of fruit, C. sanguinea 

pistil and calyx, 1.5. of fruit, flowering twig (after Baillon, 1866-1895; Le Maout and Decaisne, 

1873). CAPRIFOLIACEAE: b, Diemilla lonicera flower, part of corolla showing insertion of stamen, 

views of anther, stigma, Sambucus australis 6 flower, corolla and stamens of same, 0 flower; c, 

pistil with carpels opened showing the ovules, C.S. of ovary, fruit, 3 views of mericarp, I.s. of 

same showing seed; d, Lonicera caprifolium floral diagram, fruit, seed, Is. of same, L. xylosteum 

pair of flowers, L. ciliata I.s. of 2 united fruits (after Johnson; Baillon, 1866-1895; Martius, 

1840- 1906). 

react strongly with representatives of Caprifoli- CAPRIFOLIACEAE (Figure 1 POb-d).-Shrubs 

aceae, Rubiaceae, and Dipsacaceae, but not Sax- and small trees, rarely herbs, sometimes climbifragaceae. In this order sesquiterpenoids seem ing; xylem vessel perforation plates simple, also 

to occur only in Valerianaceae. Sambucus and scalariform, or exclusively scalariform with more 

Viburnum are set off chemically from the rest of than 20 bars (Diemilla, Viburnum); leaves oppothe Caprifoliaceae. site, simple, entire or toothed, rarely lobed or 
232 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

deeply divided, sometimes pinnately compound 

(Sambucus), exstipulate or rarely stipulate; inflorescences cymose, a dichasium lacking the middle 

flower or monochasium; flowers bisexual, actinomorphic or zygomorphic; sepals 5 (4), connate, 

the calyx small; petals 5 (4), connate, the lobes 

imbricate in bud, the corolla sometimes gibbous; 

stamens 5 (4), 2 in Carlemannia, Silvianthus, inserted on the corolla, rarely didynamous; anthers 

dorsifixed or basifixed; pollen usually 3-colporate or 3-colporoidate; pistil 1, the carpels 3 

(2-8), style 1 (-several), elongated or very short, 

the stigma(s) 1-5, apical; ovary inferior, the locules 1-3 (-8), the ovules 1 (-several) per locule, 

axile, unitegmic, usually tenuinucellar, sometimes crassinucellar (Viburnum), anatropous; 

fruit a berry, drupe, achene, or capsule; seeds 

often with a bony testa; embryo minute or small, 

straight, 0.1-0.35 (0.75-0.9 in Sambucus) the 

length of the endosperm; cotyledons 1-2 times 

the width of the radicle, 0.25-0.5 the length of 

the embryo; endosperm copious, fleshy. 

Composition: 13 genera, 450 species. 

Distribution: Centered in temperate eastern 

North America and eastern Asia; almost all Eurasia; cool temperate North America to temperate 

South America, in the mountains in the tropics; 

eastern Australia, New Guinea, New Zealand; 

only present in a small area of northwestern and 

eastern Africa. 

VALERIANACEAE (Figure 1 Zla).-Herbs, 

rarely shrubs or lianas, dried specimens usually 

malodorous; xylem vessel perforation plates simple, rarely also some scalariform (Centranthus 

ruber); leaves opposite, sometimes rosulate, often 

much divided, exstipulate; inflorescence mostly 

a thyrse, sometimes a head; flowers usually zygomorphic, bisexual or the plants rarely dioecious; sepals 5 (2-4 or merely a ring), connate, 

the calyx minute, but sometimes plumose in fruit; 

petals 5 (-3), connate, the lobes imbricate, the 

corolla often saccate or spurred, rarely bilabiate 

(Centranthus); stamens 4-1, inserted on the corolla, the filaments long or the anthers sessile, 

dorsifixed; pollen 3(4)-colporoidate; pistil 1, the 

carpels 3, style 1, elongated, the stigmas 1-3, 

apical; ovary inferior, 3-l-locular, the ovule 1, 

axile-subapical, unitegmic, tenuinucellar, anatropous; fruit an achene or nutlet; embryo 

straight, the cotyledons moderately thick, flattened, 0.7-0.9 the length of the embryo, 3 times 

as wide as the radicle; endosperm 0. 

Composition: 13 genera, 400 species. 

Distribution: Centered in temperate Eurasia 

and the mountains of South America; almost all 

Eurasia except the southeast; most of temperate 

America, only a small area of northern and southern Africa and the East Indies; absent from Australia. 

ADOXACEAE (Figure 12 1 b,c).-Herbs; xylem 

vessel perforation plates simple; leaves opposite, 

some radical, compound, the leaflets toothed, 

the scale leaves on the rhizome alternate; inflorescence a terminal dichasial cyme, capitate; 

flowers bisexual, actinomorphic, the lateral ones 

weakly zygomorphic, with musky smell; sepals 

2-3 (1-5), connate; petals 5 (4-6), connate, the 

corolla rotate; stamens 5 (4-6), inserted on the 

corolla, the anthers and filaments bifid; pollen 3- 

colporoidate; disk 0; pistil 1, the carpels 3-5 (2), 

styles 3-5, the stigmas apical, capitate; ovary 

semi-inferior, the locules 3-5, ovules 1 per locule, axile-subapical, unitegmic, tenuinucellar, 

anatropous; fruit a drupe with 1-5 pyrenes; embryo minute, 0.1-0.3 the length of the endosperm; cotyledons not broadened, 0.3-0.4 the 

length of the embryo; endosperm copious, hard. 

Composition: 1 genus, 2 species. 

Distribution: Temperate Northern Hemisphere, especially cool, moist regions. 

DIPSACACEAE (Figure 12 1 d).-Herbs, rarely 

subshrubs (Scabiosa spp.); xylem vessel perforation plates simple, rarely also scalariform; leaves 

opposite or rarely verticillate, simple, pinnately 

dissected, sometimes toothed or with a prickly 

margin, rarely entire, exstipulate; inflorescences 

often cymose heads, sometimes subspicate, rarely 

in verticillasters (Morina), usually involucrate; 

flowers bisexual, mostly zygomorphic, an involucel present; sepals 5-4, small, connate, sometimes divided into segments like a pappus; petals 

5-4, connate, the lobes imbricate in bud; stamens 
NUMBER 58 233 

FIGURE 12 1 .-VALERIANACEAE: a, Valerianu scandens d flower, 1.s. of flower, 0 flower, floral 

diagram, fruit, V. gilgiana corolla and stamens, C.S. of fruit, V. eichleriana fruit (after Martius, 

1840- 1906). ADOXACEAE: b, Adoxa moschatellina plant in flower, corolla and stamens of 5- 

merous flower, bud from below, 5-merous and 4-merous buds, views of half anther; c, I.s. of 

ovary, fruit and calyx, I.s. of fruit, seed, I.s. of same showing the minute embryo (after Le 

Maout and Decaisne, 1873). DIPSACACEAE: d, Dipsacus asper inflorescence, flower, I.s. of ovary 

with calyx and involucel, calyx, achene invested by involucel, C.S. of same, achene, D.fullonum 

flower, 1,s. of same (after Oliver, Lecomte). 
234 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

4 (2-3), equal or sometimes didynamous, inserted on the corolla, the filaments free or united 

in pairs; anthers dorsifixed; pollen 3(2-6)-colpate, 4-rupate, colporate (Morina), or porate; 

pistil 1, the carpels 2, style 1, elongated, the 

stigma(s) 1-2, capitate; ovary inferior, unilocular, the ovule 1, axile-subapical, unitegmic, tenuinucellar, anatropous; sometimes there is an 

increase in the number of antipodal nuclei in the 

embryo sac; fruit an achene or nutlet; embryo 

straight, 0.9 (0.3 in Morina) to as long as the 

endosperm; cotyledons moderately or markedly 

thick, 1.7-4.0 times wider than the radicle, 0.35- 

0.9 the length of the embryo; endosperm usually 

scanty, sometimes moderate, fleshy. 

Composition: 10 genera, -250 species. 

Distribution: Centered in the Mediterranean 

region and Near East; temperate Eurasia and 

India; northern, eastern, southern, and a small 

part of western Africa. 

RUTALES 

Trees and shrubs, less commonly herbs, rarely 

climbing, the bark often bitter; xylem vessel perforation plates simple, rarely also scalariform, 

very rarely with more than 20 bars; secretory 

canals, cavities or cells containing aromatic oil or 

resin often present; leaves alternate, less commonly opposite, usually pinnately compound, 

rarely simple, the rachis sometimes winged, the 

leaflets entire or toothed, sometimes gland-dotted, exstipulate, rarely stipulate; flowers small, 

often abundant, bisexual or unisexual; sepals 

5-3 (-8, 0), free or shortly connate; petals 5-3 

(-14, O), free or rarely connate, rarely adnate to 

a stamina1 tube; stamens 3-10 (2 to -loo), usually in 1-2 series; staminodes rarely present; filaments free, less commonly connate, rarely inserted on the corolla, sometimes with a basal 

ligular scale; anthers dorsifixed or basifixed, occasionally apiculate; pollen 3(2-8)-colporate, less 

frequently ruporate, poroid or rugoid; glandular 

disk usually surrounding the base of the ovary; 

pistils 1 (-6), the carpels 2-5 (1-20), styles 1-5 

(0-1 2), sometimes basal, stigmas apical, very 

rarely ventral; ovary superior, very rarely semiinferior or inferior, 1 -5(-20)-locular, the ovules 

1-2 (-several) per locule, axile, less commonly 

basal or subapical, very rarely parietal; fruit a 

berry, drupe, drupelets, capsule, cocci or samara, 

the seeds sometimes winged; embryo straight, 

arcuate, folded or convolute, usually as long as 

the endosperm; endosperm copious to absent. 

Distribution: Mostly pantropical and -subtropical, extending into warm temperate regions, 

in various habitats. 

Many triterpenes occur in the order. The Rutaceae have many monoterpenes in 

many genera, and several sesquiterpenes occur 

in several genera. Citrus oil terpenes inhibit some 

insects. Burseraceae and Meliaceae also have various terpenes. The Rutaceae stand out from the 

rest of the order as a maker of coumarins and 

alkaloids (of at least 10 groups). Otherwise the 

order is not alkaloid-rich. Rutaceae have some 

lignans. Chromones in Spathelia (Rutaceae) and 

Neochamaelea (Cneoraceae) may be indicative of 

relationship of their families. Degraded triterpenes with a seco-ring A are commonly found in 

Rutaceae. The only other family with these compounds is Meliaceae. Mangiferin occurs in Anacardiaceae and Melianthaceae (Bersama). 

RUTACEAE (Figure 122a-d).-Trees and 

shrubs, rarely herbs, the bark sometimes bitter; 

xylem vessel perforation plates simple, rarely also 

a few scalariform plates, rarely some multiperforate plates in a few species; aromatic oil or 

resin secreting cavities or cells nearly always present in the tissues; leaves alternate or opposite, 

very rarely verticillate, compound or simple, the 

rachis sometimes winged, entire or toothed, 

mostly with pellucid dots, exstipulate; inflorescences terminal or axillary, usually cymose, sometimes racemes, rarely a spike or head, rarely 

epiphyllous (Erythrochiton); flowers bisexual, 

rarely the plants polygamo-dioecious (Zanthoxy 

Eum), dioecious (Skimmia) or monoecious (Empleurum); sepals 4-5 (0, 3), free or basally connate; petals 4-5 (0, 3), free or rarely connate; 

stamens 4-10 (Z), rarely more (50-100 in Clymenia), obdiplostemonous or rarely diplostemonChemistry: 
NUMBER 58 235 

e 

FIGURE I22.-RUTACEAE: a, Zanthoxylum chiloperone 6 flower, 1,s. of same, views of stamen, ? 

flower, pistil, 1,s. of ? flower, C.S. of ovary; b, Pilocarpus macrocarpus flower, Is. of same; c, pistil 

and calyx, P. grandzjlorus C.S. of ovary, P. macrocarpus seed; d, Angostura odoratissima flower, 

1.5. of mericarp, fruit, 1,s. of flower, views of stamen (after Martius, 1840-1906). CNEORACEAE: 

e, Cneorum tricoccum flowering and fruiting shoot, part of same, floral diagram, flower, 1,s. of 

same;f; essential organs, fruit, I.s. of a fruiting carpel, seed, 1,s. of same (after Baillon, 1866- 

1895; Le Maout and Decaisne, 1873). 

ous, or in several series, the filaments free, rarely 

connate or inserted on the base of the corolla, 

sometimes with bifid pubescent scales basally 

(Dictyoloma); anthers about as long as wide, the 

connective sometimes apiculate, sometimes apically glandular; pollen 3(2-8)-colporate; disk usually present below the ovary, or adnate to the 

calyx tube; gynophore sometimes present; pistils 

1-5, sometimes the styles connate and ovaries 

free, carpels 2-5 (1-1 6; Clymenia IS), the styles 
236 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

1-5, free or connate, often basal, stigma(s) apical; 

ovary superior, rarely semi-inferior or inferior, 

2-5( 1 - 16)-locular, the ovules 1 -several per locule, axile or very rarely parietal (Feronia), bitegmic, crassinucellar, anatropous or semi-anatropous; fruit a berry, drupe or rarely 1-4 drupelets, a loculicidal or rarely septicidal capsule, or 

cocci, rarely a samara, sometimes the endocarp 

separates from the epicarp; embryo straight or 

arcuate; linear, or the radicle invested, 0.9-1 .0 

times the length of the endosperm; cotyledons 

thin or thick, 1-4 times the width of the radicle, 

0.4-0.9 the length of the embryo; endosperm 

copious to 0, fleshy. 

Composition: 

Distribution: Mostly pantropical, extending 

into warm temperate regions; centered in Australia and South Africa; United States to Argentina; southern Eurasia to Australia and New Zealand; Africa and Madagascar. 

CNEORACEAE (Figure 122ef). Shrublets with 

oil cells in the bark; xylem vessel perforation 

plates simple; leaves alternate, simple, entire, 

with secretory cells containing oil or resin, exstipulate; inflorescences adnate to the petiole, cymes, 

corymbs or the flowers solitary; flowers bisexual, 

the parts hypogynous; sepals 3 (-4), shortly 

connate; petals 3 (-4), free; stamens 3(-4), free, 

the filaments elongated; anthers dorsifixed, 

about as long as wide; pollen 3(4-6)-colporate; 

receptacle elongated, disk present; pistil 1, some150 genera, - 1000 species. 

sional multiple perforations in Ailanthus; secretory canals in some genera, secretory cells containing oil, resin, or mucilage are less common; 

leaves alternate, or rarely opposite or absent, 

compound or rarely simple, the leaflets entire or 

toothed, never gland-dotted, but some glandular 

teeth in Ailanthus, exstipulate or rarely stipulate; 

axillary and terminal panicles, racemes or cymose 

spikes; flowers rarely bisexual, the plants usually 

dioecious or polygamous, the parts hypogynous; 

sepals 3-5 (-8 in Holacantha), connate, rarely 

free; petals 3-5 (7-8 in Holacantha), free, rarely 

connate or 0; stamens 3- 10, rarely more (1 2- 18 

in Castela, Quassia sp.), mostly obdiplostemonous, rarely opposite the petals when equal to 

them in number, the filaments long, filiform, 

free, often with a basal ligular scale, sometimes 

5 staminodes present; anthers oblong, dorsifixed; 

pollen mostly 3-colporate, also 3-colporoidate, 3- 

colpate(?); disk under the ovary annular or cupular; gynophore sometimes present; pistils 1-5, 

the carpels (1) 2-5 (8 in Castela), often the ovaries 

free and the styles or stigmas connate; styles 1- 

5, often basal, the stigmas apical, and sometimes 

ventral (Castela); ovary l-5-locular, the ovules 1 

(2-several) per locule, axile, bitegmic, crassinucellar, anatropous, amphitropous or semi-anatropous; fruit drupelets or drupe, schizocarp, capsule or fruitlets samaras; embryo straight or arcuate, as long as the endosperm; cotyledons thin 

or thick, 3-5.8 times wider than the radicle, 0.8- 

times with septa1 glands, the carpels 3 

stigmas apical; ovary 3(-4)-locular7 the ovules 

bitegmic, campylotropous or amphitropous, 2 

per locule, axile; fruit 1-4 drupaceous cocci; seed 

arcuate, the embryo arcuate, peripheral, 1-2 

times the length Of the endosperm; cotyledons 

not broadened, 0.5-0.7 the length Of the embryo; endosperm moderate or copious, fleshy. 

(-4)9 style 3(-4)-10bed and FIGURE 123.-sIMAROUBACEAE: a, Suriana maritima floral 

diagram, 1,s. of pistil, pistil with basilar style, embryo, Quassia 

sp. floral diagram, Q. nzgrescens views of stamen, pistil; b, 

flower, I.s. of same, C.S. of ovary, fruit, I.s. of same, 1,s. of 

seed (after Le Maout and Decaisne, 1873; Martius, 1840- 

1906). BURSERACEAE: c, Protium brasiliense flower, Is. of 

same, C.S. of ovary and calyx, pistil and some of the stamens, 

views of stamen, view of seed, I.s. of same, fruit (above) 

(after Martius, 1840- 1906). ANACARDIACEAE: d, Spondias 

macrocarpa flower, calyx and C.S. of ovary, views of petal, 

pistil, 1.5. Of same, Tapirira marchandii essential organs, IS. 

of pistil, I.s. of fruit; e, Schinus lentiscifolius essential organs, 

1,s. and C.S. of ovary; f; Anacardium pumilum d flower, 0 

flower; g, views of stamen, fruit and pedicel, Is. of fruit, 

views of embryo, androecium of d flower, essential organs, 

I.s. of pistil (after Martius, 1840-1906). 

Composition: 2 genera, 3 species. 

~i~t~ib~ti~~: Western Mediterranean region, 

SrMARoUBACEAE (Figure 23a9b)*-Trees and 

shrubs, usually very bitter; xylem vessel perforation plates exclusively simple, except for occaCanary Islands; Cuba. 

238 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

0.9 the length of the embryo, embryo not divided 

in Aeschrion; endosperm moderate, scanty or 0, 

fleshy. 

Composition: 20 genera, 120 species. 

Distribution: Pantropical and -subtropical, extending into warm temperate regions in Japan 

and Korea. 

BURSERACEAE (Figure 123c).-Trees and 

shrubs with resin or aromatic oil in lysigenous or 

schizogenous canals; xylem vessel perforation 

plates simple; leaves alternate, rarely opposite, 

pinnately compound or decompound, the rachis 

often winged, rarely simple (I-foliolate), toothed, 

usually not punctate, exstipulate; inflorescences 

terminal or axillary racemes or panicles; flowers 

bisexual or the plants polygamodioecious, minute, the parts hypogynous, rarely perigynous; 

sepals 3-5, basally connate; petals 3-5 (0), free 

or rarely connate; stamens 3-10, in 1-2 series, 

obdiplostemonous, the filaments free (connate in 

Canarium spp.); anthers slightly longer than 

wide; pollen 3-colporate, rarely 3-colporoidate; 

disk surrounding base of ovary as in Rutaceae; 

pistil 1, the carpels 2-5, the style 1, short, stigma 

usually 2-5-lobed, apical; ovary superior, 2-5- 

locular, the ovules 2 (1) per locule, axile, 2(1?)- 

tegniic, crassinucellar, semi-anatropous or campylotropous; fruit a berry or drupe, sometimes a 

tardily dehiscent capsule; embryo usually folded 

or convolute, rarely straight; cotyledons thin, or 

thick and plano-convex, 12-1 5 times wider than 

the radicle, 0.7-0.9 the length of the embryo, 

the radicle sometimes invested; endosperm 0. 

Composition: 18 genera, 550 species. 

Distribution: Pantropical in a variety of habitats from rain forest to desert and mangrove; 

best represented in tropical America and northeastern Africa. 

ANACARDIACEAE (Figures 123d-g, 124a-e).- 

Trees and shrubs, rarely climbers, often with 

resin-canals in the phloem; xylem vessel perforation plates exclusively simple in most genera, a 

few scalariform plates in some species, fewer than 

20 bars except in one genus; leaves alternate, 

very rarely opposite, compound, the rachis sometimes winged, or simple, often toothed, exstipulate, very rarely obscurely stipulate; inflorescences terminal or axillary panicles or thyrses; 

flowers bisexual or the plants polygamo-dioecious or dioecious, the parts hypogynous, rarely 

perigynous or epigynous; sepals 5 (3-7), free or 

connate; petals 5 (0, 3-7), free or rarely basally 

connate; stamens 5 (3 to -15), the filaments 

elongated, free or rarely connate basally (Anacardium), sometimes all but one staminodal; anthers about as long as wide or slightly longer than 

wide, dorsifixed, rarely basifixed (Buchanania), 

occasionally apiculate; pollen (2) 3-colporate, less 

frequently 4-ruporate or 3-8?-poroid, colpoid, 

rugoid or foraminoid; disk usually present 

around the base of the ovary; gynophore rarely 

present; pistils 1 (4-6), the carpels 1-3 (5-12 

Pleiogynium), the styles 1 (5-1 2), often distant, 

rarely basal, the stigmas apical; ovary unilocular 

(2-1 2), usually superior, rarely inferior, the 

ovules 1 per pistil or carpel, axile, basal or subapical, 1 -2-tegmic, crassinucellar, anatropous, the 

funicle often long; fruit mostly a drupe; embryo 

bent or arcuate, 1.5-1.8 times longer than the 

endosperm; cotyledons thin or moderately thick, 

4.5-6.5 times wider than the radicle, 0.6-0.9 the 

length of the embryo; endosperm scanty or 0, 

fleshy. 

Composition: 70 genera, 600 species. 

Distribution: Pantropical, extending into temperate America, Africa and Eurasia; equally well 

represented in South America, Africa, and Malaysia. 

MELIACEAE (Figure 125a,b).-Trees and 

shrubs, very rarely subshrubs, with secretory cavities in the cortex and pith of a few genera, 

sometimes bitter, wood scented; xylem vessel 

perforation plates simple; leaves alternate, rarely 

opposite, mostly pinnately compound or decompound, the leaflets entire or toothed, more rarely 

simple, rarely glandular punctate, with secretory 

cells containing resin, exstipulate; inflorescences 

thyrses, panicles, racemes or umbels, sometimes 

cauliflorous; flowers mostly bisexual, rarely the 

plants dioecious; sepals 5 (3-4), free or shortly 

connate; petals 5 (3-14), free or rarely shortly 

connate, sometimes adnate to the stamina1 tube; 
NUMBER 58 239 

Q 

b 

e 

C 

FIGURE 124.-ANACARDIACEAE: a, Amphipterygzum adstringens part of d inflorescence, d flower, 

a pair of 0 flowers, one with the ovary opened to show the solitary ovule, a pair of 0 

inflorescences, one with 1 flower the other with 2; b, C.S. through 2 fruits, seed, embryo; c, 

Orthopterygzum huaucuz stamen, fruit; d, Dobinea delavuyi floral diagram of d flower, d flower, 

1,s. of same, petal, pistillode, part of 6 inflorescence, 0 inflorescence; e, 0 flower on bract, I.s. of 

same, fruit on bract, embryo (after Hemsley, Franchet). 
240 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

d p 

FIGURE 1 P!j.-%fELIACEAE: a, i2lelia azedarach floral diagram, flower, same with part of perianth 

removed to show the stamina1 tube, 1,s. of flower (upper part of petals cut off), views of anther, 

C.S. of ovary; b, part of infructescence, C.S. of fruit, seed, 1.s. of same, embryo, Trichilia 

richardiana dehiscent fruit, seed with aril (after Martius, 1840-1 906). AKANIACEAE: c, Akania 

hillii small part of inflorescence, floral diagram, bud, I.s. of flower; d, views of anther, 1,s. of 

ovary, small part of leaf, dehiscent fruit (after Stapf, Maiden). 

stamens 8 or 10 (3-25), the filaments mostly half 

way to completely connate, rarely free, (Cedrela, 

Toona), hypogynous; anthers about as long as 

wide, dorsifixed or basifixed, rarely apiculate; 

pollen 3-5-colporate (slightly ruporate, or porate); usually a disk surrounding the base of the 

ovary; pistil 1, the carpels 3-5 (2-20), style 1 or 

stigma sessile, stigma apical, often disciform or 

capitate, sometimes lobed; ovary superior 3-5( 1- 

20)-locular, the ovules bitegmic, crassinucellar, 

anatropous, campylotropous or orthotropous, 2 

(1-12) ovules per locule, axile; fruit a capsule, 
NUMBER 58 24 1 

berry, or rarely drupe, the seeds often winged; 

embryo straight, as long as the endosperm; cotyledons thin or thick, sometimes conferruminate, 

3-3.2 times wider than the radicle, 0.8-0.9 the 

length of the embryo; endosperm moderate or 

0. 

Composition: 

Distribution: Pantropical and -subtropical, extending into warm temperate regions, particularly common as understory trees in rain forest. 

50 genera, - 1400 species. 

SAPINDALES 

Trees and shrubs, sometimes lianas usually 

with tendrils, rarely subherbaceous; xylem vessel 

perforation plates simple, scalariform with few 

to many bars, or both; tissue or sap sometimes 

with resin or latex-like secretions; leaves alternate, less commonly opposite, usually compound, 

sometimes simple, usually exstipulate; flowers 

bisexual or unisexual, occasionally zygomorphic, 

the parts hypogynous to slightly perigynous; sepals 5 (3-6), usually free, sometimes basally connate; petals 5 (0, 3-6), free, rarely shortly connate, sometimes with a basal ligulate scale; stamens 8 (4-20), usually in 2 series, the filaments 

elongate, free, inserted internal or external to a 

glandular disk, very rarely basally adnate to the 

petals; anthers dorsifixed, sometimes apiculate; 

pollen 3(2-4)-colporate, 3-colpate, 3-4-porate, 

rarely 4-rugorate or 4-ruporate; glandular disk 

usually present; pistil 1, the carpels 2-3 (-5), 

styles 1-2 (-4), the stigmas apical, less frequently 

decurrent; ovary superior, 3( 1 -5)-locular, the 

ovules 1-2 (-several) per locule, axile, rarely 

parietal, subbasal or subapical; fruit usually a 

loculicidal capsule, less often a schizocarp, berry, 

drupe, samara or nut, the seeds sometimes arillate; embryo large, often bent, folded or coiled, 

sometimes straight; endosperm usually absent, 

rarely copious. 

Distribution: Pantropical, extending into temperate regions. 

Chemistry: It is obvious that Aceraceae, Sapindaceae, and Hippocastanaceae are chemically 

closely related. 

AKANIACEAE (Figure 125c,d).-Trees; xylem 

vessel perforation plates simple, also rarely scalariform with numerous bars in the young stem; 

scattered secretory cells in the leaf and cortex of 

the stem; leaves alternate, pinnately compound, 

serrate, exstipulate; inflorescences axillary or 

cauliflorous panicle or thyrse; flowers bisexual, 

the parts hypogynous; sepals 5, connate; petals 

5, free; stamens 8 (9), in 2 series, one of 5 the 

other of 3, the filaments elongated, free; anthers 

about as long as wide, apiculate; pollen 3-colpate; 

disk absent; pistil 1, the carpels 3, style 1, stigmas 

apical, not enlarged; ovary 3-locular, the ovules 

2 per locule, axile, bitegmic, crassinucellar, anatropous or semi-anatropous; fruit a loculicidal 

capsule, the seeds not arillate; embryo large, 

straight, the cotyledons thick, flattened, the 

radicle very short; endosperm copious, fleshy and 

oily, smelling of bitter almond. 

Composition: 1 genus, 1 species. 

Distribution: Eastern Australia. 

SAPINDACEAE (Figure 126b-e).-Trees, 

shrubs, and lianas usually with tendrils, rarely 

subherbaceous; xylem vessel perforation plates 

simple; tissues usually with resin or latex-like 

secretions; leaves alternate or very rarely opposite, pinnately or ternately compound, or simple, 

the leaflets entire or toothed, with secretory cells, 

rarely stipulate (in the climbing species); inflorescences terminal or axillary cymes, often cincinni, or panicles; flowers actinomorphic or 

zygomorphic, rarely bisexual, usually the plants 

polygamodioecious, the parts hypogynous; sepals 

5 (3-4), free or rarely connate; petals 5 (0, 3, 4), 

free, often with a basal ligulate scale; stamens 8 

(4, 5, 10, 12-20), inserted internal to the disk, 

the filaments elongated, free; anthers about twice 

as long as wide, sometimes apiculate, dorsifixed 

near the base; pollen 3(2-4)-colporate or 3-4- 

porate; disk usually present, unilateral or annular; pistil l, the carpels 3 (2-4), style l (2-4), 

stigmas terminal, lobed or punctiform; ovary 

3( 1-4)-locular, rarely incompletely so, the ovules 

bitegmic, crassinucellar, anatropous, campylotropous or semi-anatropous, 1 (2-several) per locule, axile or rarely parietal; fruit a capsule, berry, 
242 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 126.-MELIANTHACEAE: a, Melianthus cosmus inflorescence and leaf, M. major flower, 

same with 1 sepal reflexed, I.s. of flower, floral diagram, pistil, M. cosmus dehiscent fruit, seed 

and 1,s. of same (after Baillon, 1866-1895; Schluss). SAPINDACEAE: b, Cupania scrobiculata d 

flower, disk and base of sepals, 0 flower with petals and several stamens removed, I.s. and C.S. 

of pistil; c, fruit, dehiscent fruit, arillate seed; d, Serjania comata inflorescence and leaf, S. 

leptocarpa 6 flower, calyx showing insertion of stamens and disk (pistillode above), views of 

stamen, floral diagram; e, 2 views of lateral petal, 2 views of superior petal, S. comata fruit, 

niericarp, 1,s. of part of same showing section of seed and embryo (after Martius, 1840-1906). 
NUMBER 58 243 

nut, samara, drupe or schizocarp, the seeds often 

arillate; embryo large, the cotyledons thin, often 

plicate or coiled, or thick and plano-convex, 0.6- 

0.9 the length of the embryo; endosperm 0. 

