SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 71 

Classification, Evolution, 

and Phylogeny of the 

Families of Monocotyledons 

Aaron Goldberg 

SMITHSONUN INSTITUTION PRESS 

Washington, D.C. 

1989 
ABSTRACT 

Goldberg, Aaron. Classification, Evolution, and Phylogeny of the Families of Monocotyledons. Smithsonian Contributions to Botany, number 71, 74 pages, 41 figures, 2 tables, 1 

diagram, 1989.-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 250 monocot 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 45 and 103 monocot families in 14 to 

38 orders. I accept 57 families in 18 orders. In Table 1 I give my ordinal allocation of the 

families and that of 11 recent authors to indicate where there is agreement and where there are 

differences to be resolved. I have constructed a dendrogram to suggest relationships and degree 

of advancement of the orders. 

I have written concise, uniform descriptions of all the families of monocots 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. 

Several species are usually used to show the range of major variation within families and trends 

toward related families. 

Monocots and dicots have existed concurrently for most of their history, have been subjected 

to many of the same ecological pressures, and consequently show similar evolutionary trends. 

My approach to understanding evolutionary trends in characters is to relate them to the 

ecological factors that might be responsible for them by their selective action. The monocots 

probably originated under warm temperate or subtropical conditions favorable for growth. A 

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 and colonize generally 

unfavorable habitats. 

A second major trend has been progressively greater specialization for insect pollination. 

The primitive monocots have flowers with numerous spirally arranged parts; those having 

fIowers with few, opposite or whorled parts are derived. 

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 with a particular primitive state, the more primitive the state. 

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 a table giving the primitive and 

derived states for about 85 characters. I also indicate the extent to which I consider the states 

reversible. 

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. If a family has more than one state of a character, 

the state considered primitive for the particular family should be used 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 btitution’s annud report, Smifhsonkn Year. SERIES COVER DESIGN: Leaf 

clearing from the katsura tree Cercidiphyllum japonicum Siebold & Zuccarini. 

Library of Congress Cataloging in Publication Dats 

Goldberg, Aaron 

Classification, cvolutim, and phylogeny of the families of Monocotyledom. 

(Smithsonian contributions to botany ; no. 71) 

Bibliography: p. 

Includes -. . . - -. . index. -. -. .. .

Supr of Docs. no.: SI 1.29:71 

1. Monocotylcdcns-Classification. 2. Monocotyledons -Evolution. 3. Monocotylcdon&ylogmy. I. Title. 

11. sc;ics. 

QKlS747 no. 71 581 s [584’.012] 89-600112 [QK495.A14] 
Contents 

Page 

Introduction ........................................ 1 

Table l.'kdinal Allocation of the Monocotyledon Families by Recent Authors . . 2 

Dendrogram of Suggested Phylogenetic Relationships of the Orders of Monocotyledons ...................................... 7 

Table 2.-Evolutionary Trends in the Monocotyledons ................ 8 

Ordinal and Family Descriptions ............................. 9 

References ........................................ 71 

Index to Orders and Families .............................. 74 

iii 
Contents 

Page 

Introduction ........................................ 1 

Table l.'kdinal Allocation of the Monocotyledon Families by Recent Authors . . 2 

Dendrogram of Suggested Phylogenetic Relationships of the Orders of Monocotyledons ...................................... 7 

Table 2.-Evolutionary Trends in the Monocotyledons ................ 8 

Ordinal and Family Descriptions ............................. 9 

References ........................................ 71 

Index to Orders and Families .............................. 74 

iii 
Classification, Evolution, 

and Phylogeny of the 

Families of Monocotyledons 

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 

250 monocot 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 45 and 

103 monocot families in 14 to 38 orders. I accept 57 families 

and 18 orders. 

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 specimens in all families. 

I have written concise, uniform descriptions of all the 

families of monocots, 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. At present the only 

publications in English describing and illustrating all or almost 

all the monocot families in a single volume are Dahlgren et al. 

(1985) and Hutchinson (1973). Dahlgren et al. (1985) 

recognize twice as many families as I do. Hutchinson’s concept 

of taxa is considerably narrower than mine. Also, I believe his 

Calyciferae and Corolliferae are not natural taxa 

The illustrations have been reproduced as economically as 

possible for the purpose of illustrating major features; no 

attempt has been made to represent the detail that would be 

expected in traditional taxonomic studies published in Smithsonian Contributions to Botany. Abbreviations used in the 

Aaron Goldberg, collaborator, Department of Botany, National 

Museum of Natural History, Smithsonian Imtitution, Washington, 

D.C. 20560. 

1 

legends are as follows: 1.s. = longitudinal section; C.S. = cross 

section. 

Perusal of the references dealing with chemotaxonomy 

(Gibbs, 1974, and Hegnauer, 1962-1986) indicates how little 

is known for many of the families, so I did not consider 

chemical data under the family descriptions. However, under 

each order I have discussed those chemical data I consider 

most significant in understanding relationships of the families 

and occasionally genera. 

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 ( 1862- 1883), 

as well as those authors who have studied particular families. 

Other families accepted by the above authors: 

Cronquist, 1988. Limnocharitaceae in Alismatales; Ruppiaceae and Cymodoceaceae in Najadales; Petrosaviaceae in 

Triuridales; Acoraceae in Arales; Joinvilleaceae in Restionales; 

Hydatellaceae in Hydatellales; Heliconiaceae and Costaceae 

in Zingiberales; Cyanastraceae, Aloaceae and Hanguanaceae 

in Liliales; Geosiridaceae in Orchidales. 

Takhtajan, 1987. Thalassiaceae and Halophilaceae in Hydrocharitales; Limnocharitaceae in Alismatales; Maundiaceae in 

Juncaginales; Ruppiaceae in Potamogetonales; Cymodoceaceae 

in Cymodoceales; Melanthiaceae, Calochortaceae, Geosiridaceae, Rcophilaeaceae, Cyanastraceae, Eriospermaceae, and 

Medeolaceae in Liliales; Asphodelaceae, Dasypogonaceae, 

Aphyllanthaceae, Hyacinthaceae, Alliaceae, Hesperocallidaceae, 

Funkiaceae, Hemerocallidaceae, Phormiaceae, Blanfordiaceae, 

Doryanthaceae, and Ixioliriaceae in Amaryllidales; Convallariaceae, Ruscaceae, Asparagaceae, Dracaenaceae, Nolinaceae, 

Herreriaceae, Asteliaceae, and Hanguanaceae in Asparagales; 

Luzuriagaceae, Petermanniaceae, and Ripogonaceae in 
2 SMITHSONIANCONTRIBUTIONS TO BOTANY 

TAELE 1.4rdinal allocatim of the monocotyledon families by recent authors. 

~~ ~ 

Family 

Pandan 

Alismat 

Butom 

Hydrocharit 

Triurid 

Scheuchzeri 

Juncagin 

Lilae 

Aponogeton 

Zoster 

Potamogeton 

Zannichelli 

Najad 

Posidoni 

Junc 

Thurni 

Centrolepid 

Restion 

Flagellari 

Spargani 

TY Ph 

CYPer 

Po 

Arec 

Cyclan th 

Ar 

Lemn 

Trilli 

Lili 

Smilac 

Goldberg Cronquist Takhtajan Thorne Dahlgren et el. Hutchinion 

1988 1987 1983 1985 1973 

Pandan 

4lismat 

4lismat 

4lismat 

rriurid 

luncagin 

luncagin 

luncagin 

Yajad 

Hajad 

Xajad 

Xajad 

Najad 

Hajad 

lunc 

lunc 

lunc 

lunc 

runc (1) 

TYPh 

TYPh 

CYPer 

Po 

Arec 

Cyclanth 

Ar 

Ar 

Lili 

Lili 

Lili 

Pandan Pandan 

Alismat Alismat 

Alismat Butom 

Hydrocharit Hydrocharit 

Triurid Triurid 

Najad Scheuchzeri 

Najad Juncagin 

in Juncaginaceae Juncagin 

Najad 

Najad 

Najad 

Najad 

Najad 

Najad 

Junc 

Junc 

Restion 

Restion 

Restion 

TYPh 

TYPh 

CYPer 

CYPer 

Arec 

Cyclanth 

Ar 

Ar 

in Liliaceae 

Lili 

Lili 

Aponogeton 

Zoster 

Potamogeton 

Cymodoce 

Najad 

Posidoni 

Junc 

Junc 

Restion 

Restion 

Restion 

Restionineae 

Restionineae 

Pandan 

Alismat 

Alismat 

Alismat 

Triurid 

in Juncaginaceae 

Zoster 

in Juncaginaceae 

Zoster 

Zoster 

Zoster 

Zoster 

Najad 

Potamogetonineae 

Aponogetonineae 

Zosterineae 

Potamogetonineae 

Potamogetonineae 

Zoster 

Potamogetonineae 

Commelin 

Juncineae 

in Juncaceae 

Commelin 

Commelin 

Commelin 

Flagellariineae 

Flagellariineae 

Flagellariineae Flagellariineae 

TYPh in Typhaceae 

TYPh TYPh 

CYPer Commelin 

Juncineae 

Po Commelin 

Poineae 

Arec Arec 

Cyclanth Cyclanth 

Ar Ar 

Ar Ar 

Dioscore in Liliaceae 

Lili Lili 

Smilac Lili 

Liliineae 

Diorcoreineae 

Pandan 

Alismat 

Alismat 

Alismat 

Triurid 

Najad 

Najad 

in Juncaginaceae 

Alismat 

Najad 

Najad 

Najad 

Najad 

Najad 

CYPer 

CYPer 

Po 

Po 

Po 

TYPh 

TYPh 

CYPer 

Po 

Arec 

Cyclanth 

Ar 

Ar 

Dioscore 

Lili 

Dioscore 

Pandan 

Ali sm a t 

Butom 

Butom 

Triurid 

Alismat 

Juncagin 

Juncagin 

Aponogeton 

Aponogeton 

Potamogeton 

Najad 

Najad 

Juncagin 

Junc 

Junc 

Junc 

Junc 

Commelin 

TY Ph 

5Ph 

CYPer 

Gramin 

Palm 

Cyclanth 

Ar 

Ar 

Lili 

Lili 

Lili 
NUMBER 71 3 

TABU 1 .-Continued. 

Family 

Pandan 

Alismat 

Butom 

Hydrocharit 

Triurid 

Scheuchzeri 

Juncagin 

Lilae 

Aponogeton 

Zoster 

Potamogeton 

Zannichelli 

Najad 

Posidoni 

Junc 

Thurni 

Centrolepid 

Restion 

Flagellari 

Spargani 

TYPh 

CYPer 

Po 

Arec 

Cyclanth 

Ar 

Lemn 

Trilli 

Lili 

Smilac 

Melchior S t ebbin s Rouleau Young Benson Bmberger 

1964 1974 1981 1981 1979 1960 - 

Pandan Pandan Pandan Pandan Pandan 'andan 

lelobiae 

lelobiae 

lelobiae 

rriurid 

lelobiae 

lelobiae 

n Juncaginaceae 

lelobiae 

Potamogetonineae 

n Potamogetonaceae 

Alismatineae 

Alismatineae 

Hydrocharitineae 

Scheuchzeriineae 

Potamogetonineae 

3elobiae 

Potamogetonineae 

3elobiae 

Potamogetonineae 

Helobiae 

Potamogetonineae 

.n Potamogetonaceae 

runc 

lunc 

Commelin 

Commelin 

Commelin 

Pandan 

Pandan 

Res tionineae 

Restionineae 

Flagellariineae 

CYPer 

Gramin 

Principes 

Sy nanthae 

Spathiflorae 

Spathiflorae 

in Liliaceae 

Liliiflorae 

Liliineae 

in Liliaceae 

Alismat 

Alismat 

Hydrocharit 

Triurid 

Najad 

Najad 

__- 

Najad 

Najad 

Najad 

Najad 

Najad 

Junc 

Junc 

Restion 

Restion 

Restion 

Po 

Arec 

Cyclanth 

Ar 

Ar 

__- 

Lili 

Lili 

Alismat 

Alismat 

Hydrocharit 

Triurid 

Najad 

Najad 

Najad 

Najad 

Najad 

Najad 

Najad 

Najad 

Najad 

Junc 

Junc 

Restion 

Restion 

Restion 

5Ph 

5Ph 

CYPer 

Po 

Arec 

Cyclanth 

Ar 

Ar 

Lili 

Lili 

Lili 

Alismat 

Alismat 

Alismat 

Triurid 

Zoster 

Zoster 

in Juncaginaceae 

Zoster 

Zoster 

Zoster 

Zoster 

Zoster 

Zoster 

Junc 

in Juncaceae 

Restion 

Restion 

Restion 

in Typhaceae 

TYPh 

CYPer 

Po 

Arec 

Cyclanth 

Ar 

Ar 

--_ 

Lili 

Lili 

Alismat 

Alismat 

Hydrocharit 

Triurid 

--. 

Juncagin 

Juncagin 

Naiad 

Naiad 

Naiad 

Lili 

Lili 

Restion 

Juncineae 

Juncineae 

Restion 

Restion 

TYPh 

TYPh 

Gramin 

Gramin 

Palm 

Ar 

Ar 

Ar 

___ 

Lili 

Liliineae 

in Liliaceae 

Alismat 

Alismat 

Alismat 

Triurid 

Alismat 

Alismat 

Potamogeton 

Alismat 

Potamogeton 

Potamogeton 

Potamogeton 

Potamogeton 

Potamogeton 

Junc 

Junc 

Commelin 

Commelin 

Junc 

Pandan 

Pandan 

CYPer 

Gramin 

Palm 

Cyclanth 

Ar 

Ar 

in Liliaceae 

Lili 

in Liliaceae 
4 SMlTHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE l.-Continued. 

Family 

Agav 

Xanthorrhoe 

Philesi 

Stemon 

Dioscore 

Tacc 

Pontederi 

Amaryllid 

Vellozi 

Bromeli 

Commelin 

Mayac 

Xyrid 

Rapate 

Eriocaul 

Mu s 

Strelitzi 

Lowi 

Zingiber 

Cam 

Marant 

Haemodor 

Philydr 

Irid 

Burmanni 

Corai 

Orchid 

Goldberg Cronquist Takhtajan Thorne Dahlgren et 01. Hutchinson 

1988 1987 1983 1985 1973 

Lili Lili Amaryllid in Liliaceae 

Lili 

Lili 

Lili 

Lili 

Lili 

Lili 

Lili 

Lili 

Bromeli 

Commelin 

Commelin 

Commelin 

Commelin 

Commelin 

Zingiber 

Zingiber 

Zingiber 

Zingiber 

Zingiber 

Zingiber 

Irid 

Irid 

Irid 

Irid 

Orchid 

Orchid 

Lili 

in Smilacaceae 

Lili 

Lili 

Lili 

Lili 

in Liliaceae 

Lili 

Bromeli 

Commelin 

Commelin 

Commelin 

Commelin 

Eriocaul 

Zingiber 

Zingiber 

Zingiber 

Zingiber 

Zingiber 

Zingiber 

Lili 

Lili 

Lili 

Orchid 

Orchid 

Orchid 

Amaryllid in Liliaceae 

Smilac Lili 

Dioscore Lili 

Dioscore Lili 

Tacc Lili 

Pontederi Commelin 

Amaryllid in Liliaceae 

Dioscoreineae 

Dioscoreineae 

Dioscoreineae 

Dioscoreineae 

Pontederiineae 

Vellozi Commelin 

Bromeli Commelin 

Commelin Commelin 

Commelin Commelin 

Commelin Commelin 

Commelin Commelin 

Commelin Commelin 

Zingiber Zingiber 

Zing ib e r Zingiber 

Zingiber Zingiber 

Zingiber Zingiber 

Zingiber Zingiber 

Zingiber Zingiber 

Haemodor Commelin 

Philydr Commelin 

Lili Lili 

Burmanni Lili 

Burmanni in Burmanniaceae 

Velloziineae 

Bromeliineae 

Commelinineae Commelinineae 

Commelinineae Commelinineae 

Xyridineae Bromeliineae 

Xyridineae Bromeliineae 

Eriocaulineae Briocaulineae 

Musineae 

Musineae 

Musineae 

Zingiberineae 

Zingiberineae 

Zingiberineae 

Pontederiineae 

Pontederiineae 

Iridineae 

Iridineae 

Orchid Lili 

Orchidineae 

Asparag 

Asparag 

Asparag 

Dioscore 

Dioscore 

Dioscore 

Pontederi 

Asparag 

Vellozi 

Bromeli 

Commelin 

Commelin 

Commelin 

Commelin 

Commelin 

Zingiber 

Zingiber 

Zingiber 

Zingiber 

Zingiber 

Zingiber 

Haemodor 

Philydr 

Lili 

Burmanni 

Burmanni 

Lili 

Agav 

Agav 

Alstroemeri 

Dioscore 

Dioscore 

Haemodor 

Lili 

Amaryllid 

Haemodor 

Bromeli 

Commelin 

Commelin 

Xyrid 

Xyrid 

Eriocaul 

Zingiber 

Zingiber 

Zingiber 

Zingiber 

Zingiber 

Zingiber 

Haemodor 

Haemodor 

Irid 

Burmanni 

Burmanni 

Orchid 
NUMBER 71 5 

TABLE l.-Continued. 

Family 

Agav 

Xanthorrhoe 

Philesi 

Stemon 

Dioscore 

Tacc 

Pontederi 

Amaryllid 

Vellozi 

Bromeli 

Commelin 

Mayac 

Xyrid 

Rapate 

Eriocaul 

Mus 

Strelitzi 

Lowi 

Zingiber 

Cann 

Marant 

Haemodor 

Philydr 

Irid 

Burmanni 

Corsi 

Orchid 

Melchior Stebbins Rouleau Young Benron Emberger 

1964 1974 1981 1981 1979 1960 

iliiflorae Lili Lili 

Liliineae 

iliiflorae 

Liliineae 

n Liliaceae 

iliiflorae 

Liliineae 

iliiflorae 

Liliineae 

.iliiflorae 

Liliineae 

iliiflorae 

Pontederiineae 

iliiflorae 

Liliineae 

iliiflorae 

Liliineae 

3romeli 

Zommelin 

2ommelin 

Zommelin 

Zommelin 

2ommelin 

Scitamineae 

Commelinineae 

Commelinineae 

Commelinineae 

Commelinineae 

Eriocaulineae 

in Musaceae 

Scitamineae 

Scitamineae 

Scitamineae 

Scitamineae 

Liliiflorae 

Liliineae 

Liliiflorae 

Philydrineae 

Liliiflorae 

Iridineae 

Liliiflorae 

Burmanniineae 

Liliiflorae 

Burmanniineae 

Microspermae 

Lili 

--- 

Lili 

Lili 

Lili 

Lili 

in Liliaceae 

Lili 

Bromeli 

Commelin 

Commelin 

Commelin 

Commelin 

Eriocaul 

Zingiber 

Zingiber 

Zingiber 

Zingiber 

Zingiber 

Zingiber 

Lili 

Lili 

Lili 

Orchid 

Orchid 

Orchid 

Lili 

Lili 

Lili 

Lili 

Lili 

Lili 

Lili 

Lili 

Bromeli 

Commelin 

Commelin 

Commelin 

Commelin 

Eriocaul 

Zingiber 

Zingiber 

Zingiber 

Zingiber 

Zingiber 

Zingiber 

Lili 

Lili 

Irid 

Burmanni 

Burmanni 

Orchid 

Lili 

Lili 

--- 

Dioscore 

Dioscore 

Dioscore 

Lili 

-_- 

Lili 

Bromeli 

Commelin 

Commelin 

Commelin 

Commelin 

Eriocaul 

Zingiber 

Zingiber 

Zingiber 

Zingiber 

Zingiber 

Zingiber 

Lili 

Lili 

Lili 

Burmanni 

Burmanni 

Orchid 

in Liliaceae in Amaryllidaceae 

Lili 

Liliineae 

Lili 

Liliineae 

Lili 

Liliineae 

Lili 

Liliineae 

in Liliaceae 

Lili 

Lili 

Lili 

Lili 

Lili 

Lili 

Lili 

Mus 

Liliineae 

Commelinineae 

Commelinineae 

Commelinineae 

Commelinineae 

Commelinineae 

Commelinineae 

in Musaceae 

Mus 

Mu s 

Mus 

Lili 

Lili 

Lili 

Burmanni 

Liliineae 

Liliineae 

Liliineae 

__- 

Orchid 

in Liliaceae 

in Liliaceae 

Lili 

Dioscore 

Dioscore 

Pontederi 

Lili 

Lili 

Bromeli 

Commelin 

Commelin 

Commelin 

Junc 

Commelin 

Scitamin 

Scitamin 

Scitamin 

Scitamin 

Scitamin 

Scitamin 

Lili 

Lili 

Lili 

Burmanni 

Burmanni 

Orchid 
6 SMITHSOMANCONTRIBUTIONS TO BOTANY 

Smilacales; Trichopodaceae and Stenomeridaceae in Dioscoreales; Alstroemeriaceae in Alstroemeriales; Conostylidaceae, 

and Hypoxidaceae in Haemodorales; Heliconiaceae, and 

Costaceae in Zingiberales; Hydatellaceae in Hydatellales; 

Joinvilleaceae, Anarthriaceae, and Ecdeiocoleaceae in Restionales. 

Thorne, 1983. Trichopodaceae in Liliales; Cymodoceaceae 

in Zosterales; Heliconiaceae and Costaceae in Zingiberales. 

Dahlgren et al., 1985. Limnocharitaceae in Alismatales; 

Cymodoceaceae in Najadales; Trichopodaceae and Petermanniaceae in Dioscoreales; Luzuriagaceae, Ruscaceae, Convallariaceae, Asparagaceae, Herreriaceae, Hanguanaceae, Dracaenaceae, Asteliaceae, Nolinaceae, Dasypogonaceae, Blandfordiaceae, Calectasiaceae, Rcophilaeaceae, Ixioliriaceae, Cyanastraceae, Phormiaceae, Doryanthaceae, Eriospermaceae, Asphodelaceae, Anthencaceae, Aphyllanthaceae, Hemerocallidaceae, Funkiaceae, Hyacinthaceae, Alliaceae, and Hypoxidaceae in Asparagales; Colchicaceae, Geosiridaceae, Alstroemeriaceae, Calochortaceae, Uvulariaceae, Apostasiaceae and 

Cypripedtaceae in Liliales; Heliconiaceae and Costaceae in 

Zingiberales; Hydatellaceae in Hydatellales; Joinvilleaceae, 

Ecdeiocoleaceae and Anarthriaceae in Poales; Melanthiaceae 

and Campynemaceae in Melanthiales; Thismiaceae in Burmanniales. 

Hutchinson, 1973. Petrosaviaceae in Alismatales; Ruppiaceae in Potamogetonales; Cartonemataceae in Commelinales; 

Rmphilaeaceae and Ruscaceae in Liliales; Alstroemeriaceae 

and Petermanniaceae in Alstroemeriales; Stenomeridaceae and 

Trichopodaceae in Dioscoreales; Hypoxidaceae and Apostasiaceae in Haemodorales; Thismiaceae in Burmanniales. 

Melchior, 1964. Cyanastraceae, Hypoxidaceae and Geosiridaceae in Liliiflorae. 

Stebbins, 1974. Limnocharitaceae in Alismatales; Ruppiaceae in Najadales; Petrosaviaceae in Triuridales; Heliconiaceae and Costaceae in Zingiberales; Tecophilaeacae and 

Cyanastraceae in Liliales; Geosiridaceae in Orchidales. 

Rouleau, 1981. Limnocharitaceae in Alismatales; Elodeaceae in Hydrocharitales; Ruppiaceae and Cymodoceaceae 

in Najadales; Petrosavhceae, Aphyllanthaceae, Alliaceae, 

Alstroemeriaceae, Hypoxidaceae, Petermanniaceae, 'kcophikaceae, Cyanastraceae, Asparagaxae, Ruscaceae, CroomiaCeae 

and Trichopodaceae in Liliales; Geosiridaceae in Iridales; 

Heliconiaceae and Costaceae in Zingiberales; Apostashceae 

in Orchidales; Cartonemataceae and Abolbodaceae in Commelinales; Anarthriaceae, Ecdeiocoleaceae, Joinvilleaceae and 

Hanguaniaceae in Restionales; Hydatellaceae in Hydatellales; 

Anomochloaceae and Streptochaetaceae in Poales; Nypaceae 

in Arecales. 

Young, 1981. Cymodoceaceae in Zosterales; Cyanastraceae, 

Geosiridaceae, Almeae and Hanguanaceae in Liliales; 

Joinvilleaceae, Ecdeiocoleaceae and Hydatellaceae in Restionales; Heliconiaceae and Costaceae in Zingiberales. 

Benson, 1979. Cyanastraceae in Liliales. 

Emberger, 1960. Ruppiaceae in Potamogetonales; Phytelepha- and Nwm in palm ale^; Cyana~tra~~~, Gmiridaceae and Petermanniaceae in Liliales; Thismiaceae in 

Burmanniales; Apostasiaceae in Orchidales. 

The bibliography notes publications of several authors who 

have attempted to improve the system of classification of the 

monocots 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 (1976-1988) and the 

Kew Record of Taxonomic Literature (Royal Botanic Gardens, 

Kew, 1974-1988) can also be consulted for relevant literature 

and Stapf (1929- 193 1) for illustrations. 

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. 

Monocots have been extant nearly as long as dicots, have 

been subjected to many of the Same ecological pressures, and 

consequently show similar evolutionary trends. Like the dicots, 

I believe the earliest monocots had wind pollinated unisexual 

flowers, monosulcate pollen, and lacked vessels. Stewart 

(1983) and Muller (1981) can be consulted for a summary of 

the contribution of paleobotany to our understanding of the 

evolution of the angiosperms. 

My approach to understanding evolutionary trends in 

characters is to relate them to the 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 

developed landing platforms for insects. Some became fragrant 

or malodorous indicating the presence of food to potential 

pollinators. Some produced nectar which 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 and some flowers became 

specially adapted to the most proficient pollinators, the bees. 

The primitive insect pollinated monmts 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. 

The monmts probably originated under warm temperate 

or subtropical 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 habitats. 