Composition: 140 genera, 1500 species. 

Distribution: Pantropical, especially America, 

relatively few in temperate United States, South 

Africa, Japan, Australia, and New Zealand. 

MELIANTHACEAE (Figure 126a).-Shrubs and 

small trees; xylem vessel perforation plates simple; leaves alternate, pinnately compound, the 

leaflets toothed, stipulate; inflorescence a raceme; flowers bisexual, rarely the plants polygamodioecious, zygomorphic; sepals 5 (4)) free or 

basally connate, unequal, one sometimes spurred 

or saccate; petals 5 (4), free, subperigynous; stamens 4-5, the filaments long, inserted internal 

to the disk, free or basally connate; anthers about 

as long as wide, dorsifixed; pollen 3-colporate, 

rarely also 4-rugorate; disk unilateral; pistil 1, 

the carpels 4-5 (-2); style 1, the stigma apical, 

not enlarged; ovary superior, 4-5(-2)-locular, 

the ovules 1-5 per locule, bitegmic, crassinucellar, anatropous; fruit a loculicidal capsule, the 

seeds sometimes arillate; embryo straight, 0.5 the 

length of the endosperm; cotyledons not broadened, 0.5 the length of the embryo; endosperm 

copious. 

Composition: 2 genera, 15 species. 

Distribution: Tropical to temperate southern 

Africa. 

HIPPOCASTANACEAE (Figure 127a,b).-Trees 

and shrubs; xylem vessel perforation plates usually simple, also occasionally scalariform; secretion cells in the vegetative organs; leaves opposite, palmately compound, the leaflets serrate or 

entire, exstipulate; inflorescences terminal 

thymes, racemes or cymes (cincinni); flowers zygoniorphic, bisexual or the plants polygamomonoecious; sepals (4) 5, free or basally connate; 

petals (4) 5, free; stamens (5)-8, the filaments 

free, elongated, inserted interior to the disk; 

anthers about as long as wide, dorsifixed; pollen 

3-colporate; disk solid, annular or unilateral; pistil 1, the carpels 3, style 1, stigma apical, punctiform; ovary superior, 3-locular, the ovules 2 per 

locule, axile, bitegmic, crassinucellar, anatropous 

or anacampylotropous; fruit a loculicidal capsule, 

the seeds large, not arillate; embryo large, 

curved, the cotyledons conferruminate, much 

wider than the radicle (10 x ?), 0.8 the length of 

the embryo; endosperm 0. 

Composition: 2 genera, 15 species. 

Distribution: Temperate eastern United 

States, California; southern Mexico to northwestern South America; southeastern Europe, 

disjunct in northern India, China, and Japan. 

STAPHYLEACEAE (Figure 127c,d).-Trees and 

shrubs; xylem vessel perforation plates scalariform, usually with 20-30 bars, but up to 50 in 

species of Turpinia ; leaves opposite or alternate, 

trifoliolate or pinnately compound, rarely unifoliolate, the leaflets serrate, stipulate rarely 

exstipulate; inflorescences terminal or axillary 

panicles or racemes; flowers bisexual or the 

plants polygamous, rarely dioecious, actinomorphic; sepals (3-) 5, free or connate, petals 

(3-) 5, free; stamens 5, free, the filaments elongated, inserted on or external to the disk; anthers 

about as long as wide, dorsifixed; pollen 3(2-4)- 

colporate or 4-ruporate; disk hypogynous or 0 

(Tapiscia); pistil 1, the carpels 3 (2-4), styles 3- 

1, elongated, stigmas apical, punctiform; ovary 

superior, 3-2-locular, the ovules bitegmic, crassinucellar, anatropous sometimes becoming subcampylotropous, 1 -several (1 2) per placenta, axile or subbasal; fruit a membranous apically dehiscent capsule, follicles, berry or drupe, the 

seeds often truncate basally; embryo straight, as 

long as the endosperm; cotyledons thin, flat, 4.4 

times wider than the radicle, 0.8 the length of 

the embryo; endosperm moderate, fleshy. 

Composition: 5 genera, 50 species. 

Distribution: Southeastern Canada, eastern 

United States, California, Europe, and Asia Minor (Staphylea); Japan, China, and Indochina 

(Euscaphis); Sri Lanka, Japan, China to Malaya, 

tropical America (Turpinia); West Indies, Colombia, Peru (Huertea); China (Tapiscia). 

ACERACEAE (Figure 127e).-Trees and 

shrubs; xylem vessel perforation plates simple, 

rarely also foraminate, the sap sometimes milky; 
244 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 127.-HIPPOCAsTANACEAE: a, Aesculus hippocastanurn flower, 1.s. of same, floral diagram, stamen, pistil, l.~. of same, dehiscent fruit, seed and l.~. of same; 6, A. pavza flower, Billia 

hippocastanurn flower, petal, A. hippocastanurn petal (after Baillon, 1866-1 895; Le Maout and 

Decaisne, 1873; Pax). STAPHYLEACEAE: c, Staphylea pinnata flower, I.s. of same, views of anther, 

fruit partly opened to show seeds, flowering twig; d, I.s. of pistil, pistil, views of seed, 1,s. and 

C.S. of same (after Pax). ACERACEAE: e, Acer pseudoplatanus bisexual flower, 1,s. of same, I.s. of 

d flower, fruit separating into mericarps, embryo, A. negundo S flower, 1,s. of same, I.s. of 0 

flower (after Le Maout and Decaisne, 1873; Pax). 

leaves opposite, usually simple, lobed or toothed, 

sometimes trifoliolate or pinnately compound, 

sometimes with elongated laticiferous cells, exstipulate; inflorescences terminal or axillary panicles, racemes, corymbs, spikes, umbels or fascicles; plants polygamomonoecious or dioecious; 

sepals 5 (4), free or rarely basally connate; petals 

5 (4 or 0), free; stamens 8 (4-lo), the filaments 

free, elongated, inserted external to, on, or internal to the disk; anthers about as long as wide, 
NUMBER 58 245 

dorsifixed, rarely apiculate; pollen 3-colporate, 

3-colpate, or 3-colporoidate; disk annular, lobed 

or separate glands, rarely absent; pistil 1, the 

carpels 2, rarely 3-5 in some flowers, styles 2, 

elongated, free or basally connate, the stigmas 

decurrent ventrally or papillose all around (Acer 

negundo); ovary superior, bilocular, the ovules 

bitegmic, crassinucellar, anatropous, semi-anatropous, or orthotropous, sometimes becoming 

campylotropous, 2 (1) per locule, axile; fruit a 

samaroid schizocarp; embryo coiled or bent, 

elongated, the cotyledons thin, irregularly 

folded, 0.6 the length of the embryo, 4 times 

wider than the radicle; endosperm 0. 

Composition: 2 genera, 150 species. 

Distribution: Temperate Northern Hemisphere and mountains of the tropics, centered in 

China, Japan, and eastern North America. 

SABIACEAE (Figure 128a-c).-Trees and 

shrubs, often scandent, the leaves and flowers 

sometimes with obscure reddish glandular dots; 

xylem vessel perforation plates only simple or 

also scalariform, usually with few bars, but up to 

30 in some species, occasionally reticulate or 

forminate; leaves alternate, simple or pinnately 

compound, entire or toothed, exstipulate; inflorescences axillary, terminal or both, panicles, 

rarely racemes, or the flowers solitary; flowers 

bisexual or the plants polygamodioecious; sepals 

5 (4-6), basally connate; petals 5 (4-6), imbricate, 

free or basally connate, sometimes differing in 

size (Meliosma); stamens 5 (4-6), sometimes 3 of 

these staminodal (Meliosma), opposite the petals, 

free or basally adnate to the petals; anthers about 

as long as wide, sometimes didymous, dorsifixed 

or adnate, the connective sometimes enlarged; 

pollen 3-colporate (-colporoidate); disk annular 

or cupular, sometimes lobed or dentate, surrounding the base of the ovary; pistil 1, the 

carpels 2 (3), the styles enlongated or short, 

loosely connate or free, the stigma(s) apical, minute, rounded or capitate; ovary superior, often 

compressed, 2(3)-locular, the ovules 2 (1) per 

locule, axile, sometimes subapical, unitegmic, the 

integument failing to grow over the nucellus, 

crassinucellar, semi-anatropous; fruit a drupelet 

or schizocarp of 2 drupaceous or dry mericarps, 

sometimes 1 aborted, the seeds 1 or 1 per locule; 

embryo large, the cotyledons flat, fairly thick, 

usually folded or contorted, 0.2-0.5 the length 

of the embryo, 3.6-5.0 times wider than the 

radicle, the radicle cylindrical, bent against the 

cotyledons; endosperm very scanty or absent. 

Composition: 4 genera, 80 species. 

Distribution: Tropical America and tropical 

eastern Asia, extending into temperate Japan and 

Korea. 

GERANIALES 

Herbs and shrubs, less commonly trees, sometimes climbing; xylem vessel perforation plates 

simple, rarely also scalariform with fewer than 

25 bars; leaves alternate, less often opposite or 

whorled, simple or compound, entire, toothed, 

dissected or lobed, stipulate or exstipulate; flowers bisexual, rarely unisexual, actinomorphic or 

zygomorphic, sometimes 2-3-morphic, ie., the 

stamens and pistils varying in relative length in 

the same species; sepals 5 (3, 4), free or basally 

connate, one rarely spurred, rarely colored, 

rarely with large glands on the dorsal surface; 

petals 5 (0, 2, 4), free or shortly connate, rarely 

the laterals connate, rarely with a ligular attachment, often convolute in bud; stamens usually 

twice as many as the petals, sometimes the same 

number, rarely 3 times as many, sometimes 1-2 

series staminodal, often obdiplostemonous, the 

filaments free or basally connate, very rarely 

higher or in bundles; anthers dorsifixed, rarely 

basifixed, very rarely connate; pollen 3 (2-4)- 

colpate, 3(4)-colporate, oligo-polyforate, rarely 

6- 12-rugorate, 4-6-rupate or ruporate, or nonaperturate; glandular disk or separate glands 

often present; pistils 1, very rarely 5 (1-8), the 

carpels 3, 5 (1-12), styles 1-5, the stigmas apical, 

less commonly decurrent ventrally, rarely subsessile; ovary superior, 3, 5 (l-I2)-locular, the 

ovules 1-2 (-several) per locule, axile, rarely 

subbasal; fruit a schizocarp, capsule, less commonly a berry, drupe or samara, rarely follicular, 

the seeds sometimes arillate; embryo large, 
246 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

b 

FIGURE 128.-sABIACEAE: a, Sabia latzjCLia fruit with 2 mericarps (reduced), S. gracdis axillary 

inflorescences, S. latijdiolia fruit with 1 mericarp, S. gracilis part of inflorescence, flower, petal 

with its superposed stamen, stamen, pistil with part of calyx and disk, pistil with part of ovary 

removed to show the ovules and C.S. of ovary; b, Meliosrna arnottzana bud, flower, I.s. of same, 

petal with appendage, views of stamen, pistil and disk; c, Ophiocaryon heterophyllus floral diagram, 

stamen, pistil, 1.s. of fruit, seed, embryo, Is. of same (after Hemsley; Stapf; Baillon, 1866-1895; 

Martius, 1840-1906). 

straight, arcuate or folded; endosperm absent to 

copious. 

Distribution: Cosmopolitan in various habitats. 

Chemistry: According to Hegnauer presence 

of mustard-oil glucoside and seeds with amyloid 

and eruca acid in Tropaeolaceae and Limnanthaceae strongly indicates that they are related. Zygophyllaceae also produces mustard-oil glycosides. 

GERANIACEAE (Figure 1 29a-c).-Herbs and 

shrubs; xylem vessel perforation plates typically 

simple, occasionally reticulate or scalariform; 

leaves alternate or opposite, simple, dissected or 

compound, lobed or toothed, stipulate; inflorescences axillary, cymose, subumbellate, or the 

flowers solitary; flowers bisexual, actinomorphic 

or slightly zygomorphic; sepals 5 (4), free or 

basally connate, one sometimes spurred; petals 5 

(2-4) very rarely 0 (Rhynchotheca); stamens in 1- 

3 series of 5 (4) each, obdiplostemonous, the 

filaments elongated, usually connate basally, 

rarely connate in 5 bundles of 3 each (Monsonia); 

anthers about as long as wide, dorsifixed, rarely 

basifixed; pollen (2) 3-colpate, 3(4)-colporate or 

nonaperturate; glands usually present, alternate 

with the stamens; pistil 1, the carpels 5 (-3), styles 

1 (-5), the stigmas apical, decurrent ventrally; 
NUMBER 58 247 

Y 

/I 

d 

FIGURE 1 ~~.--GERANIACEAE: a, Geranium sanguineum floral diagram, flower, 1,s. of same 

(perianth partly removed), dehiscent fruit; b, Pelargonium inguinans floral diagram, 1,s. of 

flower, dehiscent fruit; c, Geranium albicans calyx, petal, androecium laid open, pistil, fruit, 

seed and 1,s. of same showing the embryo, Biebersteinia multzjida pistil with 1 carpel opened to 

show the ovule, views of a mericarp (above), I.s. and C.S. of a mericarp, embryo (after Baillon, 

1866-1 895; Martius, 1840-1 906; Schnizlein). VIVIANIACEAE: d, Vzviania rosea petal, stamens 

and glands, anther, pistil; e, C.S. and I.s. of ovary, I.s. of fruit, I.s. of seed, flowering stem (after 

Hutchinson, 1973). 
248 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

ovary superior, 5(-3)-locular, the ovules 1-2, 

rarely more, per locule, axile, bitegmic, crassinucellar, anatropous, campylotropous or amphitropous; fruit a schizocarp, often beaked; embryo 

folded, the cotyledons 0.5 the length of the embryo, 3.5 times wider than the radicle; endosperm 0 or sparse (Biebersteinieae). 

Composition: 10 genera, 760 species. 

Distribution: Principally temperate and subtropical; temperate North America to Argentina; 

Eurasia except the arctic; Africa except the Sahara desert; eastern and southern Australia, New 

Zealand. 

VIVIANIACEAE (Figure 129d,e).-Subshrubs 

and herbs; xylem vessel perforation plates simple; 

leaves opposite or verticillate, simple, entire, 

crenate or dentate, exstipulate; inflorescences 

axillary and terminal, subfasciculate to cymosepaniculate; flowers bisexual, the parts hypogynous; sepals (4-) 5, free or connate; petals (4-) 5 

or 0, free; stamens 10 (8), free, the filaments 

filiform, the anthers slightly longer than wide, 

dorsifixed; pollen polyforate; disk glands 5, alternate with the petals; pistil 1, the carpels (2-) 3, 

the style I, stigmas (2-) 3, linear; ovary (2-)3- 

locular, not beaked, the ovules 1-2 per locule, 

axile; fruit a loculicidal capsule; embryo linear, 

arcuate or coiled, subperipheral, 1.6 times the 

length of the endosperm; cotyledons not broadened, 0.4 the length of the embryo; endosperm 

copious, fleshy. 

Composition: 2 genera, 30 species. 

Distribution: Southern Brazil, Chile. 

LIMNANTHACEAE (Figure 1 3Oa,b).-Herbs; 

xylem vessel perforation plates simple; leaves 

alternate, pinnatisect, exstipulate; inflorescence 

terminal in a group of 3 flowers and flowers also 

solitary in leaf axils; flowers bisexual, actinomorphic; sepals 5-3, free, valvate; petals 5-3, 

free, sometimes contorted; stamens 10 or 6, the 

filaments elongated, free; anthers about as long 

as wide, dorsifixed; pollen 2-colpate (or colporate?); glands sometimes present next to the filaments; pistil 1, the carpels 5'or 3 (2), style 1, 

gynobasic, stigmas 5 or 3 (2), apical or slightly 

elongated; ovary deeply 5 or 3(2)-lobed, superior, the ovules 1 per locule, basal-axile, unitegmic, tenuinucellar, anatropous; fruit a schizocarp 

of 1-5 nutlets; embryo straight, the radicle invested; cotyledons thick, plano-convex, 0.75-0.9 

the length of the embryo; endosperm 0. 

Composition: 2 genera, 11 species. 

Distribution: Moist habitats; western United 

States and adjacent Canada, especially California, 

one species reaching the eastern United States. 

OXALIDACEAE (Figure 130c).-Herbs and 

shrubs, rarely trees (Averrhoa); xylem vessel perforation plates simple; leaves alternate, digitately 

or pinnately compound, sometimes unifoliolate, 

the leaflets entire (except some Hypseocharis), 

exstipulate; inflorescences axillary cymes, subumbellate, or cauliflorous racemes or thyrses (Averrhoa), or the flowers solitary; flowers bisexual, 

actinomorphic, heterostyled; sepals 5, basally 

connate; petals 5, free or shortly connate, contorted in bud; stamens 10 or 15, 5 sometimes 

staminodal, in 2 or 3 series, weakly obdiplostemonous, the filaments shortly connate; anthers 

about as long as wide, dorsifixed; pollen 3(4)- 

colpate, 4-rupate, 3-colporate, or 3-colporoidate; 

disk absent; pistil 1, the carpels 5, styles 5 or 1, 

free or connate (Hypseocharis), stigma generally 

capitate; ovary superior, 5-locular, the ovules 1 

or several per locule, axile, bitegmic, tenuinucellar, anatropous; fruit a loculicidal capsule, rarely 

a schizocarp (Biophytum), rarely a 5-lobed berry 

(Averrhoa); seeds sometimes arillate; embryo 

straight or cochlear (Hypseocharis), 0.9 the length 

FIGURE 1 30.-LIMNANTHACEAE: a, Limnanthes dOUglasi2 part 

of flowering shoot, floral diagram, petal, stamen and gland, 

style and stigmas, 1,s. of base of flower showing ovule in a 

carpel: 6, fruitlet, 1,s. and C.S. of same, Floerkea proserpinacoides floral diagram (after Le Maout and Decaisne, 1873). 

OXALIDACEAE: c, Oxalis elatior flower, petal, essential organ-brevistylous, intermediate and longistylous forms, androeciuni of microstylous flower laid open, fruit, pistil (microstylous form), 0. delicata seed (after Martius, 1840- 

1906). CONNARACEAE: d, Rourea cuspidata flower, 1,s. of same 

(anthers removed), views of anther (from bud); e, calyx, 

petal, gynoecium, 1,s. of 1 pistil, R. induta floral diagram;f, 

Connarus patrisii essential organs, dehiscent fruit and small 

part of inflorescence, C. suberosus fruit, calyx, 1,s. of fruit, 

seed Lvith aril (after Martius, 1840-1 906). 
NUMBER 58 249 

a 

f 
250 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

of the endosperm; cotyledons elliptic, moderately thick, 2.4-2.5 times wider than the radicle, 

0.5-0.7 the length of the embryo; endosperm 

moderate, fleshy. 

Composition: 8 genera, -900 species. 

Distribution: Cosmopolitan except arctic and 

cool temperate Northern Hemisphere; mainly 

tropical and subtropical Asia, Africa, and America, often at high elevations. 

CONNARACEAE (Figure 1 3Od-f.-Lianas, 

more rarely trees and shrubs; xylem vessel perforation plates simple; secretory cavities sometimes present in the parenchyma of the tissues; 

leaves alternate, usually compound, sometimes 

unifoliate, the leaflets entire, very rarely with 2- 

3 teeth, pulvini present, exstipulate; inflorescences terminal, axillary or at nodes of fallen 

leaves, panicles, fascicles or spikes; flowers bisexual, perhaps sometimes functionally unisexual, 

actinomorphic, the parts hypogynous, sometimes 

heteromorphic with styles and stamens of 2-3 

different lengths in the same species; rarely the 

plants dioecious; sepals 5 (4-3), free or shortly 

connate, imbricate or valvate; petals 5 (4), free, 

rarely slightly connate, rarely loriform and circinate in bud; stamens 5 (4) or 10 (8) in 2 series, 

the inner sometimes staminodial, the filaments 

elongated, often shortly connate, the anthers 

dorsifixed, about as long as wide; pollen 3-4- 

colpate, 3-colpoidate, colporate or colporoidate; 

disk present or absent; pistils 5 (1-S), unicarpellate when one, sometimes narrowed to a stipe, 

sometimes very slightly connate laterally and/or 

ventrally at the base, the style somewhat elongated, the stigma apical, capitate or obscurely 

bilobed; ovules 2 (3), ventral or subbasal, bitegmic, crassinucellar, hemianatropous; fruit 

usually follicular, sometimes dehiscent along 

both sutures or rarely indehiscent; seed 1 (2), 

often arillate or sarcotestal, the embryo straight, 

0.4-0.9 the length of the endosperm; cotyledons 

flat or plano-convex, 0.7-0.9 the length of the 

embryo, 2-7.5 times wider than the radicle, the 

latter sometimes invested; endosperm copious, 

scanty or 0, hard or fleshy, sometimes oily. 

Composition: -20 genera, -350 species. 

Distribution: Pantropical, especially Asia, extending into temperate China, Australia, and 

South Africa. 

TROPAEOLACEAE (Figure 13 la-c).-Succulent 

prostrate or twining herbs containing mustardoil; xylem vessel perforation plates usually simple 

but sometimes also reticulate; leaves alternate, or 

the lower opposite, simple, usually peltate, sometimes lobed or the margin sinuate, rarely compound, sometimes dotted, exstipulate or stipulate; flowers axillary, solitary, bisexual, zygomorphic; sepals 5, shortly connate, colored, the dorsal one produced into a spur; petals 5 (2), free, 

inserted on the calyx; stamens 8, the filaments 

elongated, free; anthers about as long as wide, 

basifixed; pollen 3-colporate; pistil 1, the carpels 

3, style 1, elongated, stigmas 3 apical-linear; 

ovary superior, 3-locular, the ovules 1 per locule, 

axile, bitegmic, tenuinucellar, anatropous; fruit 

a schizocarp of 3 cocci, rarely a berry, or samara 

(Mugallanu); embryo straight, the radicle invested; cotyledons thick, plano-convex, 0.8 the 

length of the embryo; endosperm 0. 

Composition: 2 genera, 80 species. 

Distribution: Mainly in mountains from 

southern Mexico to Chile and Argentina, but 

extending over most of the South America, in 

various habitats from cold mountain heights to 

rain forest and arid regions. 

BALSAMINACEAE (Figure 13 1 d).-Succulent 

herbs, rarely shrubs; xylem vessel perforation 

plates simple; leaves alternate, rarely opposite or 

whorled, simple, toothed, exstipulate; inflorescences axillary, subumbellate, racemiform or the 

flowers solitary; flowers bisexual, zygomorphic; 

sepals 3 (5), free, often colored, the posterior 

one produced into a nectariferous spur; petals 5, 

free or the laterals connate; stamens 5, the filaments short, broad, apically connate; anthers 

small, about as long as wide, connate into a cap 

over the stigma; pollen 3-4-colpate; disk absent; 

pistil 1, the carpels 5, stigmas 5, more or less 

obscure, apical, subsessile; ovary superior, 

5-locular, the ovules 3-several per locule, axile, 

bitegmic, tenuinucellar, anatropous; fruit a succulent capsule dehiscing elastically, rarely a 
NUMBER 58 25 1 

FIGURE 13 1 .-TROPAEOLACEAE: a, Tropaeolum brasiliense flower, I.s. of same, floral diagram, 

views of anther, views of mericarp; b, T. pentaphyllus floral diagram, 1,s. of ovary, flower; c, 

part of shoot in bud, T. brasiliense inferior petal, superior petal, T. pentaphyllum superior petal, 

I.s. of flower (after Martius, 1840- 1906). BALSAMINACEAE: d, Impatiens balsaminu 2 views of 

flower, 1,s. of same, floral diagram and dehiscent fruit, essential organs, 1,s. of same, fruit, I. 

noli-tangere dehiscent fruit (after Baillon, 1866-1 895). 

berry; embryo straight, the cotyledons moderately thick, plano-convex, 0.9 the length of the 

embryo, 3 times the width of the radicle; endosperm 0. 

Composition: 4 genera, 500 species. 

Distribution: Mainly tropical Asia and Africa, 

a few species in temperate Northern Hemisphere 

and Africa. 

STACKHOUSIACEAE (Figure 132a,b).-Small 

herbs with a woody rhizome, with a rubber-like 

substance(?); xylem vessel perforation plates simple; leaves rarely absent, alternate, simple, entire, 

exstipulate; inflorescence a terminal cyme, raceme, spike, panicle, fascicle, rarely umbel, or 

the flower solitary; flowers bisexual; sepals 5, 

basally connate into a calyx-tube, the lobes im- 
252 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

b 

FIGURE 1~~.-STACKHOUSIACEAE: a, Stackhousia monogyna flower, I.s. of same, corolla laid open 

showing separations at base, calyx laid open showing insertion of stamens, pistil, C.S. and I.s. of 

ovary, fruit, S. pulvinaris I.s. of seed; b, Stackhousia sp. inflorescence, flower laid open (corolla 

removed), Tripterococcus brunonis 1.s. of flower, part of fruit, C.S. of one of its cocci (after 

Baillon, 1866-1895; Lindley, 1853; Hutchinson, 1973; Pax). LINACEAE: c, Linum usitatissimum 

flowering shoot, bud, essential organs and 2 views of a stigma; d. L. selagznoides part of stamina1 

tube laid open, pistil, fruit from above and below, 1,s. of a hemicarpel in fruit, seed, 1,s. of same 

showing embryo; e, L. junceum three of the sepals, petal, fruit with calyx, hemicarpel in fruit 

(after Johnson; Martius, 1840- 1906). 

bricate; petals 5, free or connate, perigynous; 

stamens 5, alternipetalous, the filaments short or 

elongated, inserted at the apex of the calyx-tube; 

anthers somewhat elongated, basifixed or subbasifixed, sometimes the connective produced 

apically; pollen 3-colporate; disk lining the calyxtube; pistil 1, the carpels 3 (2-5), styles 3 (2-5), 

free or connate, sometimes with a discoid collar 
NUMBER 58 

beneath the 3 (2-5) stigma lobes, stigmas decurrent ventrally; ovary superior, 3(2-5)-locular, the 

ovules 1 per locule, basal, bitegmic, tenuinucellar, anatropous; fruit a schizocarp of 3 (2-5) 

cocci, leaving a columella, the cocci sometimes 

winged; embryo large, straight, 0.9 the length of 

the endosperm, elliptic in longitudinal section, 

the cotyledons gradually grading into the radicle; 

cotyledons 0.7 the length of the embryo; endosperm moderate, fleshy. 

Composition: 3 genera, 25 species. 

Distribution: Dry habitats, centered in Australia, extending to New Zealand, New Guinea, 

and the Philippines. 

LINACEAE (Figure 132c-e).--Trees, shrubs, 

and herbs, sometimes climbing with hooks on the 

inflorescence; xylem vessel perforation plates 

simple or scalariform with less than 25 bars; 

leaves alternate or rarely opposite (Aneulophus, 

Linum spp., Radiola), simple, entire, rarely 

toothed (Hugonia), usually stipulate; inflorescence dichasium, cincinnus, thyrse, sometimes 

appearing to be a raceme; flowers bisexual, actinomorphic, sometimes heterostyled; sepals 5 (4), 

free or basally connate; petals 5 (4), free, rarely 

with a ligular attachment, contorted in bud, caducous; stamens as many as or twice as many as 

the petals, very rarely 15 and then some staminodal (Roucheria sp.), weakly obdiplostemonous, 

the filaments basally connate, sometimes 5 or 10 

tooth-like staminodes present; anthers about as 

long as wide; pollen 3(4)-colpate, occasionally 6- 

rugate, polyforate or nonaperturate; disk 0 or an 

intra- or extra-stamina1 annulus; pistil 1, the carpels 5 (3-4), styles 5 (3-4), filiform, free or more 

rarely connate, the stigmas apical, capitate; ovary 

superior, 5(3-4)-locular or secondarily 1 O-locular, the ovules 2 (1) per carpel, axile, bitegmic, 

crassinucellar or tenuinucellar, anatropous; fruit 

a septicidal capsule or drupe (Hugonieae); seeds 

compressed, sometimes arillate; embryo straight, 

0.97 the length of the endosperm; cotyledons 

moderately thick, twice as wide as the radicle, 

0.7 the length of the embryo; endosperm copious 

or scanty, fleshy. 

Composition: 13 genera, -300 species. 

253 

Distribution: Principally temperate; cosmopolitan except for the arctic and cool temperate 

regions of the Northern Hemisphere. 

ERYTHROXYLACEAE (Figure 133a,b).-Small 

trees and shrubs; xylem vessel perforation plates 

simple, rarely a few scalariform; leaves alternate, 

simple, entire, the stipule intrapetiolar; inflorescences axillary fascicles or the flowers solitary; 

flowers bisexual or the plants rarely subdioecious, actinomorphic, usually heterostyled; sepals 

5, free or shortly connate; petals 5, free, mostly 

with a ventral ligulate bifid appendage or callosity, convolute or imbricate; stamens 10 in 2 series, obdiplostemonous, the filaments elongated, 

basally connate; anthers small, about as long as 

wide; pollen 3-colporate; glandular tissue external to filament tube; pistil 1, the carpels 3 (2 in 

Nectaropetalum), the styles 3 (1 in Nectaropetalum), free or basally connate, the stigmas apical, 

obliquely clavate or capitate; ovary superior, 3- 

locular, mostly only 1 of these fertile, with 1-2 

axile, bitegmic, crassinucellar, anatropous or 

semi-anatropous ovules; fruit a drupe; embryo 

straight, 0.9 the length of the endosperm; cotyledons thin, 2.5 times wider than the radicle, 

0.4-0.6 the length of the embryo; endosperm 

copious, fleshy, rarely 0. 

Composition: 4 genera, 250 species. 

Distribution: Centered in the Andes and Amazon basin; tropical America to part of temperate 

South America; tropical and subtropical Africa, 

Madagascar; Indochina, the East Indies, and 

northeastern Australia. 

ZYGOPHYLLACEAE (Figure 133c,d).-Shrubs 

and herbs, rarely trees, sometimes the wood 

resinous; xylem vessel perforation plates simple; 

leaves opposite or rarely alternate, pinnately 

compound or 2 (3) foliolate, the leaflets entire, 

or simple and sometimes multifid (Peganum), 

stipulate or exstipulate; inflorescences terminal 

or axillary cincinni, compound, or the flowers 

solitary; flowers bisexual or rarely the plants 

dioecious (Neoluederitzia), actinomorphic or 

rarely zygomorphic, the parts hypogynous, very 

rarely subperigynous (Sericodes); sepals 5 (4), free 

or rarely basally connate; petals 5 (4), rarely 0, 
254 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 133.-ERYTHROXYLACEAE: a, Erythroxylum coelophlebium brevistylous flower, longistylous flower, E. macrocnemium longistylous flower with petals and 2 sepals removed, E. vacciniijolium part of a flowering twig; b, E. coelophlebium scale from twig, E. macrocnemium internal 

face of petal with appendage, E. coelophlebium C.S. of ovary, E. buxus I.s. and C.S. of fruit, embryo 

(after Martius, 1840- 1906). ZYGOPHYLLACEAE: c, Kallstroemia tribuloides flower, same with 

petals removed, petal, fruit, dehiscent fruit; d, flowering shoots with 2 views of flower, views 

of stamen, dehiscent fruit with 1 remaining mericarp, 2 views of mericarp, embryo, same with 

cotytedons separated (after Martius, 1840- 1906). BALANITACEAE: e, Balanites aegyptiaca flowering twig, flower, 1,s. of same, dried fruit (after Baillon, 1840-1906). 
NUMBER 58 255 

free; stamens (3) 4-15, in 1-3 series, obdiplostemonous, the filaments elongated, free, often with 

a ventral ligular appendage; anthers about as 

long as wide, dorsifixed; pollen 3-colpate, 3(4)- 

colporate, 3-colporoidate, 4(5)-ruporate, polyforate; disk usually present; pistil 1, the carpels 

5 (2-12), style 1, short, the stigmas apical; ovary 

5(2-12)-locular, the ovules 1 to CQ per locule, 

axile, bitegmic, crassinucellar, anatropous; fruit 

usually a loculicidal and/or septicidal capsule, or 

cocci, rarely a berry (Peganum) or drupe (Nitraria); embryo straight or slightly curved, 0.7 to 

as long as the endosperm; cotyledons 1.3 times 

wider than the radicle, 0.5-0.7 the length of the 

embryo, the radicle sometimes invested; endosperm scanty or 0, hard. 

Composition: 25 genera, 250 species. 

Distribution: Mainly arid or salt habitats; 

mostly tropical and subtropical, extending into 

the temperate zones; southern United States to 

southern South America along the Andes; most 

of Africa, southern half of Eurasia, western East 

Indies, and Australia. 

BALANITACEAE (Figure 133e).-Shrubs and 

small trees with bitter bark; xylem vessel perforation plates simple; leaves alternate, 2-foliolate, 

the leaflets entire, not punctate, exstipulate; inflorescences axillary umbelliform clusters; flowers bisexual; sepals 5, free; petals 5, free; stamens 

10, the filaments elongated, inserted outside the 

disk; anthers about as long as wide, dorsifixed; 

pollen 3-colporate, occasional grains 6-rugorate; 

disk thick, surrounding the base of the ovary; 

pistil 1, the carpels 5, style 1, very short, stigma 

apical, punctiform; ovary superior, 5-locular, the 

ovules 1 per locule, axile-subapical, bitegmic, 

crassinucellar, semi-anatropous or campylotropous; fruit a drupe, the seed 1; embryo ovoid, 

green, the cotyledons thick, plano-convex, corrugated or 2-lobed; endosperm 0. 

Composition: 1 genus, 20 species. 

Distribution: Tropical Africa to Burma. 

MALPIGHIACEAE (Figure 134~-c).-Shrubs, 

lianas, and small trees; xylem vessel perforation 

plates simple; leaves opposite, very rarely alternate or ternate, simple, entire, rarely sinuateserrate or lobate, with hairs often appressed and 

medifixed and glands often present either on the 

petiole or on the lower surface, stipulate or rarely 

exstipulate; inflorescences terminal or axillary 

racemes; flowers usually bisexual, sometimes dimorphic, sometimes cleistogamous, rarely the 

plants polygamous, actinomorphic or slightly zygomorphic, the parts hypogynous or nearly so; 

sepals 5, mostly free, sometimes basally connate, 

often with 2 large glands on the dorsal surface 

of some or all; petals 5, free, convolute in bud; 

stamens mostly 10, in 2 series, obdiplostemonous, sometimes 5 reduced to staminodes, the 

filaments elongated, often basally connate; anthers 1-2 times as long as wide, the connective 

sometimes apically produced, dehiscence longitudinal or by a pore or pore-like slit; pollen 3-5- 

colporate, oligoforate, polyforate, oligoporate, 

6- 12-rugorate; disk small; pistil 1, the carpels 3 

(1-4), styles usually free, united to the apex in 

Bunchosia decussajlora, stigmas apical, punctiform 

or capitate; ovary 3( 1-4)-locular, the ovules bitegmic, crassinucellar, anatropous, semi-anatropous or campylotropous, 1 per locule, axile; fruit 

most frequently a schizocarp, often samaroid, 

sometimes a loculicidal capsule, more rarely a 

drupe; embryo straight, arcuate or rarely circinate, the radicle sometimes partly invested, the 

plumule sometimes evident; cotyledons sometimes plano-convex or one enlarged and thick 

and the other reduced, 3.3-6.0 times wider than 

the radicle, 0.5-0.8 the length of the embryo; 

endosperm 0. 

Composition: 60 genera, 800 species. 

Distribution: Forests and open areas; pantropical, extending into the warm temperate regions 

of America, Africa, and Asia; centered in South 

America. 

POLYGALALES 

Shrubs, herbs, and trees, sometimes climbing, 

very rarely saprophytes or parasites, sometimes 

resinous; xylem vessel perforation plates simple; 

interxylary or intraxylary phloem sometimes 

present; leaves alternate, less commonly oppo- 
256 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 134.-MALPIGHIACEAE: a, GaUdiChaUdkZ CongeStijh’a flower, h’falpighia ?titZda 1,s. Of 

flower, fruit, I.s. of same, Aspidopterys roxburgiana floral diagram; b, Byrsonima umbellata bud, 

sepal with 2 glands, essential organs, 4 views of stamen, 1,s. and C.S. of ovary; c, Heteropterys 

pruinosa 1,s. of flower without petals, views of anther, fruit, Tetrapterys maranhamensis hairs, 

Thryallis braszlzenszs 1,s. of seed, Byrsonima spicata embryo (after Baillon, 1866- 1895; Martius, 

1840- 1906; Niedenzu; Jussieu). 

site, simple, entire, usually exstipulate; flowers 

bisexual, usually zygomorphic; sepals 5 (4), free 

or rarely shortly connate, the inner two often 

colored, one rarely spurred; petals 1, 3-5 (0), 

free or 3 connate, 2 sometimes much smaller 

than the others, sometimes 1 larger and gibbous 

or spurred; stamens 1-12, in 1-2 series, some 

sometimes staminodal, the filaments connate or 

free, often inserted on the petals; anthers basifixed or dorsifixed, dehiscing by an apical pore 

or longitudinally, very rarely transversely; pollen 

3-polycolporate (-ruporate, 3-4-porate, rugoidate); glandular disk rarely present; pistil 1, the 

carpels 2-3 (5), style 1, the stigma(s) apical; ovary 

superior, 2-3( 1 -5)-locular, the ovules 1 -2(-several) per locule, axile, rarely apical or parietal; 

fruit a loculicidal or septicidal capsule, less commonly a nut, drupe, berry, samara, or achene, 

the seeds often hairy, sometimes winged; embryo 

large, straight, rarely folded or convolute; endosperm usually copious, sometimes absent. 

Distribution: Cosmopolitan, except the arctic, 

in various habitats. 

Chemistry: Polygalaceae are rich in triterpenoid saponin, sinapic acid (a phenolic acid), and 

polygalitol. It has xanthones. 

POLYCALACEAE (Figure 135u-f).-Herbs, 

shrubs, and climbers, rarely small trees, saprophytes or parasites; xylem vessel perforation 

plates simple; interxylary phloem sometimes 

present; leaves alternate, rarely opposite or 

whorled, simple, entire, exstipulate or rarely stipulate; inflorescences axillary or terminal racemes, spikes or panicles, or the flower solitary; 

flowers bisexual, usually zygomorphic; sepals 5, 

free or rarely connate, the 2 inner ones usually 
NUMBER 58 257 

U b 

e 

FIGURE 1 ~~.-POLYGALACEAE: a, Diclidanthera pendulajlora flower, 1,s. of same (enlarged), 

views of anther before and during dehiscence, upper part of style with stigma, I.s. and C.S. of 

ovary; b, D. laurijolia I.s. and C.S. of fruit, I.s. of seed, embryo; c, Moutabea dibotrya floral 

diagram, stamina1 tube and upper part of style within, views of anther, upper part of style and 

stigma, ,M. aculeata seed laid open, embryo; d, Polygala ligustroides flower, 1.s. of same, 

androecium (adnate to petals) laid open, views of anther, apex of style with stigma, C.S. of 

ovary; e, fruit enclosed in persistent calyx, I.s. of fruit, immature seed with aril;J P. brasiliensis 

floral diagram, P. aphylla 2 views of apex of anterior petal showing crest, P. violacea seed, I.s. 

same, embryo, Securidaca lanceolata fruits (after Martius, 1840-1 906). 
258 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 136.-KRAMERIACEAE: a, Krameria tomentosa floral diagram, flower, 1.s. of same, 

androecium with petals from a bud, androecium and 3 petals; b, views of anther, C.S. of ovary 

and I.s. of seed, 2 views of seed, fruit, I.s. of same, K. argentia bud (after Martius, 1840-1906). 

colored; petals 3-5, the outer 2 free or connate 

with the lowermost, the upper 2 free, sometimes 

minute and scale-like or absent; stamens 8 (3, 4, 

5, 7, or 10 in 2 series), monadelphous or rarely 

free (Xanthophyllum spp.), the sheath split above, 

often adnate to the petals; anthers small, basifixed, dehiscing by an apical pore, rarely nearly 

to the base, or transversely (Diclidanthera); pollen 

polycolporate, the colpi 7-28; rarely an annular 

disk within the stamens (Xanthophyllum); pistil 1, 

the carpels 2 (3, 5), one of the 2 in Monnina 

usually rudimentary, style 1, stigma(s) apical; 

ovary superior, 2( 1-5)-locular, the ovules 1 per 

locule, axile, or rarely 2-6 per placenta and 

parietal when the ovary is unilocular (Xanthophyllum), bitegmic, crassinucellar, anatropous or 

semi-anatropous; fruit usually a loculicidal capsule, sometimes a nut, drupe, berry or samara; 

seeds arillate (appendaged), often hairy; embryo 

straight, as long as the endosperm; cotyledons 

moderately or very thick and plano-convex (Xanthophyllum), 1.5-3.2 times wider than the radicle, 

0.5-0.8 the length of the embryo, the radicle 

rarely invested (Xanthophyllum); endosperm copious, moderate or rarely 0. 

Composition: 13 genera, -900 species. 

Distribution: Cosmopolitan except the arctic 

and cool temperate region of the Northern Hemisphere, New Zealand, and the central Pacific 

islands; centered in the American tropics and 

Africa. 

KRAMERIACEAE (Figure 136a,b).-Shrubs and 

herbs; xylem vessel perforation plates simple; 

leaves alternate, simple, entire, exstipulate; inflorescences axillary or terminal racemes or the 

flowers solitary; flowers bisexual, zygomorphic; 

sepals 5 (4), free; petals 5, the 3 upper longclawed, free or connate, the 2 lower usually much 

smaller, thick; stamens 4 (3), the filaments thick, 

free or adnate to the claws of the upper petals; 

anthers small, basifixed, dehiscing by a terminal 

pore; pollen 4-colporate (-ruporate); pistil 1, the 

carpels 2 (1 sterile), style 1, stigma apical, discoid; 

ovary superior, unilocular, the ovules 2, apical, 

bitegmic, anatropous; fruit a globose bristly 

achene; seed 1, the embryo straight, radicle invested; cotyledons broad and thick, 0.75 the 

length of the embryo; endosperm 0. 

Composition: 1 genus, 20 species. 

Distribution: Mainly dry regions; tropical and 

warm temperate America from southwestern 

United States to Argentina and Chile. 
NUMBER 58 259 

TRIGONIACEAE (Figure 137a,b).-Scandent 

shrubs or trees; xylem vessel perforation plates 

simple; intraxylary phloem absent; leaves opposite, rarely alternate, simple, entire, stipulate or 

exstipulate; inflorescences axillary and terminal 

racemes, panicles or thyrses, rarely 3-flowered 

cymes; flowers bisexual, zygomorphic; sepals 5, 

free or shortly connate; petals 5 or 3, free, the 

posterior one largest and gibbous or spurred, a 

gland often present opposite this petal; stamens 

3-1 2, some staminodal, the filaments connate; 

anthers about as long as wide, dorsifixed, dehiscing longitudinally; pollen 3-4-porate or 3-colporate; 1-3 glands usually adjoining the split 

stamina1 tube; pistil 1, the carpels 3, style 1, 

stigma apical, capitate or not enlarged; ovary 

superior, 3- or 1-locular, the ovules 1 to CQ per 

locule, axile, bitegmic, crassinucellar, anatropous; fruit a septicidal capsule, rarely a samara; 

seeds usually enveloped in long hair; embryo 

straight, 0.8-0.9 the length of the endosperm, 

cotyledons 5-7 times wider than the radicle, 0.8- 

0.85 the length of the embryo; endosperm 

scanty, fleshy. 

Composition: 4 genera, 35 species. 

Distribution: Lowland moist forest; tropical 

America, western Malaysia, Madagascar. 

VOCHYSIACEAE (Figure 137c,d).-Large resinous trees, shrubs, and climbers; xylem vessel 

perforation plates simple; intraxylary and interxylary phloem sometimes present; leaves opposite or verticillate, rarely alternate, simple, entire, 

stipulate or exstipulate; inflorescences racemes, 

panicle or cymose; flowers bisexual, zygomorphic; sepals 5, shortly connate, one longer than 

the others and gibbous or spurred; petals 1-3 (5, 

0), free; stamens 1 fertile, 1-4 staminodes, the 

filaments free; anthers about as long as wide, 

basifixed, rarely dorsifixed, sometimes apiculate, 

dehiscing longitudinally; pollen 3-colporate, occasionally rugoidate; pistil 1, the carpels 3, style 

1, clavate, the stigma terminal or lateral; ovary 

superior or rarely adnate to the calyx, 1-3-locular, the ovules (1) 2 to CQ per locule, axile, bitegmic, crassinucellar, anatropous or semi-anatropous; fruit a loculicidal capsule, rarely samaroid due to enlargement of sepals; seeds often 

winged, sometimes pilose; embryo straight or the 

cotyledons folded or convolute, sometimes the 

radicle invested; cotyledons thin, 6- 12 times 

wider than the radicle, 0.7-0.8 the length of the 

embryo; endosperm 0, rarely scanty (Erismadelphus). 

Composition: 6 genera, 200 species. 

Distribution: Almost entirely tropical America; one small genus in tropical western Africa. 

OLEALES 

Trees, shrubs, and lianas; xylem vessel perforation plates typically simple, rarely also scalariform with few bars; interxylary phloem rarely 

present; leaves opposite, very rarely alternate, 

simple or compound, entire or toothed, mostly 

exstipulate; flowers bisexual, rarely unisexual; 

sepals 4 (0-15), connate; petals 4 (0-l5), connate, rarely free; stamens 2 (4,5), free or inserted 

on the corolla, the filaments very rarely connate; 

anthers dorsifixed or basifixed, the loculi back 

to back; pollen 3(4)-colpate, -colporate, occasionally 4-rupate or 6-rugate; glandular disk sometimes present; pistil 1, the carpels 2, style 1, the 

stigmas apical; ovary superior, 2( 1 )-locular, the 

ovules 2 (1-10) per locule, axile (basal); fruit a 

loculicidal (circumscissile) capsule, berry, drupe, 

schizocarp or samara; embryo mostly long; endosperm copious to absent. 

Distribution: Cosmopolitan, except cool regions, in mesic to dry habitats. 

Chemistry: Gibbs considers it reasonable to 

make a separate order for the Oleaceae and to 

associate it closely with the Gentianales. 

OLEACEAE (Figure 138a,b).-Trees, shrubs 

and lianas; xylem vessel perforation plates typically simple, rarely also some scalariform with 1 

or more bars in 3 genera; leaves opposite, very 

rarely alternate (Jasminum spp.), simple or pinnately compound, entire or toothed, often with 

peltate trichomes, exstipulate; inflorescences terminal or axillary racemes, panicles or dichasia; 

flowers bisexual or rarely the plants dioecious or 

polygamodioecious; sepals 4 (0- 15), connate; 

petals 4 (0-1 5), connate or rarely free (Fraxinus 

spp.), imbricate or induplicate valvate, rarely 
260 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 137.-TRIGONIACEAE: a, Trzgonia pubescens flower, I.s. of same, essential organs, 

androecium laid open, pistil, C.S. of ovary and views of anther; b, dehiscent fruit, 1 valve of 

fruit showing exocarp, endocarp and seeds and C.S. of dehiscent fruit, 1.s. of seed, embryo and 

C.S. of seed (hair removed), Lzghtia lzcanioides flower, 1,s. of same, floral diagram, part of 

androecium (after Martius, 1840-1 906). VOCHYSIACEAE: c, Quulea cryptantha flower, 1,s. of 

same, ovary and stamen, views of anther, C.S. of ovary, dehiscent fruit; d, Vochysia haenkeana 

views of anther, flower, 1,s. of base of same, base of stamen, staminodes and part of calyx, V. 

obscura seed, embryo with convolute cotyledons and laid open (after Martius, 1840-1906). 
NUMBER 58 26 1 

FIGURE 138.--oLEACEAE: a, Oka europaea flower, 1.s. of same, floral diagram, 1,s. of fruit, seed; 

b, Jasminum azoricum flower laid open, J. ofjcinale Is. of fruit, C.S. of seed, embryo, Fraxinus 

sp. 6 flower, floral diagrams of 6 flower with 2 and 4 stamens, fruit, 0 flower, pistil and I.s. of 

calyx (after Baillon, 1866-1 895; Martius, 1840- 1906; Johnson). SALVADORACEAE: c, Salvadora 

persica bud, flower, I.s. of same, fruit, 1.s. of same, embryo (after Baillon, 1866-1895). 

convolute; stamens 2 (4 in Hesperelaea and Tessarandra and some species in a few other genera), 

free or inserted on the corolla alternate with the 

lobes; anther loculi back to back, dorsifixed or 

basifixed, the connective often produced apically; pollen usually 3(4)-colpate-colpor(oid)ate, 

occasionally 4-rupate; disk 0; pistil 1, the carpels 

2, style 1, the stigmas apical; ovary superior, 

bilocular, the ovules 2 (1-10) per locule, axile, 

unitegmic, tenuinucellar, anatropous or amphitropous; fruit a loculicidal or circumscissile (Menodora) capsule, berry, drupe, schizocarp or samara, the seeds 1-4; embryo straight, rarely contortuplicate (one species of Schrebera), 0.1 to as 

long as the endosperm, mostly long, but may be 

minute before after-ripening in Fraxinus; cotyledons thin or moderately thick, 1.3-6.0 times 

wider than the radicle, 0.5-0.86 the length of 

the embryo; endosperm copious, moderate or 0, 

fleshy. 

Composition: 28 genera, 600 species. 

Distribution: Cosmopolitan except the arctic 

and cool temperate regions of the Northern 

Hemisphere, cool temperate South America and 

New Zealand; centered in southeastern Asia and 

the East Indies. 

SALVADORACEAE (Figure 1 38c).-Trees and 

shrubs; xylem vessel perforation plates simple; 
262 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

interxylary phloem in 2 of the 3 genera; leaves 

opposite, simple, entire, small stipules sometimes 

present; inflorescences axillary and terminal 

spikes, racemes, panicles or fascicles; flowers bisexual or the plants dioecious; sepals 3-4 (2-5), 

connate; petals 4 (5), free or basally connate, 

imbricate; stamens 4 (5), hypogynous or inserted 

on the base of and alternate with the petals, the 

filaments free or basally connate; anther loculi 

back to back; pollen 3-colporoidate, exceptionally 4-rupate or 6-rug(oroid)ate; disk 0 or consisting of separate glands between the filaments; 

pistil 1, the carpels 2, style 1, short, the stigma(s) 

apical; ovary superior, 1 -2-locular, the ovules 1 - 

2 per locule, basal, bitegmic, crassinucellar, anatropous; fruit a berry or drupe, the seeds mostly 

1 ; cotyledons thick, plano-convex, 4 times wider 

than the radicle, 0.8-0.9 the length of the embryo; endosperm 0. 

Composition: 3 genera, 12 species. 

Distribution: Dry, often saline habitats; Africa 

except the west-central region; Madagascar; Arabia to India, southern China, and the western 

East Indies. 

GENTIANALES 

Trees, shrubs, and herbs, sometimes climbing, 

rarely saprophytic or parasitic and chlorophylless, often bitter; xylem vessel perforation plates 

usually simple, rarely scalariform or both; intraxylary or interxylary phloem sometimes present; 

laticiferous canals sometimes present; leaves opposite, rarely alternate or whorled, simple, very 

rarely compound, entire or very rarely toothed, 

lobed or dissected, stipulate or exstipulate; flowers bisexual, rarely unisexual, actinomorphic, 

very rarely zygomorphic; sepals 5-4 (2- 12), connate or free, sometimes with basal nectar glands; 

petals 4-5 (3-16), connate, sometimes with a 

corona, very rarely spurred; stamens 4-5 (1-16), 

the filaments free or connate, very rarely 

strongly unequal, inserted on the corolla, very 

rarely hypogynous or nearly epigynous, alternipetalous, the anthers rarely sessile, basifixed or 

dorsifixed, rarely connate; pollen 2-polycolpate, 

2-multicolporate, 3-4( 1 -6)-porate, rarely polyforate, 4-rupate or ruporate, 6-rugate or nonaperturate; pollen grains single or sometimes in 

tetrads, masses or pollinia; glandular disk usually 

present; pistil 1, the carpels 2 (3-9), style 1 (2), 

sometimes 2-4-lobed, the stigma(s) apical or ventral; ovary superior to inferior, 2( l-9)-locular, 

the locules rarely separate; ovules l-many per 

locule, axile or parietal, rarely basal or apical, 

unitegmic; fruit a septicidal, loculicidal or rarely 

circumscissile capsule, berry, drupe, schizocarp 

or follicles, rarely a nut or samara; seeds sometimes with a coma or hairy; embryo straight or 

folded, minute to large; endosperm copious to 

absent . 

Distribution: Cosmopolitan, most common in 

the tropics and subtropics, in various habitats. 

Chemistry: Alkaloids in great variety occur in 

the order. The oxindole alkaloids seem to be 

confined to Gentianales-Loganiaceae, Apocynaceae, Rubiaceae; several others are confined 

to this order. Gibbs knows nothing that sets the 

Loganiaceae or Desfontainiaceae apart; likewise 

with Gentianaceae and Menyanthaceae, both 

have gentianine, but differ in some respects, 

enough to segregate them he feels. Apocynaceae 

and Asclepiadaceae are very closely related. By 

and large the chemistry of Rubiaceae supports 

its inclusion here, but there are differences that 

set it apart fairly clearly. Loganin occurs in Mastixia (Cornaceae). The emetine group occurs in 

Rubiaceae, Alangiaceae, and Icacinaceae. The 

Convolvulaceae is the only family in “Tubiflorae” 

with ergoline alkaloids. 

LOCANIACEAE (Figure 139a-d).--Trees, 

shrubs, and herbs, sometimes climbers or epiphytes; xylem vessel perforation plates usually 

simple, rarely scalariform or both; intraxylary or 

interxylary phloem present; leaves opposite, 

rarely verticillate, simple, entire, bitter, stipulate 

or exstipulate; inflorescences terminal and axillary cymes, spikes, racemes, panicles, heads or 

rarely the flowers solitary; flowers bisexual, very 

rarely polygamo-dioecious, actinomorphic or 

very rarely not so in calyx, corolla or androecium; sepals 5-4, connate or free, rarely bipar- 
NUMBER 58 263 

FIGURE 139.-LOGANIACEAE: a, Antonia ovata l.s. of bud, flower, corolla laid open, views of

stamen, pistil, ovary, 2 views of stigma and 3 views of seed, 2 fruits, dehiscent fruit, 1,s. of fruit; 

b, Strychnos brevzf2ora cyme, corolla laid open to show insertion of stamens, calyx laid open, 

views of stamen, C.S. of ovary; c, fruit and leaf, S. brasiliensis pistil, stigma and style, C.S. of ovary 

at 3 levels, C.S. and 1,s. of seed, embryo; d, Desfontainea spinosa corolla laid open, pistil and part 

of calyx, C.S. of ovary, fruit (after Martius, 1840-1906; Hooker and Hooker, 1837-1982). 

PLOCOSPERMATACEAE: e, Plocosperma buxfoliolium flower, same laid open; f; pistil, 1,s. of ovary, 

dehiscent fruit showing the solitary seed with a tuft of hairs (after Bentham). 
264 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

tite, the lobes imbricate, very rarely 1 enlarged 

and petaloid (Usteria); petals 4-5 (- 16), connate, 

the lobes imbricate, contorted or valvate; a corona present in Scyphostrychnos; stamens 4-5 

(-16), very rarely 1 (Usteria), the filaments all 

similar, free or connate, inserted on the corolla, 

or rarely hypogynous, alternate with the lobes, 

rarely the anthers sessile; anthers usually elongated, dorsifixed or rarely basifixed, subconnate 

in Gardneria; pollen 3(2-4)-colpate, 3(4)-colporate, 3-4(-6)-porate, or 4-ruporate; disk present 

or absent; pistil 1, the carpels 2 (3-5), style 1 (2 

in Cynoctonum), the stigma+) apical, sometimes 

capitate or clavate; ovary superior or rarely semiinferior (Mitreola), 2( 1 -5)-locular, rarely 2 separate ovaries but l style (Mitreola), the ovules 

generally numerous, but 1 per locule in 1-2 

genera, mostly axile, sometimes parietal, rarely 

basal, unitegmic, tenuinucellar, anatropous, amphitropous, or semi-anatropous; fruit a mostly 

septicidal, sometimes also partially loculicidal or 

rarely circumscissile capsule, sometimes a berry 

or drupe, attachment of the seeds sometimes 

peltate; seeds sometimes minute; embryo 

straight, rarely curved, 0.3 to as long as the 

endosperm; cotyledons thin, 1-2.7 times the 

width of the radicle, 0.1-0.7 the length of the 

embryo; endosperm copious or moderate, fleshy 

to bony. 

Composition: -20 genera, -500 species. 

Distribution: Usually in dry, lowland habitats; 

pantropical, few species temperate. 