An entirely corolloid perianth was supplanted by one in 

which one series became an herbaceous calyx which provides 
NUMBER 7 1 7 

greater protection of the reproductive organs against unfavorable climatic conditions, as well as against 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 which 

enable them to resist moisture loss and prevent collapse. Also, 

in the xylem, tracheids were converted to vessels, scalariform 

perforation plates became simple and the vessels wider to 

facilitate rapid movement of water which is only periodically 

available under some 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 wcurrence among the 

families. The fewer the families showing a particular primitive 

state the more primitive the state. 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 85 

characters. I also indicate the extent to which I consider the 

states reversible. In general, character states which are most 

constant within families are least readily reversible. For 

example flower color is usually variable within families and 

commonly reversible, whereas connation of parts is much 

more constant within families and therefore difficult to reverse. 

The category “n” includes those character states which are 

Orchidales 

lridales 

7 

Zingiberales 

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 in attempting to determine 

relationships. 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 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 family should be used 

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. Collateral relatives differ 

from one another by having at least one different character from 

one another that is not in the main line of descent. Genera 

which 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 (below) suggesting 

relationships of the orders. In this treatise and in my 

Cyperales 

- Naiadales 

Suggested phylogenetic relationships 

of the orders r,f monocotyledons. 
8 

Character 

SMITHSONIANCONTRIBUTIONS TO BOTANY 

F’rimitive state Derived state Reversibiiity 

climate of origin 

Plant 

indument 

spines 

tendrils 

latex 

xylem vessels 

perforations plates 

xylem vessel and sieve tube elements 

leaves 

leaf margin 

leaf venation 

stcanates 

inflorescence 

flowers 

pollinated by 

pollinators 

involucre 

recepacle 

perianth 

perianth spur 

perianth parts 

corona 

perianth vasculature 

perianth parts. stamens, and carpels 

stamens and carpels 

nectaries 

stamens 

StaminodeS 

warm temperate or subtropical 

monoeciws or dioecious 

autotrophic 

absent 

simple 

absent 

absent 

absent 

absent 

mesophyte 

pered 

scalariform 

long, namw, angular in cross-section 

end plate slanting 

pores small 

alternate 

in more than 2 ranks 

thin 

medium 

entire 

F 

apparently parallel 

racemose 

raceme 

spike or umbel 

open cyme 

in inflorescence 

terminal 

unisexual 

OdorleSS 

green, white, or yellow 

actinomorphic 

wind 

various pollinators 

anOmOCytiC 

Cdcoptera 

short-tonwed 

absent 

convex 

flat 

monochlamydeow 

homoiochlamydews 

absent 

imbricate 

absent 

evident 

indefinite in number 

separate 

pleiomerous 

2 whorls 

isanew 

absent 

spirauy arranged 

equal to pistil in length 

equal in length 

inserted 011 receptade 

absent 

other 

hermaphroditic 

allllUal 

present 

glandular 

present 

present 

present 

present 

xerophyte 

SaPrOphfiC 

simple 

short, wide, round in cross-section 

end plate transverse 

pores large 

opposite or whorled 

distichs 

thick and sclerophyllous 

small relative to size of plant 

lobedortoothed 

closed 

obviously reticulate or pinnate 

other 

cymose 

corymb or spike 

head 

fascicle 

solitary 

dary 

bisexual 

SCented 

other cdors 

insects or birds 

particular pollinators 

Hymmoptera, Diptera, or Lepidoptera 

long-tongued 

present 

flat 

cmcave 

achlunydeous or dichlamydeous 

hetemdhnydeous 

present 

valvate 

present 

absent or obscure 

definite in number 

whorled 

connate or adnate 

isanew 

1 whorl 

oligomerous 

present 

shorter than pistil 

UliSOoStemOnouci 

adnate to perianth or epigynws 

prescnt 

Ygomorphic 

r 

r 

r 

r 

n 

r 

r 

r 

r 

r 

n (r in some 

reduced 

aquatics 1) 

n 

n 

n 

n 

r 

r 

r 

r 

r 

n 

n 

n 

n 

n 

n 

n 

r? 

C 

C 

r? 

n 

r 

r 

r 

n 

n 

n 

n 

r 

r 

r 

n 

n 

n 

r 

n 

n 

n 

n 

n 

r 

r 

r 

n 

r 

C 
NUMBER 7 1 

TABU 2.-tinued. 

9 

Character 

anthers 

connective 

pollen grains 

micrOSporangia 

cells in pollen grains at time of dispersal 

ovary 

style(s) 

carpels 

stigma(s) 

placentas in more than l-carpellate pistil 

ovules 

fruit 

dehiscence of capsule 

Seed 

embryo 

endosperm 

Primitive state Derived state Reversibility 

separate 

adnate or basifixed 

oblong 

dehiscent by slits 

exmrse or latrorse 

abundant 

4 per anther 

monaulcate 

not sticky 

free 

2 

open 

number of locules 

superia 

absent 

separate 

decurrent along ventral surface of style 

lobed 

axile 

axile 

not enlarged 

few 

crassinucellar 

bitegmic 

follicle 

ventricidal or sepicidal 

medium-sized 

small 

about as long as wide 

to number of carpels 

straight 

CopiOuS 

connate 

dorsifixed 

about as long as wide 

poricidal 

intme 

sparse or absent 

2 or 1 per anther 

pored or nonaperturate 

in tetrads or pollinia 

3 

closed 

unilocular in more than 1 -carpellate pistil 

semi-inferior to inferior 

present 

connate 

apical or style stigmatic all around 

not lobed 

parietal 

apical or basal 

enlarged and fleshy 

tenuinucellar 

unitegmic or ategmic 

other types 

loculicidal or by slits 

small or large 

curved 

much longer than wide 

scanty a absent 

sticky 

or 1 when carpels more than 1 

large 

r 

r 

r 

n 

r 

n 

n 

n 

r 

n 

n 

n 

r 

n 

r 

n 

n 

n 

n 

n 

r 

r 

n 

n 

n 

n 

r 

r 

n 

n 

n 

comparable publication on the dicotyledons (Goldberg, 1986) 

I have laid the foundation for a third volume which will be an 

elaboration of my philosophy concerning the subjects of these 

treatises and its application by giving reasons for the 

composition and placement of each taxon. It will also contain 

a synoptical key to the taxa. 

Some of my colleagues have reviewed all or part of the 

monocotyledon manuscript. I appreciate their suggestions for 

its improvement. 

Ordinal and Family Descriptions 

PANDANAUS 

The order is monotypic. 

Chemistry: Does not indicate strong relationship with any 

other order; monoterpenes are present; saponins may be absent; 

cyanogenesis is rare if present. 

PANDANACEAE (Figure I).-Trees or shrubs, often branched, 

or perennial herbs, sometimes lianas, often with aerial or pp 

roots, usually coastal or in marshes, sometimes in forests; 

raphides present, silica absent; xylem vessel perforation plates 

scalariform in all organs (except vessels absent from the stem 

of Sururungu); leaves cauline or radical, spirally arranged 

along the stem, often clustered at the apex, rarely in 2 or 4 

rows, simple, linear, sheathing, parallel-veined, leathery, the 

margins and midrib beneath spinulose, epidermal cells in 

longitudinal files, the stornates tetracytic, mostly on the lower 

surface; inflorescences terminal spadices with spathes or heads 

or panicle, or the flowers few, the bracts sometimes brightly 

colored; flowers minute, the plants dioecious, anemophilous 

or entomophilous, ornithogamous and chiropterogamous in 

Freycinetiu; perianth absent or vestigial (in dispute), short and 

gamophyllous (Sururungu); 8 flowers: stamens few to 

numerous, 9--300, variable in number, spirally arranged on a 

short or long axis, or at the Same level, the filaments free or 

connate, anthers basifixed, sometimes apiculate; pollen monoporate, 2-celled when shed; pistillode minute or absent; 9 

flowers: pistil 1 free or many connate in clusters, carpels 

1--80, the stigmas sessile or subsessile, free or connate, 

sometimes a canal leads to the ovary, the ovary superior, septal 
10 SMITHSON" CONTRIBUTIONS TO BOTANY 

a 

C 

FWUW ~.-PANDANACBAE: a, Pandanw candelabrum habit, 1.1. of 9 inflorescence, P. utilis fruiting spadix 

(reduced); b, F! candehbnun stamens, P. utilis 9 flower, 1.s. of same, cs. and 1.s. of seed, embryo; c, Freycinctia 

banksii 1.s. of a group of stamens enclosing the abortive pistil, pistil and staminodes, F. imbricafa C.S. of fruit, 

seed, 1.8. of same (after Baillon, Le Maout and Decaisne). 

nectaries reported but not confirmed, 1-many locular, the 

ovules 1 and sub-basal, or numerous and parietal, bitegmic, 

crassinucellar, anatropous, the embryo sac sometimes with 

increase in number of antipodals after fertilization, sometimes 

migration of nucellus nuclei into the embryo sac, endosperm 

formation Nuclear, later becoming cellular, staminodes absent 

or small, hypogynous or adnate to the ovary; fruits a syncarp 

or free, drupes or berries, the pericarp often with woody fibers, 
NUMBER 71 11 

the seeds 1-many, minute or medium-sized, sometimes with 

a little perisperm; embryo minute, basal, 0.1 the length of the 

endosperm, 2.5 times longer than wide; endosperm copious, 

fleshy, oily and with aleurone. Chromosomes: x = 27,30,3 1,32, 

especially 30. 

Composition: 3 genera, -500 species. 

Distribution: Primarily tropical, mostly in coastal and 

marshy habitats; tropical West Africa, India, Malaya, East 

Indies, northern Australia, New Zealand. 

hISMAT,ALES 

Herbs of wet habitats, sometimes partly or entirely 

submerged, rooted in the substrate or free-floating, often 

rhizomatous or stoloniferous, in fresh water or marine; raphides 

rare or absenr xylem vesselless throughout, or xylem vessel 

perforation plates simple and scalariform in the root; leaves 

linear or differentiated into petiole and blade with longitudinal 

and transverse veins, radical, alternate, opposite or whorled; 

inflorescence scapose or axillary, racemes, panicle, or umbels, 

often cymose, or the flower solitary; flowers entomogamous, 

hydrogamous, or autogamous, rarely anemophilous, bisexual 

or the plants polygamous, monoecious or dioecious, the parts 

hypogynous, and the torus flat to globose, or epigynous; 

perianth segments usually 6, free, sepals 3(2), green, rarely 

petaloid; petals 3(0), petaloid; stamens q-40-l), free, or rarely 

connate, or some staminodal, the anthers basifixed or 

dorsifixed, introrse or extrorse; pollen 1-sulcate, forate or 

nonaperturate, sometimes connate in short threads or in tetrads, 

3(2)-celled when shed; pistils to -100 to 1, spirally arranged 

or whorled, carpels 3-6(2-15) when connate; ovary superior 

or inferior, unilocular, the placentas rarely meeting in the 

center, the ovules many to 2, often scattered over the ovary 

wall, sometimes basal, bitegmic, crassinucellar, rarely pseudocrassinucellar, anatropous or amphitropous, rarely orrhotropous; embryo sac with 3-1 antipodal cells; endosperm 

formation Helobial; styles as many as the carpels, free or 

shortly connate, entire or 2-3-branched apically, the stigmas 

sometimes ventrally decurrenc fruit follicles, achenes or a 

berry; seeds often numerous, the embryo straight or curved, 

endosperm absent at maturity. 

Distribution: Cosmopolitan. 

Chemistry: Poorly known; cyanogenesis, tannin production, and mucilage are rare or absent; the unusual carbohydrate 

apiose is present in some members of some families, but it also 

occurs in some members of Potamogetonales and in the 

Lemnaceae; raffmose and stachyose have been reported from 

this order and Aponogetonaceae. 

ALISMATACEAE (Figure 2a-c).-Perennial, rarely annual, 

herbs of wet habitats, erect or rarely with floating leaves or 

entirely submerged, rhizomatous, latex present, raphides in 

Sagitturiu, crystals present; hairs generally absent, simple, 

unicellular or branched multicellular in a few members; xylem 

vessel perforation plates simple and scalariform in the root, 

vessels absent from the stem and leaves; leaves with an open 

sheathing petiole and blade, the latter strongly nerved, the 

venation longitudinal and transverse, the stomates paracytic 

and tetracytic, intravaginal scales present; inflorescence 

scapose, primary branches racemes or panicle, secondary 

branches often cymose, often whorls of pedicellate flowers, 

rarely the flowers subsolitary; flowers entomogamous or 

autogamous, bisexual or the plants polygamous, monoecious 

or rarely dioecious, the parts hypogynous, torus flat to globose, 

nectaries on the carpels, receptacular, or on the perigone 

(Echinodorus); sepals 3, green; petals 3(0), white or color&, 

stamens 6(3--4O), free, whorled or spirally arranged; anthers 

basifixed or dorsifmed, extrorse; pollen forate, 3-celled when 

shed; pistils to -100, spirally arranged in a head, or 6(3) in a 

single whorl, sometimes basally connate or adnate to the floral 

axis, the style terminal or basal, the stigma ventrally decurrent; 

ovules 2-00, basal or scattered over the wall of the pistils, 

bitegmic, weakly pseudocrassinucellar, anatropous, or amphitropous, the embryo sac with 1 antipodal cell, rarely 2-3; 

endosperm formation Helobial, in the large micropylar 

chamber many free nuclei are formed before the tissue becomes 

cellular; fruit achenes or follicles; seeds curved, with 

hippocrepiform embryo, endosperm absent at maturity. Chromosomes: x = 5-13, especially 7,11, large in at least some 

species. 

Composition: -15 genera, -80 species. 

Distribution: Cosmopolitan, but mostly America. 

BUTOMACEAE (Figure 2d,e).-Glabrous perennial herbs of 

wet habitats, rhizomatous, secretory canals and raphides 

absent; xylem vessel perforation plates simple and scalariform 

in the root, vessels absent from the stem and leaves; leaves 

radical, linear, not differentiated into blade and petiole, the 

stomates mostly paracytic, intravaginal scales numerous; 

inflorescence a scapose, involucrate, cymose umbel; flowers 

bisexual, entomogamous, protandrous, the parts hypogynous; 

perianth segments 6, biseriate, free, petaloid, pink, persistenr 

stamens 9, the filaments elongate, the anthers basifixed, 

introrse; pollen monosulcate, 3-celled when shed; pistils 6, the 

lower lateral surfaces nectariferous, style elongate, the stigma 

slightly decurrent ventrally, wet; ovules numerous, scattered 

over the wall of the ovary, bitegmic, pseudocrassinucellar, 

anatropous; fruit follicles, the seeds numerous, minute, 

elongate, striate; embryo straight; endosperm absent. Chromosomes: x = 8,10,11,12,13,15, especially 13, small. 

Composition: 1 genus, 1 species. 

Distribution: Europe and temperate western Asia. 

HYDROCHARITACEAE (Figure 3).-Freshwater and marine 

perennial, rarely annual, herbs, partly or entirely submerged, 

free-floating or rooted in the substrate, often stoloniferous; 

raphides and crystals absent; scalariform perforation plates in 

the roots of a few genera, the others vesselless throughout; 

leaves radical or alternate, opposite or whorled on an elongate 

stem, linear, ribbon- or strap-shaped or ensiform and submerged, or differentiated into blade and petiole, with 
12 SMFHWNIAN CONTRIBUTIONS TO BOTANY 

b 

C 

d 

e 

RGURE 2-ALISMATACEAE: 4, Hydrocleys commersonii one of the pistils, same opened to show the parietal 

placentation, H. murtii stamens, pistils, LLMocharis plwnieri seed, 1s. of same, H. commersonii plant in flower. 

b, Alisrm plantago-quatica 1.s. of flower, fruit, 1.5. of same; c, Dumosonim srellarvm floral diagram, flower, 

1.s. of same, fruit (after Baillon, Lindey. Martius). BUTOMACME: d, Bufom umbellatlcs 1.s. of flower, stamen, 

C.S. of same, pistils, inflorescence; e, dehisced fruit, seed, 1.5. of same, floral diagram (after L Maout and 

Decaisne. Bdon). 
NUMBER 71 13 

U 

C 

Y 

FIGURE 3.-HYDROCHAIUTACEAE: a, Hydrocharis morsus-raMe 8 flower, 1.s. of same, diagram of (Sflower, 

androecium, outer stamen, inner stamen with appendage; b, 1.s. of 9 flower, style opposite the sepals, style 

opposite the petals, cs. of ovary, floraldiagram of 9 flower, c, fruit, dehisced fruit, 1.s. of seed, views of embryo, 

Vallisneria spiralis (Sinflorescence (after Baillon. Le Maout and Decaisne). 

longitudinal and cross veins, and floating or emersed, entire, 

rarely toothed (Stratiom), the stornates mostly paracytic, often 

absent; often with prickle hairs, intravaginal scales present; 

inflorescence axillary, the 8 usually a cymose umbel or 

cincinnus enclosed in a spathe, the P usually solitary; flowers 

entomogamous or hydrogamous, sometimes with a nectariferous disk (Hydrocharis), rarely anemophilous (Limnobium), 

rarely bisexual, usually the plants dioecious, monoecious or 

polygamous, vestige of the opposite sex sometimes present; 

perianth rarely homochlamydeous, the sepals 3(2), free, green, 

valvate; petals 3(0), smaller or larger than the sepals, 

sometimes with a basal gland; stamens 12-1, free or 

sometimes connate, sometimes spirally arranged on an elongate 

receptacle, sometimes 3 staminodes present, the latter sometimes nectariferous (?); pollen 1-sulcate or nonaperturate, 

sometimes connate in short threads, occasionally in tetrads, 

3(2)-celled when shed; pistil 1, the carpels 3-6(2-15), styles 

as many as the carpels, often bilobed, sometimes shortly 

connate at the base, the stigmas elongate, papillose; ovary 

inferior, unilocular, rarely the placentas meet in the center of 

the ovary, the ovules usually many, sometimes few, parietal, 

sometimes appearing to be scattered over the surface, bitegmic, 

crassinucellar, usually anatropous, rarely orthotropous; embryo 

sac antipodal cells 3, persisting into early embryogeny; 

endosperm formation Helobial, division free-nuclear in the 

micropylar chamber, later becoming cellular; fruit elongate or 

globose, a berry or dry, sometimes rupturing irregularly, rarely 

regularly; seeds usually numerous, the embryo large, straight, 

radicle thick, plumule usually evident, endosperm absent at 

maturity (scanty in Otteliu). Chromosomes: x = 6-12, 

especially 6-8,ll. 

Composition: 15 genera, -90 species. 

Distribution: Cosmopolitan, but mostly tropical and absent from the arctic region. 
14 SMlTHSONIANCONTRIBUTIONS TO BOTANY 

TRIURIDALES 

The order is monotypic. 

Chemistry: Unknown. 

TRIURIDACEAE (Figure 4u,b).- Saprophytic reddish, purplish or cream herbs, sometimes rhizomatous, with a few 

alternate scales, without chlorophyll; xylem vesselless, the 

tracheary elements reduced; inflorescence a terminal bracteate 

raceme or corymb; plants monoecious, dioecious or rarely 

polygamous; flowers very small, actinomorphic, the parts 

hypogynous; perianth segments 6(3-8) in one series, sometimes acuminate, with apical uniseriate hairs, corolline, valvate, 

sometimes basally connate; 8 flowers: stamens 3 or 6(1-6), 

sometimes 3 of them staminodal, the filaments short, basally 

connate, or the anthers sessile; anthers extrorse, sometimes 

produced apically; pollen nonaperturate, 3-celled when shed, 

P flowers: pistils -15(6-50), the style terminal to basal, stigma 

decurrent or penicillate; ovule solitary, basal-mile, bitegmic, 

tenuinucellar, anampous; embryo sac antipodal cells 3, 

ephemeral; endosperm formation Nuclear, the tissue later 

becoming cellular; fruits dehisce dorsally or nutlets; embryo 

undifferentiated, endosperm present, containing aleurone and 

fat (or starch ?). Chromosomes: x = 11,12,14. 

Composition: 7 genera, -80 species. 

Distribution: Pantropical in rain forest; southern Africa. 

JUNCAGINALES 

Marsh herbs, rhizomatous or tuberous; xylem vessel 

perforation plates scalariform in the root, vessels absent from 

the stem and leaves; leaves usually basal, linear, with an open 

sheath; inflorescence a scapose raceme or spike; flowers small, 

bisexual or the plant dioecious or monoecious, anemophilous; 

perianth segments qO), biseriate, homochlamydeous, greenish; 

stamens 6 (rarely more), 3, or 1, the filaments short, anthers 

exmrse, sometimes apiculate; pollen nonaperturate, sometimes in dyads, 2-3-celled when shed; pistil 1(6), carpels 

3-6(1), the stigmas usually sessile or subsessile, the style 

rarely elongate; ovary superior, 3-locular when the carpels are 

united, the ovules 1-2(few) per carpel, basal, bitegmic, 

crassinucellar, anatropous; embryo sac antipodal cells ephemeral or undergo secondary multiplication; endosperm formation 

Helobial or Nuclear; fruit follicles, schizocarp, or nutlet; 

embryo straight, endosperm absent in mature seed 

Distribution: Cool temperate and cold Northern Hemisphere; southern tip of South America; southern Australia, New 

Zealand. 

Chemistry: Cyanogenic; Scheuchzerkxae tanniferous, Juncaginaceae not. 

SCHEUCHZERIACEAE (Figure 4c,d).-Perennial marsh herbs 

with a jointed rhizome; raphides absent, some calcium oxalate 

crystals present; xylem vessel perforation plates scalariform 

in the root, vessels absent from the stem and leaves; leaves 

often basal, linear, petiole ligulate at junction with the blade, 

the stomates tetwytic; numemus long, uniseriate hairs in axils 

of leaves; inflorescence a terminal, bracteate, few-flowered 

raceme; flowers bisexual, small, anemogamous, protogynous; 

perianth segments 6, in 2 whorls, free, similar, yellowish-green; 

stamens 6 (rarely more), free, the filaments short; anthers 

linear, basifuted, apiculate, extrorse; pollen nonaperturate, in 

dyads, 3-celled when shed, pistil 1, carpels 3(-6), shortly 

ventrally connate at the base, the stigmas sessile, papillose; 

ovary superior, the ovules 2 (few) per carpel, basal, bitegmic, 

crassinucellar, anampous; embryo sac antipodal nuclei 3, 

ephemeral; endosperm formation Helobial, at first nuclear, later 

becoming cellular in the micropylar chamber; fruit divaricate 

follicles; embryo straight, endosperm absent in mature seed. 

Chromosomes: n = 11. 

Composition: 1 genus, 2 species. 

Distribution: Cool temperate and cold Northern Hemisphere. 

JUNCAGINACEAE (Figure Sa,b).-Perennial, seldom annual, 

glabrous marsh herbs, rarely plant submerged except for one 

or two floating leaves, rhizomatous or sometimes tuberous; 

raphides absent, calcium oxalate crystals sometimes present; 

xylem vessel perforation plates scalariform in the root, vessels 

absent from the stem and leaves; leaves radical, linear, 

sheathing basally, the stomates tetracytic, intravaginal scales 

present; inflorescence scapose, an ebracteate spike; flowers 

small, anemophilous, protogynous, bisexual or the plant 

dioecious or monoecious; perianth segments 6, biseriate, 

homochlamydeous; stamens 6, 3, (4), free, the anthers 

subsessile, exmrse; pollen nonaperturate, 2- or 3- celled when 

shed, pistils 1(6), carpels 3-6, sometimes 3 aborted, the 

stigmas sessile or subsessile, papillose or plumose; ovary 

superior, 3(4)-locular in compound ovary, the ovules 1 per 

locule, basal, bitegmic, crassinucellar, anatropous; the 3 

embryo sac antipodal cells in Triglochin undergo secondary 

multiplication and persist into early embryogeny ; endosperm 

formation Nuclear and small in amount; fruit a schizocarp ur 

nutlet; embryo straight, endosperm absent in mature seed. 

Chromosomes: x = 6,8,9. 

Composition: 3 genera, -20 species. 

Distribution: Cool temperate and cold regions of Northern 

Hemisphere; southern tip of South America; southern Australia, New Zealand. 

LILAEACEAE (Figure Sc,d).-Aquatic or marsh annual, 

glabrous herbs with short rhizomes; xylem vessel perforation 

plates only in the roots; leaves basal, alternate, linear and Wte, 

with an open sheath; inflorescence scapose, bracteate, dense 

spikes, either unisexual, or 8 above, bisexual in the middle 

and 3 below; solitary flowers also occur in the axils of the 

leaves; perianth absent; stamen 1, the filament short, anther 

exmrse, about as long as wide, the sacs separate basally, the 

connective slightly produced apically; pollen nonaperturate, 

2-celled when shed; pistil 1, unicarpellate, the stigma 

subsessile in the bisexual and upper P flowers, the style long, 

filiform, with a capitate, papillose stigma in the basal axillary 

flowers; ovary superior, unilocular, uniovulate, the ovule basal, 
NUMBER 71 15 

b 

d 

FIOURB 4.-TRNRlDACEAE: a, Sciaphila albcscerrp plant in flower, 8 flower, same with perianth removed. 

flower, i pistil; b, Triwis brevistilis fruit, same opened to show the solitary led, 1s. of same showing thc 

mhute UndiffemSialed anbryo in abundant endosperm (&a Wdus). sCHEUCHZJ3RIACEAE: c, Scheuchzcria 

palwfris flower, stamm, a carpel same laid open, plant in flower. d, fruit, aecd (after Hutchinson). 
16 SMITHSONIANCONTRIBUS TO BOTANY 

a 

b 

C 

R 

d 

FIGURE 5.-JUNCACtINACEAE: a, Triglochin marithum pan of inflorescence, flower, Is. of same, T. lax@wvm 

carpel, I.s. of sme, Triglochin floral diagram; b, T. morilbnum fruit, segment of sme, I.s. of segment, T. 

palustre cs. of fruit (after Baillm, Le Maout and Decaisne). ULAEACEAE: c, Liloeo subulata upper inflorescence, 

a flower, views of stamen and perianth (bract), 9 flower from base of plans d, pistil, 1.8. of same, views of 

fruit, C.S. of same, views of embryo (after Martius). 
NUMBER 71 17 

bitegmic, crassinucellar, anatrops; the 3 antipodals of the 

embryo sac degenerate during early embryogeny; endosperm 

formation Nuclear; fruit a compressed, longitudinally ribbed 

nutlet; embryo straight, endosperm absent in mature seed. 