PLOCOSPERMATACEAE (Figure 139eJ7.- 

Shrubs and trees; leaves opposite or subverticillate, simple, entire, exstipulate; inflorescences 

axillary, with 1-4 flowers; flowers bisexual, actinomorphic; sepals 5 (6), connate; petals 5 (6), 

connate, imbricate; stamens 5 (6), inserted on the 

corolla, alternipetalous, the filaments elongated; 

anthers dorsifixed, about as long as wide; pollen 

3-colporate; disk small; pistil 1, the carpels 2, 

style 1, twice shortly bilobed apically, the stigmas 

4, clavate; ovary superior, unilocular, the placentas 2, one subapical the other subbasal, the ovules 

2 per placenta; fruit an enlongated bivalved capsule; seeds 1.2-2.3 cm. long, mostly 1, terete, 

with an apical tuft of hairs; embryo linear, 

straight; endosperm scanty, fleshy. 

Composition: 1 genus, 3 species. 

Distribution: Mexico, Central America. 

APOCYNACEAE (Figure 14Oa-e).-Lianas, 

rarely trees, shrubs, and herbs; xylem vessel perforation plates simple, very rarely also scalariform; laticiferous canals and intraxylary phloem 

present; leaves opposite or verticillate, rarely alternate, simple, entire, exstipulate or rarely with 

small interpetiolar stipules; inflorescences paniculate, often with cymose partial inflorescences, 

dichasial cymes or cincinni, or the flowers solitary; flowers bisexual, actinomorphic; sepals 5 

(4), connate, often glandular basally inside; petals 

5 (4), connate, contorted, imbricate, rarely valvate, sometimes with coronal appendages on the 

corolla; stamens 5 (4), inserted on the corolla, 

alternipetalous, the filaments short or rarely 0, 

free or rarely connate, not appendaged; anthers 

basifixed or basally dorsifixed, usually sagittate, 

connivent around the stigma, rarely connate, the 

connective often produced apically; pollen granular, rarely in tetrads, 3(-5)-colporate or 2-3(4)- 

porate; disk usually present around the base of 

the ovary, annular, cupular or of separate glands; 

pistil 1, the carpels 2 (3-5), style 1, the stigmas 

apical; ovaries often 2, superior or semi-inferior, 

the locules 1 or 2, the placentas axile in a bilocular ovary, parietal in a unilocular ovary, the 

ovules 1 or more per placenta, unitegmic, tenuinucellar, k anatropous; fruit 2 (1) follicles, 

berry, drupe, septicidal capsule or schizocarp, 

the seeds often with apical and basal tufts of 

hairs, sometimes winged, or arillate (Tabernaemontaneae); embryo straight or the radicle rarely 

recurved, 0.4 to as long as the endosperm; cotyledons thick or thin, sometimes convolute, 1-3.3 

times the width of the radicle, 0.4-0.6 the length 

of the embryo; endosperm copious, sometimes 

ruminate, scanty or 0, fleshy, sometimes firm. 

Composition: -1 80 genera, - 1500 species. 

Distribution: Pantropical, few species temperate. 

ASCLEPIADACEAE (Figure 14 1 a-c).-Lianas, 

shrubs, herbs, or rarely small trees; xylem vessel 
NUMBER 58 265 

C 

FIGURE 140.-APOCYNACEAE: a, Dipladenia gentianoides flower, D. polymorpha stigma, corolla 

laid open to show pistil and stamens, upper part of corolla opened and showing normal position 

of anthers and style, lower part of corolla tube and calyx (below), seed, embryo and same 

opened, dehiscent fruit; b, Aspidosperma subincanum bud, corolla, pistil, views of anther; c, 

Tabernaemontana reticulata bud, Ambelania tenuzyora C.S. of ovary; d, Geissospermum vellosii 

floral diagram, fruitlets, one in I.s., seed, embryo and endosperm, embryo; e, Hancornia speciosa 

calyx, fruit, I.s. of same, floral diagram (after Martius, 1840-1906). 

perforation plates simple; laticiferous canals and 

intraxylary phloem present; leaves opposite or 

verticillate; rarely alternate (Asclepias spp.), simple, entire, very rarely lobed or dentate, exstipulate; inflorescences terminal or at leaf nodes, 

mostly cymose, often umbelliform; flowers bisexual, sometimes the plants functionally dioecious 

in Periplocoideae, actinomorphic; sepals 5, connate or free; petals 5, connate, the lobes contorted, imbricate or valvate; stamens 5, inserted 
266 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

- -- 

C 

FIGURE 141 .-ASCLEPIADACEAE: a, Periploca graeca I.s. of flower, summit of style, stamen, 

Asclepias candida flower, corona segment; b, Phaeostemma riedelii part of corolla and corona 

laid open, androecium, stamen, pollinia, calyx and pistil, pistil; c, Oxypetalum guilleminianum 

bud, flower, pistil surrounded by stamens and corona, same with corona removed, single 

segment of corona, stamen, Jobinia hernandzfolia dehiscent fruit (after Martius, 1840-1 906; 

Baillon, 1866-1 895). 

on the corolla, alternipetalous, the filaments connate (free in Periploca), short, with a nectariferous coronoid appendage, the corona rarely absent; anthers basifixed, connivent or connate 

above the stigma, acute or acuminate apically; 

pollen usually in pollinia (in masses of 20 in 

Secamoneae, granular or in masses in Periplocoideae, 3-6-aperturate in the latter; disk absent; 

pistil 1, the carpels 2, united by the stigma; 

ovaries 2, superior, very rarely semi-inferior (Atherundra), the placentas ventral, the ovules usually 

numerous, unitegmic, the integument rather 

thick, pseudocrassinucellar or k tenuinucellar, 

anatropous; fruit 2 (1) follicles, the seeds usually 

compressed, mostly with an apical coma; embryo 

straight, 0.8-0.9 the length of the endosperm; 

cotyledons thin, 3-4.5 times wider than the radicle, 0.6-0.7 the length of the embryo; endosperm moderate, firm-fleshy. 

Composition: -200 genera, -2,000 species. 

Distribution: Often in dry habitats; tropical 

and subtropical, centered in south Africa and 

secondarily in South America, few species in 

temperate zones. 
NUMBER 58 267 

CONVOLVULACEAE (Figure 142a-c).-Lianas, 

herbs, shrubs, rarely small trees; xylem vessel 

perforation plates simple; laticiferous canals usually present; intraxylary phloem present; leaves 

alternate, simple, entire, lobed or subcompound, 

sometimes with extrafloral nectaries on the petiole, exstipulate; inflorescences axillary dichasia, 

cincinni or bostryx, rarely umbelliform or thyrse, 

rarely terminal racemes or panicles (Porana), or 

the flowers solitary; flowers bisexual, rarely the 

plants dioecious (Hildebrandtia), actinomorphic, 

rarely zygomorphic (Humbertia); sepals 5 (4), usually free and markedly imbricate, rarely connate 

(Wilsonia); petals 5 (4), connate, plicate-contorted, the lobes rarely imbricate; stamens 5 (4), 

inserted on the corolla toward its base, alternipetalous, the filaments elongated, the anthers 

usually elongated, basifixed; pollen (2)3-colpate 

(sometimes colporate!), polyforate, or polyrugate; disk annular, mostly lobed, surrounding the 

base of the ovary; pistil 1, the carpels 2 (3-5), 

styles 1 (0 in Erycibe, 2 in Cressa), the stigmas 2 

(-S), elongated or globose; ovary superior 2(1- 

5)-locular, sometimes deeply bilobed (Dichondra), the ovules usually 1 or 2 per locule (m in 

Humbertia), axile and subbasal, unitegmic, tenuinucellar or crassinucellar, anatropous; fruit a 

loculicidal capsule, rarely a berry, nut, or schizocarp of 1, 2, or 4 one-seeded mericarps, or 

samaroid; embryo usually folded (linear in Nephrophyllum), 2-3 times longer than the endosperm(?); cotyledons thin 1-9 times wider than 

the radicle, 0.6-0.9 the length of the embryo; 

endosperm moderate or scanty, hard and clear 

or semitransparent. 

Composition: 

Distribution: Various habitats, cosmopolitan 

except the arctic and cool temperate regions of 

the Northern Hemisphere; centered in tropical 

America and Asia, few in temperate zones. 

CUSCUTACEAE (Figure 142d).-Parasitic chlorophylless yellow or orange twining herbs; xylem 

vessel perforation plates simple; leaves 0; inflorescences fascicles; flowers bisexual; sepals 5 (4), 

free or connate; petals 5 (4), connate, the lobes 

imbricate; coronal scales alternate with the sta50 genera, -1 700 species. 

mens; stamens 5 (4), inserted on the corolla, 

alternipetalous; anthers small, basifixed?, the 

connective sometimes produced apically; pollen 

3-4-colpate, 6-rugate or 4-rupate; pistil 1, the 

carpels 2, styles 2 (1); ovary superior, completely 

or incompletely bilocular, the ovules 2 per locule, 

axile, unitegmic, tenuinucellar, anatropous; fruit 

a capsule, dehiscing transversally or irregularly, 

or berry-like; embryo linear, terete, curved or 

spirally twisted, 2-3 times longer than the endosperm?; cotyledons as wide as the radicle, 0.6 the 

length of the embryo or less; endosperm hard, 

moderate. 

Composition: 1 genus, 170 species. 

Distribution: Nearly cosmopolitan, most 

abundant in America, especially warmer regions. 

RUBIACEAE (Figure 142e,f).-Trees, shrubs, 

herbs, lianas; xylem vessel perforation plates simple, in several genera rarely also scalariform, 

usually with few bars; leaves opposite, rarely verticillate (Henriquesia), simple, entire or rarely 

toothed, very rarely sinuate-lobate or pinnatifid, 

stipulate, the stipules often interpetiolar; inflorescences terminal and axillary cymose panicles, 

heads, or small dichasia or rarely the flowers 

solitary; flowers bisexual, or rarely the plants 

dioecious or monoecious, actinomorphic or very 

rarely zygomorphic; calyx lobes (3) 4-5, often 

very small, sometimes one enlarged and petaloid; 

petals 4-5 (3-1 O), connate, the lobes contorted, 

imbricate or valvate, the corolla very rarely bilabiate, very rarely partite nearly to the base; 

stamens 4-5 (3-12), inserted on the corolla, alternipetalous, nearly epigynous in Synaptantha 

and a few other genera, the filaments very rarely 

connate, very rarely strongly unequal; anthers 

basifixed or dorsifixed, usually elongated, very 

rarely connate, the connective very rarely produced apically; pollen 3-polycolpate, 2-4-multicolporate, 3-porate, exceptionally polyforate or 

nonaperturate; disk epigynous, surrounding the 

base of the style; pistil 1, the carpels 2 (3-9), style 

1 (0 in Tetralopha), with 2 (3-4) linear or short 

apical lobes stigmatic on the ventral surface or 1 

and capitate; ovary inferior (superior in Gaertnera and Pagamea, semi-inferior in Synaptantha 
268 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 58 269 

and Henriquezia), bilocular, rarely unilocular or 

4-9 locules, the ovules 1 to ~0 per locule, mostly 

axile, sometimes apical or basal, rarely parietal 

(Gardenia), l-tegmic (0 in Houstonia), the integument massive or thin, crassinucellar or tenuinucellar, anatropous or semi-anatropous; fruit a 

septicidal or loculicidal, rarely circumscissile capsule, berry, drupe, schizocarp (Galium), the seeds 

many to 1, rarely minute, very rarely arillate or 

the apex comose; embryo straight, rarely involute, 0.2 to as long as the endosperm; cotyledons 

thick or thin, 1-4 times the width of the radicle, 

0.1-0.9 the length of the embryo; endosperm 

copious, scanty or 0, soft- to hard-fleshy, rarely 

ruminate or cartilaginous. 

Composition: -500 genera, -6500 species. 

Distribution: Cosmopolitan except arctic 

Northern Hemisphere; concentrated in the tropics and subtropics of the Eastern and Western 

Hemispheres. 

COLUMELLIACEAE (Figure 143a).--Shrubs and 

small trees, bitter in all their parts, the wood very 

hard; xylem vessel perforation plates scalariform; 

leaves opposite, simple, entire or serrate, exstipulate; inflorescences terminal and axillary cymes; 

flowers bisexual, slightly zygomorphic; sepals 5, 

free above the ovary; petals 5, connate, imbricate 

in bud; stamens 2, the filaments very short, inserted on the corolla, alternipetalous; anthers 

unilocular, twisted, the connective broad; pollen 

3-colporate; disk 0; pistil 1, the carpels 2, the 

FIGURE 1 ~~.-CONVOLVULACEAE: a, Convolvulus ottonis calyx, flower laid open, views of anther, stigmas with pollen 

grains attached, I.s. and C.S. of ovary, fruit with part of calyx, 

dehiscent fruit and calyx, seed, C.S. of same showing the 

folded cotyledons: b, Evolvulus maximilianii pistil, floral diagram, Dichondra argentea flower, corolla, part of corolla laid 

open: c, pistil and hypogynous glands, I.s. and C.S. of ovary 

(after Martius, 1840-1906; Johnson). CUSCUTACEAE: d, Cuscuta racemosa flowering shoot attached to host, flower, calyx 

laid open, corolla, same laid open showing appendages, 2 

appendages, stamen, pistils (after Martius, 1840-1 906). RuRIACEAE: e, Coffea arabica inflorescence, corolla in bud, 

flower, views of anther, 1,s. of flower (style and part of 

corolla removed), fruits;f, ovary, I.s. and C.S. of same, seed, 

same opened to show embryo, I.s. of seed, seed and C.S. at 3 

levels, embryo (after Martius, 1840-1 906). 

stigma broad, 2-4-lobed; ovary inferior, imperfectly bilocular, placentas 2, parietal, the ovules 

numerous; fruit a septicidal capsule, the seeds 

numerous, smooth; embryo minute, straight, 0.2 

the length of the endosperm; cotyledons as wide 

as the radicle, 0.4 the length of the embryo; 

endosperm copious, fleshy. 

Composition: 1 genus, 4 species. 

Distribution: Andes from Colombia to Bolivia, at k 2000-3000 m. elevation. 

GENTIANACEAE (Figure 1436-d).-Herbs, 

rarely saprophytic, very rarely shrubs; xylem vessel perforation plates simple; intraxylar phloem 

present; leaves opposite, rarely alternate (Swertia 

spp.), rarely verticillate, simple, entire, exstipulate; inflorescences usually terminal and axillary 

dichasia, sometimes monochasia or other type of 

cyme, or the flowers solitary; flowers actinomorphic, however the stamens rarely declinate, 

bisexual or the plants rarely polygamous; sepals 

4-5 (2-1 2), connate, sometimes with basal nectar 

glands; petals 4-5 (-1 2), connate, contorted or 

very rarely imbricate, rarely with coronal scales, 

rarely spurred; stamens 4-5 (-12), rarely some 

staminodal (only 1 fertile in Hoppea), inserted on 

the corolla, alternipetalous, sometimes 2 nectaries on each corolla segment; anthers elongated, 

basifixed or dorsifixed near the base, longitudinally dehiscent or poricidal, occasionally connate 

(Voyria, Leiphaimos spp. and others), rarely with 

1-4 apical or basal glands, the connective rarely 

produced apically; pollen rarely in polyads, usually 3(2-4)-colporate, exceptionally 4-ruporate 

or &rugorate, 1-3-porate in the saprophytic genera; disk annular or of 5 hypogynous glands, 

rarely absent; pistil 1, the carpels 2, style 1, bifid 

apically and the stigmas linear, or not divided 

and the stigma capitate; ovary superior, mostly 

unilocular, sometimes bilocular, the ovules numerous, mostly parietal, sometimes axile, unitegmic, tenuinucellar, anatropous or rarely & orthotropous; fruit a septicidal capsule, very rarely like 

a lantern, rarely a berry; seeds minute, numerous; embryo subglobular, linear or spatulate, 

0.2-0.9 the length of the endosperm; cotyledons 1-2 times the width of the radicle, 
270 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 58 27 1 

0.1-0.5 the length of the embryo; endosperm S A N TA LA L ES 

copious, fleshy. 

Composition: 

Distribution: Various habitats; cosmopolitan, 

most common in temperate regions and in mountains in the tropics. 

MENYANTHACEAE (Figure 143e-g).-Aquatic 

and marsh herbs; xylem vessel perforation plates 

simple or scalariform; intraxylar phloem absent; 

leaves alternate and radical, sometimes opposite 

under the flower, simple or compound, entire or 

toothed, exstipulate; inflorescence a raceme, 

panicle, cyme, fascicle, head, or the flowers solitary or paired; flowers bisexual, actinomorphic; 

sepals 5, free or connate; petals 5, connate, induplicate valvate, rarely slightly imbricate in bud; 

stamens 5, inserted on the corolla, alternipetalous; anthers mostly sagittate, dorsifixed?; pollen 

3-colporate, exceptionally 6-rugate; hypogynous 

nectaries usually present; pistil 1, the carpels 2, 

style 1, shortly bifid apically, the stigmas 2; ovary 

superior or semi-inferior, unilocular, the ovules 

numerous, parietal, unitegmic, tenuinucellar, 

anatropous; fruit a capsule, dehiscing by 2 or 4 

valves or irregularly, rarely a berry; seeds numerous to few; embryo linear, straight, 0.5-0.75 

as long as the endosperm; cotyledons as wide as 

the radicle, 0.3 the length of the embryo; endosperm copious, fleshy. 

80 genera, - 1000 species. 

Composition: 5 genera, 40 species. 

Distribution: Almost cosmopolitan. 

FIGURE 143.-cOLUMELLIACEAE: a, COlUmt?llia oblonga flowering shoot, bud, floiver, 1,s. of same, corolla laid open 

showing insertion of stamens, dehiscent fruit (after Baillon, 

1866- 1895). GENTIANACEAE: b, Centaurzum sp. inflorescence, pistil, Gentiana acaulis I.s. of flower, floral diagram, 

dehiscent fruit, seed, 1,s. of same showing the small embryo: 

c, Tapeinostemon spenneroides pistil, stigma, fruit, C.S. of same; 

d, Schultesia angustifolia flower, corolla laid open, calyx, 

views of stamen, views of pistil, C.S. of ovary, S. doniana 

pistil, stigmas (below) (after Le Maout and Decaisne, 1873; 

Martius. 1840- 1906). MENYANTHACEAE: e, Menyanthes trifoliata flowering shoot, flower, I.s. of same; f; iVymphozdes 

indica flower, stamen; g, pistil, fruit, flowering shoot (after 

Baillon, 1866-1 895; Cabrera, 1965). 

Trees, shrubs, and herbs, sometimes climbing, 

usually parasitic on roots and stems but containing chlorophyll; xylem vessel perforation plates 

scalariform, usually with fewer than 20 bars, 

simple, or both; secretory canals and intraxylary 

or extraxylary phloem sometimes present; leaves 

alternate or opposite, simple, usually entire, 

sometimes crenate, rarely lobed or reduced to 

scales, exstipulate, rarely stipulate; flowers bisexual or unisexual; sepals 3-6 (-8, 0), usually minute and connate, rarely accrescent; petals 0, 3- 

6 (-8), free or connate; stamens 3-6 (1-20) in 1 

or 2 series; staminodes sometimes present; filaments free or connate, hypogynous, epigynous 

or inserted on the corolla or calyx, the anthers 

basifixed or dorsifixed; pollen 3(2-4)-colpate, 3- 

colporoidate, 3-4(5)-porate, 4-1 2-forate; glandular disk present or absent; pistil 1, the carpels 

2-5 (l), style 1 (2-4), usually short, the stigma(s) 

apical, less commonly decurrent ventrally, sometimes sessile; ovary superior to inferior, 1(-5)- 

locular, the multilocular ovary usually unilocular 

at apex, the ovules not developed before fertilization or 1-3 (-B), 0-2-tegmic, axile, or from 

the apex of a free-central placenta, or basal; fruit 

a drupe, berry, or nutlet, rarely a samara, achene 

or coriaceous capsule; seed 1, the embryo usually 

minute or small, straight, sometimes poorly differentiated, the cotyledons sometimes 3-6 or 0: 

endosperm usually copious, oily, sometimes absent. 

Distribution: Cosmopolitan, especially pantropical and -subtropical, extending into temperate regions, in wet and dry habitats. 

Aucubin (iridoid) glycoside is absent. Gibbs says we have difficulty defining the 

order and its families from their chemistry. The 

somatic lipids of Olacaceae, Opiliaceae, Santalaceae, and Loranthaceae have similar acetylenic 

acids, lending strong support to the idea that 

these families are closely related. The seed fats 

of Olacaceae and Santalaceae have similar acetylenic acids. Santalaceae have terpenoids. 

OLACACEAE (Figure 144a,b).-Trees, shrubs, 

Chemistry: 
272 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

b 

FIGURE 144.-oLACACEAE: a, Ximenia coriacea bud, flower, views of petal, views of stamen, C.S. 

of anther before and during dehiscence, pistil, 1,s. of same, C.S. of ovary, floral diagram; b, 

Ptychopetalum olacoides I.s. of pistil, corolla laid open, views of anther, Chaunochiton loranthoides 

flower, views of anther, C.S. of anther, fruit, seed and arcuate placenta to left, I.s. of seed 

showing the minute embryo (after Martius, 1840-1 906). APTANDRACEAE: c, Aptandra Eiriosmoides axillary inflorescence, part of inflorescence, bud, flower, staminal tube and disk before 

and after dehiscence of the anthers, I.s. of staminal tube to show the pistil, 1,s. of ovary, A. 

tubzcina accrescent calyx enclosing the immature fruit, mature fruit and calyx (below) (after 

Martius, 1840-1906). 

and climbers, very rarely subherbaceous, in part inflorescences axillary, small, cymose, or a thyrse; 

hemiparasitic; xylem vessel perforation plates ex- flowers usually bisexual, the plants sometimes 

clusively scalariform, simple, or both, usually polygamodioecious; sepals 3-6, connate, rewith few bars (6-10) but more than 20 in Heis- duced, sometimes accrescent; petals 3-6, free or 

teria; leaves alternate, simple, entire, exstipulate; connate, mostly valvate; stamens 3-20, in 1-2 
NUMBER 58 

series, sometimes only opposite the petals, the 

filaments hypogynous or inserted on the corolla, 

sometimes staminodes present; anthers as long as 

wide or oblong, dehiscing longitudinally or by 

short slits, rarely by valves; pollen 3-colpate, 3- 

colporoidate, 3-4(5)-porate, 6(8)-forate, 3-4- 

diploforate; disk often annular; pistil 1, the carpels 2-5, style 1, stigma 2-5-lobed, apical; ovary 

superior, rarely semi-inferior, 1 -5-locular at the 

base, usually unilocular above, the ovules 0-1 - 

2-tegmic, tenuinucellar or pseudocrassinucellar, 

anatropous, mostly 1 per locule, axile or 1-5 

from the apex of a free-central placenta in the 

unilocular ovaries; fruit a drupe or nut, the seed 

1: embryo minute to medium-sized, straight, 0.3 

the length of the endosperm, barely differentiated; endosperm copious, oily. 

Composition: 25 genera, 240 species. 

Distribution: Pantropical, especially Africa 

and Asia, secondarily America; warm temperate 

North America to warm temperate South America; Africa except the northern part; India, Japan, China, Indochina, the East Indies, northeastern Australia, western Pacific islands. 

APTANDRACEAE (Figure 144c).-Trees; xylem 

vessel perforation plates simple; leaves alternate, 

simple, entire, exstipulate; inflorescences axillary 

racemes or panicles, ultimate divisions umbelliform; flowers bisexual or the plants dioecious; 

sepals 4-8, connate, at first small, later accrescent; petals 4-8, free, valvate; stamens 5-4, connate in a column surrounding the pistil, the anthers sessile; pollen 4(5)-porate; disk annular or 

lobed, between the petals and stamens; pistil 1, 

the carpels 2, style 1, elongated or the stigmas 

sessile; ovary superior 2- 1 -locular, the ovules 

2(?), pendulous from the apex of the free central 

placenta, ategmic; fruit a drupe, the seed 1; 

embryo minute, 0.2 the length of the endosperm, 

barely differentiated; endosperm copious, oily. 

Composition: 1 genus, 4 species. 

Distribution: Tropical America and west Africa; Malaya, Indochina. 

OCTOKNEMACEAE (Figure 145a,b).-Trees 

and shrubs; xylem vessel perforation plates scalariform, usually with fewer than 20 bars, or 

273 

simple; leaves alternate, simple, entire, exstipulate; inflorescences axillary racemes; plants dioecious; sepals 5, connate, small; petals 0 or 5; 

stamens 5, the filaments free; anthers about as 

long as wide, basifixed; pollen 3-colporoidate; 

disk lobed or 0; pistil 1, the carpels 3-5, style 1, 

short, 3-5-lobed apically, the lobes bifid; ovary 

inferior, unilocular or nearly 3(4)-locular, the 

ovules 3 (4), apical, undifferentiated, the embryo 

sac resembles that of Olacaceae; fruit a drupe, 

the seed 1, longitudinally 8(6-lO)-ribbed; embryo small, the radicle much longer than the 

cotyledons; endosperm copious, slightly ruminate. 

Composition: 1 genus, 6 species. 

Distribution: Tropical Africa. 

OPILIACEAE (Figure 145c).-Trees, shrubs, 

and lianas, all or most root-parasites; xylem vessel 

perforation plates simple; leaves alternate, simple, entire, exstipulate; inflorescences axillary racemes, spikes, umbels or panicles; flowers bisexual or the plants dioecious; calyx minute, the 

lobes obsolete; petals 4-5 (0), free or connate; 

stamens 4-5, opposite the petals, rarely alternipetalous, the filaments short or filiform, free or 

epipetalous; anthers basifixed; pollen 3-colporate 

or 3-colporoidate; disk glands alternating with 

the stamens sometimes connate; pistil 1, the carpel 1, style 1, short, the stigma apical, punctiform; ovary superior or semi-inferior, unilocular, 

the ovule solitary, basal, 0- 1-tegmic, tenuinucellar, anatropous or orthotropous; fruit a drupe; 

embryo small, linear, straight, 0.5 the length of 

the endosperm; cotyledons not broadened, 0.5 

the length of the embryo; endosperm copious. 

Composition: 8 genera, 60 species. 

Distribution: Pantropical and -subtropical, especially Asia. 

MEDUSANDRACEAE (Figure 146a,b).-Trees 

with secretory canals throughout, buds perulate; 

xylem vessel perforation plates scalariform, commonly with 20 but as many as 50 or more bars; 

leaves simple, alternate, crenulate, stipulate; inflorescences axillary slender pendulous racemes; 

flowers bisexual, actinomorphic; sepals 5, free or 

very shortly connate, open in bud, accrescent; 
274 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 145.-oCTOKNEMACEAE: a, Octoknema klaineana inflorescence, 6 flower, 1.s. of same 

with pistillode, 0 flower, I.s. of same, P flower from above, I.s. of fruit showing the minute 

embryo, C.S. of fruit; b, 0. af$nis lateral view of stigmas, style and staminodes, 0. winkleri I.s. 

and C.S. of fruit, seed, 0. affinis embryo (enlarged) (after Pilger and Krause). OPILIACEAE: c, 

Agonandra brasiliensis 0 inflorescence, part of d inflorescence, floral diagram, bud, petal (dorsal 

view), androecium and disk, views of stamen and ventral view of petal, pistil and disk, I.s. of 

pistil (after Martius, 1840- 1906). 

petals 5, free, imbricate; stamens 5, oppositipetalous, the filaments short, not connate but 

often very shortly adnate to the base of the petals, 

staminodes 5, much elongated at anthesis, becoming several times longer than the stamens; 

anthers 4-locular, about as long as wide, dorsifixed, dehiscing by dorsal and ventral valves; 

pollen 3-colporoidate; disk absent; pistil 1, the 

carpels 3 (4), styles 3 (4), short, conical, free, 

distant, the stigmas apical, minute; ovary superior, unilocular, the placenta free central, ovules 

6 (-8) apical-axile; fruit a 3(4)-valved coriaceous 

capsule, the seed solitary, large; embryo small, 

straight, 0.3 the length of the endosperm, the 

cotyledons not broadened, 0.2 the length of the 

embryo; endosperm copious, slightly ruminate. 

Composition: 1 genus, 1 species. 

Distribution: Rain forest; tropical west Africa. 

CARDIOPTERIDACEAE (Figure 146c, d).- 

Climbing twining herbs with articulated laticiferous tubes; xylem vessel perforation plates simple; 

leaves simple, alternate, entire or lobed, exstipulate; inflorescences secund axillary loose 

panicle of cymes, ebracteate; flowers minute, actinomorphic, ebracteate, bisexual or the plants 

andromonoecious; sepals 5 (4), shortly connate, 
NUMBER 58 275 

FIGURE 146.-hfEDUSANDRACEAE: a, hfedusandra richardsiana flower, bud, floral diagram, 

stamen and petal, pistil, I.s. of ovary, inflorescence, part of same enlarged; b, fruit, dehisced 

fruit, seed, section of same showing the small embryo (black), cavity (hatched) and endosperm 

(after Hutchinson and Dalziel). CARDIOPTERIDACEAE: c, Perzpterygium quznquelobum inflorescence superimposed on leaf, axillary infructescence, bud, corolla laid open, anther, 1,s. of ovary 

(note the unequal styles), fruit, P. platycarpurn pistil; d, bud of d flower, pistillode, 2 sepals, 3 

petals and a stamen (after Hutchinson, 1973; Lecomte). 

imbricate; petals 5 (4), shortly connate, imbricate; stamens 5 (4) adnate to the corolla, alterComposition: 1 genus, 3 species. 

Distribution: Assam, southeastern Asia, Manipetalous, the filaments short, anthers dorsifixed, about as long as wide to oblong, longitudinally dehiscent; pollen 3-colpor(oid)ate; disk 0; 

pistil 1, the carpels 2, styles 2 or 1 and apically 

bifid, one accrescent, the other short, the stigmas 

apical, capitate; ovary superior, unilocular, the 

ovules 2, naked(?), one sometimes aborting, apical; fruit a flat samara, the seed 1, linear; embryo 

very minute, the endosperm copious, fleshy. 

laysia, Queensland. 

SANTALACEAE (Figure 147a,b).-Trees, 

shrubs, and herbs, sometimes parasitic on roots 

or stems (Phacellaria on Loranthaceae); xylem 

vessel perforation plates simple; leaves alternate 

or opposite, simple, entire, sometimes reduced 

to scales, exstipulate; inflorescences axillary and 

terminal racemes, spikes, heads, often cymose, 

sometimes the flower solitary; flowers small, bi- 
276 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 147.-sANTALACEAE: a, Acanthosyris spinescens part of flowering branch, floral diagram, 

inflorescence in bud, flower, 1,s. of same laid open (also showing disk), Is. of ovary; b, Thesium 

aphyllum 1,s. of flower, apical and lateral views of fruit, I.s. of same showing seed with embryo 

and endosperm (after Martius, 1840-1 906). LORANTHACEAE: c, Psittacanthus glaucoma I.s. of 

bud, P. biternatus flower, views of anther, fruit, I.s. and C.S. of same; d, Struthanthus uulgaris 

pistil, petal and stamen of exterior series, same of interior series, views of stamen of interior 

series, C.S. and I.s. of fruit (embryo linear, surrounded by endosperm) (after Martius, 1840- 

1906). 

sexual or the plants monoecious or dioecious; 

sepals 3-6, free or basally connate, valvate or 

slightly imbricate, sepaloid or petaloid; petals 0; 

stamens 3-6, inserted on the calyx opposite the 

lobes; anthers about as long as wide, basifixed or 

dorsifixed; pollen 3-brevicolpate, colporoidate, 

3-colporate, 3-porate; disk perigynous or epigynous; pistil 1, the carpels (2) 3-5, style 1, the 

stigma 1 and capitate or (2) 3-5-lobed; ovary 

unilocular, inferior, semi-inferior or rarely superior (Anthoboleae), the ovules 1-3 (4, 5), not 

developed in Anthoboleae, basal, 0- 1 -tegmic, 
NUMBER 58 277 

tenuinucellar, orthotropous, anatropous or semianatropous; fruit a nut or drupe; seed 1, without 

a testa; embryo linear, terete, straight or minute 

and undifferentiated, 0.1-0.95 the length of the 

endosperm; cotyledons thin, 0.3-0.7 the length 

of the embryo; endosperm copious, fleshy. 

Composition: 30 genera, 400 species. 

Distribution: Often dry habitats; cosmopolitan except the arctic region, most common in 

tropics and subtropics. 

LORANTHACEAE (Figure 147c,d).-Shrubs, 

more rarely herbs, hemiparasites on the branches 

of trees or rarely terrestrial trees or shrubs parasitic on roots; xylem vessel perforation plates 

simple; secretory canals and intraxylary or extraxylary phloem sometimes present; leaves 

mostly opposite or whorled, rarely alternate, simple, entire, sometimes reduced to scales, exstipulate; inflorescences cymose, sometimes spikes or 

racemes, the flowers solitary or 2-3 together in 

axils of bracts or extra-axillary, rarely terminal; 

flowers bisexual or the plant dioecious; sepals 3- 

4, free above the ovary or connate and 4-6- 

toothed or truncate (0 in Antiduphne); petals 4- 

6 or 0, free or connate; stamens 3-6, epigynous 

or inserted on the perianth opposite the lobes or 

at their base; anthers dorsifixed, sometimes unilocular or transversely locellate, dehiscing longitudinally or by terminal pores or transverse slits; 

pollen 3( 2-4)-colpate, less frequently colpor(oid)ate; disk present or absent; pistil 1, the 

carpels 3(?), style 1 or the stigma sessile, apical; 

ovary inferior, unilocular, the ovules not differentiated, the placenta basal; fruit a berry, drupe 

or samaroid; seed 1, without a testa; embryo 

linear, small, straight, 0.3-1 25 times the length 

of the endosperm; cotyledons sometimes 3-6 or 

0, 0.1-0.5 the length of the embryo; endosperm 

copious, succulent or sometimes 0. 

Composition: 35 genera, 1300 species. 

Distribution: Pantropical, especially Southern 

Hemisphere, extending into the temperate 

zones. 

MISODENDRACEAE (Figure 148u-c).-Subshrubs hemiparasites on the branches of Nothofugus trees; xylem vessel perforation plates simple; leaves alternate, simple, entire or crenate, 

sometimes scale-like, exstipulate; plants dioecious; inflorescences axillary and terminal small 

spikes, racemes, head or the flowers solitary; S 

flower: perianth 0; stamens 2-3 (1-4), the filaments elongated, inserted around a disk; anthers 

basifixed, unilocular, dehiscing by 2 valves or a 

tangential slit; pollen 4-1 2-forate; Q flower: calyx adnate to ovary?; petals 0; disk epigynous; 

pistil 1, the carpels 3, style 1, short, the stigmas 

3, decurrent ventrally; ovary inferior, incompletely 3-locular, the ovules 3, basal; fruit a nutlet 

or achene distributed by 3 enlarged, feathery 

setae; seed 1, without a testa; embryo straight, 

minute, 0.1-0.3 the length of the endosperm; 

cotyledons minute, hardly differentiated, shorter 

than the radicle; endosperm copious, fleshy. 

Composition: 1 genus, 10 species. 

Distribution: Nothofugus forests of southern 

third of the Andes. 

GRUBBIACEAE (Figure 149a,b).-Small shrubs, 

not parasitic; xylem vessel perforation plates scalariform, with 20-40 or more bars; leaves opposite, simple, linear, entire, exstipulate; inflorescences axillary 3-flowered dichasia or strobiloid 

compound dichasia; flowers bisexual, minute; sepals 4, valvate; petals 0; stamens 8, in 2 series, 

the filaments free; anthers medifixed or basifixed, small, dehiscing by valves; pollen 3-colporate, 3-colporoidate; disk epigynous, annular; pistil 1, the carpels 2, style 1, short, the stigmas 2, 

apical; ovary inferior, initially bilocular, becoming unilocular, the ovules 2, unitegmic, tenuinucellar, anatropous, the placentas axile, becoming 

free-central or basal; fruit a drupe or nutlet, the 

seed 1; embryo linear, subterete, 0.7-0.8 the 

length of the endosperm; cotyledons as wide as 

the radicle, 0.3-0.4 the length of the embryo; 

endosperm copious, fleshy and oily. 

Composition: 1 genus, 5 species. 

Distribution: Cape Province, South Africa. 

TH YMELA EA L ES 

Trees and shrubs, rarely herbs or lianas, usually with tough fibrous bast; xylem vessel perfo- 
278 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

w 

FIGURE 148.-bfISODENDRACEAE: a, Misodendron punctulatum d plant, leaf with d inflorescence, 

d flower, I.s. and C.S. of anther, 0 flower, I.s. of same; b, 0 plant in fruit, fruit, 1,s. of same, seed 

with embryo at apex, embryo; c, M. brachystachyum seed germinating within the fruit and 

attaching to a branch, 0 flower, I.s. of same, C.S. of ovary (after Le Maout and Decaisne, 1873; 

Baillon, 1866-1895; Hooker and Hooker, 1837-1982). 

ration plates usually simple, rarely scalariform; 

intraxylary phloem usually present, sometimes 

interxylary phloem, occasionally mucilaginous 

secretory canals and spaces; leaves alternate or 

opposite, simple, entire, mostly exstipulate; flowers usually bisexual; sepals 4-5 (3-6), connate 

into a tube, rarely nearly free, often petaloid; 

petals 0 or 4-12, mostly scale-like, free or connate, inserted on the calyx tube; stamens 4-10 

(1 to -80) in 1 or 2 series, inserted on the calyx, 

the filaments mostly not connate; anthers basifixed, dorsifixed or adnate, rarely apiculate; pollen usually oligo-polyforate, sometimes 3-colporate; glandular disk hypogynous or adnate to the 

calyx, sometimes absent; pistil 1, the carpels 2 

(3- 12), often pseudomonomerous, often heterostylous, the styles 1 (4), often excentric, 

stigma(s) apical; ovary superior, l(2- 1 2)-locular, 

the ovules 1 (2) per locule, axile, sometimes 

subapical; embryo sac sometimes with increased 

number of antipodals; fruit usually an achene, 

nut, berry, or drupe, sometimes enclosed by a 

fleshy calyx tube, rarely a loculicidal capsule, the 

seed(s) often arillate or carunculate; embryo usually large, straight; endosperm mostly scanty or 

absent, rarely copious. 
NUMBER 58 279 

C 

FIGURE 149.-GRUBBIACEAE: a, Grubbia rosmarinzfofa flowering shoot, glomerule with its 

bracts and without them, single flower, 1,s. of same, petal and stamens; b, flower laid open, C.S. 

of ovary, fruit from 3 flowers, I.s. of same showing 1 fertile fruit and minute embryo in 

abundant endosperm, G. stricta 1.5. of fruit showing the entire seed (after Baillon, 1866-1895; 

Marloth, 1925). GEISSOLOMATACEAE: c, Geissoloma marginaturn flowering twig, floral diagram 

(nectar foveoles internal to the stamens), flower, part of perianth showing insertion of stamens, 

views of anther; d, pistil, Is. of same, C.S. of ovary, dehiscent fruit (the seeds of 2 locules 

exserted), seed with aril, 1,s. of same showing the minute embryo (after Marloth, 1925; Gilg). 

Distribution: Centered in Africa and Australia, extending to temperate and tropical Eurasia and America. 

GEISSOLOMATACEAE (Figure 149c,d).-A low 

shrub; xylem vessel perforation plates scalariform with 12-33 bars; intraxylary phloem absent; leaves simple, opposite, decussate, entire, 

coriaceous, the stipules minute, 1 mm., or absent(?); flowers solitary, bisexual, axillary, actinomorphic, closely subtended by 6 decussate 

bracts; sepals 4, very shortly connate, shortly 

acuminate, petaloid, imbricate in bud; petals 0; 

stamens 8, in 2 whorls, inserted on the base of 

the calyx, the filaments elongated, not connate; 

anthers dorsifixed, ellipsoid, small, about as long 

as wide, the connective slightly produced apically, the sacs basally separate, dehiscing longitudinally; pollen 3-colporate; 8 foveolate glands 
280 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

present between the ovary and stamens; pistil 1, 

the carpels 4, styles 4, free basally, connate apically, the stigma(s) apical, not enlarged; ovary 

superior, 4-locular, the carpels loosely connate, 

the ovules 2 per locule, apical-axile, bitegmic, 

anatropous, the embryo sac 8 nucleate (Polygonum type); fruit a 4-winged loculicidal capsule, 

the seeds often 1 per locule, carunculate; embryo 

straight, as long as the endosperm according to 

some, with foliaceous or linear cotyledons and 

short radicle, but the illustration in Marloth 

shows a small embryo, 0.2 the length of the 

endosperm and cotyledons 0.5 the length of the 

embryo; endosperm copious, fleshy. 

Composition: 1 genus, 1 species. 

Distribution: 

GONYSTYLACEAE (Figure 150a,b).-Trees, 

sometimes large, rarely shrubs; xylem vessel perforation plates simple, phloem fibers abundant, 

intraxylary phloem absent, mucilage secretory 

cells or spaces usually present in the leaves and 

stem; leaves alternate or rarely opposite or 

subopposite, simple, entire, coriaceous, the lateral veins numerous, relatively inconspicuous, 

the midrib prominent beneath, mostly glandularpunctate, exstipulate; inflorescences a terminal 

thyrse or rarely subumbel; flowers bisexual, actinomorphic; sepals 5, shortly connate, the lobes 

imbricate or rarely valvate in bud; petals minute 

(disk of some authors) 7-40, deltoid to linearsubulate or rarely an entire low annulus; stamens 

usually numerous, 8-80, the filaments free or 

rarely connate in groups (Lethedon spp.), inserted 

on the base of the calyx, the filaments short, 

filiform; anthers oblong, basifixed, sometimes 

hippocrepiform, dehiscing longitudinally; pollen 

oligoforate, with small spinulose excrescences; 

disk 0; pistil 1, the carpels 3, 5 (2-12), style 1, 

elongated, filiform, bent, much longer than the 

stamens, sometimes parastyles present, the 

stigma small, capitate, clavate or rarely obscurely 

lobed; ovary 3, 5 (2-12)-locular, the ovules 1 per 

locule, axile-subapical, bitegmic, crassinucellar, 

anatropous; fruit usually a woody loculicidal capsule, or rarely soft (indehiscent?), the seeds large, 

arillate; cotyledons thick, radicle short; endosperm 0. 

South Africa, in dry habitats. 

Composition: 1 genus, 25 species. 

Distribution: Southeastern Asia, the East Indies to Fiji. 

THYMELAEACEAE (Figure 15Oc,d).-Usually 

shrubs, sometimes trees, rarely herbs or lianas, 

with tough fibrous bast, many poisonous with 

acrid, rarely caustic sap; xylem vessel perforation 

plates simple, intraxylary phloem present except 

in Lagetta and Drapetes, interxylary phloem occurs in some genera, mucilaginous secretory canals and spaces sometimes present; leaves alternate or opposite, simple, entire, exstipulate; inflorescences terminal, axillary, at nodes of fallen 

leaves, rarely cauliflorous, spikes, racemes, 

heads, umbels, fascicles or rarely cymes, or the 

flowers rarely solitary, sometimes involucrate, 

the bracts rarely colored; flowers usually bisexual 

or sometimes the plants dioecious or polygamous, actinomorphic or rarely slightly zygomorphic, rarely dimorphic, often sweetly scented; 

sepals 4-5 (3-6), connate into a tube or rarely 

nearly free, the calyx often petaloid, the lobes 

imbricate in bud or rarely absent; petals 4-12, 

mostly scale-like, free, connate or absent, usually 

smaller than the calyx lobes, inserted on the 

calyx-tube at its mouth or slightly below; stamens 

4-1 0 (1 -2 in Pimelea), usually in 2 series, sometimes 1 and then opposite or alternate with the 

calyx lobes, inserted on the calyx-tube at its 

mouth or within the tube, the filaments short or 

long, not connate; anthers as long as to twice as 

long as wide, basifixed, dorsifixed or adnate, 

rarely the connective produced apically, dehiscing longitudinally; pollen usually ‘oligo-polyforate, usually with vestigial spinuloid excrescences 

and patterns f similar to the “croton-pattern;” 

disk hypogynous, annular, cupular, or of separate 

scales or sometimes adnate to the calyx-tube or 

0; pistil 1 , sometimes stipitate, pseudomonomerous or the carpels 2, often heterostylous, the style 

1, short or elongated, often excentric, the stigma 

capitate, discoid or ovoid, often papillose or penicillate, rarely punctiform; ovary superior, l(2)- 

locular, the ovules 1 per ovary or locule, pendulous, but vascular strand basal, or axile, bitegmic, 

crassinucellar, anatropous or rarely semi-anatropous to nearly orthotropous, the embryo sac 
NUMBER 58 28 1 

FIGURE 1 ~~.-GONYSTYLACEAE: a, Gonystylus bancanus part of inflorescence, I.s. of flower, 

calyx from below, views of stamen, seed, embryo; b, G. borneensis floral diagram, G. forbesii 

floral diagram, G. bancanus fruit, I.s. and C.S. of same (after Koorders and Valeton, Domke, 

Miquel). THYMELAEACEAE: c, Daphnopsis brasiliensis bud, 8 flower with pistillode (enlarged), P 

flower, fruit, I.s. of same, D. martii part of 8 flower laid open, with pistillode, anther before 

and during dehiscence, pistillode with hypogynous glands, Schoenobiblus daphnoides 8 flower; 

d, Linostoma calophylloides flower laid open, portion of perianth further enlarged showing the 

squamulae on the perianth, floral diagram, stigma, I.s. of ovary, views of anther, Lasiadenia 

rupestris fruit, 1,s. of same showing seed and embryo (after Martius, 1840-1906). 
282 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

sometimes with increased number of antipodals; 

fruit an achene, nut, berry or drupe, sometimes 

enclosed by a fleshy calyx-tube, rarely a loculicidal capsule (Aquilaria, Gyrinops), the seed(s) often 

arillate or carunculate; embryo straight, as long 

as the endosperm, the cotyledons thick, planoconvex, rarely flat (Gnidia balzeri), rarely narrow 

and semi-cylindrical (Lachnaea, Pimelea, Rhamnoneuron), 0.6-0.9 the length of the embryo and 

to 5.5 times wider than the radicle, the latter 

sometimes invested; endosperm mostly scanty or 

absent, rarely copious (Lachnaea, Pimelea). 

Composition: 48 genera, -500 species. 

Distribution: Centered in Africa and Australia, extending to temperate and tropical Eurasia, the East Indies and New Zealand; disjunct 

areas in temperate and tropical America. 

POLEMONKALES 

Herbs, shrubs, lianas, and small trees; xylem 

vessel perforation plates simple, occasionally also 

scalariform with few bars; leaves alternate, less 

commonly opposite, usually entire and simple, 

sometimes toothed, lobed, or compound, exstipulate; flowers bisexual, rarely unisexual, actinomorphic, rarely zygomorphic; sepals 5 (4- 

12), free or connate; petals 5 (4-12), connate, 

sometimes with coronal scales; stamens 5 (1, 4- 

15) in 1 (2) series, epipetalous, rarely inserted at 

2 levels, rarely unequal, rarely some staminodal; 

anthers dorsifixed or basifixed, sometimes acute 

apically, rarely connate; pollen 3(2- 1 2)-colporate, 6-1 O-porate, oligo-polyforate, 3-8-colpoidorate, 3-colporoidate, sometimes 3-colp(or)ate3-pseudo-colpate, 2-colpate, 6-9-rugate; glandular disk annular, hypogynous, or absent; pistil 

1, the carpels 2-3 (4), sometimes heterostyled, 

the style 1 (2), sometimes 2-4-lobed, often gynobasic, the stigmas apical or linear; ovary superior, very rarely semi-inferior, 1-3 (4-1 0)-locular, the ovules l-many per locule, axile or parietal, sometimes subbasal, unitegmic, rarely bitegmic; fruit nutlets, drupe, berry, or loculicidal 

or rarely septicidal capsule; seeds rarely winged 

or long-hairy; embryo straight or slightly curved, 

rarely folded, usually large; endosperm copious 

to absent. 

Distribution: Cosmopolitan, most common in 

western America and the Mediterranean region, 

most common in dry habitats. 

Chemistry: Boraginaceae seed-fats have some 

unusual fatty-acids. It is the only family in the 

“Tubiflorae” with pyrrolizidine alkaloids, and in 

great number. 

POLEMONIACEAE (Figure 15 la-c).-Herbs or 

rarely shrubs, lianas, or a small tree (Cantua); 

xylem vessel perforation plates simple, also scalariform with 1-10 bars in some species; leaves 

alternate or opposite, entire, rarely toothed (Loeselia) or pinnately compound, exstipulate; inflorescences terminal, rarely axillary corymbs, involucrate heads or cymes, rarely the flowers solitary (Cobaea); flowers bisexual, actinomorphic, 

rarely somewhat zygomorphic; sepals 5 (4), connate; petals 5 (4), connate, contorted in bud; 

stamens 5, epipetalous, alternate with the corolla 

lobes, the filaments long or short; anthers longer 

than wide, dorsifixed; pollen 6- 1 O-porate, oligopolyforate, or 3-8-colpoidorate; disk annular, 

surrounding the base of the ovary; pistil 1, the 

carpels 3 (2, 4), the style 1, filiform, the stigmas 

3 (2, 4), linear; ovary superior, 3(2, 4)-locular, 

the ovules 1 or more per locule, axile, unitegmic, 

tenuinucellar, anatropous; fruit a loculicidal or 

rarely septicidal (Cobaea) capsule, or rarely indehiscent; seeds rarely minute; embryo straight 

or slightly curved, 0.9 the length of the endosperm; cotyledons moderately thick, 1-2.8 times 

the width of the radicle, 0.3-0.6 the length of 

the embryo; endosperm mostly copious or moderate, fleshy or firm-fleshy, sparse or lacking in 

Cobaea, Cantua, and Bonplandia. 

Composition: 16 genera, 300 species. 

Distribution: Centered in temperate western 

North America; most of North America; Alaska 

to the southern tip of South America; temperate 

Eurasia. 

FOUQUIERIACEAE (Figure 15 1 d,e).-Shrubs 

and small trees; xylem vessel perforation plates 

simple or with occasional scalariform plates with 

a few bars; leaves alternate, simple, entire, the 
NUMBER 58 283 

FIGURE 15 1 .-POLEMONIACEAE: a, Polemonium caeruleum flowering shoot, floral diagram, 

flower, Is. of same, pistil; b, Phlox suaveolens 1,s. of flower, dehiscent fruit with calyx, seed, I.s. 

of same; c, Cobaea pendulzjlora inferior part of corolla laid open to show insertion of stamens, 

ovary and disk and part of calyx, stigma and part of style, I.s. and C.S. of ovary, 1,s. of seed, C. 

trianaei flower (after Le Maout and Decaisne, 1873; Baillon, 1866-1895; Karsten; Hooker and 

Hooker, 1837-1982). FOUQUIERIACEAE: d, Fouguieria splendens bud, flower, I.s. of same, 

flowering twig, pistil, views of anther and base of filament; e, 1,s. and C.S. of ovary, F. spinosa 

dehiscent fruit, seed (after Johnson, Niedenzu). 
284 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 1 .?I~.-HYDROPHYLLACEAE: a, Phacelia californica flowering shoot, flower, pistil and 

part of calyx; b, corolla, ovary and hypogynous disk, C.S. and 1.s. of ovary, views of stamen; c, 

P. pinnatzfide part of corolla laid open, Hydrophyllum virginianum I.s. of ovary, dehiscent fruit; 

d, Hydrolea spinosa flower, corolla laid open, floral diagram, views of stamen; e, pistil, 1,s. and 

C.S. of ovary, fruit, I.s. and C.S. of same, seed, I.s. and C.S. of same showing embryo (after 

Johnson; Brand; Le Maout and Decaisne, 1873; Martius, 1840-1906). 

petioles becoming spines, exstipulate; inflorescences terminal racemes or panicles; flowers bisexual, the parts hypogynous; sepals 5, unequal, 

free, markedly imbricate; petals connate high, 

the corolla tubular, the lobes 5, imbricate in bud; 

stamens 10-15 (-23), in 1-2 series, the filaments 

filiform, free or shortly connate; anthers about 

twice as long as wide, dorsifixed; pollen S-colporate; base of the ovary glandular; pistil 1, the 

carpels 3, the style 1, filiform or short and stout, 

divided shortly or to the middle, stigma(s) 3, 

linear or not divided; ovary unilocular, the placentas 3, parietal, septiform, each with 4-6 bitegmic, tenuinucellar, anatropous ovules; fruit a 

3-valved capsule; seeds compressed, winged all 

around or long-hairy; embryo straight, as long as 

the endosperm; cotyledons thin, 2.8 times wider 

than the radicle, 0.8 the length of the embryo; 

endosperm scanty, fleshy. 

Composition: 1 genus, 11 species. 

Distribution: Arid southwestern United 

States and Mexico. 
NUMBER 58 285 

HYDROPHYLLACEAE (Figure 152a-e).-Herbs, 

rarely subshrubs; xylem vessel perforation plates 

simple; leaves alternate, sometimes radical, rarely 

opposite, entire, lobed, or compound, the hairs 

often rough, exstipulate; inflorescences axillary 

and terminal, cymose, sometimes cincinni, dichasia or umbellate, or the flowers rarely solitary; 

flowers bisexual, actinomorphic; sepals 5 (6-8, 

10- 12 in Codon), free or basally connate; petals 

5 (6-8, 10-12 in Codon), connate, the lobes 

imbricate, rarely contorted; stamens 5 (4, 6-8, 

10-1 2 in Codon), often inserted on the corolla 

towards its base, sometimes differing in length 

and inserted at different levels, alternipetalous, 

often with a scale-like appendage at the base of 

the filament; anthers dorsifixed; pollen 3-colporoidate, sometimes 3-pseudocolpate-3-colpate or 

6-9-rugate; disk hypogynous or absent; pistil 1, 

the carpels 2, (3 in Romanzofja), styles 2, or 1 

and bifid apically, the stigmas apical, capitate; 

ovary superior or semi-inferior (Nama spp.), unilocular, with 2 parietal placentas or 2(3)-locular 

with 2 (3) axile placentas, the ovules often 

numerous, sometimes only 2 per placenta 

(Hydrophyllum), unitegmic, tenuinucellar, f anatropous; fruit a capsule, loculicidal or rarely septicidal; seeds sometimes minute, sometimes carunculate; embryo straight, 0.2-1.0 as long as 

the endosperm; cotyledons 1-3 times as wide as 

the radicle, 0.2-0.4 the length of the embryo; 

endosperm copious or scanty, fleshy or hard. 

Composition: 18 genera, 250 species. 

Distribution: Centered in dry habitats of western United States; Alaska to temperate South 

America; northeastern Siberia; disjunct areas in 

tropical and southern Africa, Madagascar; India 

to Indochina, the western East Indies, and a small 

area in northern Australia. 

BORACINACEAE (Figure 153a-e).-Herbs, 

shrubs, and small trees, rarely lianas, often scabrid; xylem vessel perforation plates simple, 

rarely also a few reticulate (Cordia); leaves alternate or very rarely opposite, simple, usually entire, sometimes toothed, rarely lobed, exstipulate; inflorescences cymes, usually a coiled 

(scorpioid) cincinnus, rarely the flowers solitary; 

flowers bisexual, rarely the plant gynodioecious 

(species of Echium) or dioecious (Rochefortia), actinomorphic, rarely zygomorphic; sepals 5 (4-7), 

free or basally connate; petals 5 (4-7), connate, 

the lobes contorted or imbricate in bud, many 

with coronal scales covering the entrance to the 

corolla-tube; stamens 5 (4-7), rarely 1 fertile and 

the others staminodal, inserted on the corolla, 

rarely at 2 levels, rarely unequal, alternipetalous; 

anthers rarely coherent, basifixed or basally dorsifixed, often acute apically, sometimes apically 

produced; pollen often 3 (2- 1 2)-colporate, rarely 

3-colporoidate, 3-colporate-3-pseudocolpate, 2- 

colpate; disk annular, hypogynous, or absent; 

pistil 1, the carpels 2, style 1, rarely 2 (Pteleocarpu) or bilobed, sometimes twice bilobed (Cordia), sometimes heterostyled, terminal or usually 

gynobasic, the stigmas 1-4, apical, various; ovary 

superior, 2 or 4( 1O)-locular, the ovules usually 4, 

subbasal or axile, unitegmic, tenuinucellar, or 

pseudo-crassinucellar (Heliotropium), anatropous 

or semi-anatropous; fruit 4 (8- 10 in Trigonotis) 

nutlets, a drupe or berry, the seed rarely with an 

elaiosome; embryo straight or slightly curved, 

0.7-1 .O the length of the endosperm; cotyledons 

moderately thick or flat, rarely folded (Cordia), 

or 2-partite (Amsinckia), 1-(Ehretia) or 1.8-2.7 

times wider than the radicle, 0.4-0.9 the length 

of the embryo, the radicle sometimes partly invested; endosperm mostly 0, sometimes scanty or 

moderate, fleshy. 

Composition: 100 genera, 2000 species. 

Distribution: Cosmopolitan; Boraginoideae 

centered in western North America and the Mediterranean region; Ehretioideae and Cordioideae 

are tropical and subtropical. 