Chromosomes: n = 6. 

Composition: 1 genus, 1 species. 

Distribution: Western America from southern Canada to 

southern South America. 

NAJALMLE 

Freshwater or marine rhizomatous or tuberous submerged 

or emergent herbs: xylem usually vesselless, sometimes 

scalariform perforation plates in the root: leaves distichous or 

basal, alternate or opposite, the blade linear to orbicular, the 

veins longitudinal, sometimes also transverse, the sheath open, 

stomates absent; inflorescence spikes, terminal or axillary, or 

flowers solitary or few in axillary cymes; flowers small or 

minute, bisexual or the plants monoecious or dioecious, the 

parts hypogynous, anemogamous or hydrogamous; perianth 

absent or of 3-4 scales; stamens 4-1(-12), the filament 

elongate or the anthers sessile, often extrorse, sometimes 

apically produced; pollen monocolpate or nonaperturate, 

globose or filiform, 2-3-celled when shed; pistils 3-4(1-9), 

the styles 1-4, elongate, or the stigmas sessile, sometimes 

decurrent ventrally; ovary unilocular, the ovules 1-8, basal, 

apical or ventral, bitegmic, crassinucellar or pseudocrassinucellar, anatrops, orthotropous or campylotropous; embryo sac 

antipodals ephemeral or undergoing secondary multiplication 

to form 4-5 cells; endosperm formation Helobial or Nuclear: 

fruit usually nutlets, achenes or drupelets, sometimes follicuk, 

embryo straight or curved, endosperm absent in mature seed. 

Distribution: Cosmopolitan. 

Chemistry: No distinctive characters have been uncovered. 

APONOGET~NACEAE (Figure 6u-c).-Freshwater perennial 

herbs with tuberous rhizome; raphides absent, calcium oxalate 

crystals present; xylem vesselless; leaves radical, usually 

floating, sometimes submerged, long petiolate, the blade 

elliptic to linear, with some longitudinal and numerous 

transverse veins, the stomates paracytic, intravaginal scales 

present, otherwise glabrous; inflorescence a scapose ebracteate 

spike, sometimes branched, flowers small, bisexual or rarely 

the plants dioecious, the parts hypogymus; perianth parts 0-3, 

petaloid or bract-like, abaxial when 2; stamens 6-12, in 2-4 

cycles, the filaments elongate, anthers small, extrorse; pollen 

monocolpate, 3-celled when shed; pistils 3-6, nectariferous, 

the style grading into the ovary, the stigmas somewhat 

decurrent ventrally; ovules 2-8, basal, bitegmic, the integuments free or f connate, crassinucellar, anatrops; embryo 

sac with 3 antipodals which persist into early embryogeny; 

endosperm formation Helobial; fruit 3-6 follicles, the seeds 

with straight embryo, endosperm absent at maturity. Chromosome~: x = 8,12,13,24,26,38,46, especially 8. 

Composition: 1 genus, -25 species. 

Distribution: Centered in tropical Africa and Madagascar; 

southern 2/3 of Africa: India, southeast Asia, few islands of the 

East Indies; northern Australia. 

ZOSTERACEAE (Figure 6d).-Submerged marine perennial, 

glabrous herbs on rocks, with short creeping rhizome, 

sometimes tuberous, and erect branches; crystals absent; xylem 

vesselless, the tracheary elements reduced; leaves alternate, 

linear, sometimes minutely denticulate, the apex of the sheath 

with 2 lateral projections, stomates absent, intravaginal scales 

present; inflorescence on one side of a flattened spadix with a 

row of bracts on each margin, at fist enveloped by a leaf 

sheath, the plants monoecious or dioecious; perianth absent; 

8 flowers consist of a single, sessile, dorsifixed, elongate 

anther, pollen filamentous, about 2500 pn long, 4 pn wide, 

2-celled when shed, 3 flowers consist of 1 pistil and a vestigial 

anther, the stigmas 2, large, flat; ovary unilocular with 1 

pendulous, bitegmic, pseudocrassinucellar, orthotrops ovule; 

the 3 antipodal cells of the embryo sac persist into early 

embryogeny; endosperm formation Nuclear; fruit a nutlet, 

achene or drupelet, the seed longitudinally ribbed, endosperm 

absent at maturity. Chromosomes: x = 6,7,8,10. 

Composition: 3 genera, 18 species. 

Distribution: Mostly temperate, few tropical or arctic; 

North America; Eurasia; western and southern Australia, New 

Zealand; eastern Africa, Madagascar. 

P~TAMOGETONACEAE (Figure 7a-c).-Aquatic perennial, 

glabrous herbs with creeping rhizome and erect stems; xylem 

vessel perforation plates sometimes scalariform in the root, 

vessels absent from the stem and leaves: leaves distichous, 

alternate or opposite, immersed or floating, ribbon-like, linear, 

oblong, lanceolate, ovate or orbicular, entire or rarely sermlate, 

sometimes differentiated into blade and petiole, the stomates 

paracytic, intravaginal scales present: inflorescences axillary 

spikes: flowers small, bisexual, the parts hypogymus, anemogamous or hydrogamous; sepals 4, valvate, free, or 0 (Ruppia); 

petals 0; stamens 4, sessile, adnate to claw of sepal, or 2 

(Ruppia), the anthers extrorse; pollen nonaperturate, round 

(Potamgeton) or tubular (Ruppiu), 3-celled when shed; pistils 

4(1, rarely more than 4, to 10 in Ruppiu), the stigma sessile 

or subsessile, slightly decurrent ventrally, the ovule 1, 

bitegmic, crassinucellar, orthotropous or campylotropous; 

embryo sac antipodal cells 3, soon degenerate; endosperm 

formation Helobial, free-nuclear division occurs in the 

micropylar chamber, but later the tissue becomes cellular, fruit 

achenes, nutlets or drupelets; embryo curved, endosperm 

absent in mature seed. Chromosomes: x = 8,10,13-15, 

especially 13,lO. 

Composition: 2 genera, -100 species. 

Distribution: Cosmopolitan, in fresh or somewhat brackish water. 

ZANNICHELLIACEAE (Figure 7d).-Submerged, slender, branching perennial glabrous herbs of fresh to salt water, with a 

creeping slender rhizome: raphides absent (?), crystals absent; 

xylem vesselless, the tracheary elements reduced: leaves linear, 
18 

a 

d 

SMlTHSONIANCONTRIBUTIONS TO BOTANY 

b 

FIc3vRe 6.-AwplroaETONACEAE: a, Aponosclon fenesfrdu inflorescmce and leaf, A. diptochyvs part of pht 

and inflorecrcena, part of infloresmce, Aponogcton flowet, 6, A. bernieriunvr flower, piatil laid open; c, A. 

dirlochyuc 1.s. of reed, geminating embryo (after Le Macut and Decaisne. Bailla). ZOSTERACBAB: d, Zostera 

tmrh anther, the same dehiscing, pistil, 1.s. of same, fruit, I.s. of same, sed. embryo, inflorescence and part 

of spathe (after Bdon). 
NUMBER 71 19 

FIGURE 7.-mOQEToNACEAE: a, POramOgetOn crkpvs flowering branch, flower, 1.5. of same, P. 

slenmtachys flower, views of stamen and puianth segmen; b, gynoecium. 1.s. of a pistil. fruit, 1.s. and C.S. of 

same; c, Ryppio maritima infloresancea, anthers before and after dehiscma, pollen grain, pistil, 1s. of same, 

fruit, 1.5. of same, embryo (after Madus, Baillm). ZA"ICHEUCEAB: d, Zonnichellia palvsrris intlorescmce, 

stamen and pollen grain, 1.s. of pistil, fdtlets, 1,s. of a fruitlet showing the folded embryo (after Martius). 
20 SMITHSONIANCONTRJBUTONS To BOTANY 

alternate, opposite or whorled, sometimes minutely denticulate, 

with a short sheath and mostly ligulate, stomates absent, 

intravaginal scales filiform; plants monoecious or dioecious; 

flowers minute, solitary or few in cymes in the leaf axils; 

perianth of 3 free scales or cupular (Zannichellia) or absent; 

stamens 3-1, rarely connate, the filament elongate or the 

anthers sessile; anthers elongate, basifixed, the connective 

sometimes apically produced; pollen nonaperturate, globose 

(Zannichellia) or filiform (Cymodoceae) and 2000 pm by 10 

p, 2-3-celled when shed; pistils 3-4(1-9), the style 1, 

mostly elongate, with a capitate, peltate or ~mbonate stigma, 

or 2-4-lobed and the stigmas fiiform; ovary superior, 

unilocular, the ovule solitary, apical, bitegmic, crassinucellar, 

anatropous; the 3 antipodals of the embryo sac may undergo 

secondary multiplication to form 4 or 5 cells; endosperm 

formation Helobial; fruits sessile or stipitate nutlets or achenes; 

embryo coiled, endosperm absent in mature seed. Chromosomes: x = 6-8. 

Composition: -6 genera, -20 species. 

Distribution: Cosmopolitan. 

NAJADACEAE (Figure 8a,b).-Small annual, submerged, 

glabrous herbs of fresh or brackish water, the stem filiform, 

much branched; raphides and crystals absent; xylem vesselless, 

the tracheary elements reduced; leaves linear, usually toothed, 

with a short sheath, sub-opposite or verticillate; plants 

monoecious or rarely dioecious; flowers minute, 1-2 sessile 

in the axils of leaves at base of branches; 8 flower consists of 

a single 4(-l)-locular sessile anther enclosed in a bilabiate 

close-fitting perianth and a spathe, the pedicel elongating prior 

to anther dehiscence; pollen ellipsoidal, nonaperturate, 3-celled 

when shed; ? flower naked or surrounded by a membranous 

perianth or spathe; pistil 1, the styles 3(2), stigmas 3(2), ventral, 

elongate; ovary unilocular with 1 basal, bitegmic, crassinucellar, anatropous ovule; the 3 antipodal cells of the embryo sac 

persist into early embryogeny; endosperm formation Nuclear, 

later becoming cellular throughout; fruit a nutlet, the seed 

smooth or sculptured; embryo straight, endosperm absent in 

mature seed. Chromosomes: x = 6,7, especially 6. 

Composition: 1 genus, -35 species. 

Distribution: Cosmopolitan except the arctic region. 

POSIDONIACEAE (Figure 8c).-Submerged marine, perennial, glabrous herbs on rocks, with a short thick rhizome; xylem 

vesselless, the tracheary elements reduced; crystals absent; 

leaves subbasal, alternate, liguliform, entire or serrulate, 

rounded at the apex, the sheath open and ligulate, the persistent 

fibrous leaf-bases cover the stem, stomates absent, intravaginal 

scales present; inflorescences spikes in mils of foliaceous 

bracts at the apex of a scape, the lower flowers bisexual, the 

upper generally 8; perianth absent or of 3 caducous scales; 

stamens 3(4), hypogynous, the anthers large, sessile, extrorse, 

the anther sacs separated by connective which is also apically 

produced; pollen filiform; pistil 1, carpel 1, the stigma sessile, 

fimbriate; ovary superior, unilocular, the ovule 1(2), ventral; 

fruit a drupe; embryo straight; endosperm absent. Chromosomes: n = 10. 

Composition: 1 genus, 3 species. 

Distribution: Southern and western Australia; Mediterranean. 

TYPHALES 

Herbs of wet habitats, rhizomatous; raphides commonly 

present; xylem vessel perforation plates scalariform; leaves 

alternate, linear to strap-shaped, distichous; plants monoecious; 

inflorescences usually dense, scapose, the flowers numerous, 

small or minute, anemophilous; perianth absent or 3-6 

sepaloid scales; stamens 2-6, the filaments free or connate, 

anthers elongate, basifixed, pollen 2-3-celled when shed, 

single or in tetrads, 1-ulcerate; pistil 1, carpels 1-2, the styles 

1-2, stigmas decurrent ventrally; ovary superior, 1(2)-locular, 

ovule 1(2), bitegmic, crassinucellar, anatropous; embryo sac 

antipodals sometimes multiplying to as many as 150 cells; 

endosperm formation Nuclear; fruit achenes or nutlets; embryo 

straight, elongate; endosperm copious. 

Distribution: Nearly cosmopolitan, most commonly in the 

Northern Hemisphere. 

Chemistry: Tanniferous; sometimes cyanogenic; not saponiferous. 

SPARGANIACEAE (Figure 9a,b).-Perennial herbs of wet 

habitats, rhizomatous; raphides and calcium oxalate crystals 

commonly present; xylem vessel perforation plates scalariform 

in mot and stem and perhaps leaves; leaves linear, basally 

sheathing, distichous, usually emergent, sometimes floating, 

the stomates paracytic; plants monoecious, the inflorescences 

panicles or spikes of spherical heads, the 8 above the 3; 

flowers small, with 3-6 sepaloid scales, protogynous, anemophilous; stamens 3-6, the filaments free or partially connate, 

anthers oblong, basfixed, extrone; pollen 3-celled when shed, 

ulcerate, the ulcus f pore-like; pistil 1, carpels 1(2), very rarely 

3, the style(s) elongate, stigma(s) 1(2), decurrent ventrally; 

ovary superior, l(2)-locular, ovule 1 per locule, bitegmic, 

crassinucellar, anatropous; embryo sac antipodal cells undergo 

postfertilization multiplication, forming as many as 150 cells; 

endosperm formation Nuclear (Helobial ?); fruit a nutlet; 

embryo straight, 0.8-0.9 the length of the endosperm, 8-13 

times longer than wide; endosperm copious, mealy, with starch, 

protein, and oil. Chromosomes: n = 15. 

Composition: 1 genus, -15 species. 

Distribution: Throughout temperate and cold Northern 

Hemisphere; few in the East Ides, southern Australia, New 

Zealand. 

TYPHACEAE (Figure 9c,d).-Perennial emergent herbs of 

wet habitats, aerial stems jointless, erect, with a thick rhizome; 

raphides and calcium oxalate crystals commonly present; 

xylem vessel perforation plates scalariform in root, stem and 

leaves; leaves alternate, distichous, strap-shaped, basally 

sheathing, the stomates paracytic; plants monoecious, the 
NUMBER 71 21 

\Wl 

C 

LJ 

FIQURE I.-NMADACEAE a, Najas major part of plant showing a flower in the axil of a leaf, 8flower. 1.8. and 

C.S. of same showing the solitary stamen, 3 flower, b, 1.s. of ovary showing the solitary basal ovule, fmit, 1.8. 

of same. N. guadelupenriS seed, 1.s. of same showing the straight embryo (after Martius). POSIDONIACEAE: c, 

Posidonia oceunim flower, 1.s. of same, embryo, 1.s. of same, plant swn after germination (after Baillon). 

inflorescence a scapose, dense cylindrical spadix with numerous minute flowers, the 8 above the 3; flowers without a 

perianth, merely subtended by bristles, protandrous, anemophilous; stamens 2-5, the filaments free or connate, anthers 

linear, basifixed, the connective usually apically produced; 

pollen 2-celled when shed, single or in tetrads, ulcerate, the 

ulcus single, k pore-like; pistil 1, stipitate, carpel 1, the style 

elongate, stigma decurrent ventrally; ovary superior, ovule 1, 

bitegmic, crassinucellar, anatropous; embryo sac antipodals 

degenerate won after fertilization (Davis), or increase in 

number (Engler Syllabus); endosperm formation Nuclear; fruit 

achenes or nutlets with a tuft of long hairs (wind dispersed), 

the seed striate; embryo straight, 0.5-1.2 times the length of 

the endosperm, 6-9.7 times longer than wide; endosperm 

copious, mealy, with aleuron and oil. Chromosomes: x = 15. 

Composition: 1 genus, -10 species. 
22 SMITHSONIANCONTRIBUTONS TO BOTANY 

d 
NUMBER 71 23 

Distribution: Nearly cosmopolitan, but most widely distributed in the Northern Hemisphere. 

JUNCALES 

Herbs of wet to dry habitats, usually rhizomatous, the stem 

sometimes jointed, solid or the internodes hollow: raphides 

absent, silica rarely present; xylem vessel perforation plates 

usually scalariform and simple, sometimes only scalariform, 

vessels sometimes absent from the leaves; leaves alternate, 

sometimes distichous, often basal, usually elongate and 

narrow, sometimes reduced to a sheath, the latter usually open, 

sometimes ligulate; inflorescences usually terminal and compound; flowers small or minute, almost always anemophilous, 

bisexual or the plants dioecious or monoecious, the parts 

hypogynous; perianth segments 6-3(0), usually glumaceous, 

rarely scarious, free or rarely shortly connate; stamens 6 or 

3(-l), the filaments elongate, anther basifixed or dorsifixed, 

sometimes apically produced; pollen 1-aperturate, ulceroid or 

sulcoid, sometimes shed in tetrads, 2-3-celled; pistil 1, carpels 

3-1, the styles 3-1(0), usually long, the stigmas sometimes 

decurrent ventrally, sometimes plumose: ovary 3-l-locular, 

the ovules 1-many per locule, axile-basal or -apical, or 

parietal, bitegmic, crassinucellar or tenuinucellar, anatropous 

or orthotrapous; embryo sac antipodal cells ephemeral or 

persistent into early embryogeny, sometimes multiplying to 

15 cells; endosperm formation Helobial or Nuclear; fruit a 

loculicidal capsule or nutlet, the seeds small, 1 to many; 

embryo small, axile or lying against the endosperm; endosperm 

copious. 

Distribution: Cosmopolitan. 

Chemistry: Usually tanniferous, sometimes cyanogenic 

(not Restionaceae); usually not saponiferous; alkaloids absent; 

seed fats of some Juncaceae, Liliaceae and Philesiaceae contain 

eicosenoic acid. 

JUNCACEAE (Figure lOa,b).-Perennial or seldom annual 

herbs often of moist habitats, rarely a shrub (Prionium), 

rhizomatous, usually glabrous but Luzula with long multicellular hairs; raphides, crystals and silica absent; xylem vessel 

perforation plates usually scalariform and simple in all organs 

or sometimes only scalariform in the stem and leaves; leaves 

filiform, flat or terete, mostly basal, alternate, often ligulate, 

the sheath usually open, rarely closed, rarely with a spinulose 

margin (Prionium); epidermal cells and stomares usually 

elongate longitudinally, stomates paracytic; inflorescence 

FICJURE 9.4PAROANIACEAE: a, Sprganium simpkx upper pen of plant with 

4 8 inflorescences above and 2 9 below, S. ramapum 1 .s. of 6 intlo~~scence, 

9 flower, 1.s. of same, Sparganiwn pistil with 2 styles; b, Sparganim 1.s. 

and C.S. of ovary, S. natans 8 flower, C.S. of fruit with 4 bracts. seed, embryo 

(after Le Maout and Decaisne, Barn). ~PHACEAE: c, 7jpha latifolio upper 

part of plant in flower (much reduced), I: angwtifolia part of 8inflorescmce, 

part of 9 inflarescence, Tpha 6 flower, pollem in tetrads; d, 1.s. of ovary, 

diagram of 9 flower, fruit, 1 .s. of seed, embryo (after Le Msout and Decaisne. 

Baillon, Mamus). 

terminal and axillary cymes, panicles, corymbs or heads, 

sometimes solitary; flowers small, nectarless, almost always 

anemophilous, bisexual or rarely the plants dioecious, the parts 

hypogynous; perianth segments 6(3) in 2 whorls, glumaceous, 

rarely scarious, usually greenish or reddish brown, rarely white 

or yellowish; stamens 6(3), free, the anthers basifixed, rarely 

apically produced, pollen ulcer(oid)ate, shed in tetrads, each 

grain 3-celled; pistil 1, carpels 3, styles 1 or 3, short or long, 

stigmas 3, decurrent: ovary 1- or 3-locular, the ovules 3 to 

many, axile-basal or patietal, bitegmic, crassinucellar, anatropus; embryo sac 3 antipodal cells may persist into early 

embryogeny; endosperm formation Helobial, free-nuclear 

divisions in the mimpylar chamber are followed by cell 

formation; fruit a loculicidal capsule, the seeds small, 

sometimes tailed, only Voladeria fruit 1-seeded (?Juncaceae); 

embryo axile, small, 0.2-0.3 the length of the endosperm, 

1.3-2.0 times longer than wide; endosperm copious, starchy. 

Chromosomes: x = 3-36, especially 6,lO; diffuse centromere 

in chromosomes of Luzula. 

Composition: 9 genera, -350 species. 

Distribution: Cosmopolitan, chiefly cold temperate and 

montane, rare in tropical regions. 

THURNIACEAE (Figure lOc).-Coarse rhizomatous glabrous 

herbs of wet habitats; raphides and calcium oxalate crystals 

absent, silica bodies present; xylem vessel perforation plates 

scalariform and simple in the mot, scalariform in the stem and 

leaves: leaves basal, linear-acuminate, leathery, entire or with 

spinulose margins, keeled, the sheath short, open, the stomates 

paracytic, sometimes tetracytic; inflorescences dense globular 

heads, subtended by several leafy bracts, at the apex of a stout, 

3-bgonous or subterete scape; flowers small, bisexual, 

anemophilous; perianth of 6 free, (or shortly connate ?), 

similar, oblanceolate, membranous parts; stamens 6, the 

filaments long-exserted, adnate to the base of the perianth; 

anthers about 3 times longer than wide, basifixed; pollen united 

in tetrads, grains with an obscure roundish aperture; pistil 1, 

carpels 3, connate part of the style short, style arms 3, fdiform, 

the stigmas decurrent ventrally; ovary superior, 1(3)-1ocular, 

the ovules axile-subbasal, 1-few per locule, anatropous; fruit 

a loculicidal capsule, the 3 seeds elongate, pointed at both 

ends; embryo axile in the endosperm, 0.3 the length of the 

endosperm, 7 times longer than wide; endosperm copious, 

mealy, starchy. 

Composition: 1 genus, 3 species. 

Distribution: Guayana, Venezuela, Amazon basin of Brazil, and rarely Colombia. 

CENTROLEPIDACEAE (Figure 1 la,b).-Small, tufted, annual 

or perennial herbs of wet habitats: raphides, crystals and silica 

bodies absent; sometimes with multicellular unbranched or 

branched filamentous hairs, xylem vessel perforation plates 

scalariform in all organs: leaves basal, more rarely imbricate 

along the stem, alternate, sometimes distichous, with a short 

open sheath; the stomates paracytic, somewhat similar to those 

in Poaceae; inflorescences scapose spikes or heads, or rarely 
24 SMITHSONIANCONTRIBUTIONS TO BOTANY 

FIoURE ~~.-JUNCACE%E: a, Juwvp bdbosvs plant in flower, 1.s. of flower, floral diagram, Juncvp one of the 

fruit valves showing the numerous seeds, seed, 1,s. of same deprived of its cellular testa; b, Lvncla campcsrrir 

flower, 1.s. of same, dehisced fruit, seed, 1.1. of same (after Baillon, Le Maout and Decaisne). THURNIACUE: 

c, Thurnia sphaerocephala part of plant showing globular inflorescence. 'I: jennmnii flower, 1.s. of sac, pistil, 

'I: sphaerocephala seed, 1.s. of same, 1.s. of lower part of seed showing small embryo in abundant endosperm 

(after Hooker, Baillon, Pilger). 

the flower solitary, mostly subtended by relatively large bracts; 

flowers minute, anemogamous, bisexual or the plants momecious; 

perianth 0, the flower subtended by 1-3 bracteoles; stamens 

1(2), the filaments filiform; anthers 1-2-locular, elongate, 

dorsifKed or basifixed, pollen 1-aperturate, sulcoid or ulceroid, 

3-celled when shed, pistil 1, the carpels 1-3, styles 1 per carpel, 

free or connate, filiform, the stigmas venrrally decumnc ovary 

superior, 1-3-locular; ovules 1 per locule or carpel, apical, 
NUMBER 71 25 

FIOURE 11 .-CWTROLEPIDACEAE: a, Centrolepis divmmondii plant in flower, inflorescence, pseudanthium, 

seed, 1.s. of same showing the basal anbryo; b, Gaimardia aurtralis plant in flower, flower, 1s. of same, 

dehisced fruit, Aphelia cyperoidcs flower, 1.s. of same (after Baillon). RESTIONACEAE: c, Rcsrw wginatur upper 

half of plant in flower, R. triticcvp 6 flower, floral diagram of 9 flower, 1.s. of 9 flower, Lanyvocaados 

grandis 9 flower, 1.8. of same; d, Restio tetraphyh 6 flower, 3 flower joined to its bract, 1.s. of pistil. Restw 

dehisced fruit, seed, 1.s. of same showing the basal embryo and abundant endosperm (after Le Maout and 

Decaisne, Baillon). 
26 SMITHSONIANCONTRIBUTIONS TO BOTANY 

bitegmic, crassinucellar, anatrops, the antipodals of the 

embryo sac increased in number; endosperm development 

Nuclear; fruit a membranous loculicidal capsule, or rarely 

indehiscent; embryo small, marginal, lying against the 

endosperm, 0.25 the length of the endosperm, as long as wide; 

endosperm or perisperm copious, starchy, mealy. Chromosomes: x = 10-14. 

Composition: 5 genera, -30 species. 

Distribution: Centered in southern Australia and New 

Zealand; Hainan, Indochina, East Indies; southern tip of South 

America. 