S CR OPH ULA R IA LES 

Herbs, shrubs, lianas, and trees, sometimes 

root parasites; xylem vessel perforation plates 

simple; intraxylary or interxylary phloem sometimes present; leaves opposite or alternate, simple, less commonly compound, entire or toothed, 

lobed, or dissected, sometimes reduced to scales, 

rarely with insectivorous bladders, rarely with a 
286 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 153.-BORAGINACEAE: a, Ehretia serrata flower, I.s. of same, C.S. of ovary, E. elliptica 

Is. of ovary, 1,s. and C.S. of fruit, seed, embryo; b, Coda calocephala corolla, same laid open, 

views of anther, fruit, C.S. of same, C. leucocalyx part of corolla laid open to show insertion of 

stamens; c, C. excelsa pistil, Is. and C.S. of ovary, embryo, same opened, C. trachyphylla flower; 

d, Cynoglossum amabile part of inflorescence, Antiphytum cruciatum flower, same opened, fruit 

and calyx; e, Heliotropium clausseni pistil and calyx, coherent anthers, fruit, corolla opened to 

show position of anthers, pistil (after Baillon, 1866-1895; Sargent; Martius, 1840-1906; Smith, 

1970). 

terminal tendril, exstipulate, very rarely stipu- spurred, gibbous, ventricose, saccate, personate, 

late; flowers bisexual, zygomorphic, less com- campanulate or funnelform; stamens 4-5 (2-8), 

monly actinomorphic; sepals 5-4 (2-6), usually epipetalous, equal or often didynamous, rarely 

connate, the calyx rarely truncate, bilabiate or connate in pairs; sometimes 1-3 staminodes presspathaceous; petals 5-4 (6-8), connate, the co- ent; anthers basifixed or dorsifixed, sometimes 

rolla regular or often bilabiate, sometimes connivent or connate in pairs or in a group of 4, 
NUMBER 58 287 

usually 2-locular, sometimes 1 locule reduced, 

sometimes the loculi divaricate; pollen 2- 13-colpate or -colporoidate, less often 6-rugate, 4-rupate, zone- or spiraperturate, polyforate, 2-4- 

porate, or nonaperturate; glandular disk usually 

surrounding the base of the ovary; pistil 1, the 

carpels 2 (3-5), style 1, usually elongate, the 

stigma(s) apical; ovary superior, rarely semi-inferior or inferior, 2- 1 (-5)-locular, the locules 

sometimes incomplete, ovules often numerous, 

sometimes few to 1 per locule, more commonly 

axile than parietal, rarely basal-axile, basal, or 

subapical, unitegmic, tenuinucellar; fruit a septicidal, loculicidal or infrequently circumscissile 

capsule or berry, uncommonly nutlets or a 

drupe, the seeds often numerous, minute, and 

sculptured, sometimes winged; embryo straight, 

arcuate or bent, often elongate, the cotyledons 

narrow or broad, minute and undifferentiated in 

Orobanchaceae; endosperm copious to absent. 

Distribution: Cosmopolitan, in various habitats. 

Chemistry: Solanaceae is known for its alkaloids. Aucubin (iridoid) occurs in Scrophulariaceae, Buddlejaceae, Globulariaceae, Bignoniaceae, Lentibulariaceae, and Pedaliaceae(?). 

Orobanchin is present in Buddlejaceae, Gesneriaceae, Orobanchaceae, and Verbenaceae but 

not Solanaceae or Lamiaceae. Catalpinoside and 

stachyose occur in Catalpa and Paulownia; the 

latter is also present in some Scrophulariaceae. 

Plantaginaceae has verbascase, present in at least 

2 species of Scrophulariaceae; it also has aucubin 

(iridoid) glycosides. Gibbs believes Buddlejaceae 

seems to fit well in the Tubiflorae. 

SOLANACEAE (Figure 154a-e).--Herbs, 

shrubs, small trees, and lianas; xylem vessel perforation plates simple, intraxylary phloem present; leaves alternate, rarely opposite (Sclerophylax), sometimes an opposite pair beneath the 

inflorescence, simple, entire, lobed, rarely 

toothed or dissected, exstipulate; inflorescences 

axillary cymes, dichasia, sometimes cincinni, or 

the flowers solitary; flowers bisexual, mostly actinomorphic, rarely zygomorphic; sepals 5 (4-6), 

connate; petals 5 (4-6), connate, plicate, contorted or rarely valvate, variously shaped; stamens 5 (2, 4, 6), inserted on the corolla, alternipetalous, usually equal and all fertile, rarely differing in length, rarely only 2 or 4 fertile and 

didynamous, then a staminode sometimes present; anthers basifixed or dorsifixed, often connivent, the loculi parallel, dehiscence longitudinal or poricidal; pollen 3-5 (2-6)-colpate, -colporoidate, -colporate or rugor(oid)ate or nonaperturate; disk surrounding the base of the ovary; 

pistil 1, the carpels 2 (3-5), style 1, the stigma 

apical, capitate or very shortly 2(-5)-lobed; ovary 

superior, the locules 2 (1-5), the ovules usually 

numerous (2-4 in Henoonia, Sclerophylax), axile, 

unitegmic, tenuinucellar, anatropous, semi-anatropous, or campylotropous; fruit a septicidal, 

rarely circumscissile capsule or berry, rarely a 

drupe (Grabowskia has 4 pyrenes); seeds usually 

numerous, rarely 1-2 (Henoonia, Sclerophylax, 

Lycium, Cestrum), minute, often sculptured; embryo arcuate, coiled or rarely straight, 0.8-3.0 

times the length of the endosperm; cotyledons 

1-3 times the width of the radicle, 0.2-0.6 the 

length of the embryo; endosperm copious, moderate, scanty or 0 (Goetzeinae), fleshy and semitransparent . 

Composition: 90 genera, -2300 species. 

Distribution: Cosmopolitan except the arctic 

and cool temperate regions of the Northern 

Hemisphere; mainly tropical and subtropical; 

centered in South America. 

NOLANACEAE (Figure 154f,g).-Herbs or small 

shrubs; xylem vessel perforation plates simple; 

intraxylary phloem present; leaves alternate, 

sometimes paired where branches meet, simple, 

entire, exstipulate; flowers usually solitary, axillary, sometimes in a raceme, bisexual, actinomorphic; sepals 5, connate; petals 5, connate, the 

corolla campanulate or funnelform, plicate in 

bud; stamens 5, equal or 3 long and 2 short, 

inserted low on the corolla, alternipetalous; anthers about as long as wide or oblong, the sacs 

slightly divergent basally; pollen 3-colporate; disk 

surrounding the base of the ovary, sometimes 

lobed; pistil 1, the carpels typically 5, style 1, the 

stigma capitate or obscurely 2-5-lobed; ovary 
288 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

W 0 

b 

C 

LJ 

FIGURE 1 ~~.-SOLANACEAE: a, Nicandra physaloides 1.s. of flower, floral diagram, "cotiana 

tabacum flower, A'. glauca I.s. of flower, 1,s. of lower part of flower, C.S. of ovary; b, views of 

anther, iV. tabacum stigma; c, seed, 1,s. of same, dehiscent capsule; d, Solanum tuberosum flower, 

1,s. of same, anther, C.S. of fruit, seed, 1,s. of same, Petunia axillaris part of corolla laid open 

showing different lengths of stamens; e, Brunfelsia obovata 1,s. of corolla showing pistil and 

stamens, dehiscent fruit, stamen, stigma (after Baillon, 1866-1 895; Johnson; von Wettstein, 

1935; Smith and Downs; Martius, 1840-1 906). NOLANACEAE: f; "Volana paradoxa flowering 

shoot, Alona coelestis 1,s. of flower; g, Nolana prostrata floral diagram, views of anther, pistil, 

fruit, drupelet, I.s. of same showing sections of 4 seeds, I.s. of seed, A'. rupestris base of style 

and ovary, fruit, A'. atriplicifolia pistil, Alona phylicifolia pistil (after Le Maout and Decaisne, 

1873; Baillon, 1866-1895; yon Wettstein, 1935). 
NUMBER 58 289 

superior, 5-locular, the ovules 4-1 per locule, 

basal-axile, unitegmic, tenuinucellar, semi-anatropous; fruit a schizocarp of 3 to w nutlets, the 

endocarp sometimes stony; embryo terete, arcuate or coiled, 1.6 times the length of the endosperm or longer; cotyledons as wide as the 

radicle, 0.6-0.7 the length of the embryo; endosperm copious or scanty. 

Composition: 2 genera, 80 species. 

Distribution: Mostly along the seashore; temperate and subtropical Chile and Peru. 

SCROPHULARIACEAE (Figure 1 55a-d).-Herbs 

and shrubs, rarely small trees or lianas or greenleaved root parasites; xylem vessel perforation 

plates simple; intraxylary phloem absent; leaves 

alternate, opposite or rarely verticillate, simple, 

usually toothed, sometimes entire or dissected, 

exstipulate; inflorescences axillary or terminal 

spikes, racemes, panicles, or cymes and then usually dichasia, or the flowers solitary; flowers bisexual, zygomorphic or rarely nearly actinomorphic; sepals 5 (4-8), usually shortly connate; 

petals 5 (4-8), very rarely 0 (Besseya spp.), connate, the corolla often bilabiate, sometimes 

spurred, gibbous or saccate, the lobes imbricate 

in bud; stamens inserted on the corolla, usually 

4, didynamous, often 2 (3-8), one staminode 

sometimes present; anthers sometimes connivent 

in pairs, the sacs often divergent basally or sometimes united or reduced to 1, dehiscence longitudinal, rarely poricidal; pollen 2-7-colpate, 

-colporoidate, or -colporate, occasionally spiraperturate or k irregular; disk at the base of or 

below the ovary, annular or to one side, sometimes lobed; pistil 1, the carpels 2 (3), style 1, the 

stigmas 2, flattened, or 1 and capitate, apical; 

ovary superior, bilocular, rarely unilocular (Rehmannia), the ovules numerous, rarely few or 1 

per locule (Selagineae), axile, unitegmic, tenuinucellar, anatropous, semi-anatropous or rarely 

campylotropous; fruit a capsule, usually septicidal, sometimes loculicidal or poricidal, very 

rarely a berry or nutlet (Selagineae, Tetrachondra); seeds usually minute, usually numerous, 

often sculptured; embryo straight or slightly arcuate, often terete, 0.4-1.0 the length of the 

endosperm; cotyledons 1-3 times the width of 

the radicle, sometimes thick, 0.1-0.6 the length 

of the embryo; endosperm copious or scanty, 

rarely 0, fleshy. 

Composition: 200 genera, 3000 species. 

Distribution: Cosmopolitan; centered in the 

North Temperate Zone; mainly in mountains in 

the tropics, poorly represented in rain forest. 

BUDDLEJACEAE (Figure 156a,b).-Trees and 

shrubs, very rarely herbs, indument when present glandular, stellate or lepidote; xylem vessel 

perforation plates simple; intraxylary phloem absent; leaves opposite or verticillate, very rarely 

alternate, entire or toothed, sometimes stipulate; 

inflorescences thyrses, racemes, spikes, heads or 

cymes; flowers bisexual, actinomorphic or 

slightly zygomorphic; sepals 4 (j), connate; petals 

4 (5), connate, imbricate; stamens 4 (5?), inserted 

on the corolla; anthers longer than wide, the sacs 

not, or slightly separate, parallel; pollen usually 

3 (4)-colporate (-colporoidate) or 4-ruporate; disk 

sometimes present; pistil 1, the carpels 2, style 1, 

the stigma enlarged, shortly bilobed; ovary superior, bilocular, sometimes incompletely so, 

rarely 4-locular, the ovules numerous, axile, unitegmic, tenuinucellar, semi-anatropous; fruit 

usually a septicidal capsule, rarely a drupe or 

berry, the seeds minute, often winged or produced at each end; embryo straight, 0.5 the 

length of the endosperm; cotyledons not broadened, 0.4 the length of the embryo; endosperm 

moderate, copious or scanty. 

Composition: 8 genera, 150 species. 

Distribution: Tropical to warm temperate; 

centered in eastern Asia; southwestern United 

States to warm temperate South America; Madagascar, tropical, southern, and eastern Africa to 

southern Arabia, India, eastern and southern 

Asia to the East Indies. 

GLOBULARIACEAE (Figure 156c).-Small 

shrubs or herbs; xylem vessel perforation plates 

simple; leaves alternate, often also in a rosette, 

simple, entire, exstipulate; inflorescences terminal, sometimes also axillary, usually involucrate, 

chaffy heads or dense spikes; flowers bisexual, 

zygomorphic; sepals 5, connate; petals 5, con- 
290 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

b 

C 

d 

FIGURE 155.-sCROPHULARIACEAE: a, Gerardia communis corolla laid open showing stamens 

and pistil, views of anther, sectons of fruit, I.s. of seed, Scrophularia leporella 3 views of corolla, 

pistil and hypogynous disk and part of calyx, staminode, upper part of style, stigma, C.S. of 

ovary, fruit, and calyx; b, Penstemon sp. pistil and calyx, pistil, C.S. of ovary, dehiscent fruit, 

Veronica buxqolia corolla and stamens, I.s. of corolla, face view of corolla, stamen, C.S. of ovary; 

c, Selugo corymbosa flowering shoot, floral diagram, S. stricta flower, I.s. of same, S. cinerea calyx, 

stamen, S. corymbosa 1,s. of seed; d, Hebenstretia dentata flower, I.s. of same, C.S. of fruit, 2 views 

of seed, 1,s. of same (after Martius, 1840-1906; Johnson; Baillon, 1866-1895; Le Maout and 

Decaisne, 1873). 
NUMBER 58 29 1 

FIGURE 156.-BUDDLEJACEAE: a, Buddleja anericana flowering twig, flower, 1,s. of same, 

dehiscent fruit, seed; b, B. colvilei corolla laid open, views of stamen, pistil and part of calyx, 

C.S. of ovary (after Rendle, 1938; Hooker and Hooker, 1837-1982). GLOBULARIACEAE: c, 

Globularia vulgaris flowering shoot, flower without calyx, floral diagram and Is. of flower, 

pistil, I.s. of ovary, corolla laid open showing insertion of stamens, calyx laid open, 1.s. of seed 

in 2 planes (after Le Maout and Decaisne, 1873; Baillon, 1866-1895). 

nate, the corolla bilabiate, the upper lip bilobed 

to completely reduced, the lower lip trilobed, the 

lobes imbricate in bud; stamens 4 (2), didynamous, epipetalous, the anthers dorsifixed, dehiscing by a single slit; pollen (2)3-colporate; disk 

hypogynous, annular, or an anterior gland, or 0; 

pistil 1, the carpels 2, style 1, elongated, the 

stigmas 2, acute, or 1 and capitate; ovary superior, unilocular at maturity, the ovule 1, subapical, unitegmic, tenuinucellar, anatropous; fruit a 

nutlet, the pericarp membranous; embryo 

straight, often terete, 0.8-0.9 the length of the 

endosperm, the cotyledons 0.5-0.7 the length of 

the embryo, 1-2 times as wide as the radicle; 

endosperm moderate to copious, fleshy. 

Composition: 2 genera, 30 species. 

Distribution: Centered in the Mediterranean 

region, extending into most of Europe and to 

the Caspian Sea; northeastern Africa. 

LENTIBULARIACEAE (Figure 157a,b).-Herbs 

of wet habitats; xylem usually much reduced; 

leaves alternate, often in basal rosettes, or re- 
292 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 58 293 

duced to scales, sometimes with bladders, insectivorous, the submersed sometimes finely dissected, exstipulate; inflorescence a scapose raceme or spike or the flower solitary; flowers 

bisexual, zygomorphic; sepals 5 or 4, free or 

shortly connate, in 1 or 2 series or united to form 

2; petals 5, connate, the corolla spurred, gibbous 

or personate, the lobes imbricate; stamens 2, 

inserted on the base of the corolla, sometimes 2 

staminodes present; anthers basifixed or dorsifixed, 1 -2-locular, the loculi divaricate; pollen 

apertures 3- 1 8-colporate or colporoidate, occasionally ruporate or k irregular; disk absent; 

pistil 1, the carpels 2, style 1, short, or the stigma 

sessile, 2- 1 -lobed; ovary superior, unilocular, the 

ovules numerous or rarely 2, basal, unitegmic, 

tenuinucellar, anatropous; fruit a capsule, dehiscing by 2 or 4 valves or circumscissile, rarely an 

achene, the seeds numerous, rarely 1, sculptured; 

embryo straight, the cotyledons as wide as the 

radicle(?), 0.2-0.4 the length of the embryo; 

endosperm 0. 

Composition: 4 genera, -200 species. 

Distribution: Cosmopolitan. 

OROBANCHACEAE (Figure 157c).-Chlorophylless herbs parasitic on roots; xylem vessel 

FIGURE 157.-LENTIBULARIACEAE: a, Utriculariajlaccida calyx, corolla, lf. tricolor stamens, U. Jaccida 1,s. of fruit, U. 

cornuta face of flower, a view of stamens and pistil, another 

view of pistil and 1 stamen, 2 views of pistil; b, stigma, I.s. 

and C.S. of ovary, habit showing bladders, Polypomopholyx 

laciniata calyx, Genlisea ornata calyx, Utricularia cucullata 

seed (after Johnson; Martius, 1840- 1906). OROBANCHACEAE: c, Orobanche ramosa flowering plant attached to host, 

flower, 1,s. of same, dehiscent fruit, seed, 0. eryngii floral 

diagram, stamen, 0. hederae 1,s. of seed (greatly enlarged) 

showing the minute embryo in copious endosperm (after 

Baillon, 1866-1899; Le Maout and Decaisne, 1873; Beck). 

ACANTHACEAE: d, Arrhostoxylon glabrum bud, flower, I.s. of 

corolla showing insertion of stamens, views of anther, pistil, 

and calyx; e, Justicia longepetiolata views of anther, stigma, 

fruit and calyx, fruit with calyx removed, valves of fruit with 

and tvithout seeds, seed (after Martius, 1840-1906). BIGNONIACEAE: J; Cybistax antisyphilitica flower, 1,s. of same, 

corolla laid open showing the stamens and staminode, floral 

diagram, stigma, 1,s. of ovary and disk, I.s. of seed showing 

embryo, part of dissepiment (above) and part of dehiscent 

fruit (after Martius, 1840-1906). 

perforation plates simple; leaves alternate, reduced to scales, exstipulate; inflorescence a terminal raceme or spike or the flower solitary; 

flowers bisexual, zygomorphic, some cleistogamous; sepals (2) 4-5, connate; petals 5, connate, 

the corolla bilabiate or nearly regular, the lobes 

imbricate in bud; stamens 4, didynamous, inserted on the corolla, a staminode present or 

absent; anthers often connivent in pairs; pollen 

apertures 3-colpoidate, 3-colp(oroid)ate, or k irregular; pistil 1, the carpels 2 (3), style 1, elongated, the stigma apical, capitate or 2-4-lobed; 

ovary superior, unilocular (basally bilocular in 

Christisonia), the ovules numerous, parietal, unitegmic, tenuinucellar, anatropous or semi-anatropous, sometimes the placentas meeting in the 

center of the ovary; fruit a loculicidal capsule 

dehiscing by 2 valves; seeds minute, numerous, 

sculptured; embryo minute, globular, undifferentiated, 0.1 the length of the endosperm; endosperm copious, fleshy. 

Composition: 13 genera, 150 species. 

Distribution: Centered in temperate Eurasia; 

few species in America, southern Africa, and 

Australia, few tropical. 

ACANTHACEAE (Figure 157d,e).-Herbs, 

lianas, and shrubs, rarely small trees; xylem vessel 

perforation plates simple; intraxylary or interxylary phloem sometimes present; leaves opposite, very rarely alternate (Elytraria), simple, usually entire, sometimes lobed and spiny, exstipulate; inflorescences axillary dichasia or monochasia, sometimes a terminal raceme or the flowers solitary; flowers bisexual, zygomorphic, 

nearly regular in Thunbergioideae; bracts often 

colored and conspicuous; sepals 4-5 (3), sometimes truncate (Thunbergia); petals 4-5, connate, 

the corolla bilabiate, rarely unilabiate, the lobes 

imbricate or contorted in bud; stamens 4, didynamous or equally long, sometimes 2, rarely 5 

(Pentstemonacanthus), inserted on the corolla, the 

filaments free or partially connate in pairs; 1-3 

staminodes frequently present; anthers 2- 1- 

locular, the loculi confluent or separate, basifixed 

or dorsifixed; pollen 3-9-colpate, 3-colpoidorate, 

2-4-porate, zone-or spiraperturate; disk below 
294 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

the ovary, sometimes annular or cupular; pistil 

1, the carpels 2, style 1, elongated, the stigmas 

2, one often smaller than the other; ovary superior, bilocular (1-locular in Elytraria and Mendoncia), the ovules 2 (1- 10) per locule, axile (parietal 

in Elytraria), unitegmic, the integument thick, 

tenuinucellar, anatropous, campylotropous or 

amphitropous; fruit a loculicidal capsule, rarely 

a drupe (Mendoncia), the seeds few, funicle usually indurate; embryo large, the cotyledons usually foliose and thin, sometimes moderately thick, 

usually slightly to markedly bent, 0.9 the length 

of the endosperm, 3.5-9.0 times wider than the 

radicle, 0.7-0.9 the length of the embryo, the 

radicle sometimes invested; endosperm usually 

0, sometimes moderate (Nelsonioideae) or 

scanty, fleshy. 

Composition: 250 genera, 2600 species. 

Distribution: Various habitats; mainly pantropical, centered in southeastern Asia, Africa, 

and tropical America; few temperate; United 

States to warm temperate South America. 

BICNONIACEAE (Figure 157f).-Trees, shrubs, 

and lianas, very rarely herbs; xylem vessel perforation plates typically simple, but with a few 

multi-perforate plates in some genera; interxylary phloem sometimes present; leaves opposite, rarely alternate or verticillate, mostly compound, sometimes simple, often toothed, sometimes entire, sometimes the terminal leaflet(s) 

converted to a branched tendril, exstipulate; inflorescences usually dichasia, sometimes tending 

to become cincinnal, sometimes a panicle, or the 

flowers solitary; flowers bisexual, somewhat zygomorphic, sometimes nearly actinomorphic; sepals 5, connate, the calyx 5-toothed, rarely truncate or bilabiate; petals 5 (sometimes 6-7 in 

Catophractes), connate, the corolla usually campanulate or funnelform, the lobes imbricate in 

bud, rarely valvate; stamens 4, didynamous, or 2 

(5-7), inserted on the corolla; staminodes 1 or 3; 

anthers usually connivent in pairs, the sacs divergent 180" (only 1 sac of each anther fertile in 

Millingtonia); pollen 3-colporoidate, nonaperturate, 2- 12-colpate, 4-rupate, 6-rugate or spiraperturate; disk usually present, hypogynous, annular 

or cupular; pistil 1, the carpels 2, style 1, the 

stigma apical, bilobed, ventrally papillose; ovary 

superior, bilocular, rarely 1- or 4-locular, the 

ovules numerous, axile or rarely parietal, unitegmic, tenuinucellar, anatropous or semianatropous; fruit an often elongated loculicidal or septicidal capsule, or a large berry with a hard 

surface; seeds numerous, rarely few (Argylia), 

often flattened and winged; cotyledons spread 

out, straight, large, thin, rarely thick (Argylia), 

3-4 times wider than the radicle, 0.8 the length 

of the embryo, the radicle invested; endosperm 

0. 

Composition: 120 genera, 700 species. 

Distribution: Mainly pantropical, few temperate America and Africa; centered in tropical 

South America. 

GESNERIACEAE (Figure 158a).-Herbs, 

shrubs, or rarely small trees or lianas, many epiphytes; xylem vessel perforation plates simple; 

leaves opposite, rarely alternate or whorled, 

sometimes radical, simple, entire or toothed, 

rarely pinnatisect, exstipulate; inflorescences 

usually cymose, sometimes racemose, or the flowers solitary; flowers bisexual, usually zygomorphic, rarely nearly actinomorphic (Ramonda); sepals 5 (4-6), free or basally connate; petals 5 (4- 

6), connate, the corolla often bilabiate, often 

ventricose, rarely spurred, the lobes imbricate in 

bud; stamens 4, didynamous, rarely equal, or 2, 

rarely 5 (Ramonda, Sinningia spp.), inserted on 

the corolla, often near the base; sometimes 1 or 

3 staminodes present; anthers connate or connivent in pairs or in a group of 4, rarely free, 

FIGURE 1 ~~.-GESNERIACEAE: a, Sinningia pendulina corolla, same laid open showing the stamens and staminode, 

pistil with hypogynous glands and calyx, fruit and calyx, C.S. 

of fruit, views of seed, embryo, Codonanthe devosiana fruit 

(after Martius, 1840- 1906). PEDALIACEAE: b, Martynia lutea 

flower, 1,s. of same, floral diagram, fruit, C.S. of same, seed, 

I.s. of same; c, Sesamum orientale flowering shoot, pistil, C.S. 

of ovary, dehiscent fruit, seed, C.S. of same, embryo (after 

Baillon, 1866-1893; Le Maout and Decaisne, 1873). MYOPORACEAE: d, Myoporum acuminatum flower, Is. of same, M. 

parvijlorum floral diagram; e, flowering twig, corolla laid 

open, C.S. of ovary, I.s. and C.S. of fruit, seed, 1.s. of same 

(after Baillon, 1866-1 895; Le Maout and Decaisne, 1873). 
NUMBER 58 295 

e 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

bilocular; pollen 3 (2-4)-colporoidate or colporate, occasionally 4-ruporoidate or 6-rugoroidate; 

disk annular or cupular or of separate glands or 

scales, or to one side of the ovary, rarely absent; 

pistil 1, the carpels 2, style 1, elongated, stigma 

apical, simple and capitate or bilobed; ovary superior, semi-inferior or inferior, unilocular, 

rarely bilocular or incompetely 2- or 4-locular, 

rarely only 1 carpel fertile (Hemiboea), the ovules 

numerous, usually parietal, rarely axile, unitegmic, tenuinucellar, anatropous; fruit a 2- or 4- 

valved, usually loculicidal, rarely septicidal (Ramonda) or circumscissile, often elongated capsule, or a hard or soft berry; seeds numerous, 

minute, sculptured; embryo straight, 0.8-0.9 the 

length of the endosperm; cotyledons as broad as 

the radicle, 0.3-0.5 the length of the embryo; 

endosperm copious or moderate, rarely 0. 

Composition: 125 genera, 1,800 species. 

Distribution: Mostly pantropical in cloud forest, few in temperate South America, Africa, 

Australia, New Zealand, and Eurasia. 

PEDALIACEAE (Figure 158b,c).-Herbs, rarely 

shrubs or small trees; xylem vessel perforation 

plates simple; leaves opposite or alternate, simple, usually irregularly lobed and toothed, sometimes entire, exstipulate; inflorescences axillary 

cymes or solitary flowers, or terminal racemes; 

flowers bisexual, zygomorphic; sepals 5, connate 

or sometimes nearly free, sometimes spathaceous 

or bilabiate; petals 5, connate, the 5 lobes imbricate in bud, the corolla sometimes ventricose or 

spurred; stamens 4 and didynamous, or 2 (Trapella), inserted on the corolla; staminodes 1-2; 

anthers often connivent, the sacs parallel and 

dorsifixed, or divaricate; pollen 5- 13-colpate, 

-colporoidate, polyforate, or non-aperturate; 

disk annular, hypogynous; pistil 1, the carpels 2, 

style 1, elongated, the stigmas apical, 2 or 1 and 

punctiform; ovary superior or rarely inferior 

(Trapella), 1-2- or 4-locular, the ovules 1 to 

numerous per locule, axile or parietal, unitegmic, 

tenuinucellar, anatropous; fruit a loculicidal capsule, nut or subdrupe, often horned, hooked, or 

winged; seeds 1 -many, sometimes sculptured; 

embryo straight, 0.9-1.0 the length of the endosperm; cotyledons moderately thick, 2-2.8 

times wider than the radicle, 0.6-0.8 the length 

of the embryo; endosperm scanty or 0. 

Composition: 15 genera, 60 species. 

Distribution: Tropical and subtropical, few in 

temperate regions; mostly in dry and coastal habitats; Pedalioideae in most of Africa, part of 

India, Indochina, the East Indies, and northern 

Australia; Martynioideae in tropical and subtropical America. 

LAMIALES 

Shrubs, herbs, trees, and lianas, often with 

marked characteristic aromatic fragrances; xylem vessel perforation plates simple, very rarely 

also a few scalariform with few bars; intraxylary 

phloem very rarely present (Oftia and Avicennia); 

leaves opposite, rarely alternate, simple, rarely 

compound, usually toothed, sometimes entire or 

dissected, often gland-dotted, exstipulate; flowers bisexual, rarely unisexual, mostly zygomorphic; sepals 5-4 (-8), connate, the calyx 

subactinomorphic or bilabiate; petals 5-4 (-1 6), 

connate, the corolla mostly bilabiate, sometimes 

actinomorphic; stamens 4, 2 (-16), rarely 1-3 

staminodal, inserted on the corolla, usually didynamous, the filaments very rarely basally connate (Coleus); anthers dorsifixed, the sacs often 

divaricate, sometimes 1 sac of the anther 

aborted; pollen 3 (2-5)-colpate, 3-colporate, 

rarely 3 (4)-colporoidate or 6-rug(or)ate; pistil 1, 

the carpels 2 (3-5 in Geunsia, Duranta), style 1, 

terminal or gynobasic, the stigmas apical; ovary 

superior, 1-2 (3-5)-locular, the ovules 2 (-8) per 

carpel, usually basal, rarely axile, very rarely the 

placenta free-central (Avicennia), unitegmic, tenuinucellar; fruit nutlets, drupe, very rarely a 

capsule; embryo usually straight, nearly as long 

as the endosperm; endosperm absent or scanty, 

very rarely moderate. 

Distribution: Cosmopolitan, in various habitats. 

Chemistry: Verbenaceae and Lamiaceae are 

chemically very similar. They are rich in monoterpenoids and share a number of terpenoids. 

Furans occur most often in Lamiaceae and My- 
NUMBER 58 297 

oporaceae, but they are present in several unrelated families too. 

MYOPORACEAE (Figure 158d,e).-Shrubs or 

small trees; xylem vessel perforation plates simple; secretory cavities present in the leaf and 

stem; intraxylary phloem present in Oftia; leaves 

alternate or rarely opposite, simple, entire or 

toothed, often gland-dotted, exstipulate; inflorescences axillary cymes, or the flowers solitary; 

flowers bisexual, usually zygomorphic sometimes 

actinomorphic; sepals 5, connate; petals 5, connate, the corolla subactinomorphic, 5-lobed, to 

bilabiate, the lobes imbricate in bud; stamens 4, 

didynamous, rarely 5, inserted on the corolla; 

sometimes 1-3 staminodes present; anther sacs 

divergent basally; pollen 3-colpate, 3-colporate, 

3 (4)-colporoidate, rarely 6-rugorate; pistil 1, the 

carpels 2, style 1, elongated, stigma apical, obscurely bilobed; ovary superior, the locules 2 (3- 

lo), ovules 1-8 per locule, axile-apical, unitegmic, tenuinucellar, anatropous; fruit a drupe or 

1-8 nutlets; embryo straight or slightly curved, 

subterete, 0.9 the length of the endosperm; cotyledons plano-convex, as wide as the radicle, 0.4 

the length of the embryo; endosperm scanty or 

0, fleshy. 

Composition: 4 genera, 100 species. 

Distribution: Centered in Australia; all Australia, New Zealand, New Guinea; few in eastern 

China, Japan, and the West Indies. 

VERBENACEAE (Figure 1 59a-d).-Shrubs, 

trees, herbs, and lianas; xylem vessel perforation 

plates simple except for rare reticulate plates and 

a few scalariform plates in some species of Vitex; 

intraxylary phloem present in Avicennia ; leaves 

opposite or whorled, rarely alternate (some 

Chloanthoideae and Ammonia), usually simple, 

rarely compound, toothed, entire or dissected, 

exstipulate; inflorescences terminal, rarely axillary spikes, heads, corymbose-paniculate, dichasial cymes with a cincinnal tendency, or the flowers solitary; flowers bisexual, or the plants polygamous, zygomorphic or actinomorphic; sepals 

4-5 (4), connate; petals 4-5 (-16), connate, the 

lobes imbricate in bud; stamens 4 (2-16), didynamous, inserted on the corolla, rarely 1 or 3 

staminodes present; anthers dorsifixed, the sacs 

often divaricate; pollen 3 (2-5)-colpate, 3-colporate, rarely 6-rugate or 3-colporoidate; disk annular or absent; pistil 1, the carpels 2 (3-5 in 

Geunsia, Duranta), style 1, terminal, the stigma 

usually bilobed, sometimes punctiform (Stilbe) or 

3-5-lobed (Geunsia, Duranta); ovary superior, 

the locules 2 (only 1 fertile in Stilbe) (4-5), sometimes incomplete, the ovules unitegmic, tenuinucellar, anatropous or suborthotropous (Vitex, 

Avicennia), 1-2 per locule, usually basal, rarely 

axile, sometimes the placenta free-central (Avicennia); fruit a drupe, nutlet(s), rarely a capsule 

(Campylostachys, Caryopteris, Nyctanthes, Dimetra, 

Avicennia); embryo straight, 0.9 to as long as the 

endosperm if present; cotyledons moderately 

thick, rarely conferruminate or plicate, 1.3-3.0 

times wider than the radicle, 0.5-0.8 the length 

of the embryo, the radicle rarely invested; endosperm 0 or scanty, rarely moderate (Avicennia, 

Stilbe, Chloanthes), fleshy. 

Composition: 75 genera, 2600 species. 

Distribution: Pantropical and subtropical, few 

in warm temperate zones, lacking in arctic and 

cool temperate zones. 

PHRYMACEAE (Figure 1,59e).-Herbs; xylem 

vessel perforation plates simple; leaves opposite, 

simple, toothed, exstipulate; inflorescences terminal and axillary spikes; flowers bisexual, zygomorphic; sepals 5, connate, the calyx bilabiate; 

petals 5, connate, the corolla bilabiate; stamens 

4, didynamous, inserted on the corolla; anthers 

about as long as wide, the sacs not divergent; 

pollen 3-colpate; pistil 1, the carpels 2, style 1, 

terminal, the stigma very shortly bifid; ovary 

superior, unilocular, the ovule 1, basal, unitegmic, tenuinucellar, suborthotropous; fruit a 

nutlet, the pericarp membranous; embryo convolute or longitudinally folded, the cotyledons 

flattened, broad, 0.8 the length of the embryo; 

endosperm 0. 

Composition: 1 genus, 2 species. 

Distribution: Temperate eastern Asia and 

eastern North America. 

LAMIACEAE (Figure 159f-j).-Herbs, rarely 

shrubs, very few small trees (Hyptis), a few climb- 
298 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

0 

4 

d 

r 
NUMBER 58 299 

ers (Scutellaria), usually with marked characteristic aromatic fragrances; vessel perforation 

plates simple, except some scalariform plates with 

few bars at right angles to their usual direction 

in Cymaria elongata; leaves opposite, rarely 

whorled, very rarely alternate (Zcomum), often 

glandular-punctate, simple, usually toothed, 

rarely pinnately dissected (Cedronella), sometimes entire, exstipulate; infloresences terminal 

or axillary, usually a dichasial cyme, becoming 

cincinnal and glomerate in its later branchings, 

rarely a raceme, spike or head, rarely the flowers solitary; flowers bisexual, rarely the plants 

gynodioecious, zygomorphic, rarely subactinomorphic; sepals 5 (4 in Preslia), connate, the calyx 

subactinomorphic or bilabiate; petals 5 (4), connate, the corolla often bilabiate, rarely subactinomorphic; stamens 4, often didynamous, or 2, 

inserted on the corolla, the filaments separate 

(basally connate in Coleus), sometimes 1-2 staminodes present; anther sacs usually divaricate, 

sometimes only 1 sac of each anther fertile; pollen usually 3 (4)- or 6-colpate, rarely 3-colporoidate; disk mostly 4- or 2-lobed, often to one 

side of the ovary; pistil 1, the carpels 2, style 1, 

usually gynobasic, sometimes terminal, the 

stigma usually bifid and the branches linear, 

rarely shortly 4-fid (Cleonia); ovary superior, the 

locules 2, eventually 4-lobed, the ovules 2 per 

FIGURE 1 .~~.-VERBENACEAE: a, Verbena hastata flower, 1 .s. 

of barlie, stamen, pistil, developing fruit invested by calyx 

and bract and C.S. of same (below); b, Amasonia lasiocaulos 

corolla in bud, flower !\.ithout the calyx, corolla laid open, 

pistil, views of anther; c, calyx and fruit, fruit, C.S. of same, 

2 view5 of ii ntitlet, C.S. of same, seed, embryo; d, Avicennia 

nitida floral diagram, flower laid open, 1 .s. of pistil, placental 

column and ovules, 1 .s. and C.S. of fruit, embryo laid open 

(after Johnson; Martius, 1840- 1906; Sargent). PHRYMACEAK e, Phryma leptostachya flower, 1,s. of same, calyx (reflexed in fruit), 1.s. of calyx and fruit showing the solitary 

seed, C.S. of fruit showing folding of the cotyledons (below) 

(after Johnson, Briquet). LAMIACEAE:~; Hyptis vzrgata flower, 

corolla laid open; g, ovary and disk, 2 views of nutlet, 1 .s. of 

same in 2 planes, C.S. of same: h, calyx laid open showing 

ovary. pistil, views of anther; i, C.S. and 1 .s. of fruit showing 

gymbasic style;j, Salvia hilarii bud, flower, corolla laid open 

shouing stamens and staminodes, pistil, stamen (after Martius, 1840-1 906). 

carpel, basal-axile, unitegmic, tenuinucellar, 

anatropous or semianatropous; fruit 4 (-1) 

nutlets, rarely a drupelet (Stenogyne, Gomphostemma, Prasium); embryo usually straight, sometimes bent (Scutellaria, Catopheria), 0.9 to as long 

as the endosperm if present; cotyledons moderately thick, or thin, 2.5-5.5 times wider than the 

radicle, 0.5-0.7 the length of the embryo, the 

radicle usually invested, rarely accumbent or incumbent on the cotyledons; endosperm usually 

0, sometimes scanty (Prosthanthera), fleshy. 

Composition: 180 genera, 3000 species. 

Distribution: Cosmopolitan; centered in the 

Mediterranean region to central Asia, less frequent in the Southern Hemisphere; various habitats, mostly open areas, few in rain forest. 

C A MPA N ULA L ES 

Herbs, rarely shrubs, small trees, or climbers; 

xylem vessel perforation plates simple, rarely also 

a few scalariform with few bars: laticiferous canals sometimes present; medullary bundles or 

intraxylary phloem sometimes present: leaves alternate, rarely opposite or radical, simple, 

toothed or entire, very rarely lobed, exstipulate; 

flowers bisexual, very rarely unisexual, actinomorphic or zygomorphic; sepals 5 (3-lO, O), basally connate, rarely free: petals 5 (3- lo), connate, 

very rarely free, the lobes valvate in bud, rarely 

imbricate, the corolla campanulate, bilabiate or 

tubular, a paracorolla rarely present; stamens 5 

(2-lo), free or connate, inserted on the corolla 

or on a disk, rarely adnate to the style, the 

anthers basifixed, free or connate, very rarely 

didymous; pollen 3- 1 O-colpate, 3-6 (2)-colporate, 3 (2)- or 6-7-colporoidate, 4-8-poroid, occasionally 4-rupate, sometimes spinulose; glandular disk usually present; pistil 1, the carpels 2- 

5 (-lo), style 1, sometimes 2-3-fid, elongate, 

sometimes indusiate, the stigmas apical, sometimes slightly elongate; ovary inferior, semi-inferior, or rarely superior, the locules 2-5 (1-lo), 

ovules numerous to 1, axile, basal or apical, 

rarely parietal or the placenta free-central, unitegmic, tenuinucellar: fruit a capsule dehiscent 

by pores, lateral slits, apically, loculicidally, sep- 
300 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

ticidally or circumscissile, rarely a berry, drupe, 

nutlet, or achene, the seeds often minute; embryo minute to nearly as long as the endosperm, 

straight, the cotyledons usually 1-2 times the 

width of the radicle and up to half the length of 

the embryo; enhsperm usually copious or moderate, rarely absent. 

Distribution: Cosmopolitan, in various habitats. 

Chemistry: Gibbs says Campanulaceae and 

Goodeniaceae hang together, but the Asteraceae 

seem to differ rather markedly from the other 

members of the order, perhaps enough to warrant separation from it. The chemistry of Campanulales and Cucurbitaceae fits quite well together, but the cucurbitaceous amino-acids, seedfats and abscence(?) of inulin, would all mark off 

the Cucurbitaceae from the Campanulales. 

CAMPANULACEAE (Figure 1 6Oa,b).-Mostly 

herbs, a few shrubs and small trees (Clermontia 

spp.), rarely climbers (Cyphia); xylem vessel perforation plates simple, but a few scalariform bars 

in Platycodon, Lightfootia, Roella, and Lobelia; 

laticiferous canals present; medullary bundles 

present in a number of genera; sometimes intraxylary phloem present; glandular hairs absent; 

leaves simple, alternate, rarely opposite or 

whorled, toothed (entire in Sphenoclea), exstipulate; inflorescences terminal or axillary, rarely 

epiphyllous (Ruthiella), racemes, spikes or heads 

or cymose, sometimes thyrsiform or the flowers 

solitary; flowers usually bisexual, the plants very 

rarely dioecious, actinomorphic or zygomorphic; 

sepals 5 (3- lo), basally connate, very rarely free, 

usually adnate to the ovary; petals 5 (3-10, 0 

when cleistogamous), connate, rarely free (Dialypetalum and 2 other genera), valvate, the corolla 

campanulate, tubular or bilabiate; stamens 5 (3- 

lo), inserted basally on the corolla or on the disk, 

the filaments free or connate; anthers free or 

connate; pollen 4- 1 0-colpate, (2) 3-6-colporate, 

(2) 3 or 6-7-colporoidate, sometimes spinulose; 

disk epigynous, sometimes cupular or tubular; 

pistil 1, the carpels 2-5 (-lo), style 1, elongated, 

the stigmas 2-5 (-lo), apical and sometimes 

slightly elongated; ovary inferior, semi-inferior, 

or rarely superior (Cyananthus), the locules 2-5 

(1 -lo), the ovules mostly numerous (1 per locule 

in Mezleria), axile, basal or apical, rarely parietal, 

unitegmic, tenuinucellar, anatropous; fruit a capsule dehiscent by pores or by lateral slits, or 

apically loculicidal, or circumscissile (Lysipomia, 

Siphocodon, Sphenoclea), rarely a berry; seeds minute, the embryo straight, 0.1-0.7 the length of 

the endosperm; cotyledons 1-2 times the width 

of the radicle, 0.06-0.5 the length of the embryo; 

endosperm copious or moderate, fleshy. 

Composition: 65 genera, 2000 species. 

Distribution: Cosmopolitan; Campanuloideae 

centered in north temperate zone, only one genus being confined to the tropics; few in the 

Southern Hemisphere but several in South Africa; Lobelioideae centered in America, many 

tropical, often at higher elevations; southern 

Canada to the southern tip of South America; 

southern 2h of Africa, Madagascar; Japan, northeastern Siberia to India, Indochina, the East Indies, Australia, New Zealand; western Europe 

and the Mediterranean region. 

GOODENIACEAE (Figure 160c,d).-Herbs and 

small shrubs; xylem vessel perforation plates simple; laticiferous canals absent; medullary bundles 

present in 3 genera; leaves alternate or rarely 

opposite, sometimes rosulate, simple, toothed or 

entire, exstipulate; inflorescences cymes, racemes, spikes, panicles or heads or the flowers 

solitary in the leaf axils; flowers bisexual, zygomorphic to subactinornorphic; sepals 5 (3, 0), 

connate or free; petals 5, connate, the corolla 

bilabiate or rarely unilabiate, the lobes valvate in 

bud; stamens 5, free or rarely shortly adnate to 

the corolla; anthers basifixed, usually free, sometimes connivent or connate around the style; 

pollen usually 3-colporate, 4-8-poroid, occasionally 4-rupate, sometimes minutely spinulose; pistil 1, the carpels 2, style 1, or 2-3-fid, elongated, 

indusiate apically, the stigma bilobed; ovary usually inferior, sometimes semi-inferior, 1-2 (4)- 

locular, the ovules 1 or more per locule, mostly 

basal, sometimes axile, unitegmic, tenuinucellar, 

anatropous or campylotropous; fruit a capsule, 

drupe or nut, the seeds flat, the embryo usually 
NUMBER 58 30 I 

C 

FIGURE 160.-CAMPANULACEAE: a, Wahlenbergia brasiliensis flower, 1 .s. of same, C.S. of ovary, 

1,s. of fruit, seed, U’. linarioides C.S. of ovary, views of stamen; b, Centropogon surinamensis 

flower, 1,s. of lower part of same, C.S. of ovary, upper part of stamina1 tube showing united 

anthers and stigma, fruit, seed, 1 .s. and C.S. of same (after Martius, 1840-1 906). GOODENIACEAE: 

c, Scaevola plumier2 bud, flower, corolla laid open, stamens showing anthers in dorsal view, 2 

stamens showing anthers in ventral view, stigma surrounded by indusium; d, style, 1 .s. of ovary, 

1 .s. of fruit, pyrene, embryo (after Martius, 1840-1906). 
302 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

straight, 0.8 the length of the endosperm; cotyledons thin, twice as wide as the radicle, 0.4-0.5 

the length of the embryo; endosperm usually 

copious, sometimes moderate, fleshy. 

Composition: 14 genera, 300 species. 

Distribution: Centered in Australia, especially 

the southwest; shores of southeastern Asia, eastern, southern and western Africa, Madagascar, 

eastern tropical America and temperate western 

South America. 

BRUNONIACEAE (Figure 16 1 a,b).-Herb; xylem vessel perforation plates simple; leaves simple, rosulate, entire, exstipulate; inflorescences 

scapose cymose heads, bracteate; flowers bisexual, subactinomorphic; sepals 5, connate; petals 

5, connate, the lobes valvate in bud; stamens 5, 

the filaments not connate, inserted on the corolla 

near the base, the anthers basifixed, connate 

around the style; pollen 3-colporate; pistil 1, the 

carpels 2, style 1, indusiate apically, the stigma 

small, not divided; ovary superior, unilocular, 

ovule 1, basal, unitegmic, tenuinucellar, anatropous; fruit an achene or nutlet; embryo straight, 

the cotyledons grading gradually into the radicle; 

cotyledons 4 times wider than the radicle, 0.5- 

0.8 the length of the embryo; endosperm 0. 

Composition: 1 genus, 1 species. 

Distribution: Eastern and western Australia, 

with a narrow southern connecting strip. 

CALYCERACEAE (Figure 16 1 c,d).-Herbs, 

rarely suffrutescent; xylem vessel perforation 

plates simple; secretory cavities and external 

glands absent; phloem strands absent from cortex; leaves alternate, sometimes rosulate, simple, 

entire or pinnately lobed, exstipulate; inflorescence an involucrate head; flowers bisexual, 

rarely the central flowers of a head 8 by abortion, 

rarely cleistogamous, actinomorphic or zygomorphic; sepals 5 (4-6), the calyx-tube adnate to the 

ovary; petals 5 (4-6), connate, the lobes valvate 

in bud; stamens 5 (4-6), the filaments free or 

connate, inserted high on the corolla; anthers 

free or connate around the style; pollen 3-colporate; disk absent; pistil 1, the carpels 2, style 

1, elongated, not indusiate, the stigma apical, 

capitate; ovary inferior, unilocular, the ovule 1, 

apical, unitegmic, tenuinucellar, anatropous; 

fruit an achene or nutlet; embryo straight, 0.75- 

0.9 the length of the endosperm; cotyledons as 

wide as the radicle, 0.25-0.3 the length of the 

embryo; endosperm copious or scanty, fleshy. 

Composition: 4 genera, 50 species. 

Distribution: Southern half of South America, most commonly in the Andes, mainly in dry 

habitats of scrub and steppe vegetation. 

STYLIDIACEAE (Figure 16 1 e,f).-Herbs or 

rarely undershrubs; the sap not milky; xylem 

vessel perforation plates simple; leaves simple, 

entire, alternate, sometimes rosulate or in whorls 

on the stem, exstipulate; inflorescence terminal 

raceme, spike, corymb, or cymes, sometimes scapose; flowers bisexual or unisexual and then the 

plants usually monoecious, mostly zygomorphic; 

sepals 5 (4-9), the calyx-tube adnate to the ovary, 

or the sepals free or connate; petals 5 (4-9), 

often with a paracorolla, connate, the lobes imbricate in bud, the lowest one often smaller or 

larger than the others; stamens 2, the filaments 

connate and adnate to the style, but not to the 

corolla; anther sacs sometimes separate; pollen 

3-6-colpate, spinuliferous; disk present or absent; pistil 1, the carpels 2, style 1, divided apically, not indusiate; ovary inferior, bilocular or 

unilocular, the ovules numerous per locule, axile 

or free-central, unitegmic, tenuinucellar, anatropous; fruit a septicidal capsule or rarely indehiscent; seeds small, the embryo minute, barely 

differentiated, 0.06 the length of the endosperm; 

endosperm copious, fleshy and oily. 

FIGURE 16 1 .-BRUNONIACEAE: a, Brunonia australis plant in 

floiier, flower, 1 .s. of same, same laid open, stamen, stigma 

and indusium: b, fruit, embryo, 1 .s. of same (after Baillon, 

1866-1 89.5; Le Maout and Decaisne, 1873). CALYCERACEAE: 

c, Boopis bupleuroides inflorescence, 1 .s. of bud, flower, 

Acicarpha procumbens flower, anthers and apex of filament 

tube: d, Boopis bupleuroides views of fruit (after Martius, 

1840- 1906). STYLIDIACEAE: e, Stylidiurn adnatun flowering 

shoot, 1,s. of flower, floirer, corolla laid open, 2 views of 

androecium and stigma, dehiscent fruit, 1 .s. of seed showing 

the minute embryo;J S. graminfolium flower, 1,s. of same, 

Phyllacne magellanica plant in flower, 1 .s. of flower, Levenhookia pusilla flower, 1 .s. of same (after Baillon, 1866-1895; 

Le Maout and Decaisne, 1873). 
NUMBER 58 303 
3 04 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Composition: 5 genera, 140 species. 

Distribution: Centered in Australia; few species in New Zealand, eastern India, southern 

parts of the East Indies, and southern tip of South 

America. 

ASTERA LES 

The order is monotypic. 

ASTERACEAE (Figure 162a-f).-Mostly herbs, 

sometimes shrubs, rarely trees or climbers, often 

with a taproot; xylem vessel perforation plates 

simple, rarely also a few scalariform or reticulate 

plates; medullary and cortical bundles sometimes 

present; intraxylary phloem present in 3 genera; 

glandular hairs sometimes present; laticiferous 

vessels present in 2 tribes; oil tubes often present; 

leaves more often alternate than opposite, rarely 

whorled, sometimes rosulate, simple, toothed to 

deeply dissected, more rarely entire, exstipulate; 

inflorescences terminal racemose heads, sometimes in short dense spikes, involucrate, very 

rarely individual flowers surrounded by bracts; 

flowers actinomorphic or zygomorphic, bisexual 

or the plants monoecious, rarely dioecious; sepals 

absent or represented by a pappus of scales, 

bristles or hairs; petals 5 (4), very rarely the 9 

flower apetalous (Ambrosia), connate, the lobes 

valvate in bud, when zygomorphic usually ligulate, rarely bilabiate; stamens 5 (4), inserted on 

the corolla, the filaments not connate or very 

rarely connate (Ambrosia); anthers basifixed or 

rarely attached above the base (Arctotis, Gazania), 

sometimes tailed or produced apically, connate 

into a tube surrounding the style, very rarely 

free; pollen usually 3-colporate, sometimes to 12 

apertures, colpoidorate apertures form a transition to pores, sometimes 3-4 (6)-colpate, occasionally rugate, usually spinulose; disk annular, 

surrounding the base of the style, rarely absent; 

pistil 1, the carpels 2, style 1, elongated, bifid 

apically, very rarely 3-5-fid in some flowers of a 

specimen, stigmas 2 (I), often linear, rarely quite 

hairy (Artemisia), indusium absent; ovary inferior, unilocular, ovule 1, basal, unitegmic, tenuinucellar, anatropous, the embryo sac often with 

increased number of antipodal nuclei; fruit an 

achene (cypsela), very rarely a drupe (Chrysanthemoides); embryo straight, the cotyledons usually 

moderately thick and broad, plano-convex, 

rarely semicylindrical or convolute, 4 times wider 

than and grading gradually into the radicle, 0.3- 

0.8 the length of the embryo; endosperm 0. 

Composition: 

Distribution: Cosmopolitan; best represented 

in temperate and subtropical regions, in almost 

all habitats, but especially dry ones; uncommon 

in wetlands and tropical rain forest; Eupatorieae, 

Vernonieae, Heliantheae, and Liabeae are centered in the subtropics of America, the second 

also in Africa and Asia; the Cardueae and Lactuceae are centered in the Mediterranean region 

and Near East, the Anthemideae in those regions 

and South Africa; the Calenduleae and Arctoteae 

are centered in South Africa; the Mutiseae are 

centered in temperate regions of South America, 

especially the Andes and Guayana Highlands, 

and in Africa. 

-900 genera, - 14,000 species. 

INCERTAE SEDIS 

I cannot suggest a position for the following 

families. 

TREMANDRACEAE (Figure 163a-c).-Xeromorphic shrublets, rarely twining; xylem vessel 

perforation plates simple, perhaps sometimes 

also a few scalariform plates present; leaves alternate, opposite, or verticillate, rarely absent, 

simple, entire or toothed, often glandular, 

sometimes stellate tomentose, exstipulate; flowers solitary, axillary, long pedicillate, bisexFIGURE 162.-ASTERACEAE: a, C'ernonia macrophylla inflorescence, 1,s. of same, flower, 1,s. of same and corolla laid 

open, Erigeron maximus 1,s. of flower, fruit with pappus, 

stigmas and 1 .s. of fruit; b, Senecio erisithalifolius disk flower, 

1,s. of same: c, anther tube laid open, stigmas, base of ray 

flower; d, Mikania scandens inflorescence, flower, Trichocline 

macrocephala ray flower, disk flower, C.S. of fruit; e, anther 

tube laid open, stigmas;f; Aspilia setosa 1,s. of inflorescence, 

A. foliosa disk flower, 1 .s. of same, 1 .s. of base of ray flower 

(sterile), Agrianthus empetrijolius flower, 1 .s. of same, views 

of anther (after Martius, 1840- 1906). 
NUMBER 58 305 
306 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

a 

FIGURE 163.-TREMANDRACEAE: a, Tetratheca hirsuta essential organs, flowering twig, pistil 

with one of the ovary locules laid open; b, T. verticzllata flowering twig, floral diagram, T. 

procumbens flower, 1 .s. of pistil, stamen, 1 .s. of seed showing the minute embryo; c, T. verticillata 

flower, same with corolla removed, 1,s. of same, pistil (after Lindley, 1853; Le Maout and 

Decaisne, 1873). ELAEAGNACEAE: d, Elaeugnus angust~oliu part of flowering twig, 1 .s. of flower, 

1 .s. of ovary and part of perianth, floral diagram, d flower; e, 1 .s. and C.S. of fruit, stone, 1 .s. 

of seed, embryo, radicle and pluinule (after Le Maout and Decaisne, 1873). 

ual, actinomorphic, the parts hypogynous; 

sepals 4-5 (3), free, rarely basally connate, valvate in bud; petals 4-5 (3), free, induplicatevalvate; stamens 8, 10 (6), free, in 1 or 2 series, 

the filaments very short, or elongated; anthers 

elongated, sometimes beaked, the beak sometimes longer than the locules, dehiscing by a 

single apical pore; pollen 3-colporate or 3-colporoidate; glandular, annular, lobate disk sometimes present between the petals and stamens; 

pistil 1, the carpels 2, (an occasional pistil has 3 

carpels), style 1, rarely bifid for 0.3 its length, 

elongated, slender, the stigma(s) apical punctiform or slightly capitate; ovary bilocular, the 

ovules 1-2 (-4) per locule, axile, sometimes subapical, bitegmic, crassinucellar, anatropous; fruit 

a compressed capsule, dehiscence loculicidal and 

sometimes also septicidal; seeds pilose or glabrous, often with a conspicuous chalaza1 appendage; embryo straight, cylindrical, 0.7. the length 
NUMBER 58 307 

FIGURE 164.-PITTOSPORACEAE: a, Pittosporum revolutum flowering twig, flower, 1 .s. of same, 

floral diagram, P. tobira dehiscent fruit, I.s. of seed showing the minute embryo; b, Sollya 

heterophylla part of flowering twig, essential organs, 1 .s. of pistil, C.S. of ovary, infructescence, 

1 .s. of seed; c, Bzllardiera longzjlora flowering twig, C.S. of ovary, part of fruiting twig, Citriobatus 

multLJorus flowering branch, 1,s. of pistil, c. s. of ovary, Pronaya elegans flower, 1,s. of same 

(after Baillon, 1866-1 895; Pax). 

of the endosperm, the cotyledons not broadened, 

0.2 the length of the embryo; endosperm copious, fleshy or subcartilaginous. 

Composition: 3 genera, 30 species. 

Distribution: Extratropical Australia, especially southern Western Australia. 

ELAEAGNACEAE (Figure 163d,e).-Shrubs and 

small trees, sometimes the branchlets ending in 

thorns, sometimes producing gum when traumatized; root nodules containing nitrogen-fixing 

organisms present; tannins abundant; xylem vessel perforation plates simple; parenchyma diffuse; leaves alternate or rarely opposite, simple, 

entire, the indument lepidote, rarely also stellate, 

exstipulate, stomata of ranunculaceous type; inflorescences axillary or at defoliated nodes of 

previous year’s growth, fascicles, spikes, racemes, 

corymbs, cymes or the flowers solitary; flowers 

actinomorphic, bisexual or the plants dioecious 

or polygamous; sepals 4 (2-8), connate, the calyx 

tubular or rarely saucer-shaped (8 HippophuE), 

the lobes valvate in bud, often colored; petals 0; 

stamens 4 (3-5), alternisepalous or 8, the filaments inserted at throat of calyx or rarely free, 

short or elongated (Hippophae),, or rarely the 

anthers sessile; anthers oblong or as long as wide, 

dorsifixed or subbasifixed (Hippophue); pollen 3 

(2)-colporate, or rarely 4-ruporate; disk usually 

present, inserted at throat of calyx-tube, conic, 

cup-shaped or of 8 separate glands; pistil 1, unicarpellate, style elongated, the stigma laterally 

decurrent, linear or rarely subcapitate; ovary 
308 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

superior, ovule 1, basal, bitegmic, crassinucellar, 

anatropous; fruit an achene but drupe- or berrylike due to development of the closely investing 

calyx-tube; embryo large, straight, 0.9 the length 

of the endosperm, the cotyledons thick, planoconvex, 0.7-0.8 the length of the embryo, 2.0- 

3.5 times wider than the radicle, the latter partly 

invested; endosperm scanty or 0. 