RESTIONACEAE (Figure 1 lc,d).-Perennial herbs, usually 

xerophytic but sometimes in wet habitats, often covering large 

areas like grasses and sedges, tufted or with an elongate 

rhizome covered with scale-like sheaths, the stems often 

jointed, solid or the internodes hollow, quadrangular, flattened 

or terete; raphides absent, crystals generally absent, silica 

bodies present; xylem vessel perforation plates simple in the 

root, simple or scalariform or both in the stem, rarely simple 

and scalariform in the leaves, but vessels usually absent from 

the last; leaves usually reduced to a mostly open sheath, rarely 

with a linear blade, very rarely with a ligule, sometimes 

somewhat ensiform, distichous, rarely basal, the epidermis 

with long and short cells, stomates paracytic, similar to those 

in Poaceae, often sunken; hairs uncommon, unicellular or 

malticellular, simple or branched filaments or flattened, 

fan-shaped and closely appressed to the epidermis; inflorescences 1- to many-flowered spikelets arranged in spikes or 

panicles, rarely solitary or pared, sometimes differing between 

the sexes; flowers small, anemophilous, the parts hypogynous, 

very rarely bisexual (some Phyllocomos, Lepyrodia hermuphrodita), the plants usually dioecious or rarely monoecious; 

perianth 3-6 glumes, in 2(1) series, or rarely absent fmm the 

3 flower, free or rarely the inner ones basally connate, or 

different fmm the outer; 8 flower: stamens 3 or 2(1), the 

filaments linear, free or rarely connate, the anthers oblong 

1(2)-locular, dorsifixed, the sacs separate at base and apex, the 

connective often apically produced, pistillode present or 

absent; pollen 1-ulcerate, -ulceroidate or -sulcoidate, often 

graminoid, 2-3-celled when shed, 3 flower: pistil 1, carpels 

3-1, the styles 3-l(0 in Chondropetalm), free or connate, 

usually linear, sometimes plumose, the stigmas sometimes 

decurrent ventrally, rarely surrounded by a disk (Hypodiscus), 

3 small staminodes present or absent; ovary 3-l-locular, the 

ovules 1 per locule, axile-apical, bitegmic, crassinucellar or 

tenuinucellar, orthompous; the 3 antipodah of the embryo sac 

usually ephemeral, but in a few species secondary multiplication occurs and up to 15 cells are formed but degenerate soon 

after fertilization; endosperm formation Nuclear; fruit a 

loculicidal capsule or nutlet; embryo lenticular, lying against 

the endosperm, less than 0.1 the length of the endosperm, 4-5 

times wider than long; endosperm copious, mealy, starchy. 

Chromosomes: x = 6-13, especially 6. 

Composition: -28 genera, -350 species. 

Distribution: Centered in South Africa and Australia, New 

Zealand; Madagascar, Malawi; Indochina, Malay peninsula; 

southwestern South America. 

FLAGELLARIACEAE (Figure 12).-Erect herbs (Joinvillea), 

or somewhat woody climbers (Flugellaria) with a rhizome; 

stem internodes solid (Flagellaria) or hollow (Joinvillea); 

raphides absent; xylem vessel perforation plates simple and 

scalariform in the stem and leaves; leaves alternate, elongate, 

lanceolate or linear, entire, grass-like, distichous in at least 

Flagellaria, the apex ending in a cirrhose tip in the latter, with 

an elongate closed (Flagellaria) or open (Joinvillea) sheath, 

the epidermal cells often in longitudinal files, stomates 

paracytic, similar to those in Poaceae, silica present (Joinvillea) 

or absent (Flagellaria), secretory cells present (Flagellaria) 

or absent (Joinvillea), glabrous (Flagellaria) or with pricklehairs or multicellular hairs (Joinvillea); inflorescence a 

terminal panicle or compound spike; flowers bisexual, small, 

apparently anemophilous; perianth of 6 parts, free or shortly 

connate, dry (Joinvillea) or somewhat corolloid (Flagellaria); 

stamens 6, rarely 1 aborts, hypogynous or adnate to the base 

of the perianth segments, the filaments free, elongate, the 

anthers rather small, oblong, sagittate at base, retuse at apex, 

basifixed or dorsifixed near base; pollen 1-ulcerate, the aperture 

f circular, graminoid, 2-celled when shed; pistil 1, carpels 3, 

the styles 1, with 3 linear, shortly connate arms, or 3, ventrally 

stigmatic, plumose and papillate; ovary superior, 3-locular, the 

ovules 1 per locule, axile, orthotropous or anatrops; fruit a 

small drupe, the seeds 1-3, globose, embryotega present; 

embryo small, lenticular, lying against the endosperm, less 

than 0.1 the length of the endosperm, 8 times wider than long; 

endosperm copious, mealy, starchy. Chromosomes: Joinvillea 

x = 18; Flagellaria n = 19, 18? 

Composition: 3 genera, 7 species. 

Distribution: Tropical and subtropical; southern 2/3 of 

Africa, Madagascar, southern India, Indochina, the East Indies, 

northeastern Australia. 

CYPERALES 

The order is monotypic. 

Chemistry: Sometimes tannifems; seldom cyanogenic or 

Saponifwous; many silica rich; mucilage absent; rarely 

producing simple indole alkaloids and sometimes ethereal oils. 

CYPERACEAE (Figure 13).-Perennial, or infrequently annual herbs or very rarely woody, usually of wet habitats, often 

tufted, sometimes rhizomatous or tuberous, stem solid or rarely 

hollow, often aigonous, rarely compressed, very rarely 

articulated; prickle-hairs It ubiquitous, silica-bodies often 

present, raphides absent; xylem vessel perforation plates most 

often simple, also scalariform or both in roots, stems and 

leaves; leaves usually basal or subbasal, cauline ones usually 

3-ranked rarely distichous (Coleochloa), linear, grass-like, 

rarely terete, rarely reduced to a sheath, the sheath closed, 

rarely open (Coleochlw), a ligule rarely present, the stomates 
NUMBER 71 27 

b 

FIGURE ~~.-FWE~.LARIACEAE: u, Flugclluriu irrdicu inflorescence and leaves showing the characteristic 

circinate apex. flower (the anthers fallen), 1.s. of same, fruit, 1.s. of same showing the minute embryo at the 

apex of abundant endosperm; b, flower, pistil and C.S. of same, fruit, 1.s. of same with 2 aborted loculu, part 

of infructesm~e (after Engler and Dmde, Baillon). 

paracytic or rarely tetracytic, usually in longitudinal rows, 

guard cells dumbbell-shaped; inflorescences simple or compound spikes, arranged in umbels or panicles, usually 

subtended by 1 or more leafy bracts; flowers small, each 

subtended by a glume, bisexual or the plants monoecious, very 

rarely dioecious, distichously or spirally arranged, rarely the 

3 spikelets reduced to 1 bract and 1 flower; anemophilous, or 

very rarely entomophilous (Dichroomenu); perianth absent, 

rarely scales, bristles or hairs in its place, very rarely 

subpetaloid; lodicules absent; stamens 3( 1-22), hypogynous, 

the filaments free, anthers oblong or linear, basifixed, rarely 

apiculate; pollen 3-celled when shed, 1-4-aperturate, the 

apertures ulceroid, poroid or elongate; pistil 1, the carpels 3-2, 

style 1 with 2-3 branches, the stigmas papillate, usually 

decurrent; ovary superior, unilocular, uniovulate, the ovule 

basal, bitegmic, crassinucellar, anatropous; embryo sac antipodal cells ephemeral; endosperm formation Nuclear, the tissue 

later becoming cellular; fruit an achene, very rarely a drupe, 

often uigonous, sometimes lenticular; embryo axile, 0.2-0.5 

the length of the endosperm, 1-1.5 times as long as wide; 

endosperm copious, starchy, oily, and with a protein layer, 

mealy or fleshy. Chromosomes: x = 35-45 or more; diffuse 

centromere in chromosomes of some species. 

Composition: -80 genera, -3500 species. 

Distribution: Cosmopolitan, centered in temperate regions. 

POALES 

The order is monotypic. 

Chemistry: A variety of fructosans present; rarely tanniferous; occasionally cyanogenic; sometimes produce isoquinoline, simple indole or pyrmlizidine alkaloids; sometimes 

saponiferous; mucilage absent. There is very close similarity 

between the chemistries of Poaceae and Cyperaceae according 

to Hegnauer. Harbome points Out the presence of the 

characteristic leaf flavmids (glycoflavones, tricin) of the 

Poaceae in Cypemxae; flavone, flavanols and leucoantho- 
28 SMlTHX"ICONTRIBUTI0NS TO BOTANY 

d 
NUMBER 71 29 

cyanins are fairly numerous in Poaceae and Arecaceae, but 

there is only one report of flavanol and no leucoanthocyanins 

from Araceae; coumarin and d-mannitol are present in Poaceae 

and Arecaceae. 

POACEAE (Figures 14, 15).-Perennial or infrequently 

annual herbs, more rarely shrubs or trees, the stems terete, 

rarely flattened or angled, jointed, the nodes solid, often 

swollen, internodes usually hollow, rarely solid; silica present, 

raphides and calcium oxalate crystals absenc xylem vessel 

perforation plates simple, sometimes also less commonly 

scalariform, in roots, stem and leaves; leaves usually linear, 

occasionally broad, flat, rarely terete, ovate, elliptic, lanceolate 

or setaceous, alternate, distichous, very rarely spirally arranged 

(Micruiru), the sheath usually open, rarely closed, a ligule 

usually present, sometimes a petiole between blade and sheath, 

the epidermis with long and short cells, stomates paracytic, in 

longitudinal rows, guard cells dumbbell-shaped; 2( 1)-celled 

micro-hairs common, sometimes unicellular macro-hairs form 

an indument readily visible to the naked eye, hairs rarely 

uniseriate; inflorescences panicles, racemes, spikes or heads 

of spikelets; flowers (florets) small, usually bisexual, rarely the 

plants monoecious or dioecious, anemophilous, very rarely 

entomophilous, usually consisting of the reproductive organs 

enclosed in 2 bracts (lemma and palea), the floret@) in a 

spikelet@) usually subtended by 2 bracts (glumes), 2-3(0- 

many) hyaline scales (lodicules) internal to the lemma, 

sometimes the lemma, more rarely glumes, awned; florets 

l-many per spikelet, distichous; perianth absens stamens 3, 

6(1-120), the filaments filiform or rarely short, free or rarely 

connate basally; anthers oblong, basifixed but the sacs usually 

partly separate basally and apically, rarely apiculate; pollen 

3-celled when shed, l-ulcerate; pistil 1, the carpels 2-3, styles 

2( 1,3), free or connate, the stigmas 2( 1-3) generally plumose, 

sometimes only papillate, rarely ventral (Chusqueu guudichaudii); ovary superior, unilocular, uniovulate; ovule apical 

to basal, 2( 1 ,O)-tegmic, usually pseudocrassinucellar, sometimes tenuinucellar, anatropous, hemitropous or campylotropous; secondary multiplication of the antipodal cells is 

characteristic and as many as 300 cells may be produced; 

endosperm formation Nuclear; fruit a caryopsis, rarely a drupe, 

nut, berry or utricle, sometimes hairy and wind dispersed, 

embryo with a relatively large haustorium (scutellum), basal 

to lateral, lying on the abaxial surface of the endosperm, 

F~UW ~~.-CYPERACEAE: a, Cyprus longus floral diagram, spikeld, flower, 

1 .s. of ovary, 1.s. of fruit showing small basal embryo and abundant endosperm, 

C. stricta inflorescences; b, Schanoxiphivm rufum spikelet with 6 flowers 

above and P flower below, spikelet reduced to the 3 condition plus a 

glumigerous rhachilla, Carexstricta dflower, flower, 1.s. of same showing 

pengone enclosing ovary and style, Scleria lvcida fruiq c, Rhynchospra 

sparguntoides inflorescences, 6 flower, anther. perfect flower, fruit, 1.s. of 

same, R. recyryata stigma and upper pert of style; d, R. alba part d 

infloresrmce, 3 flower, Oreobolus pctinatw fruit, flower, Ewndra arktata 

I.s. of inflorescence (after Bdm. Martius). 

0.1-1.1 times the length of the endosperm, 3.3-8.0 times 

longer than wide; endosperm copious or rarely lacking in the 

mature seed (Melocannu), starchy, sometimes also oily, usually 

with a protein layer too. Chromosomes: x = 3-21, especially 

7-10, monocentric. 

Composition: -600 genera, -8OOO species. 

Distribution: Cosmopolitan and widespread, dominant in 

prairie, savanna and other open formations, less common in 

shady, wooded habitats, more common in dry than wet habitats. 

I4RECAL.B 

The order is monotypic. 

Chemistry: Noteworthy for their seed-fats. Lauric acid 

seems to be the most abundant fatty acid in seed-fats of most 

genera of palms. Lauric and myristic acids are also most 

abundant in the seed-fats of Lauraceae and Myristicaceae. 

Among other families Poaceae and Arecaceae have n-hexadec9-enoic acid (palmitoleic acid). Several produce waxes; very 

often tanniferous; various sorts of polyphenols present; rarely 

producing pyridine alkaloids; occasionally saponiferous; seldom cyanogenic. See Poales for additional comparison with 

that order. 

ARECACEAE (Figures 16-18).-Perennial trees, shrubs and 

climbers, sometimes rhizomatous, sometimes spiny; raphides 

and silica-bodies common; hairs sometimes absent, but often 

present particularly in the inflorescence and on young leaves, 

typically with a multicellular or unicellular base of sclerotic 

or cutinized cells and a distal uniseriate filamentous, or 

shield-like group of thin-walled cells, less commonly microscopic prickle-hairs; xylem vessel perforation plates scalariform or simple and scalariform, rarely vessels absent from the 

stem; leaf traces diverging toward center of stem before turning 

to exit in leaves; leaves alternate, clustered at the apex of the 

stem, scattered along the stem in climbing species, rarely 

radical, with a closed sheath and often a tubular ligule between 

the sheath and petiole, large to very large, pinnately or 

palmately divided or entire, folded in bud, induplicate or 

reduplicate, often prickly, the stomates temytic or hexacytic; 

inflorescence usually large, compound, rarely simple, spikes 

or a panicle, sometimes in dichasia, rarely heads, axillary or 

below the leaves, rarely terminal, subtended by one or more 

spathes; flowers entomophilous or anemophilous, usually 

bracteate, usually small, usually actinomorphic, the parts 

hypogynous, sometimes embedded in the spadix, bisexual or 

the plants monoecious, dioecious or polygamous; perianth 

usually in 2 whorls, rarely in a single whorl or the parts (to 

10) spirally arranged, homochlamydeous or more rarely 

heterochlamydeous, often leathery, rarely glumaceous, usually 

green, sometimes white, tan or yellow to red; sepals 3(2), free 

or connate, imbricate or open in bud; petals 3(2), free or 

connate, valvate or imbricate; stamens usually 6, more rarely 

numerous or few (3--400), the filaments free or connate, 

rarely adnate to the petals, the anthers as long as wide to linear, 
30 SMlTHSONIAN CONTRIBUTIONS TO BOTANY 

n 

C 
NUMBER 71 31 

1' b 

FIOURE ~~.-POACEAB: a, ZLa mys plant in flower, B flower, part of 8 infloresance, 9 inflorescence without 

the bracts, canpound fruit, 9 flower, b, Tripcwn daciyloidcs Bflower with palea, 9 flower, 1.s. of fruit, T. 

angvlatwn inflorescence; c, Imperata brasilielrpis inflorescence, spikelet, essential organs (after Baillon, 

Martius). 

FIGURE 14.-POACEAE: a, AWM sativa inflorescences, spikelet, flower with dorsifaed or basifixed, the sacs sometimes separate at thew, 

one glue removed, fruit, 1.s. of same showing the basal lateral embryo, and sometimes rarely separate from apex to m, 

spresbding from a bifid connective and geniculate with the embryo, 1.s. of same (below); b, Oryur sativa diagram of spikelet, spikelet, 

same with glumes removed, inflorescence, Zizoniopsir microstachy 1.s. of 

pan of leaf showing ligule at juncture of blade and sheath, Digitaria lanuginow fihWIt (pies of Geomm); @en 2-celled when shed, 

pan of infloreescence, Park infermedia stamens; c, Glyceria nervaka usually monosulcate, trichotomo-sulcate, or 2-sulcate, 2- 

essential organs, Brm mollis pistil, 1.s. of same showing the solitary bad sulculate (-z~isulcula&) in Nyp, usually smooth, sometimes 

ovule, strePto&yw crinata stamen, Pistil, floral diagram, base ofph showing %him; sometimes an &.& present; 1(3), 

the carpels 3(1-lo), sometimes with the ventral suture open, 

distichous condition and open sheaths; d, Bambwa bambos part of 

inflorescence, spikelet, Churqueu Ieptophylla part of stem showing swollen 

joints and hollow internode., Bambura bvnbos gynoedum and glumes, 

Merosiachys plurifroru floral diagram, Streptochaeta spicata floral diagram absent, sometimes filiform with 4-10 very long Stigmas 

(after Martius, Baillon). (Phyrelephas); ovary superior, 1-3(4-1O)-locular, septal 

the styles and stigmas usually 1-39 sometimes the styles 
32 SMlTHSONIANCONTRIBUl'IONS To BOTANY 

d 

e 

FIOIIRE 16.--A~eclr~~m: a, Phoenixdactylgera 1.8. of aflower, 1.8. of 9 flower, 1.5. of fruik seed, 1.s. and 

C.S. of same showing the minute anbryo in abundant endosperm, embryo mlarged, same with the ootyledon 

detached, b, Cocos nvcjfera 1.s. of fruit showing the minute embryo in abundant endosperm, fruit with husk 

remwed showing the 3 carpels, Arenga gowutus 1.8. of 6flower; c, Caryota wenv branch of inflorescence, C. 

soboljfcra plant in flower, C. wens 6 flower, 9 flower in bud, same at anthcsis, 1.s. of same, Areca curechu 

fruit, 1.s. of same; d, Bactrir bahnophora 9 flower, 1.8. of same, Raphia nJfF. fruit and &inflorescence, Arenga 

hngkab 1s. and cs. of fruit e, Trithrinax acanthocoma flower, calyx, petal, androecium, gynoecium. 1.s. of 

pistil (after Bdm, Le Maout and Decaisne, Martius). 
NUMBER 71 33 

C 

d 

ROW ~~.-ARECACEAE: a, Chamaerops hwnilu part a€ inflorescence in spathe, 9 flower, Gflower, diagram 

of 8 and 9 flowers; b, Borossusflabelljfcr 1.s. of pan of 6 inflorescence, 9 flower, part of 6 inflomcenoe; c, 

Moximiliana marijm 8 flower, petal with 2 stamens. Micrmoelwn pulchra 1.s. of d flower, 1.s. of 9 flower, 

Astrocaryum weddcllii dflower, petal with 2 stamens, fruit; d, Alhgoptera maritimum androecium and a perianth 

segment, 6 flower, Cah ciliarip 1.8. d flower, 9 flowers, 1.s. of 9 flower, pistil, c, Gconoma rubescens 

9 flower, pistil end staminode, staminode. pistil, Attdea phalerata 9 flower, pistil and staminodal cup, d 

flower, stamens, stamen (enlarged) (after Le Maout and Decaisne, Baillon, Manius). 
34 SMITHSONIANCONTIUBUTONS TO BOTANY 

a 

FIciUW 18.-ARBCACEAE: 0, Ehck mclcrnococcu &lower, sepal, petal, androecium, 1.s. of same, anddum 

with all stamens but one moved, 2 views of anther, flower, pistil; b, Phyrclcphus nvlcrocorpa 8 

inflorescence, 1.s. of P inflorescence, pistil, 1.8. of same, Nypufruticortr dflower (3 mate stamens), C.S. of 

fruit (after Baillon, Martius). 

nectaries often present; ovules axile (subbasal to apical), 1 per 

ovary or locule, bitegmic, crassinucellar, anatropous, rarely 

semi-anatropous or orthotropous, embryo sac antipodal cells 

usually ephemeral, rarely persisting into embryogeny, endosperm formation Nuclear, later cellular; fruit a berry or 

drupe; seeds 1-6, large, the embryo minute, typically 0.2-0.5 

the length of the endosperm, 1-3.5 times longer than wide, 

lateral or oblique, basal or above the middle; endosperm 

copious, often very hard, with cellulose, oil and aleuron, but 

starch absent. Chromosomes: x = (8-10),13-18,(19), especially 14,16,18. 

Composition: -220 genera, -3000 species. 

Distribution: Mainly pantropical, especially Asia and then 

America, extending into the warm temperate regions, in various 

habitats, including the understory of lowland rain forest, 

mangrove, deserts, but with accessible ground water, mountain 

thickets. 

CYCLANTHALES 

The order is monotypic. 

Chemistry: Sometimes tanniferous; probably not saponiferous; mucilage present. 

CYCLANTHACEAE (Figure 19).-Rhizomatous perennial 

herbs or shrubs, or lianas with aerial roots, rarely trees, rarely 

epiphytic; juice watery or milky; raphides commonly present, 

calcium oxalate crystals sometimes present, silica cells absent; 

xylem vessel perforation plates scalariform in the roots and 

leaves, vessels absent from the stem; leaves spirally arranged 

or distichous, large, sometimes fan-shaped, entire or bilobed 

apically, parallel veined, the petiole sheathing, the stomates 

tetracytic; inflorescence solitary, terminal or axillary, a spadix, 

sometimes screwlike, subtended by conspicuous green, white, 

yellow or red spathes; the plants monoecious, the flowers 

FIciUW 19.--cYc~TH~~: a, Cyclmrhus bipartitus inflorescence before 

anthesis and at anthesis. 1.s. of part of inflorescence, 1.8. of 2 and 1 8 

flower, upper part of leaf (greatly reduced); b, Curlvdovim luncijoliu 

inflaescence. C. plmotu plant in flower, C. lutifoliu spadix with fruits and 

8 flowers; c, C. funifcru 8 flower. 1.s. of same, C. plicuto 8 flower from 

above, lateral view of same, 1.s. of same; d, C.S. of spedix. C.funiferu flower, 

1.s. of same; c, C. plicuto C.S. of flower, C.S. of ovary, C. pulmtu group of 

fruits SM latenlly and fran above, Curlvdovim 1.s. of seed showing the 

minute embryo in abundant endosperm (after Bdon, Madus, Le Maout and 

Decaisne). 
NUMBER 71 35 

C 

d 
36 SMlTHSONIANCONTRIBUTIONS TO BOTANY 

small, spirally arranged, nectaries absent, the receptacle 

sometimes concave and the parts perigynous, more rarely 

epigynous, the female surrounded by 4 males, or the sexes in 

superposed whorls; pollinated by beetles; 8 flowers: perianth 

in one whorl, cupular and several toothed or lobed, or rarely 

absent; stamens 6-150, the filaments basally connate and 

sometimes adnate to the perianth, the anthers as long as wide 

or linear, basifixed or dorsifixed; pistillode absent; pollen 

2-celled when shed, 1-sulcate or ulceroidate; 3 flowers: free 

or connate; perianth absent or of 4 segments, free or basally 

connate, staminodes 4 and often fdiform; pistil 1, the carpels 

4(1,3), the style 1 or 0, stigmas 1, 2 or 4, often elongate and 

divergent; ovary unilocular, superior or often embedded in the 

spadix, the ovules numerous, bitegmic, crassinucellar, anatropous, parietal or apical; embryo sac antipodal cells 3; 

endosperm formation Helobial; fruit a syncarp or free bemes; 

seeds numerous, small, sometimes with a sarcotesta, sometimes sculptured; embryo minute, basal, straight or curved, 

0.1-0.2 the length of the endosperm, 2-4 times longer than 

wide; endosperm copious, fleshy or horny, with oil, aleurone, 

hemicellulose, rarely starch. Chromosomes: n = 9,15,16. 

Composition: 11 genera, -150 species. 

Distribution: Mostly in moist forests; tropical America; 

West Indies. 

ARALES 

Large herbs with a rhizome or corm to a minute thallus, 

rarely woody, sometimes climbers, often of wet habitats; 

raphides nearly universal (absent from Acorus and some 

Lemnaceae); xylem vessel perforation plates scalariform in the 

root, vessels absent in the minute plants; leaves simple to 

compound, radical or alternate, sometimes distichous, linear 

to nearly orbicular, the major veins longitudinal and parallel 

or more often palmate or pinnate, the minor ones reticulate, 

usually petiolate and with an open or closed sheath, 

inflorescence a spadix and spathe, bracteoles absent; flowers 

small, bisexual or the plants monoecious, rarely paradioecious; 

perianth homochlamydeous, the segments 0,4,6,(-9), free or 

connate; stamens 4, 6(1-8). free or connate, the anthers 

basifixed, rarely apically produced, pollen sulcate, porate, or 

nonaperturate, 2-3-celled when shed, pistil 1, the wpels 

3(1-9), style 1, long or short, the stigma apical, ovary 

3( 1 -9)-locular, superior or embedded in the spadix, the ovules 

l-many, axile, basal, apical or parietal, bitegmic, crassinucellar, rarely tenuinucellar, anatropous or hemianatropous, rarely 

orthotropous; embryo sac antipodals ephemeral, rarely multiplying secondarily; endosperm formation ab initio Cellular, 

fruit usually a berry, sometimes a utricle; seed coat sometimes 

fleshy; embryo large or small, usually axile, endosperm 

copious to absent in the mature seed. 

Distribution: Mostly pantropical and subtropical, few 

temperate. 

present; mucilage fairly common, some have latex; many 

Chemistry: bucoanthocyanins, anthocyanins and flavonols 

cyanogenic; amines common in the inflorescence, attracting 

carrion flies, also produced by Aristolochiaceae and Asclepiadaceae; some tannins and saponins common in Araceae but not 

Lemnaceae; the phenolic acids of Araceae and Lemnaceae are 

quite similar; alkaloids especially the indole group in Araceae 

but not Lemnaceae; the phenolic acids do not strongly support 

relation of Araceae and Pandanaceae. According to Gibbs the 

little we know of Cyclanthaceae is not against relation with 

Araw and in general the chemistry of Arecaceae is in line 

with relation to the Arales. However there are differences. 

Gibbs has never found HCN in Araceae, but others have found 

it. 