Composition: 3 genera, 50 species. 

Distribution: Mostly coastal regions and 

steppes in temperate regions and mountains of 

subtropical regions; temperate North America 

and temperate to tropical Eurasia, one species 

extending to northeastern Australia. 

PITTOSPORACEAE (Figure 164a-c).-Trees, 

shrubs, and climbers; xylem vessel perforation 

plates simple, very rarely scalariform with few 

bars (1 - 10); resin abundant in canals in the bark; 

leaves simple, alternate, or whorled, usually entire, very rarely sinuate-toothed, juvenile leaves 

of Pittosporum anomalum are serrate, sometimes 

pellucid-dotted (Citriobatus), exstipulate; inflorescences axillary or terminal, mostly thyrses, 

rarely simple cymes, corymbose, or the flowers 

solitary; flowers bisexual, rarely the plants polygamous, actinomorphic, rarely slightly zygomorphic (Cheiranthera); sepals 5, free or connate; 

petals 5, free or basally connate, imbricate, the 

claws often connivent; stamens 5, alternipetalous, 

free, sometimes connivent, hypogynous; anthers 

lanceolate-oblong, basifixed or subbasally dorsifixed, rarely connivent around the style, dehiscing longitudinally or by 2 apical pores; pollen 

usually 3-colporate, occasionally 4-ruporate or 4- 

rupate; disk absent, rarely present at the base of 

a stipe (Citriobatus); pistil 1, the carpels 2 (3-5), 

style 1, short or long, the stigmas apical, capitate, 

almost entire or obscurely 2-5-lobed; ovary superior, unilocular, rarely 2-3-locular or incompetely 2-5-locular, the ovules several per locule, 

parietal or basal, rarely axile, unitegmic, tenuinucellar, anatropous and campylotropous?, the 

funicles sometimes long (Bursaria); fruit a loculicidal and sometimes also septicidal capsule or 

berry, rarely dry and indehiscent, the seeds 1- 

numerous, rarely winged; embryo undifferentiated, minute, linear, straight, 0.1 the length of 

the endosperm; endosperm copious, firm-fleshy. 

Composition: 9 genera, 220 species. 

Distribution: Centered in Australia, especially 

southern Western Australia, extending into tropical regions; Pittosporum also in tropical and subtropical Africa, Madagascar, Asia, and East Indies, and New Zealand. 
References 

Axelrod, D.L. 

1960. The Evolution of Flowering Plants. In S. Tax, 

editor, The Evolution of Lqe, pages 227-305. Chicago. 

The Potentialities and Limitations of Wood Anatomy in the Study of the Phylogeny and Classification of Angiosperms. Journal of the Arnold Arboretum, 38:243-254. 

Degeneriaceae, a New Family of Flowering Plants 

from Fiji. Journal of the Arnold Arboretum, 

23:356-365, 5 plates. 

Bailey, I.W. 

1937. 

Bailey, I.W., and X.C. Smith 

1942. 

Baillon, H.E. 

Bedell, H.G., and J.L. Reveal 

1866-1 895. 

1982. 

Histoire des plantes. Volumes 1-1 1. Paris. 

Amended Outlines and Indices for Six Recently 

Published Systems of Angiosperm Classification. 

Phytologia, .51:65-156. 

Plant Classijkation. 2nd edition. Lexington, Mass. 

Benson, L.D. 

Benthani, G., and J.D. Hooker 

Bessey, C.E. 

1979. 

1862-1 883. 

19 15. 

Boivin, B. 

19.56. 

Genera Plantarum. 3 Volumes. London. 

Phylogenetic Taxonomy of Flowering Plants. Annals of hlissouri Botanical Garden, 2: 109-1 64. 

Les Families des Tracheophytes. Bulletin de la 

Societei Botanique de France, 103:490-305. 

Flora de la provincia de Buenos Aires. Buenos Aires, 

Argentina. 

Foundations of Plant Geography. New York and 

London. 

Cabrerd, A.L. 

1963. 

Cain, S.A. 

1944. 

Capuron, R. 

1963a. Diegodendon. Adansonia (new series), 3(3):385- 

1963b. Rhopalocarpacees. In Flore de Madagascar et des 

392. 

Comores, number 127. Paris. 

The Systematics of Angiosperms. In A Century of 

Progress in the A'atural Sciences, 1853-1953, pages 

405-483. San Francisco: California Academy of 

Science. 

1964. Systematic Botany-An Unending Synthesis. 

Taxon, 13:257-273. 

Constance, L. 

19.55. 

Corner, E.J.H. 

1976. The Seeds of Dicotyledons. 2 volumes. Cambridge, 

England. 

The Evolution and Classzjication of Flowering Plants. 

Boston. 

An Integrated System of Classtjication of Flowering 

Plants. New York. 

Brunelliaceae. In Flora h'eotropica. Darien, Connecticut. 

Cronquist, A. 

1968. 

198 1. 

Cuatrecasas, J. 

1970. 

Curtis, U'. 

1787-1983. The Botanical Magazine. Volumes 1-182. 

London. 

A System of Classification of the Angiosperms to 

be Used to Demonstrate the Distribution of Characters. Botaniska A'otiser, 128:l 19-147. 

A Revised System of Classification of the Angiosperms. Botanical Journal of the Linnean Society, 

General Aspects of Angiosperm Evolution and 

Macrosystematics. Nordic Journal of Botany, 

3:119-149. 

The Bases of Angiosperm Phylogeny. Annals of 

the Missouri Botanical Garden, 62:5 15-834. 

Dahlgren, R. 

1975. 

1980. 

80:91-124. 

1983. 

Davidse, G., editor 

1975. 

Davis, G.L. 

1966. Systematic Embryology of the Angiosperms. New 

York. 

Principles of Angiosperm Taxonomy London. 

Morphology of the Angiosperms. New York. 

New Evidence of Relationships and Modern Systems of Classification of the Angiosperms. Nordic 

Journal of Botany, 3:3-155. 

Phylogenetic Patterns and the Evolutionary Process. 

New York. 

Les Vegetaux vasculaires. In M. Chadefaud and 

L. Emberger, Traite de Botanique Systematique. 

Paris. 

Davis, P.H., and V.H. Heywood 

1963. 

Eames, A. 

1961. 

Ehrendorfer, F., and R. Dahlgren, editors 

1983. 

Eldredge, N., and J. Cracraft 

1980. 

Ernberger, L. 

1960. 

Engler, A. 

188 1 - 1984. Botanische Jahrbiicher fur Systematik. Pflan309 
310 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

zengeschichte und P’anzengeographie. Leipzig. 

1900-19.53. Das P’anzenreich. 107 volumes. Leipzig 

and Berlin. 

1926. Angiospermae: Kurze Erlauterung der Bluten 

und Fortpflanzungsverhaltnisse. In A. Engler and 

K. Prantl, Die naturlichen Pjlanzenfamilien. 2nd 

edition, volume 14a. Leipzig. 

Engler, A,, and E. Gilg 

Syllabus der P’anzenfamilien. 9th and 10th editions. Berlin. 1924. 

Engler, A,, and K. Prantl 

1887-191 3. 

1924-1 980. 

Die naturlichen Pfanzenfamilien. 1st edition, 

Die natiirlichen Pfanzenfamilien. 2nd ediparts 3 and 4. Leipzig. 

tion, volumes 16b-2 1, 28b. Leipzig. 

Pollen ‘Worphology and Plant Taxonomy: Angaosperms. Stockholm. 

Anatomy ofSeed Plants. 2nd edition. New York. 

The Principles of Pollination Ecology. 3rd edition. 

Oxford and New York. 

Erdtnian, G. 

1952. 

Esau, K. 

1977. 

Faegri, K., and L. van der Pi1 

1978. 

Gibbs, R.D. 

1974. Chemotaxonomy of Flowering Plants. 4 volumes. 

Montreal and London. 

Good, R. 

1974a. Features of Evolution in the Flowering Plants. New 

1974b. The Geography of the Flowering Plants. 4th edition. 

York. 

London. 

Families of Dicotyledons. U’altham, Massachusetts. 

L’Origine et le systkme phylktique des Angiosperms. Archives Neerlandaise des Sciences Exactes et 

h’aturelles, series 3B, 1: 146-234. 

Hawkes, J.G., R.N. Lester, and A.D. Skelding, editors 

The Biology and Taxonomy of the Solanaceae. London and New York. 

Gundersen, A. 

1950. 

Hallier, H. 

1912. 

1979. 

Hegnauer, R. 

1962-1973. Chemotaxonomie der Pjlanzen. 6 volumes. 

Basel and Stuttgart. 

The Biology and Chemistry of the Umbellijerae. London. 

The Biology and Chemistry of the Compositae. London 

and New York. 

Heywood, V.H., D.M. Moore, I.B.K. Richardson, and W.T. 

Flowering Plants of the World. New York. 

1837-1 982. Hooker’s Icones Plantarum. Volumes 1-38. 

Heywood, \’.H., editor 

197 1. 

Heywood, V.H, J.B. Harborne, and B.L. Turner, editors 

1977. 

Stearn, editors 

1978. 

Hooker, W.J., and J.D. Hooker 

London. 

Hutchinson, J. 

1964, 1967. 

1969. 

1973. 

The Genera of Flowering Plants. 2 volumes. 

Oxford, England. 

Evolution and Phylogeny of Flowering Plants. London and New York. 

The Families of Flowering Plants. 3rd edition. Oxford, England. 

Hutchinson, J., and J.M. Dalziel 

1954-1 972. Flora of West Tropical Africa. 2nd edition (3 

volumes iii 4 parts). London. 

A Glossary of Botanic Terms. 4th edition. London 

and New York. 

Taxonomy of the Flowering Plants. New York and 

London. 

An Introduction to the System of Classification 

Used in the Flora of Australia. In Flora of Australia, 1:77-111. Canberra. 

Insects as Flower Visitors and Pollinators. Annual 

Review of Entomology, 28:407-453. 

Flowering Plants, Evolution and ClasszJication of 

Higher Categories: Plant Systematics and Evolution. 

Supplement 1. \’ienna and New York. 

Taxonomy of Vascular Plants. New York. 

A General Sjstem of Botany. London, Boston, and 

New York. 

Jackson, B.D. 

1928. 

Johnson, A.M. 

1931. 

Kanis, A. 

1981. 

Kevin, P.G., and H.G. Baker 

1983. 

Kubitzki, K., editor 

1977. 

Lawrence, G.H.M. 

Le Maout, E., and J. Decaisne 

195 1. 

1873. 

Lemke. A. 

1929-1 93 1. Dictionaire descriptifet synonymique desgenres 

de plantes phanerogames. 9 volumes. Brest, France. 

The Vegetable Kingdom. 3rd edition. London. 

An Introduction to the Embryology of the Angiosperms. 

New York. 

The Flora of South Africa. Volume 2, part 2, 210. 

Capetown. 

Comparative Internal Morphology of Seeds. American Midland Naturalist, 36:5 13-660. 

Lindley, J. 

1853. 

Maheshwari, P. 

1950. 

Marloth, R. 

1925. 

Martin, A.C. 

1946. 

Martius, C.F.P. 

1840- 1906. 

Meeuse, A.D.J. 

Flora Brasiliensis. 15 volumes (in 40 parts). 

Munich, Vienna, and Leipzig. 

The Descent of the Flowering Plants. Acta Botanica ,Veerlandica, 19:61-72, 133-140. 

A Contribution to the Taxonomy and Biology of 

Rafflesia arnoldi in West Sumatra. Annales Bogoriensis, 3( 1):33-44. 

1970. 

Meijer, W. 

1958. 
NUMBER 58 31 1 

Melchior, H., editor 

1964. Engler’s Syllabus der Pfanzenfamilien. 12th edition, volume 2. Berlin. 

Anatomy of the Dicotyledons. 1st edition, 2 volumes. 

Oxford, England. 

Anatomy of the Dicotyledons. 2nd edition, 2 

volunies. Oxford, England. 

Fossil Pollen Records of Extant Angiosperms. The 

Botanical Review, 47:l-142. 

Sinopsis de la Flora Chilena, claves para la identijiracion de familias y generos. Santiago, Chile. 

Advances in Legume Systematics. 2 volumes. Kew, 

England. 

The Pollination of Flowers. London. 

Compendium van de Terminologie, Nomenclatuur en 

Systematiek der Zaadplanten. 3rd edition. Utrecht. 

Vuscular Plant Systematics. New York, Evanston, 

San Francisco, and London. 

1983. Pollination Biology. Orlando. 

1938. 

Ricardi, %I. 

Metcalfe, C.R., and L. Chalk 

1950. 

1979-1983. 

Muller, J. 

198 1. 

Munoz-Pizarro, Carlos 

1959. 

Polhill, R.M., and P.H. Raven, editors 

198 1. 

Proctor, M.> and P. Yeo 

1973. 

Pulle, A..4. 

19.52. 

Radford, A.E., W.C. Dickison, J.R. Massey, and C.R. Bell 

1974. 

Real, I>., editor 

Rendle, A.B. 

The Classijication of Flowering Plants. 2nd edition. 

Cambridge, England. 

1967. Revision taxononiica de las Malesherbiaceaes. 

Guyana, 16, 139 pages. 

Guide to the Generic Names Appearing in the Index 

Kewensis and Its Fqteen Supplements, Pars IV: Conspectus Systematis. Cowansville, Qukbec, Canada. 

Rouleau, E. 

198 1. 

Royal Botanic Gardens, Kew 

1974-1 985. Kew Record of Taxonomic Literature Relating 

to Vascular Plants (for 1971-1980). 10 volumes. 

London. 

An Appraisal of the Orders and Families of Primitive Extant Angiosperms. Journal of the Indian 

Botanical Society. 50:2 15-226. 

Boraginaceas. In Flora Illustrada Catarinense. Itajai, Brazil. 

Sneath, P.H., and R.R. Sokal 

h‘umerical Taxonomy: The Principles and Practice of 

‘Vumerical Classzjlcation. San Francisco. 

Present Aspects of Evolutionary History of Telomophyta. Annales Wniuersitatis Scientiarum Budupestinensis: Sectio Bzologica, 4: 167-1 78. 

Smith, A.C. 

197 1. 

Smith, L.B. 

1970. 

1973. 

Sob, C.R. de 

1961. 

Sporne, K.R. 

1975. The Morphology of Angiosperms: The Structure and 

Evolution of Flowering Plants. New York. 

1976-1985. Taxonomic Literature. 2nd edition, 5 volStafleu, F.A., and R.S. Cowan 

umes. Utrecht and The Hague. 

Stapf, 0. 

1929-1 93 1. Index Londinensis to Illustrations of Flowering 

Plants, Ferns and Fern Allies. 6 volumes (additional 2-volume supplement by W.C. Worsdell 

issued in 1941). Oxford, England. 

Flowering Plants, Evolution above the Species Level. 

Cambridge, Massachusetts. 

Manual of Vascular Plants of the Lower Yangtze 

Vulley, China. Corvallis, Oregon. 

Flowering Plants: Origin and Dispersal. Edinburgh, 

Scotland. 

Outline of the Classification of Flowering Plants. 

The Botanical Review, 46:225-359. 

The Systematic Arrangement of Dicotyledonous 

Families. In C.R. Metcalfe and L. Chalk, Anatomy 

of Dicotyledons. 2nd edition, volume 2, pages 180- 

201. 

Thanikaimoni, G. 

1972-1 980. 

Stebbins, G.L. 

1974. 

Steward, A.N. 

1958. 

Takhtajan, A. 

1969. 

1980. 

1983. 

Index Bibliographique sur la morphologie 

des pollens d’Angiospermes. In Trauaux de la 

section scientijlque et technique de l’lnstitut Francais 

de Pondichery, India, volumes 12 (parts 1 and 2), 

13, and 18. 

A Phylogenetic Classification of the Angiosperms. 

In M.K. Hecht, W.C. Steere, and B. Wallace, 

Evolutionary Biology, 9335-106. New York and 

London. 

Proposed New Realignments in the Angiosperms. 

Nordic Journal of Botany, 3:85-1 17. 

Die Areale und Arealtypen der Angiospermen-Familien. Leipzig. 

Handbuch der systematischen Botanik. 4th edition. 

Leipzig and Vienna. 

A Dictionary of the Flowering Plants and Ferns. 8th 

edition. Cambridge, England. 

Thorne, R.F. 

1976. 

1983. 

\.‘ester, H. 

1940. 

Wettstein, R. 

1935. 

Willis, J.C. 

1973. 

Young, D.A., and D.S. Seigler, editors 

198 1. Phytochemistry and Angiosperm Phylogeny. New 

York. 

Young, D.J., and L. Watson 

The Classification of Dicotyledons: .4 Study of the 

Upper Levels of the Hierarchy. Australian Journal 

of Botany, 18:387-433. 

1970. 
Index to Families and Orders 

Acanthaceae, 293 

Aceraceae, 243 

Achariaceae, 13 1 

Achatocarpaceae, 164 

Actinidiaceae, 103 

Adoxaceae, 232 

Aextoxicaceae, 18 1 

Agdestidaceae, 167 

Ai7oaceae, 160 

Akaniaceae, 24 I 

Alangiaceae, 126 

Amaranthaceae, 170 

Amborellaceae, 48 

Anacardiacede, 238 

Ancistrocladaceae. 1 14 

Annonaceae, 41 

Apiaceae, 226 

Apiales, 225 

Apocynaceae, 264 

Aptandraceae, 273 

Aquifoliaceae, 158 

Araliaceae, 226 

Aristolochiaceae, 54 

A r i s t o I oc h ia I es, 5 4 

Asclepiadaceae, 264 

Asteraceae, 304 

Asterales, 304 

Asteranthaceae, 194 

Austrobailevaceae, 43 

Balanitaceae, 255 

Balanopaceae, 73 

Balanopales, 73 

Balanophoraceae, 15 1 

Balanophorales, 15 1 

Balsaminaceae, 250 

Barbeuiaceae, 164 

Barbeyaceae, 83 

Barringtoniaceae, 194 

Basellaceae, 169 

Bataceae, 99 

Batales, 99 

Baueraceae, 2 1 2 

Begoniaceae, 225 

Begoniales, 224 

Berberidaceae, 59 

Betulaceae, 79 

Bignoniaceae, 294 

Bixaceae, 139 

Bombacaceae, 187 

Bonnetiaceae, 106 

Boraginaceae, 285 

Brassicaceae, 95 

Bretschneideraceae, 99 

Brunelliaceae, 2 15 

Bruniaceae, 2 12 

Brunoniaceae, 302 

Buddlejaceae, 289 

Burseraceae, 238 

Buxaceae, 33 

Byblidaceae, 222 

Cabombaceae, 70 

Cactaceae, 160 

Caesalpiniaceae, 86 

Callitrichaceae, 6.5 

Calycanthaceae, 50 

Calyceraceae, 302 

Campanulaceae, 300 

Campan u lales , 2 99 

Canellaceae, 45 

Zannabaceae, 83 

Zapparaceae, 93 

Zaprifoliaceae, 23 1 

Zardiopteridaceae, 274 

Zaricaceae, 13 1 

Zaryocaraceae, 1 16 

Zaryophyllaceae, 168 

Zaryophyllales, 158 

::asuarinaceae, 8 1 

Zasuarinales, 81 

Zelastraceae, 153 

Zelastrales, 15 1 

Zephalotaceae, 2 19 

,eratophyllaceae, 70 

>ercidiphyIlaceae, 30 

:henopodiaceae, 17 I 

Zhloranthaceae, 70 

Zircaeasteraceae, 62 

=istaceae, 138 

Zistales, 137 

Zlethraceae, 14 1 

Zlusiaceae, 1 10 

=neoraceae, 236 

;ochlospermaceae, 139 

Zolurnelliaceae, 269 

Conibretaceae, 203 

Connaraceae, 250 

Convolulaceae, 267 

Coriariaceae, 19 1 

Cornaceae, 230 

Cornales, 228 

Corynocarpaceae, 189 

Crassulaceae, 208 

Crossosomataceae, 189 

Crypteroniaceae, 200 

Ctenolophonaceae, 15 1 

Cucurbitaceae, 137 

Cunoniaceae, 2 15 

Cuscutaceae, 267 

Cbrillaceae, 146 

Daphniphyllaceae, 37 

Datiscaceae, 225 

Davidiaceae, 228 

Davidsoniaceae, 2 10 

Degeneriaceae, 43 

Dialypetalanthaceae, 194 

Diapensiaceae, 146 

Dichapetalaceae, 133 

Didiereaceae, 162 

Didymelaceae, 38 

Diegodendraceae, 1 19 

Dilleniaceae, 101 

Dilleniales, 10 1 

Dioncophyllaceae, 220 

Dipsacaceae, 232 

Dipsacales, 230 

Dipterocarpaceae, 1 14 

Donatiaceae, 2 14 

Droseraceae, 220 

Droserdles, 220 

<benaceae, I22 

Zbenales, 122 

heagnaceae, 307 

<laeocarpaceae, 183 

Clatinaceae, 1 I2 

Snipetraceae, 144 

Xpacridaceae, 144 

Xreniosynaceae, 208 

Cricaceae. 14 1 

Cricales, 139 

Crvthropalaceae, 156 

312 
NUMBER 58 313 

Erythroxvlaceae, 253 

Escalloniaceae, 2 15 

Eucornmiaceae, 83 

Eucryphiaceae, 108 

Euphorbiaceae, 180 

Euphorbiales, 178 

Euponiatiaceae, 43 

Eupteleaceae, 30 

Fabaceae, 87 

Fabaks, 8.5 

Fagaceae, 79 

Fagales, 75 

Flacourtiaceae, 127 

Fouquieriaceae, 282 

Francoaceae, 208 

Frankeniaceae, 173 

Funiariaceae. 9 1 

Garryaceae, 230 

Geissoloniataceae, 279 

Gentianaceae, 269 

Gentianales, 262 

Geraniaceae, 246 

Geraniales, 245 

Gesneriaceae, 294 

(;lobulariaceae, 289 

Gomortegaceae, 52 

Gonystvlaceae, 280 

Goodeniaceae, 300 

Goupiaceae, 155 

Greyiaceae, 2 18 

Grossulariaceae, 2 1.5 

Grubbiaceae, 277 

Gunneraceae, 64 

Gyrostenionaceae, 163 

Haloragaceae, 62 

Haloragales, 62 

Harnamelidaceae, 35 

Hamamelidales, 33 

Hernandiaceae, 54 

Himantandraceae, 43 

Hippocastanaceae, 243 

Hippuridaceae, 64 

Hoplestigmataceae, 134 

Humiriaceae, 1 14 

Hydnoraceae, 148 

Hydrangeaceae, 2 10 

Hydrophyllaceae, 285 

Hydrostachyaceae, 222 

Hypericaceae, 1 12 

Icacinaceae, 1.38 

Idiosperniaceae, 30 

Illecebraceae, 168 

Illiciaceae, 45 

Irvingiaceae, 153 

I teaceae, 2 1 2 

Ixonanthaceae. 15 1 

Juglandaceae, 77 

Krameriaceae, 258 

Lacistemataceae, 127 

Lactoridaceae, 68 

Lamiaceae, 297 

Lamiales, 296 

Lardizabalaceae, 57 

Lauraceae, 52 

Laurales, 52 

Lecythidaceae, 192 

Leeaceae, 156 

Leitneriaceae, 75 

Lennoaceae, 146 

Lentibulariaceae, 291 

Lirnnanthaceae, 248 

Linaceae, 253 

Lissocarpaceae, 126 

Loasaceae, 134 

Loganiaceae, 262 

Loranthaceae, 277 

Lythraceae, 197 

Magnoliaceae, 38 

Magnoliales, 38 

Malesherbiaceae, 129 

Malpighiaceae, 255 

Malvaceae, 185 

Malvales, 18 1 

Marcgraviaceae, 1 16 

Medusagynaceae, 108 

Medusandraceae, 273 

Melastomataceae, 200 

Meliaceae, 238 

Melianthaceae, 243 

Menispermaceae, 55 

Menyanthaceae, 27 1 

Mimosaceae, 87 

Misodendraceae, 277 

Molluginaceae, 168 

Monimiaceae, 48 

Monotropaceae, 144 

Moraceae, 85 

Moringaceae, 99 

Myoporaceae, 297 

Myricaceae, 77 

Myristicaceae, 43 

Mvrothamnaceae, 33 

Myrsinaceae, 175 

Myrtaceae, 192 

Myrtales, 19 1 

Nandinaceae, 59 

Nelumbonaceae, 73 

h'epenthaceae, 65 

Nepenthales, 65 

Neuradaceae, 189 

Nolanaceae, 287 

Nyctaginaceae, 167 

Nyrnphaeaceae, 7 1 

Nymphaeales, 70 

Nyssaceae, 228 

Ochnaceae, 1 16 

Octoknemaceae, 273 

Olacaceae, 27 1 

Oleaceae, 299 

Oleales, 259 

Oliniaceae, 200 

Onagraceae, 203 

Opiliaceae, 273 

Orobanchaceae, 293 

Oxalidaceae, 248 

Paeoniaceae, 6 1 

Pandaceae, 18 1 

Papaveraceae, 9 1 

Papaverales, 89 

Paracryphiaceae, 108 

Parnassiaceae, 208 

Passifloraceae, 129 

Pedaliaceae, 296 

Pellicieraceae, 108 

Penaeaceae, 206 

Pentadiplandraceae, 93 

Pentaphylacaceae. 106 

Peridiscaceae, 134 

Petiveriaceae, 164 

Philadelphaceae, 2 10 

Phrymaceae, 297 

Phytolaccaceae, 164 

Piperaceae, 67 

Piperales, 67 

Pittosporaceae, 308 

Plantaginaceae, 178 

Plantaginales, 178 

Platanaceae, 33 

Plocosperrnataceae, 264 

Plumbaginaceae. 173 

Podophyllaceae, 59 

Podostemaceae, 2 2 2 

Polenioniaceae, 282 

Polenioniales, 282 
314 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Polygalaceae, 256 

Polygalales, 255 

Polygonaceae, 177 

Polygonales, 177 

Portulacaceae, 162 

Priniulaceae, 171 

Priniulales, 17 1 

Proteaceae, 73 

Proteales, 73 

Pterostenionaceae, 2 10 

Punicaceae, 197 

Py rolaceae, 142 

Quiinaceae, 1 12 

Rafflesiaceae, 148 

Rafflesiales, 148 

Ranunculaceae, 61 

Ranunculales, 54 

Resedaceae, 93 

Rhaninaceae, 156 

Rhaninales, 155 

Rhizophoraceae, 197 

Rhoipteleaceae, 77 

Rosaceae, 188 

Rosales, 187 

Rubiaceae, 267 

Rutaceae, 234 

Rutales, 234 

Sabiaceae, 245 

Salicaceae, 8 1 

Salicales, 8 1 

Salvadoraceae, 261 

Santalaceae, 275 

Santalales, 27 1 

Sapindaceae, 24 1 

Sapindales, 24 1 

Sapotaceae, 124 

Sarcolaenaceae, 122 

Sargentodoxaceae, 57 

Sarraceniaceae, 65 

Sarraceniales, 63 

Saurauiaceae, 104 

Saururaceae, 67 

Saxifragaceae, 206 

Saxifragales, 206 

Schisandi-aceae, 45 

Scrophulariaceae, 289 

Scrophulariales, 285 

Scyphostegiaceae, 134 

Scytopetalaceae, 1 19 

Simaroubaceae, 236 

Simniondsiaceae, 75 

Siphonodontaceae, 155 

Solanaceae, 287 

Sonneratiaceae, 195 

Sphaerosepalaceae, 1 19 

Stachyuraceae, 134 

Stackhousiaceae, 25 1 

Staphpleaceae, 243 

Stegnosperniataceae, 167 

Sterculiaceae, 183 

Strasburgeriaceae, 1 19 

Stylidiaceae, 302 

Styracaceae, 124 

Styracales, 124 

Syinplocaceae, 108 

Tamaricaceae, 173 

Tetracarpaeaceae, 2 14 

Tetracentraceae, 30 

Tetranieristaceae, 1 10 

Theaceae, 106 

Theales, 104 

Theligonaceae, 162 

Theophrastaceae, 175 

Thymelaeaceae, 280 

Thyinelaeales, 277 

Tiliaceae, 183 

Tovariaceae, 9 1 

Trapaceae, 203 

Tremandraceae, 304 

Trigoniaceae, 259 

Trimeniaceae, 48 

Trochodendraceae, 30 

Trochodendrales, 29 

Tropaeolaceae, 250 

Turneraceae, 13 1 

Clmaceae, 85 

Crticaceae, 83 

Urticales. 81 

Vahliaceae, 212 

Valerianaceae, 232 

Verbenaceae, 297 

Violaceae, 13 1 

Violales, 126 

l’itaceae, 156 

L’ivianiaceae, 248 

Vochysiaceae, 259 

Winteraceae, 39 

Zygophyllaceae, 253