ARACEAE (Figures 20, 2la).-Perennial herbs, rarely 

woody, sometimes climbers with adventitious roots, or 

epiphytes, often of wet habitats, with a rhizome or corm, some 

with latex, very rarely aromatic (Acorus), raphides nearly 

universal (absent from Acorus), generally glabrous (uniseriate 

hairs in Pistia); xylem vessel perforation plates scalariform in 

the mot, scalariform or vessels absent in the stem; leaves 

radical or alternate on elongate stems, sometimes distichous, 

simple, entire, divided, or compound, linear to nearly orbicular, 

the major veins parallel or more often palmate or pinnate, the 

minor ones sometimes evidently reticulate, usually petiolate 

and with an open or closed sheath, inaavaginal scales rarely 

present, the stomates paracytic, tetracytic, or subsidiary cells 

more than four; inflorescence a scapose or axillary spadix, the 

flowers rarely few, subtended by and often enclosed in a 

sometimes colored spathe, individual floral bracts absent, very 

rarely vestigial bracteoles present (Pothos), the flowers sessile, 

very rarely pedicellate; flowers small, often fetid, the parts 

hypogynous, bisexual or usually unisexual, sometimes with a 

rudiment of the opposite sex, and the plants monoecious, males 

on the upper part of the spadix, often striking sterile flowers 

above OT below the males, the females below, or rarely 

paradioecious, sometimes flowers absent from the apex of the 

spadix; perianth homochlamydeous, of 0, 4, 6,(-9) free or 

connate segments in 1 or 2 whorls, nectarifemus in Anthurium; 

stamens 4, 6(1-8), free or connate, rarely completely so in 

synandria, the filaments often flattened, the anthers usually 

basifixed rarely dnate, or the locules immersed in an 

FICWRE 20.-AluCr~~: a, Acorus cah spadix (naturally not enclosed in 

spathe), flower, 1,s. of part of ovary showing the apical ovules, fruit, 

Gymostachys anceps flower from above, 1.s. of pistil with a single apical 

ovule; b, Zantedeschia C.S. of lower and upper part of ovary, Arum maculatum 

stamen, 1.s. of pistil, fruit, 1.s. of seed, embryo, spadix with the spathe removed 

showing 6 flowers above the 9; c, Orontinm aquaticwn flower from above, 

stamen, 1s. of pistil showing the single basal ovule, Calla palustrip spadix and 

spathe, flower, 1 .s. of same (the flower normally lacks a perianth); d, Anfhwiwn 

scandens part of spadix, lateral view of flower, Pistia stratwtes spathe opened 

showing the single pistil below and the androecium above. 1,s. of same. fruit 

adnate to the spthe, 1.8. of seed, c, Philcdendrm sqnamifcrwn 1.s. and C.S. 

of fruit, seed, 1.s. of same with exterior integument removed, Ambrosino 1.8. 

of inflorescence with pistil separated from androecium by a diaphragm, pistil 

and diaphragm from above, androeciUm; f, Caladium striatips synmdrium 

lateral view and from above, C.S. of same, pistil. 1s. and C.S. of same (after 

Baillon, Le Maout and Decaisne. Madus). 
NUMBER 71 37 
38 SMITHSOMANCONTRIBUTIONS TO BOTANY 

b 

C 

d 

)I\ 0 *- *..- 

FwUm 21.-ARACEAB: a, Arkam vdgarc spadix showing dflowers above and ? below, dflower, 1.s. of 

? flower, Awwrphophallvs hpc group of 6 flowers, 1.3. of 9 flower, Symplocarpvs 1.s. of seed 

(exalhmiuous), Cytusperma 1.8. of d (after Baillon, Le Maout and Decaisne). LWACEAE: b, Lemaa 

minor in flower, inflorescencein bract, 1.9. of pistil. fruit, L. paucicostata fruiting frond; c, L. wldiviana stamen, 

L. rrudca seed. same with testa removed. 1.s. of same. anbryo, Spiro&Ia polyrrhizu essential organs, stamen, 

fruiting fd (after Le Maat and Decaiane, Martius). TRILLUmE: d, Paris qw&.ifdia plant in flower, flower. 

1.r. of same, flaral diagram; c, R pol~hylla C.S. of wary, plant in flower, floral diagrpm, Trilliwn crcctwn floral 

diagram (after Baillon, Le M.out and Deuisne). 
NUMBER 71 39 

undifferentiated stamen, the connective rarely apically produced, dehiscence poricidal or longitudinal; pollen 1-sulcate, 

3(2-4)-sulculate, zonisulculate, 3(4)-porate, oligoforate, polyfe 

rate or nonaperturate, the type correlated with pollinator, rarely 

in tetrads (Xunthosoma), 2-3-celled when shed, pistil 1, the 

carpels 3(1-9), the style long, short or absent, the stigma(s) 

apical, wet; ovary 3(1-9)-locular, superior or embedded in the 

spadix, the ovules 1-many, axile, basal, apical or parietal, 

bitegmic, crassinucellar, rarely tenuinucellar, anampous or 

hemianatropous, rarely orthotropous; embryo sac antipodals 

usually 3 and ephemeral, rarely some secondary multiplication 

occurs; endosperm formation Cellular from the start; fruit a 

berry; seeds sometimes with a sarcotesta; embryo large or 

small (Pistiu), usually mile in the endosperm, curved when the 

latter is absent, often 0.7-0.8 the length of the endosperm, 

7-10 times longer than wide; endosperm copious, oily and 

sometimes also starchy, to absent in the mature seed, 

sometimes perisperm also present (Acorus). Chromosomes: x 

= 6?,7-9,ll-17,21,22, especially 8,12-16. 

Composition: -1 10 genera, -2500 species. 

Distribution: Pantropical and subtropical, mostly on the 

forest floor or vines on trees, few in the temperate zones. 

LFMNACEAE (Figure 2lb,c).-Small or minute aquatic 

perennial herbs, a mere thallus, often rootless, the stomates 

anomocytic; raphides sometimes present; xylem vessels 

absent; inflorescence consists of 1 3 flower and 1-2 8 flowers 

on the surface of the thallus at its margin, sometimes with a 

small membranous spathe, the plants monoecious rarely 

dioecious; perianth absent; 8 flowers: stamens 1-2, filament 

present or the anthers sessile; anther of 2 separate sacs, 

transversely or apically dehiscent; pollen 1-aperturate, ulceroid, 3-celled when shed; 3 flowers: pistil 1, the carpel 1, 

style 1, short, the stigma apical; ovary superior, unilocular, the 

ovules 1 -few, basal, bitegmic, crassinucellar, hemianatropous 

or anatropous; embryo sac antipodals usually ephemeral, rarely 

persisting to early endosperm formation; endosperm formation 

Cellular from the start; fruit a utricle, the seed with a fleshy 

outer seed coat; embryo axile, straight, 0.8 the length of the 

endosperm, 2.4 times longer than wide, endosperm scanty or 

absent in the mature seed, fleshy, with starch and other 

reserves. Chromosomes: x = 5,8,10,11, especially 10,ll. 

Composition: 6 genera, -30 species. 

Distribution: Cosmopolitan, except much of the arctic 

region, in quiet fresh water. 

LIUM 

Usually herbs, less commonly shrubs, rarely trees, sometimes climbing, with rhizome, corm, bulb or tuber; raphides 

present in some Liliaceae, Agavaceae, Stemonaceae, Dioscoreaceae, Xanthorrhoeaceae, and Velloziaceae; xylem 

vessel perforation plates scalariform, simple or both, vessels 

sometimes absent from the stem or leaves; leaves linear to 

orbicular, basal or cauline, alternate or whorled, rarely 

opposite, the veins extending from the base to the apex, in the 

broader leaves transverse veins also evident; inflorescence a 

raceme, spike, umbel, panicle, thyrse, cyme, head, or the flower 

solitary; flowers usually conspicuous, bisexual, or less 

commonly the plants dioecious or polygamo-dioecious, 

actinomorphic, rarely zygomorphic, usually entomogamous; 

perianth segments usually 6, free or connate, corolline, rarely 

the outer whorl sepaloid; outer whorl 3(2-10) segments, inner 

3(0-7) segments; stamens 6(1--48, in bundles when many), 

hypogynous, epigynous or adnate to the perianth, the filaments 

free or connate, anthers basiiixed or dorsifiied, rarely adnate, 

the connective sometimes apically produced; pollen usually 

1-sulcate, sometimes 2-sulculate, trichotomosulcate, or 2- 

porate or nonaperturate, 2(3)-celled when shed; nectaries in the 

septa of the ovary or on the base of the perianth segments; 

pistil 1, the carpels 3(2-lo), rarely nearly free, the styles 

1(-lo), the stigmas apical or ventrally decurrent; ovary 

superior, semi-inferior or inferior, 3( 1 - 10)-locular, the ovules 

usually numerous, less commonly few to 1 per locule, axile, 

rarely parietal, basal or apical, 2(0-l)-tegmic, crassinucellar 

or tenuinucellar, anatropous, hemianatropous, campylotropous 

or orthotropous; embryo sac antipodals ephemeral or persistent 

into endosperm formation, rarely increased to 11; endosperm 

formation Helobial or Nucieat, fruit a loculicidal or septicidal, 

rarely ventricidal, capsule, less commonly a berry, rarely a 

samara, utricle or nutlet; seeds 1-many, various in size, shape 

and consistency, sometimes longitudinally ribkd; embryo 

small to large, straight or curved, axile; endosperm copious, 

hard or fleshy, very rarely floury. 

Distribution: Cosmopolitan. 

Chemistry: Steroidal saponins and sapogenes are characteristic of Liliaceae, Agavaceae and Dioscoreaceae. The order 

is not rich in alkaloids. The Liliaceae are notable for their 

steroidal alkaloids and mpolones, cardenolides and scilladienolides, some produce inulin and fructosans. Steroidal 

alkaloids also occur in Ranunculaceae. Sremonu has a unique 

pup of alkaloids. Amaryllidaceae has unique alkaloids which 

set it off from all other Liliales; however a few members of the 

Menispermaceae have one or two of the same alkaloids. 

Taccaceae has many alkaloids, dioscorin occurs in Dioscoreu. 

The seed fats of LiliaCeae, Xanthorrhoeaceae, Agavaceae and 

Hypoxis are rather similar. Gibbs knows of no chemical 

character that really distinguishes Dioscoreaceae from some, 

at least, of the other families of the order. No group of 

chemicals is unique to the Agavaceae; its chemistry is very 

like that of Liliaceae. According to Gibbs, chemistry does not 

favor segregating Smilacaceae from Liliaceae and the Wurmbamideae may wmt family status. A family Colchicaceae 

has been suggested, its outstanding chemical character is 

tropolone alkaloids, including colchicine. The Allieae lack the 

characteristic alkaloids of the Amaryllidaceae. Velloziaceae 

lack saponins, are often tannifmus, commonly produce resin 

or gum, are not cyanogenic. Some Xanthorrhoeaceae produce 

napthaquinones. 

TRILLMCEAE (Figure 2ld,e).-Herbs with simple erect stem, 

rhizomatous, raphides present; xylem vessel perforation plates 
40 SMlTHSC"ICO"I'RIBUTIONS TO BOTANY 

scalariform in the mts, vessels absent from the stem and 

leaves; leaves entire, whorled or opposite, usually fairly broad, 

with midrib and one major pair of longitudinal veins, the 

transverse veins between the midrib and longitudinal veins 

quite evident, the stem also with a few short basal leaf-sheaths, 

the young leaves sometimes floccose; inflorescence a terminal 

solitary flower or umbel; flowers bisexual, actinomorphic, the 

parts hypogynous; sepals 3-5(2-lo), free, usually green, 

sometimes brownish, purplish or white; petals 3-5(0-7), free, 

usually colored, frequently differing in size and shape from the 

sepals; stamens 6(2-20), the filaments usually short, free, the 

anthers elongate, basifixed, connective usually produced 

apically and usually between the sacs; pollen 1-sulcate; pistil 

1, the carpels 3(-lo), the styles linear, 3(1-lo), usually deeply 

lobed, the stigmas ventrally decurrent; ovary 3(-lO)-locular, 

with septal nectaries, the ovules numerous, axile, bitegmic, 

crassinucellar, anatropous; embryo sac antipodal cells 3, 

usually ephemeral, occasionally persisting into early embryogeny; endosperm formation Helobial (sometimes Nuclear ?), 

division at first free-nuclear, later the tissue becomes cellular: 

fruit usually a berry, sometimes a fleshy loculicidal capsule; 

seeds several, globular, the embryo small, 0.1 the length of the 

endosperm, 1-2 times longer than wide; endosperm copious, 

hard or fleshy, starchy. Chromosomes: x = 57, especially 5. 

Composition: 3 genera, -50 species. 

Distribution: Rmperate North America and Eurasia. 

LILIACEAE (Figures 22-24a).-Perennial or rarely annual 

herbs, or rarely somewhat woody, rarely climbing, with 

rhizome, corm, bulb or tuber; velamen present on roots of 

several genera; raphides present in some members; xylem 

vessel perforation plates scalariform, simple or both, vessels 

sometimes absent from the stem or leaves; leaves linear to 

orbicular, basal or cauline, alternate or whorled, rarely redud 

to scales, the veins when conspicuous mostly extending from 

the base to the apex, in broader leaves cross veins also evident 

and sometimes the venation is pinnate (Lilium spp.), usually 

entire, rarely the margin prickly, the stornates mostly 

anomocytic, rarely paracytic (e.g., Astelia, Hosta), rarely with 

various kinds of hairs; inflorescences raceme or spike, rarely 

an umbel (Allium), subcapitate, cyme or solitary, rarely 

spathamus (Allium), terminal, axillary or scapose; flowers 

actinomorphic, rarely slightly zygomorphic, usually bracteate, 

usually entomogamous, bisexual or rarely the plants polygamodioecious; perianth petaloid, the segments free or connate, 

6(4,8,10 in Antherolophus) in 2 similar whorls, rarely basally 

saccate; stamens 6(3,4,8,9,12), hypogynous or adnate to the 

perianth, the filaments elongate, free or connate; rarely 3 

staminodes present (Erinna); anthers basitixed or dorsifixed, 

dehiscing longitudinally, rarely poricidal, usually introrse or 

lmrse, more rarely extrorse; pollen 2(3)-celled when shed, 

usually 1 -sulcate, sometimes 2-sulculate, trichotomosulcate, 

zonisulculate, 2-porate, 3-aperturate, irregularly apertumte or 

nonaperturate; nectaries in the septa of the ovary or on the 

base of the perianth segments, seldom absent; pistil 1, the 

carpels 3(2,4), rarely nearly free (Tofwldia, Neodregea, 

Dipidax),the style 1(3), stigmas 3, apical or decurrent ventrally; 

ovary 3( 1,2,4)-locular, superior, rarely semi-inferior, the ovules 

many, rarely 1-few, axile, very rarely parietal (species of 

Asteliu), bitegmic, crassinucellar or tenuinucellar, anatropous, 

hemianatropous or orthotropous; embryo sac antipodals 3, 

ephemeral or persistent into endosperm formation, rarely 

increased up to 11; endosperm formation Helobial, (Nuclear?); 

fruit a loculicidal or septicidal, rarely ventricidal capsule, more 

rarely a berry; seeds 1-many, the embryo axile in the 

endosperm, straight or curved, usually small, 0.1-2.0 the 

length of the endosperm, 1.6-16 times longer than wide; 

endosperm copious, fleshy or cartilaginous, never floury, with 

protein, oil, hemicellulose, rarely any starch. Chromosomes: x 

= 3-202337, especially 6-8,12. 

Composition: -230 genera, -3500 species. 

Distribution: Cosmopolitan, except much of the arctic 

region, most abundant in temperate regions. 

SMILACACEAE (Figure 24b,c).-Perennial herbs and shrubs, 

usually climbing, often with a large tuberous rhizome, the stem 

often with prickles; raphides generally present; xylem vessel 

perforation plates scalariform in the root, stem and leaves, 

vessels sometimes absent from the stem and leaves; leaves 

alternate, more rarely opposite, often leathery, usually fairly 

broad and petiolate, with a midrib and 1-2 pairs of arcuate 

veins from the base to the apex and conspicuous cross-veins, 

the petiole sometimes tendriliferous, the stomates anomocytic; 

inflorescences axillary umbels, racemes or spikes, rarely a 

terminal panicle; flowers small, actinomorphic, the parts 

hypgynous, rarely bisexual, the plants usually dioecious, 

staminodes present in 3 flower but pistillode absent from d 

flower; perianth segments usually 6, rarely the petals absent, 

usually greenish or yellowish, free or rarely connate; stamens 

6(3,9), the filaments free or connate, adnate to base of perianth, 

the anthers about 3 times longer than wide, basifixed, introrse; 

pollen nonaperturate, 1-sulcate, 1-sulcoidate ?; pistil 1, the 

carpels 3, styles 3(-1), short, the stigma 3(-1)-lobed, decurrent 

ventrally; ovary 3(-l)-locular, the ovules 1-2 per locule, axile; 

fruit a berry, the embryo small, 0.2 the length of the endosperm, 

3 times longer than wide; endosperm copious, hard, with 

hemicellulose, lipid, protein, and rarely some starch (Ripogonum). Chromosomes: x = 13-16, especially 16. 

ETaURB 22.-mCEAE: a, Tofieldia calyculota flower, 1.s. of same, apex of 

stamen, C.S. of wary, seed (enlarged), floral diagram, Veratrwn floral diagram; 

b, Veratrwn apex of stamen, cs. of ovary, 1.8. and C.S. of seed. embryo, 

ZygadanlLp gkaberrk 1.1. of flower, c, Anguillarka dioica 1.8. of 8 flower, 

1.s. of ? flower, Narlhecium ossifragum 1,s. of flower, d, inflorescence, 

dehisd fiUit, 1.8. of seed showing the small embryo, part of tails cut off 

(gnatly enlarged), Colchicwn autwMole anddum and perianth-limb laid 

open, stamen, apex of a style showing sanewhat decurrent stigma, 1,s. of seed 

showing minute anbryo in abundant endosperm, Is. of flower, c, wary and 

base of styles, 1.s. of same, C.S. of ovary, dehiscing fruit, C.S. of same, mrm; 

f, seed and 1.s. of same, fld diagram, Makanthemwn biforia flower, 1.s. of 

same (after Baillm, Le Macut and Decaisne). 
NUMBER 71 41 

b 

e 

f 
42 SMITHSONIANCONTRIBUTIONS TO BOTANY 

Composition: 4 genera, -375 species. 

Distribution: Mainly pantropical and subtropical, extending into temperate regions. 

AGAVACEAE (Figure 246).-Usually large perennial herbs, 

or shrubs or trees, rarely scandent, of dry habitats, usually 

rhizomatous, sometimes tuberous; raphides commonly present; 

xylem vessel perforation plates simple or scalariform in the 

roots, sometimes scalariform in the leaves, vessels absent from 

the stem and sometimes leaves; leaves elongate, crowded, often 

basal, sometimes spirally arranged along the stem, entire or 

toothed, usually thick, fibrous and leathery or fleshy, usually 

not sheathing, or the sheath open, the stomates usually 

anomocytic or tetracytic, sometimes paracytic (Doryunthes); 

inflorescence usually a large scapose panicle, thyrse, raceme 

or spike, rarely a head or the flower solitary; flowers usually 

bisexual, rarely the plants polygamo-dioecious or dioecious, 

actinomorphic or rarely slightly zygomorphic; perianth segments 6, corolline, free or connate; stamens 6, usually adnate 

to the corolla, more rarely epigynous or hypogynous, the 

filaments elongate, anthers oblong, usually dorsifiied, rarely 

basifxed (Dolyanthes); pollen 1-sulcate or 2-sulculate, provided with a thick reticulate or retipilate sexine, usually single, 

rarely tetrads, 2-celled when shed, pistil 1, the carpels 3, style 

1, usually elongate, slender, rarely short or absent, the stigma 

usually enlarged and shortly 3-lobed; ovary superior (e.g., 

Yucca) or inferior (e.g., Agave), septal nectaries usually 

present, 3-locular, the ovules numerous to 1 per locule, axile, 

or rarely unilocular (Daylirion) with 3-6 basal ovules; ovules 

bitegmic, crassinucellar, anatropous; embryo sac antipodal 

cells rarely show secondary multiplication to 5 cells (Doryunthes); endosperm formation Helobial, perhaps sometimes 

incorrectly reported as Nuclear; fruit a loculicidal capsule or 

berry, the seeds many to one, usually compressed; embryo 

within the endosperm, 0.4 to as long as the endosperm, 5 times 

longer than wide; endosperm copious, fleshy, or hard, with 

hemicellulose, oil, protein, but no starch; perisperm present in 

Yucca. Chromosomes: x = 8,10,12,15-21,24,30, especially 

19,30(5 large, 25 small). 

Composition: -20 genera, -600 species. 

Distribution: Pantropical and subtropical, extending into 

the temperate zones. 

XANTHORRHOEACEAE (Figure 25).-Usually large xerophytic perennial herbs and shrubs, often with a thick, 

sometimes branched, woody stem, or rhizome, sometimes 

resinous, raphides commonly present; xylem vessel perforation 

plates simple or scalariform in the root, rarely scalariform in 

the leaves, vessels absent fmm the stem and usually leaves; 

leaves usually linear, sometimes liguliform, usually tufted, 

basal or apical, rarely scattered dong the stem, alternate, not 

sheathing or with a short open sheath, sometimes with a prickly 

margin, the stomates anomocytic, less often paracytic; 

inflorescence a panicle, dense cylindrical spike, globular head, 

often scapose, or rarely the flowers solitary and axillary or 

terminal; flowers small or rarely large, bisexual or the plants 

dioecious; perianth mostly glumaceous rarely large and 

colored, the segments 6 in 2 whorls, free or shortly connate, 

the inner whorl sometimes petaloid; stamens 6 in 2 whorls, the 

inner attached to the base of the inner perianth segments, the 

outer free and hypogynous; anthers about as long as wide, 

2-locular, basifixed or dorsifiied; pollen 1 -sulcate, 2-sulculate, 

zonisulculate or spiraperturate; pistil 1, carpels 3, the styles 

usually elongate, 3 or 1 and 3-lobed, stigmas 3, apical, 

sometimes capitate; ovary superior, 3-locular, with 1-several 

axile ovules per locule, or 1 -1ocular with 3 basal ovules; ovules 

bitegmic, crassinucellar, anatrops; embryo sac with 3 large 

antipodal cells; fruit a loculicidal capsule or rarely a nutlet; 

embryo straight, endosperm copious, hard, horny, with protein, 

oil, and hemicellulose. Chromosomes: x = 7-9,11,17,24, 

especially 11,8. 

Composition: 8 genera, -60 species. 

Distribution: Endemic to Australia except for 3 species in 

New Guinea and 1 in New Caledonia. 

PHILE~IACEAE (Figure 26u).-Perennial shrubs, subshrubs, 

sometimes thin-stemmed climbers, rhizomatous; raphides 

sometimes present; xylem vessel perforation plates simple or 

scalariform or both in the roots, scalariform or vessels absent 

from the stem, vessels absent from, or perhaps plates rarely 

scalariform in the leaves; leaves alternate, sometimes distichous, usually petiolate, ovate or oblong with only the midrib 

conspicuous or often 1-2 pairs of arcuate lateral veins 

extending from the base to the apex or 4 pairs from the midrib 

to apex (Petermanniu), reticulations between lateral veins 

mostly evident, rarely some leaves reduced to sheaths, the 

stomates anomocytic; inflorescences terminal or axillary 

(leaf-opposed in Petermanniu), cyme, raceme, fascicle or the 

flower solitary; flowers bisexual, actinomorphic, often large, 

sometimes medium-sized, perianth nectaries present; perianth 

segments 6, corolline and similar or the inner much larger than 

the outer, free or rarely connate (Behniu, Petermunniu); 

stamens 6, hypogynous or rarely adnate to the perianth, the 

filaments free or basally connate, anthers elongate, large, 3-6 

times longer than wide, dorsifixed, introrse or latrotse (exmrse 

in Petermunniu); pollen 1-sulcate, sometimes aichotomosulcate (both in Geitonoplesium), or nonaperturate (Lupgeriu, 

Philesiu), spinulose in the latter 2 genera, 2-celled when shed; 

pistil 1, the carpels 3, style 1, elongate, the stigma terminal, 

shortly 3-lobed or capitate; ovary superior or rarely inferior 

R~UW 23.--LImm: a, PolygoMtwn vulgare plant in flower, flower, 1.8. 

of same, Aspidhra elolior flower, 1.s. of same, pistil, plmt in flower, b, 

EriarpcrmvAl parvifolivm flower, 1s. of same, dehisoed fruit, 1.s. of seed, c, 

Aloe Vera 1.s. of flower, Asphodclw ramxw lde of ovary with dmd 

surface qxmed, 1.s. of flower, Peliosanthes teka ruptured fruit showing the 

seeds, P. campnulalo 1.s. of flower, 1.8. of seed; d, Asparagw ominah 1.s. 

of 8 flower. 1.s. of perfect flower, fruiting branch, 1.8. of B&. Tricyrtir 

stigmas, floral diagram; e, Rwcw 8 flower, 9 flower, 1.1. of 9 flower, 1s. 

of fruit, Allivm cepa seed, 1 .s, of same (after Baillq Le Maout pnd Decaisne). 
NUMBER 71 43 

b 

d 

h 

e 
-3 
NUMBER 71 45 

a 

FIc3URE ~.-XANTHORRHOBACEAB: a, Xanth.orrhoea hurtilk flower, 1.s. of same, dehisced fruit, seed. 

Lomandra longiforia 6 flower, 1.s. of same; b, Calectrrsia cyaneu part of plant in bloom, flower, 1.s. of same, 

Darypogon bromeliifolius inflorescence, flower. 1.8. of same (after Baillon). 

(Petermnniu), 3- or 1-locular, the ovules usually numerous, 

sometimes few, axile or parietal, bitegmic, crassinucellar, 

campylotropous; embryo sac antipodals large; fruit a berry, 

sometimes large, rarely later becoming dry and dehiscent 

(Eusfrephus), the seeds globose, embryo straight or slightly 

curved, 0.3 the length to as long as the endosperm; endosperm 

hard, with protein and oil. Chromosomes: x = 10, also n = 

15,19. 

Composition: 8 genera, 10 species. 

Distribution: Peru to the southern tip of South America; 

eastern Australia, New Zealand, the East Indies to New 

Caledonia and Fiji; southeast Africa. 

STEMONACEAE (Figure 26b).--Glabrous perennial herbs or 

rarely subshrub (Stichoneuron), erect or sometimes climbing, 

rhizomatous, or with tuberous roots; raphides sometimes 

present; xylem vessel perforation plates scalariform in the roots 

and sometimes in the stem, vessels absent from the leaves and 

sometimes stem; leaves alternate, opposite or in whorls of 3-5, 

often distichous, basal or along the stem or subapical, entire, 

petiolate, midrib and several pairs of lateral arcuate veins 

extending from the base of the blade to the apex or from the 

midrib toward the apex, with numerous fine straightish 

cross-veins between the arcuate veins, rarely with a short open 

sheath (Pentastemom), sometimes 2-3 sheaths at the base of 

the stem, the stomates anomocytic; inflorescences axilhry, the 

flowers solitary and pedicellate or few-flowered cincinnae in 

to moderate in size, nectaries absent (?); perianth petaloid, the 

segments free or connate, 4, 2-seriate or 5 in one whorl 

(Pentastemom); stamens 4 (5 in Pentastemom), the filaments 

short, free or connate (Pentusternnu), or adnate to the very 

base of the pehth; anthem elongate 0~ subglobose (Pentastemom), basifixed oT donfix& oT sate and be anther sacs 

separate, the connective sometimes produced apically; pollen 

I-Sulcate, m1Y nonaperm (Pentastemom); Pistil the 

carpels 2 (3 in Pentastemom), style elongate or the stigma 

FIOURE 24.--LILIACME: a, Danai racemosus flowering branch with cladodes 

and leaf scales, flower, 1s. of same, Convailaria majaiip 1s. of flower, stamen 

inserted on base of corolla, pistil. inflorescence against pt of leaf (after 

Baillon, Le Ma,,,,t and De,sne). sMILAcAcBAE: b, smilru brrrsiliclrrip 

branchlet with tendrils and spines and inflorescence in bud, s. mauritanica 

flowering P branch, S. pwpwata 1.s. of ? flower showing the dmmt 

stigmas; C, s. hcrboCc0 ? flower, s. rOtundifo1k fruit. s. aspera 1.8. of seed, 

embtyo, S. brrrsilklrris stamen, views of aflower, Smilax Gfloral diagram 

(after Baillon, Martius, Le Maout and Decaisne). A~AVACEAE: d, Agave 

americana flower, 1.s. of same, plant in bloom (much reduced), Yucca 

treculeana 1.s. of flower, 1.s. and C.S. of a fruit, C.S. and 1.s. of seed showing 

linear embryo in abundant endosperm (after Baillon, Sargent). 

racemes or subumbel; flowem bisexual, actinomwhic, small 
46 SMITH!3O"IANCONTRIBUTIONS TO BOTANY 

C 
NUMBER 71 47 

sessile; ovary superior, semi-inferior or inferior, unilocular, the 

ovules numerous to 2, basal, apical or parietal (Pentustemonu), 

bitegmic, crassinucellar, anatropous; embryo sac antipodal 

cells very ephemeral; endosperm formation Nuclear, the tissue 

later becoming cellular; fruit a capsule dehiscing by 2 valves, 

or indehiscent (Pentustemonu), the seeds 1-many, broadellipsoid, longitudinally grooved or ribbed, with a basal, lobed 

aril, the funicle sometimes very long; embryo usually small, 

0.1-0.7 the length of the endosperm, 1.5-7.0 times longer 

than wide; endosperm copious, fleshy or hard, starchy. 

Chromosomes: x = 7. 

Composition: 4 genera, -25 species. 

Distribution: Southeastern Asia, the East Indies, northern 

Australia; southeastern United States. 

DIOSCOREACEAE (Figure 26c-e).-Climbing perennial herbs 

or shrubs often with tuberous rhizomes, rarely spiny, glabrous, 

or hairs uni- or bicellular, stellate, or glandular; raphides 

present; xylem vessel perforation plates scalariform at least in 

the root and stem, vascular plexus in the nodes distinctive and 

consistent; leaves petiolate, alternate, rarely opposite and 

whorled, the blade usually broad, usually entire, sometimes 

palmately lobed or compound, often with a cordate base, the 

main veins extending from the base to the apex of the leaf and 

cross-veins quite evident, the stomates mostly anomocytic; 

inflorescences axillary cymes, spikes, racemes, panicles or 

heads, the plants dioecious; flowers small, inconspicuous, 

actinomorphic, entomophilous, perianth nectaries common; 

perianth segments 6 in 2 series, connate; 8 flowers: stamens 

6 or 3, sometimes 3 staminodes present, adnate to the base of 

the perianth, the filaments elongate, free or shortly connate, the 

anthers basifmed, extrorse or introrse, less than twice as long 

as wide, the sacs sometimes separate, the connective often 

apically produced, a pistillode sometimes present; pollen 

1-sulcate or 2(3)-sulculate, 2-celled when shed, 3 flowers: 

pistil 1, the carpels 3, styles 1 or 3, the stigmas 3, each 

sometimes 2-parted; ovary inferior, septal nectaries common, 

3-locular, the ovules 2 or 4 per locule, axile, bitegmic, 

crassinucellar, anatropous; staminodes often present; embryo 

sac antipodal cells degenerate soon after fertilization; endosperm formation Nuclear, the tissue later becoming celluk, 

fruit usually a capsule, sometimes samara or berry, the seeds 

FIGURE Zd-PHILESWCaAa: a, Philesia bvz~olia flowering twig, Lapageria 

rosea flowering branch, 1.s. of flower, anther, stigma, C.S. of way, fld 

diagram (after Lindley, Bailla, Le Maout and Decaisne). STEMONACEAE: b, 

Stemom tuberare stamen, C.S. of same and C.S. of anther (above), flower, 1.1. 

of same, S. glo*wsoides flowering twig, dchisced fruit with pendent seds on 

long funicles, seed, 1s. of same (atla Bdq Lindlcy). DIOSCOREACBAB: c, 

Trichopus zeyhnicw flower, 1,s. of same, views of stamen. style lad stigmas, 

C.S. of ovary. fruit, seed, 1.8. and cs. of same; d, Dioscorea batatas part of 8 

plant in bloom. part of 9 plant in flower end fruit, D. adenoccupo 6 flower, 

1s. of same, views of stamen; e, Dwscopea 9 flower, D. sativa fruit, D. 

adenocarp dehisced fruit, winged sad, Tcynvp ca~uurir 9 flower, 1,s. of 

same, fruits, Rujania hastaka fruit (after Knuth. Bdon, Le Maout and 

Decpisne, Martius). 

often winged, embryo small, 0.3 the length of the endosperm, 

as long as to 3 times longer than wide; endosperm copious, 

horny, with protein, oil, and hemicellulose. Chromosomes: x 

= 9,10,12,14,18, especially 10. 

Composition: -6 genera, -600 species. 

Distribution: Pantropical and subtropical, a few species 

extending into temperate regions. 

TACCACEAE (Figure 27u).-Perennial herbs with creeping, 

sometimes tuberous rhizomes; raphides present; xylem vessel 

perforation plates scalariform in the root, vessels absent from 

the stem and leaves; leaves basal, large, broad, entire to deeply 

lobed, with a midrib and fairly numerous lateral veins 

extending from it, the petiole often long, the stomates 

anomocytic; inflorescence scapose, the flowers in a cymose 

umbel, often subtended by a combination of linear and broad 

bracts; flowers bisexual, actinomorphic; perianth homochlamydeous, the segments 6 in 2 series, connate, campanulate, 

petaloid, dark-colored (brownish); stamens 6, adnate to the 

perianth, the filaments very short, broad, concave, the anthers 

inflexed, about as long as wide, adnate or basifmed, anther 

sacs separated by connective, the connective shortly produced 

apically; pollen 1-sulcate, 2-celled when shed, pistil 1, the 

carpels 3, intercarpel nectaries sometimes present, the style 1, 

short, style arms broad, usually bilobed, relexed, ovary inferior, 

unilocular, the ovules numerous, parietal, bitegmic, crassinucellar, anatropous; embryo sac antipodal cells ephemeral; 

endosperm formation Nuclear, the tissue later becoming 

cellular; fruit a berry or rarely loculicidal capsule; seeds 

numerous, longitudinally ridged, the embryo minute, included 

in the endosperm, basal or lateral, less than 0.1 the length of 

the endosperm, twice as long as wide; endosperm copious, 

somewhat cartilaginous, with aleurone, lipid, but no starch and 

little or no hemicellulose. Chromosomes: x = 15. 

Composition: 2 genera, -30 species. 

Distribution: Pantropical, centered in southeastern Asia, 

extending into Polynesia, limited to Amazon basin in America 

PONTEDERIACEAE (Figure 276,c).-Freshwater perennial or 

rarely annual (Hydrothrix) herbs with a short or creeping 

rhizome, vertical stems short, rarely submerged (Hydrothrix); 

raphides present; xylem vessel perforation plates scalariform 

in the mot, scalariform or vessels absent in the stem, vessels 

absent from the leaves; leaves mostly in a rosette, sometimes 

distichous, rarely spirally mged along an elongate stem, the 

blade broad or narrow with primary veins extending fmm base 

to apex, petiole distinct, sheath mostly closed, intravaginal 

scales sometimes present, the stornates paracytic; inflarescence 

a raceme, panicle, thyrse or rarely the flower solitary; flowers 

bisexual, actinomorphic or zygomorphic; perianth petaloid, 

marcescent, the segments 6 (4 in Scholleropsis), bisexiate, 

connate, rarely free (Monochoriu species); stamens 6 or 3 (1 

in Hydrothrix), mostly adnate to the perianth but free from one 

another, sometimes dissimilar and unequal in length, sometimes 1-2 staminodal; anthers oblong, rarely one larger than 

the others, basifixed or dorsifixed, dehiscence longitudinal, 
~~.-TA~cIwJ: a, Tacca cristata inflorescence, flower, 1.s. of same, T. lcontopctaloides fruit. seed, 

1.s. of same showing the minute embryo in abundant endosperm (after Baillm). PONTEDEUCIWJ: 6, Pontcderia 

cordaka flower. 1s. of same, flowering stem; c, Eichhornia fricolor put of inflorescence, views of fruit, same 

dehiscing, seed, Monochoria vaginalip seed, 1.s. of same showing the long embryo in abundant endosperm (after 

Baillm, Martius). AMARYUIDACEAE: d, Hypxis crecfa inflorescence wd pat of leaf, 1.s. of bud, H. decwnberrp 

flower, views of stamen, Hypoxip C.S. of ovary, dehisced fruit, C.S. of same, seed, 1.s. of same, embryo; c, 

Alstrocmeria monticoka perianth segmmts, A. plurtaglua upper part of style and stigmas, apex of stamen, A. 

vcrsicdw seed and 1s. of same showing the small embryo in abundant endosperm, A. pritkacim 1.s. of flower 

(after Le Maout wd Decaisne, Martius, Baillm). 
NUMBER 71 49 

0 

a 

horn ~~.-AMAI~YLLIMCBAB: a, NarciWvr tazetta &his& fruit, seed, 1.8. of same showing the minute 

embryo. 1.s. of flower showing corata and stamens inserted on the perianth. pistil, Hymemcallis cahthina 1.8. 

of flower showing the stsminodal conma (after Bdm). VEULYZIACENE b, Vellozia epidendroidcs part af stem 

showing the persistent leaf bases and numerous adventitious r00cs (in cs.) within them, V. caruncularis scapse 

flower and parts of leaves, V. tenelh part of periantb and stamens, V. hemisphaeruu 1.s. of flower, Barhcenia 

vandellii lateral view of stamen, B. luzul~olia flower opened out; c, B. pwpwca flower, stamen, 1.s. of flower, 

Burbncenia floral diagram, B. gradis dehisced fruit, seed, 1.s. of same showing the minute embryo in abundant 

endosperm, 8. am'ca seed, I.s. of same (lower pair) (after Martius, Baillon). 
50 SMITHSOMAN CONTRIBUTIONS TO BOTANY 

rarely by pores; pollen elongate, 2( 1,3)-sulculate, 2-celled 

when shed; pistil 1, the carpels 3, style 1, often trimorphic, the 

stigma apical, punctiform or shortly 3- or 6-lobed; ovary 

superior, internal septal nectaries usually present (absent in 

Heteruntheru), 3-locular, the ovules numerous, axile or 

1-locular and uniovulate, bitegmic, crassinucellar, anatropous; 

embryo sac antipodals ephemeral; endosperm formation 

Helobial, the tissue later becoming cellular; fruit a loculicidal 

capsule or utricle with one fertile cell; seeds small, longitudinally ribbed; embryo cylindrical, axile in the endosperm, as 

long as the endosperm, 5-7.8 times longer than wide; 

endosperm copious, floury, starchy and with an outer layer of 

aleurone. Chromosomes: x = 8,14,1536, especially 8. 

Composition: -7 genera, -30 species. 

Distribution: Primarily pantropical, extending into temperate America, Asia and Africa. 

AMARYLLIDACEAE (Figures 27d,e, 284. -Perennial herbs 

usually of dry habitats, with a tunicated bulb, very rarely a 

rhizome; raphides usually present; xylem vessel perforation 

plates scalariform in the roots, vessels absent from the stem 

and leaves; leaves entire, often linear or lorate, basal, usually 

with inconspicuous veins, sometimes with a closed sheath, the 

stomates anomocytic, generally glabrous (except Hypoxis); 

inflorescence a scapose cymose umbel, rarely subcapitate, 

rarely a solitary flower, often enclosed in bud by 1 or more 

membranous bracts; flowers bisexual, actinomorphic rarely 

zygomorphic, usually showy; perianth 6 segments in 2 whorls, 

free above the ovary or connate, corolline, sometimes with 

nectaries, a corona sometimes present; stamens 6(3,--20 in 1 

species of Gethyllis and 6 bundles with about 8 anthers each 

in another), epigynous or adnate to the perianth, the filaments 

free or connate, long or short, the anthers elongate, dorsifixed 

or occasionally basifmed, introrse, dehiscence longitudinal, 

rarely by apical pores; pollen usually 1-sulcate or 2-sulculate, 

2-celled when shed, pistil 1, the carpels 3, style slender, the 

stigmas apical, capitate or 3-lobed. ovary inferior, with septal 

nectaries, 3-locular, rarely 1-locular by abortion (Calostem), 

the ovules mostly numerous, rarely 1-few per locule, axile or 

rarely parietal (Leontochir, Empodium), 1- or 2-tegmic 

(ategmic in Crinum), pseudocrassinucellar, anatropous or 

semi-anampous; embryo sac antipodals often persist into early 

embryogeny; endosperm formation Helobial, the tissue later 

cellular, Nuclear reported but needs confirmation; fruit a 

loculicidal capsule or more rarely a berry; embryo 0.6 the 

length of the endosperm, 6 times longer than wide; endosperm 

copious, with cellulose or starch, also aleutone, and oil (?), 

enclosing the embryo. Chromosomes: x = 6-12,14324, 

especially 7-9,11,12. 

Composition: -75 genera, -1 100 species. 

Distnbutwn: Pantropical and subtropical, extending into 

the temperate zones, especially of the Southern Hemisphere. 

VELLOZIACEAE (Figure 28b,c).-F’erennial herbs and sparsely 

branched shrubs, usually of dry habitats; raphides sometimes 

present; xylem vessel perforation plates simple in the root, 

scalariform or absent in the stem and leaves (Tulbotiu); leaves 

in a basal rosette or apical tufts, linear-acuminate, with midrib, 

sometimes with a saw-toothed margin, the sheath persistent, 

the stomates paracytic, rarely tetracytic; flowers usually 

solitary on each scape, pedicellate, terminal, bisexual, rarely 

functionally unisexual and the plants dioecious (Burbucenwpsis), actinomorphic; perianth segments 6 in 2 whorls, petaloid, 

free or basally connate, sometimes with a small corona behind 

the anthers; stamens 6(--48) in 6 bundles when more than 6, 

the filaments epigynous (Tulbotiu) or adnate to the perianth, 

sometimes flattened, the anthers sometimes subsessile (Bmbaceniopsis), introrse, linear, basifixed or dorsifixed, pollen 

1-sulcate or nonaperturate, sometimes in tetrads (Velloziu); 

pistil 1, the carpels 3, style 1, the stigma capitate or shortly 

3-lobed; ovary with internal septal nectaries, inferior, 3-locular, 

the ovules very numerous, axile on stalked placentas, bitegmic, 

pseudocrassinucellar, anampous; fruit a loculicidal capsule, 

sometimes rupturing irregularly; seeds numerous, the embryo 

small, 0.2-0.3 the length of the endosperm, 2.5-4.0 times 

longer than wide; endosperm copious, hard, with hemicellulose, protein, oil, and starch. Chromosomes: n = 9,16 (Goldblatt 

in Raven), -2436. 

Composition: 6 genera, -270 species. 

Distribution: Tropical South America (1 species in Panama) and Africa, Madagascar, southern Arabia. 

BROMEWALES 

The order is monotypic. 

Chemistry: Steroid saponins sometimes present; tannin 

and mucilage common; alkaloids absent. 

BROMELIACEAE (Figures 29,30).-Perennial herbs or rarely 

woody plants often of dry habitats, and often epiphytic in moist 

habitats, sometimes rhizomatous or stoloniferous; hairs characteristically peltate shield-like scales with a uniseriate stalk, 

functioning to conserve and absorb water, rarely other types 

also present; raphides and silica present; xylem vessel 

perforation plates usually scalariform, sometimes simple in the 

roots, vessels sometimes absent from the stem and leaves; 

leaves mostly in a rosette, spirally arranged, dy distichous, 

liguliform. without a midrib (except Pitcairniu spp.), entire or 

often with spiny teeth, the sheath open, the stomates tetracytic 

or sometimes hexacytic; inflorescence a terminal, rarely 

axillary, panicle, raceme, spike or head, rarely the flower 

solitary, often with brightly colored bracts; flowers almost 

FrouRs ~~.-BROMKL.IACBAB: a, Navia acadis pistil, dchisrmt fruit and 

calyx, seed, 1.8. of same, Brocchinia panicdata C.S. of ovary, Pitcairnk 

incrmic 1s. of pistil showing the 6-inferior ovary, Dyckja cncholirioihs 

1.8. d flower, b, 8ep.l pd, views d stamm, pistil, d, 1.8. of me, 

Accha aquilcga indument scale greatly enlarged; c, Vricsea gigantea flower, 

1.8. of sme, V platynctm d grutly darged, Tilland~ia streptoatpa 

iuflorercence, puts d luf, Bromclia wbaniana part of luf; d, Tilladtia 

Bromclia whaniana flower, sepal, petal and stamens. 1.s. of wary, rpu d 

style MI showing the decurrent stigma, Acchmca aquilcga 1.1. of flower, the 

perianth and stamens removed (after Mutius. Baillon). 

SpUUbM EUUIQ, 1.8. d OVW. 1.8. Of d Showing the hd rmbryo, 
NUMBER 71 51 

b 

C 
52 SMlTHSONIANCONTRIBUTIONS TO BOTANY 

C 

FIGURe 30.-BROMELIACEAE: a, Ananas comosvs multiple fruit, Bromlia pingVin fruit, Billbcrgia spcciosa 

plant in flower, floral diagram, 1.8. of flower. b, B. nutans C.S. of wary, 1.s. of seed showing the small embryo 

in abundant endosperm. Poptea pctroplifana lower part of petal with apendages. sepal. stamen, upper part of 

style with stigmas; c, Streptocalyx Iongifolivs petal and stamen, 1.6. of wary, Araemoccvs parvijlow calyx, 

Acantharlachy strobilocea flower, 1.s. of same, fruit, seed, 1.s. of same (after Mdus, Baillm). 
NUMBER 71 53 

always actinomorphic, rarely weakly zygomorphic, bisexual 

or more rarely the plants polygamous or dioecious, entomogamous, ornithogamous, rarely anemogamous; perianth heterochlamydeous, but sometimes the sepals colored (Sodiroa, 

Mussungeu); sepals 3, free or basally connate; petals 3, free or 

connate, usually considerably larger than the sepals, sometimes 

with a pair of basal appendages; stamens 6, free or basally 

adnate to the petals, the filaments usually separate, sometimes 

basally connate, often long-exserted, the anthers linear, usually 

dorsifixed, pollen usually 1-sulcate or 1-sulcoidate, sometimes 

2-polyaperturate, then the apertures slightly elongate, circular 

or irregular, 2-celled when shed, rarely in tetrads; pistil 1, the 

carpels 3, style 1, elongate, the stigmas 3, usually linear, 

sometimes spirally twisted around each other; ovary superior, 

semi-inferior or inferior, 3-locular, with internal septal 

nectaries, the ovules usually numerous, axile, bitegmic, 

crassinucellar, anatrqus, rarely campylowopous; embryo sac 

with 3 small antipodal cells; endosperm formation Helobial, 

the tissue at first free-nuclear, later becoming cellular, fruit a 

septicidal, more rarely loculicidal, capsule, berry or syncarp; 

seeds usually numerous, small, sometimes appendaged or 

winged; embryo lateral to the endosperm or rarely embedded 

in it, 0.2-1.8 times the length of the endosperm, 2-10 times 

longer than wide; endosperm copious, mealy, starchy. Chromosomes: x = 8,9,17,25,27,28, especially 25,8. 

Composition: -50 genera, -2000 species. 

Distribution: Tropical and warm temperate America, 

except for one West African species. 

COMMELINALES 

Herbs, often of wet habitats, or mesophytic to somewhat 

xerophytic, sometimes succulent; raphides in Commelinaceae 

(except Curtonema), absent from the other families; xylem 

vessel perforation plates simple, scalariform or both, vessels 

sometimes absent from the leaves, very rarely from the stem; 

leaves spirally arranged or distichous, sometimes basal, usually 

elongate; flowers actinomorphic or zygomorphic, often ephemeral and mescent, bisexual or rarely the plants polygamomonoecious, more rarely monoecious, very rarely dioecious, 

the parts mostly hypogynous; paianth usually heterochlamydeous; sepals 3(2,0), green or hyaline, very rarely 

colored, rarely connate; petals 3(0), cmlline, rarely scarious, 

rarely connate; stamens 6(4-l), sometimes 3(4) staminodes 

present, the filaments free or connate, very rarely epipetalous; 

anthers basifixed or dorsifixed, dehiscent longitudinally or by 

apical pores; pollen l(2)-sulcate or nonaperturate or spiraperturate, 2-3-celled when shed; nectaries absent; pistil 1, the 

carpels 3(2), style 1, rarely appendaged, the stigma apical, 

usually unlobed, rarely mid; ovary superior, 3(-l)-locular, 

the ovules 1-many per locule, axile or parietal, rarely basal, 

bitegmic, crassinucellar or tenuinucellar, orthotropous to 

anatrqus; fruit a loculicidal capsule, rarely a berry; seeds 

often sculptured, the embryo small, lying against the endosperm; endosperm copious, mealy or floury, starchy. 

Distribution: Primarily pantropical and subtropical, relatively few in temperate zones. 

Chemistry: Sometimes saponiferous; sometimes tanniferous; phenolic acids, sinapic and ferulic acid common and 

sometimes abundant; mucilage in Commelinaceae at least. 

According to Gibbs, there is rather close agreement between 

the chemistries of Bromeliaceae and Commelinaceae at least. 

COMMELINACEAE (Figure 3lu-c).-Perennial or less commonly annual herbs, often succulent, erect to decumbent or 

rarely climbers, very rarely epiphytes, the stems usually 

articulated and the nodes swollen, sometimes tuberous, 

stoloniferous, rarely rhizomatous or bulbiferous; raphides 

present (not in Curtonema), silica bodies somtimes present; 

xylem vessel perforation plates simple in all organs, rarely also 

scalariform, or vessels absent from the stem and leaves 

(Curtonema); leaves spirally arranged or distichous along the 

stem, rarely in a rosette, often elongate, with a prominent 

midrib, more rarely the blade ovate to suborbicular, entire, the 

sheath closed, the stomates commonly tetra- or hexacytic, 

sometimes paracytic; sometimes a variety of hairs, e.g., 

3-celled glandular microhairs much larger than those of 

grasses, and macrohairs present; inflorescences terminal or 

axillary, rarely leaf-opposed or basal, simple or compound 

cincinnus, thyme, or fascicle, rarely 1-flowered, rarely spikelike (Curtonema), sometimes within a protecting bract; flowers 

usually actinomorphic, sometimes zygomorphic, bisexual or 

rarely the plants polygamo-monoecious, the parts mostly 

hypogynous; perianth heterochlamydeous, but sometimes the 

sepals are colored, sepals 3, free or rarely connate; petals 3, 

usually free, rarely connate, ephemeral and deliquescent; 

stamens q3-l), sometimes 2-4 converted to attractive 

staminodes, the filaments free or very rarely basally connate, 

very rarely basally epipetalous, sometimes bearded with 

brightly colored hairs, anthers usually less than twice as long 

as wide, rarely 1 larger than the others, basifixed, dorsifixed, 

or versatile, dehiscing longitudinally, or rarely by an apical 

pore; pollen 1-sulcate, rarely with additional apertures, 2-celled 

(3 in Floscopu scandens) when shed; nectaries absent; pistil 

1, the carpels 3 (2 in a few genera), style 1, the stigma usually 

1, acute or capitate, rarely trifia ovary 3(2)-locular, the ovules 

few to 1, rarely many per locule, uniseriate or biseriate, axile, 

bitegmic, crassinucellar, rarely tenuinucellar, orthotropous, 

hemianatropous, (or anatropous ?); embpyo sac antipodals 

ephemeral but persistent in Tinuntiu erectu; endosperm 

formation Nuclear, the tissue usually later becoming cellular; 

fruit a loculicidal capsule, rarely berry; seeds often sculptured, 

sometimes with an aril, rarely winged; embryo marginal, its 

position indicated by an embryotega, 0.3 the length of the 

endosperm, 1.3-1.7 times longer than wide; endosperm 

copious, mealy, starchy. Chromosomes: x = 4-20, especially 

6,8,9,12,14. 

Composition: -50 genera, -700 species. 

Distribution: Primarily pantropical and subtropical, ex- 
54 SMITHSOMAN CONTRIBUTIONS TO BOTANY 

d 

f 
NUMBER 71 55 

tending into the temperate zones, usually in mesic open or 

forest habitats. 

MAYACACEAE (Figure 3ld--.-Herbs of freshwater and 

marshes, often submerged, the stems filiform, sometimes 

branched; raphides, calcium oxalate crystals and silica absent; 

xylem vessel perforation plates scalariform in root, stem and 

leaf; leaves numerous, linear, spirally arranged along the stem, 

apically minutely bidentate, not sheathing, the stomates 

paracytic, sometimes uniseriate hairs present in the leaf axils; 

inflorescence subumbellate or a solitary flower, at the apex of 

the stem, the pedicels long, the flowers subtended by a 

membranous bract; flowers bisexual, actinomorphic, the parts 

hypogynous, nectaries absent; sepals 3, free, green; petals 3, 

free, corolline, clawed, stamens 3, free, the anthers basifixed, 

4-locular, dehiscing by an apical pore or short slit; pollen 

monosulcate; pistil 1, the carpels 3, style 1, the stigmas apical, 

1 or shortly trilobed, ovary unilocular, the ovules several, 

parietal, orthotropous; fruit a loculicidal capsule, the seeds 

small, subglobose and reticulately sculptured, the embryo 

apical, lying against the endosperm, underlying an embryotega, 

0.2-0.3 the length of the endosperm, 1.6 times wider than long 

to 1.3 times longer than wide; endosperm copious, mealy, 

starchy (?). Chromosomes: n = 8. 

Composition: 1 genus, -5 species. 

Distribution: Tropical America, West Indies, southeastern 

United States; one species in Angola. 

XYIUDACEAE (Figure 32u,b).-Perennial, less commonly 

annual herbs generally of wet habitats, with a short rhizome, 

rarely a bulb; raphides absent, but calcium oxalate crystals 

sometimes present; plant glabrous or with uniseriate mucussecreting glandular hairs on inner surface of leaf sheath, rarely 

with branched hairs, rarely entirely pilose; xylem vessel 

perforation plates simple in the root, stem and leaves; leaves 

mostly basal, distichous or spirally arranged, linear, or 

sometimes terete or flattened, the sheath short, open, the 

stomates paracytic or anomocytic; inflorescence scapose, rarely 

terminal on a leafy stem (Arutitiyopeu), head or dense spike, 

rarely loose (Achlyphilu), rarely paniculate, or the flower 

solitary, the flowers subtended by imbricate, leathery, usually 

rigid bracts; flowers bisexual, ephemeral, nectaries absent; 

sepals 3 (sometimes 2 in Abolbodu), all similar or two bractoid, 

the other membranous, usually free, petals 3, colored, 

FIGURE 31.40MMELINAcEAE: a, Dichoruandra thyrsgora flower from 

front and behind, pistil and calyx, stamens and petals, essential organs, pistil. 

D. tcjvccmir stamens; b, led showing the dd sheath, Comrnclina hlbcrosa 

1.8. of flower, Tredcscan~ia virgiica 1.s. d flower, dehisad fruit, embryo; c, 

Cmlina erecta inflorescence. C. -a flower with petala mnoved, 

stsminode, short stamen, long stamen, seed, 1.s. of same showing the mall 

embryo at margin of abundant mdoaperm, emtnyotega (after Martius, Bailloo). 

hhYACACm: d, Mapcafluviafih flower. 1,s. of same showing the parietal 

placentaion, seed. I.s. of same showing mall embryo in abundant endosperm; 

c, Mayca floral diagram, M. sellowhna views of mther showing apical pore, 

C.S. of same; fruit, same dehiscing and &ad, M. longipcs apex d led 

(greatly enlarged). part of plant in flower and fruit (&a Martius, Baillm). 

actinomorphic or zygomorphic (Orectunthe), connate, sometimes only slightly so; stamens 3 fertile, and 3 staminodal or 

lacking (Xyris luceru), very rarely all fertile (Xyris nigromucronutu), &ate to the perianth; anthers linear, extrorse or introrse, 

dehiscing longitudinally; pollen usually elongate, 1-sulcate, 

2(3)-sulculate or nonaperturate, 2-3-celled when shed, pistil 

1, the carpels 3, style 1, sometimes with 2-3 basal glandular 

appendages (Abolbodu, Orectanthe, Arutitiyopeu), the stigma 

apical, unlobed or 3-lobed, ovary superior, usually 1-locular 

and the ovules parietal, rarely 3-locular, sometimes imperfectly 

so, and the ovules axile (Abolbodu, Orectunthe, Arutitiyopeu); 

ovules usually numerous, sometimes few, bitegmic, tenuinucellar, orthotropous or anatropous (Abolboda, Orectunthe); 

embryo sac antipcdals sometimes soon degenerate; endosperm 

formation Nuclear, the tissue later becoming cellular; fruit a 

loculicidal capsule, the seeds numerous, small, usually 

longitudinally striate, embryo small, lying against the endosperm, 0.2 the length of the endosperm (0.5 in Abolboda, 

Orectunthe), 0.5 as long as wide; endosperm copious, mealy, 

starchy and sometimes also oily. Chromosomes: x = 8,9,13,17. 

Composition: 5 genera, -250 species. 

Distribution: Mainly pantropical and subtropical, mostly 

America, extending into temperate America and Africa. 

RAPATEACEAE (Figure 32c,d).-Perennial herbs often of 

wet habitats, rarely epiphytic, often large, with a short, thick 

rhizome; raphides and calcium oxalate crystals absent; silica 

bodies present; xylem vessel perforation plates usually 

scalariform in root and stem, rarely simple in them and leaves, 

vessels usually absent from the leaves; leaves distichous, linear, 

ensiform, rarely terete, basal, equitant, the sheath short, open, 

stomates paracytic, anomocytic or tetracytic; uniseriate mucusproducing hairs present; inflorescence scapose, head or 

subumbellate, often subtended by 1-2 large sometimes white 

bracts and the individual flowers subtended by several 

sometimes colored bracteoles; flowers bisexual, actinomorphic, nectaries absent; sepals 3, hyaline, rigid, free or connate; 

petals 3, cmlline, connate; stamens 6, sometimes adnate to 

the corolla, the filaments short or long, free or connate, the 

anthers linear, basifixed and sometimes apically produced, 

dehiscent by pore(@ or an apical cleft; pollen l(2)-sulcate ar 

zonisulculate, pistil 1, the carpels 3, style 1, the stigma apical, 

simple, often punctate; ovary superior, 3( 1)-locular, sometimes 

incompletely so above, the ovules several to one per locule, 

axile or basal, anatropous; fruit a loculicidal capsule, the 

seed(s) sometimes longitudinally striate, rarely carunculate; 

embryo lying against the endosperm, small, 0.1 the length of 

the endosperm, 3 times wider than long; endosperm copious, 

mealy, starchy. Chromosomes: n = 11 (Muschalocephalus). 

Composition: 16 genera, 80 species. 

Distribution: Tropical South America, 1 genus extending 

into Panama; 1 genus in tropical West Africa 

ERI~CAULACEAE (Figure 33).-Perennial, seldom annual 

herbs usually of wet habitats, emergent or rarely submerged, 

the stem short, rarely stoloniferous; raphides absent but 
56 SMITHSOMANCONTRIBUTIONS TO BOTANY 

d 

RoUm 32.-Xm=: a, Xyric hrifofrio inflmscmce showing the numerous imbricnte hcts, X. indica 

1.s. of flower, floral diagrun showing different typed of seprls, seed (greatly enl.rged), 1.s. of same showing 

the minute embryo in abundant endosperm; b, X. &x$ofia sepals, style and stigmas, C.S. of ovuy, fruit, one valve 

showing the numerous minute ads, Abolbo& pppigii flower, A. pmrchon views of anther, A. pppigii 

pistil showing the stylar appendages, C.S. of ovary, dehisd fruit (after Martius, Bai.llon). RAPATEACEAE: c, 

Schocnocephalium mariianum inflorescence, flower bud with bmcts, part of flower showing the inner perianth 

parts and stamens and one external perianth part, C.S. of perianth, perianth parts and stamens, stamen, perisnth 

part and base of filaments (enlarged), pistil, d, stigma, fNit, same dehisd, Rapafeo pafadma flower, I.s. of 

same (after Martius, Bdon). 
NUMBER 71 57 

, 

FTauRE 33.-ERIocAULAcBAE: a, Paepalanthw lanato-albur inflorescence, 6 flower, calyx of 6 flower, 6 

flower opmed out, 0 flower, sepal of 0 flower, papl of 3 flower, pistil showing the stylar appendages; b, 

Eriocadon cadescenr plant with many heads, E. septangdare 6 flower, 3 flower opmed, 1.s. of ovary, seed 

(enlarged), 1.s. of ssme showing minute anbryo in abundant endosperm. 6 floral diagram, 0 floral diagram 

(after Manius. Baillon, Le Maout and Deuisne). 

calcium oxalate crystals present, silica very rarely present; 

xylem vessel perforation plates usually simple and scalariform 

in root, stem and leaves; leaves usually basal, sometimes 

clustered at the end of the stem or scattered along it, spirally 

arranged, rarely distichous, linear, rarely ensiform, the sheath 

short or absent, the stomates paracytic; hairs 3-celled, 

glandular, unequal-armed T-shaped, or filamentous uniseriate 

ones often forming dense axillary clusters; inflorescence often 

hairy scapose heads, subtended by an involucre which is 

sometimes white; flowers very small, numerous, actinomorphic 

or zygomorphic, unisexual, the plants monoecious, very rarely 

dioecious; nectaries absent; perianth scarious, often white, 

segments 4 or 6, in 2 whorls, outer segments free or connate, 

the inner free or connate, sometimes with an apical gland, inner 

segments and sometimes also outer rarely absent and replaced 

by hairs; stamens 4 or 6(1-3), the filaments free or rarely 

connate or Anate to the base of the perianth; anthers slightly 

longer than wide, the sacs separate at the base and apex, 

longitudinally dehiscent; pollen spiraperturate, 3-celled when 

shed, pistil 1, the carpels 2-3, style 1, often appendaged, 

2-3-lobed, the lobes often bifid at the apex; ovary superior, 

2-3-locular, the ovules 1 per locule, axile-subapical, bitegmic, 

tenuinucellar, orthompous; embryo sac antipodal cells ephemeral; endosperm formation Nuclear or perhaps Helobial, the 

tissue later becoming cellular; fruit a thin-walled, loculicidal 

capsule; embryo lying against the endosperm, 0.2-0.4 the 

length of the endosperm, 0.5-0.8 as long as wide, endosperm 

copious, mealy or floury, starchy. Chromosomes: x = 8-10, 

especially 8. 

Composition: 13 genera, -1 150 species. 

Distribution: Mainly pantropical and subtropical, centered 

in South America, few temperate; eastern North America; 

Great Britain, Ireha eastern Asia. 

ZINGIBERALES 

Usually large herbs, usually supported by rigid leaf sheaths 

rather than the stem, rarely small herbs or trees, rhizomatous 

or tuberous; raphides in Musaceae, Strelitziaceae and Lowiaceae, 

absent from Zingikmceae, Cannaceae and Marantaceae; 

xylem vessel perforation plates simple or scalariform in the 

roots, rarely in the stem, vessels usually absent from the stem 

and leaves; leaves usually large, convolute in bud, spirally 

arranged, sometimes distichous or basal, the blade usually with 

a midrib and numerous lateral parallel veins extending from it, 

the petiole usually long, sheath usually open; inflorescence 

bracts often colored, flowers bisexual or the plants monoecious 
58 SMITHSONIANCONTRIBUTIONS M BOTANY 
NUMBER 71 59 

or rarely polygamous, zygomorphic or irregular, entomogamous or ornithogamous; perianth segments 6 in 2 whorls, 

homochlamydeous or heterochlamydeous often differing in 

size, connate or free; stamens 5(6)-1, epigynous or adnate to 

the corolla, 1-4 often petaloid staminodes, the filaments long 

or short, anthers linear, basifixed, rarely dorsifixed; pollen 

usually nonaperturate, sometimes forate or spirapemte, 

2-3-celled when shed, nectaries epigynous or septal; pistil 1, 

the carpels 3(2), style 1, the stigma apical, 3-lobed, bilabiate 

or not divided; ovary inferior, 3(2,l)-locular, the ovules usually 

numerous, rarely few-1 per locule, axile, parietal or basal, 

bitegmic, crassinucellar, anatropous or semi-anatropous; embryo sac antipodah ephemeral, rarely persisting after fertilization or multiplying secondarily; fiuit usually a loculicidal 

capsule or berry, rarely a schizocarp, the seeds often globose, 

sometimes arillate; embryo small to large, marginal or within 

perisperm; perisperm copious. 

Distribution: Pantropical and subtropical, few extending 

into temperate regions. 

Chemistry: Rich in phenolic acids; not alkaloid rich, 

sometimes producing indole alkaloids, sometimes tanniferous 

and producing flavonoids, mucilage sometimes present; 

sometimes aromatic. Characterization of individual families is 

difficult. Only Zingiberaceae stands out a little with very 

unusual flavonoids with no B-ring substitution, and a great 

variety of mono- and sesquiterpenoids. 

MUSACEAE (Figure 34u,b).-Large rhizomatous glabrous 

perennial herbs; raphides and silica bodies present; xylem 

vessel perforation plates simple and scalariform in the root, 

vessels absent from the stem and leaves; leaves large, convolute 

in bud, spirally arranged the blade entire, with a midrib and 

numerous parallel veins extending from it toward the margin, 

the sheath open, the stomates with several subsidiary cells only 

weakly differentiated from the other epidermal cells, guard 

cells symmetrical; inflorescence a raceme or spike of cymose 

clusters of flowers covered by colored bracts; flowers 

entomogamous and ornithogamous, zygomorphic or irregular, 

sometimes bisexual but usually the plants momious, c3 

flowers within the upper bracts, P within the lower bracts; 

perianth segments 6 in 2 petaloid whorls, 5 connate, 1 free, 

stamens 5, sometimes a staminode also present, the filaments 

filiform, free of the perianth; anthers linear, basifixed, 

2-locuk, pollen nonaperturate; pistil 1, the carpels 3, style 1, 

ROURE 34.-MUSACEAE: a, Mwa sinemis infloresance, young fruit and part 

of leaves (greatly reduced), Emefe edule flower, fruit, seed fran below and 

C.S. of same showing the pit a€ the Mum; b, Mwa sqpientvm flaral diagmn, 

flower, the free petal, M. coccinea flower. stamem and style. the free pe.tal, 

stigma (after b Macut and Decsisne, Martius). STRBLPIPACEAB: c, Rawnah 

(Phenakospcmuun) guyanemis flower, seed with hairy ad, 1.s. of same 

showing the modeme embryo, Ravenah ma&gascariensis seed, 1.s. of same, 

Heliconicr formosa I.s. of lower part of flower, H. metallic0 cs. d base d 

flower, abortive stamen; d, H. brasiliemis part of inflomcu~ce m flower and 

fruit. comlla opened out with lateral sepals. intermediate sepal, stamen, style, 

apex of style and stigma (after Martius, Le Maout and Dwaiane). 

elongate, the stigma 3-lobed, apical, ovary inferior, with septal 

nectaries, the nectar copious, 3-locular, the ovules axile, 

numerous, bitegmic, crassinucellar, anatropous; embryo sac 

antipodah ephemeral; endosperm formation Nuclear, the tissue 

later becoming cellular; fruit a berry, the seeds with an 

embryotega; embryo at the margin of the perisperm, 0.3 the 

length of the latter, twice as wide as long; perisperm copious, 

mealy, starchy, endosperm also present. Chromosomes: x = 

9-1 1,16,17. 

Composition: 2 genera, -40 species. 

Distribution: Tropical Africa, Ivladagascar; southeast Asia, 

the East Indies, Queensland. 

ST-ACEAE (Figure 34c,d).-Perennial rhizomatous 

usually glabrous herbs or trees; raphides and silica bodies 

present; xylem vessel perforation plates scalatiform and 

sometimes also simple in the roots, scalariform or vessels 

absent in the stem, vessels absent from the leaves: leaves 

distichous, large to very large, the blade with a midrib and 

numerous lateral parallel veins, the petiole long, with an open 

sheath, the stomates paracytic or with numerous subsidiary 

cells only weakly differentiated from the epidermal cells, guard 

cells symmeirical, some species with branched, uniseriate, 

candelabra-like hairs on the petiole and parts of the inflorescence (Heliconiu); inflorescences cincinni, terminal or in the 

axil of cymbiform bracts, the latter sometimes colored 

(Heliconiu); flowers large, zygomorphic, bisexual; perianth 

segments 6, often differing in size, shape and sometimes color, 

free or 5 connate and 1 free (Heliconiu); stamens epigynous 

or adnate to base of perianth, 5(6), the sixth often a petaloid 

staminode; filaments linear, the anthers 2-celled, linear, 

basifixed, pollen nonaperturate or aperture indistinctly 

f porelike (Heliconiu); pistil 1, the carpels 3, style 1, fdiform, 

the stigmas 3, fdiform or capitate; ovary inferior, with se.ptal 

nectaries, 3-locular, the ovules many or 1 per locule 

(Heliconiu), axile or basal (Heliconiu), bitegmic, crassinucellar, anatropous; embryo sac antipodals soon degenerate; 

endosperm formation Nuclear; fruit a loculicidal capsule or 

schizocarp (Heliconiu), the seeds globose, arillate or not arillate 

(Heliconiu), with an embryotega; embryo marginal or within 

the perisperm, 0.35-0.9 the length of the perisperm, 1.4 times 

wider than long to 7.5 times longer than wide; perisperm 

copious, mealy and starchy. Chromosomes: x = 7-13, 

especially 11,12. 

Composition: 4 genera, -60 species. 

Distribution: Moist tropical America; South Africa, 

Madagascar; New Guinea to Samoa. 

LOWIACEAE (Figure 35u).-Perennial rhizomatous, glabrous 

herbs; raphides and silica cells present; xylem vessel perforation plates scalariform in the roots, vessels absent from the stem 

and leaves; leaves basal, distichous, fairly large, with a midrib 

and numerous parallel veins extending apically from it, also 

conspicuous transverse veins, the petiole long, sheath open, the 

stomates paracytic, guard cells asymmetrical, inflorescence 

directly from the rhizome, a few-flowered cyme, the bracts 
60 SMllXSO~ANCONTRIBUTIONS TO BOTANY 

FIOURB 35.-LOWIACEAE: a, Orchkhntha hngijlora, flower, stamen, 0. maxillarwidos interior tepals, stamens 

and style. 1.s. of OVW, seed with aril (after Schumann). ZJNOIBERACBAE: b, Alpinia flower, 1.s. of same, C.S. 

of fruit, He+hiwn gardurianum floral diagram; c, flower, fruit, seed. 1.s. of same, Elctiaria cardamtnnwn 

dehiscing fruit, Rcnealmia racemosa flower, labellum; d, 1.5. and C.S. of ovary, fruit, views of seed, Rcnealmia 

adlate seed, 1.s. of same showing the embryo enclosed by little endoeperm (white) and much perisperm 

(stippled), C.S. of seed at level of embryo. embryo; c, Kaenpferia pandnrata inflorescence, the 3 inner perianth 

segments, anther enclosing the apex of the style betwem its lobes, style with 2 staminodes at its base, cs. of 

ovary. Costus spiralis stamen and apex of style, C. malortieanus cs. of anther. floral diagram (after Le Maout 

and Decaisne, Martius, Lindley). 
NUMBER 71 61 

colored but not very large; flowers large, zygomorphic, 

bisexual, malodorous; sepals 3, linear, connate basally into a 

slender tube; petals 3, the middle one large and colored, the 

laterals small; stamens 5, epigynous, unilateral, the filaments 

short, anthers 2-locular, linear, basifixed, the connective 

sometimes apically produced, pollen smooth, spheroidal, 

nonaperturate; pistil 1, the carpels 3, style fairly short, the 

stigmas 3, laciniate, wet; ovary inferior, the ovules numerous, 

axile, anatropous; fruit a loculicidal capsule, the seeds globose, 

arillate, their food reserve starchy; embryology unknown. 

Chromosomes: n = 9. 

Composition: 1 genus, 2 species. 

Distribution: Southern China, Hainan, Malay Peninsula, 

Borneo. 

ZINGIBERACEAE (Figure 35b-e).-Perennial herbs with 

elongate or tuberous rhizomes, often aromatic; plants not often 

conspicuously pubescent, but micro-hairs common, usually 

unicellular, rarely forked or stellate or 2-celled, commonly 

thick and lignified, others uniseriate, long, thin-walled and 

ephemeral except for base; raphides absent but calcium oxalate 

crystals and silica cells present; xylem vessel perforation plates 

in the roots usually scalariform, sometimes also simple, rarely 

scalariform in the stem, vessels usually absent from the stem 

and leaves; leaves basal or cauline, distichous or spirally 

arranged (Costoideae), often large, often divided into blade, 

petiole and sheath, with a ligule between the latter two, the 

blade with a midrib and numerous fine parallel veins extending 

obliquely upward from it, the sheath open or closed 

(Costoideae), the stomates mostly temytic, sometimes 

paracytic or with more than 4 subsidiary cells, guard cells 

asymmetrical; inflorescence terminal, sometimes scapose, 

raceme, spike, head, cyme, or the flower solitary, the bracts 

often colored; flowers zygomorphic, bisexual or rarely the 

plants polygamous; sepals 3, calycine, connate; petals 3, 

connate, one usually larger than the others, colored and showy, 

sometimes pouched; stamen 1, adnate to the corolla, the 

filament long or short, sometimes petaloid, the anther adnate, 

rarely dorsifixed, elongate, 2-locular, sometimes spurred, the 

connective sometimes appendaged and sometimes separating 

the locules, 2 or 4 petaloid staminodes usually present; pollen 

usually nonaperturate, forate or spiraperturate or k intermediate 

between both in Costoideae, 2-celled when shed; nectaries 

epigynous or septal (Costoideae); pistil 1, the carpels 3(2), style 

1, the upper part usually enveloped by the anther, the stigma 

capitate or 2-lipped, or rayed, wet; ovary inferior, 3(2)-locular, 

the ovules numerous, axile, or 1-locular and the ovules parietal, 

or rarely basal and the ovules few, bitegmic, crassinucellar, 

anatropous or semi-anatropous; embryo sac antipodak ephemeral; endosperm formation Helobial, the tissue ultimately 

cellular; fruit usually a loculicidal capsule, sometimes a berry, 

the seeds often covered by an aril; embryo 0.8 to as long as the 

perisperm, 3.6-7.0 times longer than wide; endosperm small, 

hard or mealy, perisperm large, mealy or hard, starchy. 

Chromosomes: x = 7-14,16,17,21,26, especially 11,12. 

Composition: 49 genera, -1OOO species. 

Distribution: Centered in Indomalaysix northern 2/3 of 

South America; southern 2/3 of Africa, Madagascar, East 

Indies, northeastern Australia. 

CANNACEAE (Figure 36u,b).-hge perennial herbs, sometimes of wet habitats, glabrous, usually with tuberous 

rhizomes; raphides absent, but calcium oxalate crystals and 

silica cells present; xylem vessel perforation plates both simple 

and scalariform in the root, vessels absent from the stem and 

leaves; leaves cauline, spirally arranged, large, broad, the blade 

with a strong midrib and numerous lateral veins extending from 

it, the sheath open, ligule and pulvinus absent, the stomates 

with 2, 4, or more subsidiary cells, guard cells usually 

symmetrical, inflorescence terminal, composed of cincinni 

arranged in racemes or panicles; flowers bisexual, irregular, 

mostly large and showy; sepals 3, free, herbaceous; petals 3, 

one usually smaller than the others, basally connate; stamen 

1, adnate to the corolla, the anther unilocular, adnate to one 

side of the petaloid filament; staminodes 3(1,4), petaloid, 2 

shortly basally connate and adnate to the petals; pollen 

nonaperturate, 3-celled when shed; pistil 1, the carpels 3, style 

1, petaloid, the stigma apical, linear; ovary inferior, with septal 

nectaries, 3-locular, the ovules numerous, axile, bitegmic, 

crassinucellar, anatropous; endosperm formation Nuclear 

(possibly Helobial), the tissue ultimately cellular, fruit a 

capsule, often warty, sometimes indehiscent; seeds round, the 

embryo straight, 0.8-0.9 the length of the perisperm, 2.4-4.3 

times longer than wide; endosperm sparse, perisperm copious, 

very hard, starchy. Chromosomes: x = 9. 

Composition: 1 genus, -50 species. 

Distribution: Southeastern United States, tropical and 

subtropical America, West Indies; according to some authors 

1-3 species are indigenous to Africa and Asia. 

MARANTACEAE (Figure 36c,d).-Perennial herbs, rarely 

shrubby, often with a tuberous rhizome, often of moist or 

swampy forests; raphides absent but calcium oxalate crystals 

and silica cells present; xylem vessel perforation plates simple 

or scalariform in the root and stem; leaves distichous along the 

stem or basal, differentiated into an asymmetrical blade, 

petiole, and open sheath, with a pulvinus at the apex of the 

petiole, with a distinct midrib and numerous lateral parallel 

veins, the stomates with 2,4, or more subsidiary cells, guard 

cells asymmetrical; plants generally glabrous to the naked eye, 

the hairs usually unicellular, typically sunk in the epidermis; 

inflorescence terminal, sometimes scapose, sometimes on a 

separate shoot (Thymocarpus, some Calathea), sometimes a 

sympodium of racemes (Thymocarpus), cymose, spike- or 

panicle-like or capitate, the bracts sometimes colored, flowers 

bisexual, irregular; sepals 3, free, petals 3, unequal, connate; 

stamen 1, the anther unilocular, adnate to one side of the 

petaloid filament; staminodes 2-4, petaloid, adnate to the 

corolla, sometimes 2 connate; pollen nonaperturate; pistil 1, 

the carpels 3, two usually aborting, the style stout, curved, 

stigma apical; ovary inferior, with septal nectaries, 1(3)-locular, 
62 SMITHSOMANCONTRIBUTIONS TO BOTANY 

0 

d 

ROUW ~~.~XTNACBAB: a, Canna &ucCr flower, C. dcnvdota labellurn, style and stamen, C. coccinca 

inflorescmce on pnrt of leaf, 1.8. and cs. of immature fn& b, floral diagram, C.Jaccida stamen (pan petaloid), 

C. indica fruit, seed. 1.s. of same (enlarged). embryo, Cam 1.s. of anbryo (after Martius, Le Maout and 

Decaisne, Lindley). hlU”MaR c, Calatka villosa flower, sepal, petal, stamen, cs. and 1.s. of ovary, C. 

cichlcri stamm; staminode, stamen and style; mlla, calyx and ovary; d, C. micam views of seed. 1.s. of sme 

ahowing the curved embryo. C. brchcmi0M base of blade and enlarged apex of petiole, Thfia deafbata fruit. 

C.S. d same with only one fertile cell floral diagram (after Mutius. Le Maout and Decaisne, Lindley). 
NUMBER 71 63 

FroURe ~~.-HAEMODORACEAE: a, Haemodonon spuatwn flower spd out, apex of stamen. C.S. of ovary, H. 

teretij'diwn 1s. of flower, fruit, Haemo&rwn seed. sdm of endosperm showing the small embryo in abundant 

endosperm, embryo; 6, ConoJtylip serratij'olia 1s. of flower, PhMocatya)il~olia I.s. of flower, Anigosanthw 

flower, 1 .s. of same. seed. 1.s. of same, stamen, floral diagram (after Baillon, Le Maout and Decpisne, My). 

PHILYDRACEAE: c, Pritzclh pygmaea #ant in bloom, flower, flaal diagram, Philydivm lanuginoswn petah and 

stamen, dehisced fruit, sd, 1.s. of same showing the moderate embryo in abundant endosperm (after Bdm). 

IRIDACEAE: d, Iris germanica habit showing the quitant, distichous leaves and Ihiume, flower, 1.s. of ssme 

showing parts of 4 perianth segments, one broadened entire style arm and an&r in I.s. with a stamen heath 

it, flower with perianth and stamens moved. showing the style arms and ovary (after Le Maout and Decaisne). 
64 SMlTHSONIANCONTRIBUTIONS TO BOTANY 

the ovules 1 per locule, mile-basal, bitegmic, crassinucellar, 

anmopous or anacampylotropous; embryo sac antipodals 

ephemeral or persisting at least until fertilization or multiplying 

secondarily; endosperm formation Nuclear; fruit a loculicidal 

capsule or berry, sometimes muricate (Truchyphynium, 

Thymocurpus) or papillose (some Culutheu), the seeds 

sometimes arillate; embryo curved or folded, 1.4-1.6 times 

longer than the perisperm, 7.3-25 times longer than wide; 

perisperm copious, mealy, starchy. Chromosomes: x = 

4,6,7,9,11-13, especially 9,12,13. 

Composition: -30 genera, -350 species. 

Distribution: Pantropical, centered in America, extending 

into temperate America. 

IRIDALES 

Herbs, very rarely low shrubs or saprophytic, with rhizomes, 

corms, tubers or stolons, rarely bulbs; xylem vessel perforation 

plates simple or scalariform in the roots, scalarifom or vessels 

absent in the stem and leaves; leaves mostly basal, linear to 

ensiform, distichous, often equitant, the sheath short, open; 

flowers bisexual, actinomorphic or zygomorphic; perianth 

segments 6(3,4), petaloid, connate or free; stamens 3(6,1), 

epigynous or adnate to the perianth, the filaments free or 

connate, anthers basifixed or dorsifixed; pollen usually 

1-sulcate, less often 2(3)-sulculate, spirapertur(oid)ate, nonaperturate, polyrugoidate, 2(-4)-porate, or 1(-3)-ulcerate, 

2(-3)-celled when shed, pistil 1, the carpels 3, styles 1 or 3, 

the stigmas apical or decurrent ventrally; ovary usually inferior, 

less commonly semi-inferior or superior, 3(-1)-locular and the 

ovules mile or rarely 1-locular and the ovules parietal, usually 

numerous, rarely few or only 1 per locule, bitegmic, 

crassinucellar, rarely tenuinucellar, anatropous, rarely hemianatropous; embryo sac antipodal cells soon degenerate or 

sometimes persisting into early endosperm formation; endosperm formation Nuclear or Helobial; fruit a loculicidal 

capsule, very rarely dehiscing by slits or circumscissile, very 

rarely a berry, the seeds usually numerous; embryo enclosed 

in the endosperm, small to moderate in size; endosperm usually 

copious, rarely scant or absent, usually hard. 

Distribution: Cosmopolitan, centered in the tropics, South 

Africa and Australia. 

Chemistry: Often tanniferous; sometimes saponiferous; 

rarely cyanogenic; alkaloids absent; Iridaceae, fiilydraceae 

and Burmanniaceae (except Thimiu) lack raphides, Haemodoraceae usually has them. Iridaceae often produce mangiferin 

and sometimes naphthaquinones or anthraquinones. Isophysis 

is chemically in line with membership in Iridaceae. Haemodoraceae usually have a characteristic polyphenolic red 

pigment (phenalones) in the roots and rhizomes. 

HAEMODORACEAE (Figure 37u,b).-Perennial herbs with 

tubers or short rhizomes, or sometimes stolons, stems and roots 

often with red sap; raphides usually present; xylem vessel 

perforation plates simple and scalariform in the roots, 

scalariform or vessels absent in the stem and leaves; leaves 

basal, linear to ensiform, distichous, equitant, the sheath short, 

open or closed, the stomates paracytic, or nearly anomocytic 

(bphiolu); inflorescence cymose, arranged in racemes or 

panicles, or subcapitate, often densely villous; flowers 

bisexual, actinomorphic or slightly zygomorphic; perianth 

segments 6, in 1 or 2 series, connate or free, often woolly; 

stamens 3 or 6, sometimes 2-3 staminodes present, 1 plus 2 

staminodes in Pyrrorhizu, the filaments free from one another, 

adnate to the perianth; anthers elongate, basifmed or dorsifixed, 

introrse, dehiscing longitudinally; pollen 1 -sulcate, 2(-4)- 

prate, 2-celled when shed; pistil 1, the carpels 3, style 1, 

elongate, the stigma apical, acute or capitate; ovary superior, 

semi-inferior or inferior, with septal nectaries, 3-locular or 

1-Zlocular by abortion, the ovules numerous to 1 per locule, 

axile, bitegmic, crassinucellar, hemianatropous to anatropous; 

embryo sac antipodals soon degenerate; endosperm formation 

Helobial, the tissue ultimately cellular, fruit a loculicidal 

capsule, rarely indehiscent (Bwberettu); embryo 0.1 the length 

of the endosperm, as long as wide, rarely rather large; 

endosperm copious, fleshy, mostly starch, some protein, oil 

and hemicellulose. Chromosomes: x = 4-8,10,15, especially 

6,8. 

Composition: -15 genera, -75 species. 

Distribution: Centered in eastern and western Australia; 

New Guinea; South Africa; tropical America, eastern United 

states. 

PHILYDRACEAE (Figure 37c).-Perennial herbs of wet 

habitats, often floccose, the hairs uniseriate, sometimes 

glandular, sometimes with a short rhizome; raphides present 

(?), styloid crystals sometimes present; xylem vessel perforation plates scalariform in the roots, vessels absent from the stem 

and leaves; leaves basal with a short open sheath, distichous, 

equitant, linear or ensiform, the stomates paracytic or 

tetracytic; inflorescence a simple or compound spike, sometimes woolly; flowers bisexual, zygomorphic, covered by a 

bract in bud; perianth corolline, the segments 4, biseriate, the 

inner much smaller than the outer, free or basally connate, 

marcescent; stamen 1, free or adnate to the inner tepals, the 

filament elongate, flattened; anther straight, arcuate, or 

longitudinally coiled; pollen sometimes in tetrads, 2-celled 

when shed, 1-sulcate, rarely 3- or zonisulculate ?; pistil 1, the 

carpels 3, style 1, elongate, persistent, the stigma capitate, dry; 

FmUW ~~.--IRIDAcEAE: a, Iris dehisced fruit, 1.s. of seed, floral diagram 

showing the stigmas opposite the stamens, Bekamcanda fld diagram showing 

the stigmas alternate with the stamens, Tigridio mcleagris essential organs, 3 

stamens forming a column and 6 fUom style arms, lateral view and fmn 

above; b, Sisyrinchivm sellowianum inflorescence, Geosiris aphylla 2 plants 

in flower, inflorescence, flower with 2 spathes, anther, essential organs, pistil, 

c, fruit, cs. of same, bmch of placenta with seeds, floral diagram (after Le 

Macut and Decaisne. Jder, Martius). BURMANNUCEAE: d, Bw~laMicr 

bkolor plant, flower, 1.11. of same, Apteria setacea plant. flower laid open; e, 

Bwmannio sellovianu part of inflorescence, flower opened out, floral diagram, 

Dictymtega floral diagram (after Martius, Baillon, Le Maout and Decaisne). 
NUMBER 71 65 

a 

d 

e 

I 
66 SMITHSONIANCONTRIBUTIONS To BOTANY 

ovary superior, without septal nectaries, 3-locular and the 

ovules axile, or incompletely 3-locular and the ovules parietal; 

ovules numerous, bitegmic, crassinucellar, anatropous; embryo 

sac antipodal cells degenerate soon after fertilization; endosperm formation Helobial, the tissue ultimately cellular; fruit 

mostly a loculicidal capsule, rarely indehiscent; seeds numerous, minute, sculptured, tailed at both ends; embryo straight, 

in the axis of the endosperm, 0.8 the length of the latter, 5.5 

times longer than wide; endosperm moderate, starchy, sometimes also oil and protein present. Chromosomes: x = 8,17, 

especially 8. 

Composition: 4 genera, 5 species. 

Distribution: Eastern and southwestern Australia; southeastern China, Indochina, New Guinea. 

IRIDACEAE (Figures 37d, 3&-c).-Perennial, rarely annual 

(Sisyrinchium spp.) herbs or very rarely low shrubs, with 

corms, rhizomes, tubers or rarely bulbs, very rarely a colorless 

saprophyte (Geosiris); raphides absent, but often calcium 

oxalate crystals present, rarely hairy; xylem vessel perforation 

plates simple or scalariform or both, but vessels frequently 

absent from the stem and leaves; leaves basal, often ensiform, 

or linear, equitant and distichous, very rarely scales (Geosiris); 

inflorescences terminal, cymose, arranged in spikes, racemes 

or panicles, or the flower solitary, the spathes very rarely 

colored (Oenostuchys); flowers bisexual, actinomorphic or 

zygomorphic: perianth petaloid, marcescent, the segments 6, 

2-seriate, free or connate, rarely with a spur or sac at the base 

(Kentrosiphon); stamens 3, rarely one of them staminodal 

(Diplurrhenu), epigynous or adnate to the perianth, the 

filaments free or connate, anthers basifixed or dorsifixed, 

extrorse; pollen usually 1-sulcate, less often 2-sulculate, 

spiraperau(oid)ate, nonaperturate, or polyrugoidate, 2(3)- 

celled when shed, pistil 1, the carpels 3, styles 1 or 3, the 

stigmas 3, sometimes divided at the apex, apical or decurrent 

ventrally, sometimes the style branches petaloid; nectaries on 

the perianth, frlaments or septa of the ovary; ovary inferior, 

very rarely superior (Isophysis), 3-locular and the ovules axile 

or very rarely 1-locular and the ovules parietal (Hermodactylus); ovules usually numerous, rarely few or one per locule, 

bitegmic, crassinucellar, anatrops; embryo sac antipodah 

sometimes persistent into early endosperm formation: endosperm formation Nuclear; fruit a loculicidal capsule; embryo 

enclosed in the endosperm, 0.4-0.7 the length of the 

endosperm, 4-7 times longer than wide; endosperm copious, 

hard, with hemicellulose, protein and oil, rarely also starch 

(Rudinosiphon). Chromosomes: x = 3-19,223, especially 

8-10.15. 

Composition: -60 genera, -1500 species. 

Distribution: Cosmopolitan, except the arctic, centemi in 

South Africa and tropical America, to a lesser extent in eastern 

Meditenanean region. 

BURMANNTACEAE (Figures 38d,e, 39u,b).--Sapmphytic chlorophylless or green autotrophic annual or perennial herbs, 

sometimes with a creeping rhizome or small tuber: raphides 

sometimes present; xylem vessel perforation plates scalariform 

in the root, stem and leaves; leaves reduced to scales or linear 

or lanceolate, entire, sometimes equitant, basal or alternate all 

along the stem, the sheath short, open, the stomates mostly 

anomocytic, sometimes absent; inflorescence a terminal 

dichasial or monochasial cyme, or a solitary flower, sometimes 

with colored bracts; flowers bisexual, actinomorphic, or the 

corolla rarely zygomorphic (Ophiomeris, Afrothism’u); perianth corolline, of 6(3) segments connate in a tube, sometimes 

winged, the outer often different in size or shape from the 

inner, sometimes appendiculate, sometimes persistent; stamens 

3 or 6, usually adnate high, rarely low, on the perianth-tube, 

subsessile, sometimes connate in a ring or deflexed tube, the 

connective often broadened and apically produced, and the 

anther sacs separate, dehiscence latrorse or introrse; pollen 

nonaperturate or with I(-3)-ulcerate, circular or slightly 

elongate apertures, 1-sulcate(?), 2-3-celled when shed; pistil 

1, the carpels 3, style 1, the stigmas 3, apical, usually enlarged, 

often capitate, sometimes each bifid; ovary inferior, with septal 

or epigynous nectaries, 3- or 1-locular, the ovules very 

numerous, axile or parietal, bitegmic, tenuinucelk, embryo 

sac with 3 small antipodal cells; endosperm formation 

Helobial, the tissue cellular at time of zygote division; fruit 

usually a capsule, sometimes fleshy, sometimes winged, 

dehiscent by 3 longitudinal slits or by numerous transverse 

slits, or circumscissile, the seeds numerous, elongate, minute 

but larger than those of most orchids, the funicle often rather 

long; embryo arrested in 4-10-celled stage, further development occurring after the seeds are shed, endosperm scant or 

absent in the mature seeds. Chromosomes: x = 6,8,9. 

Composition: -17 genera, -125 species. 

Distribution: Panaopical, mainly in rain forest, some in 

savanna or grassy fields, few in temperate regions. 

ORCHIDALES 

Herbs, rarely saprophytic, often epiphytic or climbing in the 

tropics, sometimes with short rhizomes, tubers or swelling of 

the stem; raphides probably universal, silica cells sometimes 

present; xylem vessel perforation plates usually simple in the 

roots, sometimes also scalariform, vessels present or absent in 

the stem and leaves; leaves simple, entire, rarely plicate, often 

distichous, reduced to scales in saprophytes, the sheath nearly 

always closed; inflorescence racemose; flowers zygomorphic, 

bisexual or very rarely the plants polygamous, monoecious or 

dioecious; perianth segments 6 in 2 whorls, mostly petaloid, 

free or connate in each whorl, frequently one segment with a 

nectariferous spur; stamens 1 or 2, rarely 3 or 6, adnate to the 

style; pollen 1-sulcate, 2-sulculate(?), 3-4-poroid or nonaperturate, mostly in pollinia, 2-celled when shed; pistil 1, the 

carpels 3, the stigma short, apical or subapical; ovary inferior, 

unilocular, the ovules very numerous, parietal, very rarely 

3-locular and the ovules axile, usually bitegmic, rarely 

unitegmic, tenuinucellar, anatrops; embryo sac antipodals 
NUMBER 71 67 

\I 

d 

FICIIJRE BURMAN" MAN"^ a, Bvrm~nia dirticha plant, dehiscing atsmen, style and stigmas, fmit, C.S. of 

same, sed, 1.1. of same, mbvo at base of endosperm; b, Geomitra clavigera flower (greatly drged). 1.s. of 

same (after IL Maout and Decairne, Bailloo). C~RSIACBAE: c, Corsk ornafa flower, 1 .s. d same (after Bdon). 

ORCHIDACEAE: d, Aposratia odorota flower, stamens and style, GI. of ovary, seed, Apratia floral diagram, 

Neuwiedia floral diagram (after Lindley, Le Maout and Decdsne). 
68 SMITHSONIANCONTRIBUTIONS TO BOTANY 

ephemeral and frequently not all 3 produced, endosperm 

formation absent or Nuclear, the nuclei few; fruit a capsule 

mostly dehiscing by longitudinal slits, very rarely somewhat 

berried, seeds very numerous, minute, the embryo immature 

when the seeds are. shed; endosperm absent, very rarely scanty 

in the mature seeds. 

Distribution: Cosmopolitan, most diverse and abundant 

in the tropics. 

Chemistry: Many have alkaloids, including alkaloid mines, 

a group of indole alkaloids peculiar to orchids, indolizidines, 

pyrrolidines and pyrrolizidines; saponins uncommon; not 

cyanogenic; no leucoanthocyanins; mucilage in many; occasionally tanniferous. 

CORSIACEAE (Figure 39c).-Chlorophylless saprophytic 

perennial herbs with a short rhizome or small tubers; leaves 

reduced to scales alternate all along the stem; inflorescence a 

solitary terminal flower, flower bisexual or unisexual, zygomorphic; perianth parts 6, free, corolline, the median one of 

the outer series enlarged, usually with nectariferous tissue at 

its base, and enclosing 5 linear tepals; stamens 6, the filaments 

short, close to the style and adnate to it, or perhaps some 

epigynous, or adnate to the base of the large tepa, anthers as 

long as wide, subbasally dorsifixed, extrorse; pollen 1-sulcate, 

the sulcus not very distinct; pistil 1, the carpels 3, style short, 

the stigmas 3, short, not enlarged; ovary inferior, the ovules 

numerous, parietal; fruit a capsule dehiscing by 3 valves; seeds 

numerous, minute, elongate, with a long funicle, the testa 

obscurely longitudinally grooved; embryo undifferentiated, 

endosperm scanty. Chromosomes: x = 9. 

Composition: 2 genera, -10 species. 

Distribution: Centered in New Guinea; Queensland, Solomon Islands; one species in Chile (Aruchnitis unifrora). 

ORCHIDACEAE (Figures 39d-4 I).-Perennial herbs (one 

species annual), rarely half-shrubs, obligately mycorrhizal 

(except some Arethuseae), rarely saprophytic, rarely climbers 

(Vanilla), the stem mostly sympodial, sometimes monopodial, 

often thickened basally and with aerial roots often with 

2-many cell layers of velamen, rhizomes or roots often 

tuberous; plants usually appearing glabrous, but sometimes 

various kinds of hairs present in different species; raphides 

present, silica cells sometimes present; xylem vessel perforation plates usually simple in the roots, sometimes also 

scalariform, vessels present or absent in the stem and leaves; 

leaves simple, entire, rarely plicate, alternate, rarely opposite 

or whorled, often distichous, rarely deciduous in the dry 

season, or absent, often fleshy, reduced to scales in the 

saprophytes, various in shape, rarely reticulate-veined, the 

sheath nearly always closed, smmates paracytic, less often 

anomocytic, seldom tetracytic; inflorescence terminal or 

axillary, sometimes scapose, spikes, racemes or panicles or a 

solitary flower, rarely capitate; flowers zygomorphic, usually 

resupinate, rarely the corolla nearly actinomorphic (Apostasia), 

bisexual or very rarely the plants polygamous, monoecious or 

dioecious; perianth segments 6 in two whorls, usually petaloid, 

more rarely the outer whorl sepaloid; the segments often differ 

in size, shape and color, free or connate in each whorl, 

frequently a petal or sepal is prolonged into a nectariferous 

spur, sometimes extraflod nectaries present; stamens 2 (the 

third sometimes a staminode covering the anthers), or 1 

(sometimes with 2 staminodes), rarely 3 abaxial, 2 of the inner 

cycle and 1 of the outer (Neuwiedia), adnate to the style and 

the latter sometimes adnate to a petal, pollen 1-sulcate, 

2-sulculate, 3-4-poroid or nonaperturate, in pollinia often 

elaborated by caudicles (formed from differentiated pollen 

grains) and viscidia or stipes (formed from part of the stigma 

or style), more rarely single, in tetrads or masses, the individual 

grains 2-celled when shed; pistil 1, the carpels 3, stigmas 3 

fertile, or 2 and the third partially to completely transformed 

into a rostellum; ovary inferior, sometimes with nectaries in 

the wall between the carpels, unilocular, very rarely ailocular 

(Apostasia, Neuwiedia, Selenipedium, Phragmipedium), the 

ovules very numerous, minute, parietal or rarely axile, usually 

bitegmic, rarely unitegmic, tenuinucellar, anatropous; embryo 

sac antipodals ephemeral and frequently not all 3 are produced; 

endosperm formation, when it occurs, Nuclear, and then only 

few nuclei are produced; fruit a capsule, mostly opening by 3 

or 6 longitudinal slits, rarely a berry (Neuwiediu curtisii and 

some Vanilla); seeds very numerous, minute, slightly larger 

and black in Apostusiu and Vanilla, the embryo always 

immature when the seeds are shed; endosperm absent in the 

mature seed. Chromosomes: x = 6-16,18,20,21,24, especially 

10,16,18,20,21. 

Composition: -700 genera, -15000 species. 

Distribution: Cosmopolitan, in almost all habitats, most 

diverse and abundant in the tropics where the majority are 

epiphytes in forests; almost all temperate and all arctic genera 

are terrestrial. 

l%m ~O.-ORCHIDACEAE: a, Orchis fld diagram, flower, smne enlarged 

without ovary (ST = stigma, L = anther cell, P = pollinim. R = retinadun), 

Skanhopea ocdaia flower, b, Habenoria achalermis infiarescence, flower, 

column, 1 .s. of ssune (below) showing the entrance to the spur, pollinium, c.a. 

of ovary, H. areckrvoletae C.S. of ovary; c, Selenipediwn palmfolium sepal, 

petal, labellum, S. kabelicmum 3 views of column, 1.8. of same, S. palm#olium 

C.S. of wary; d, Epktephiwn sclerophyllwn lateral sepal, ped, labellum and 

column, 2 views of column, wary, Stenorrhynchos argentinus labellum, 

column @oth in lateral view); c, pollinia of Malaxis paludoscl, Phaius 

tankruillii. Brawia maculata, Opb apifera, Orchis, pprt of Orchis 

pollinium gnotly enlarged, pollen of Stenorrhynch sjxcwsus, EpidcndrMl 

anther without the pollinia, pollinia (after Martius, Lindley, Le Maout and 

Decaisne). 
NUMBER 71 69 
70 SMITHSONIANCONTRIBUl"I0NS TO BOTANY 

RCWRR 41.-oRCHIDACBAB: a, vaftiiia chamissonis labehnn, appendiced of same. column, 1.s. of same, cs. 

of anther, Vmiila cs. of fruit, dehisced fruit; b, column a4 Arethupo, Stenorrhynchos, Brassia maculota, Orchis 

IIYU&, Ophrys apifcra C.S. of fruit, seed (enlarged), Erythrdes peterionvs seed (greatly enlarged), 

Stenorrhynchar argenlinlu reed (greatly adarged); c, Orchis debisced fruit leaving in place the 3 medim nerves 

of the carpels, Fermndczia acuta dchisced fruit, diagram of same; d, Angraecwn dehisced fruit, diagram of 

same, Plewothallis ckalua dehiaced fruit, diagram of same; e, Epidcndrwn dehisced fruit, Coryanthes speciaro 

flower, Biphih polysyka lateral sepal (after Martius, Le Maout and Decpisne, Mdley). 
References 

Arber, A. 

1925. Monocofyledonr: A Morphologigal Study. Cambridge, England. 

Axelrod, D.L. 

1960. The Evolution of Flowering Plants. In S. Tax, editor, The Evolution 

of Life, pages 227-305. Chicago. 

Bailey, I.W. 

1957. 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. 

BaiUon, H.E. 

Bedell, H.G., and J.L. Reveal 

1894-1895. 

1982. 

Histoire des Plantes. Volumes 12, 13. Paris. 

Amended Outlines and Indices for Six Recently Published Systems 

of Angiosperm Classification. Phytologia, 51x55-156. 

1979. Plant Classification. 2nd edition. Lexington, Mass. 

1862-1883. Genera Plantarwn. 3 Volumes. hdon. 

1915. Phylogenetic Taxonomy of Flowering Plants. Annals of the Missouri 

Botanical Garden, 2:109-164. 

Heresy Revived The Monomt Theory of Angiosperm Origin. 

Evolutionary Theory, 5~189-225. 

1986. A Phylogenetic Analysis of the Orchidaceae. Smithsonian Contributions to Botany, 61: 79 pages. 

Cain, S.A. 

1944. Foundations of Plant Geography. New York and London. 

Cave, M.S., editor 

1958-1%5. Index to Plant Chromosome Numbers for 1956-1964. 2 

Bmson, L.D. 

Bentham. G., and J.D. Hooker 

Bessey, C.E. 

Burger, W.C. 

1981. 

B~s-Balogh, P., and V.A. Funk 

volumes. Chapel Hill, North Carolina. 

1943. The Origin and Certain Trends of Specialization in the Monocotyledoneae. Ameruan Jownal of Botany, 3O:ll-17. 

1953. Independent Origin of Vessels in the Monocotyledons and 

Dicotyledons. Phytomorphology, 322-44. 

1955. The Systematics of Angiosperms. In A Century of Progress in the 

Natural Sciences, 1853-1953, pages 405-483. San Francisco: 

California Academy of Sciences. 

1964. Systematic Botany-An Unending Synthesis. Tpron, 13957-273. 

1981. An Integrated System of Classfiation of Flowering Plants. New 

Yolk. 

1988. The Evolution and Class#cation of Flowering Plants. Second 

edition. New York. 

Curtis, w. 

1787-1983. 

Dahlgren, R.M.T. 

1975. 

Cheadle, V.I. 

Constance, L. 

Cronquist, A. 

The Botanical Magazine. Volumes 1-182. London. 

A System of Classification of the Angiospenns to be Used to 

Demonstrate the Distribution of Characters. Botaniska Notiser, 

1980. A Revised System of Classification of the Angiosperms. Botanical 

Journal of the Linnean Society, 80.91-124. 

1983. General Aspects of Angiosperm Evolution and Macrosysteanatics. 

Nordic Journal of Botany, 3:119-149. 

128~119-147. 

Dahlgren, R.M.T.. and H.T. Clifford 

Dahlgren, R.M.T., H.T. Clifford, and P.F. Ye0 

1982 The Monocotyledonr: A Comparative Study. hdon and New York. 

1985. The Families of the Monocotyledom: Structure, Evoluiwn, and 

Taxonomy. Berlin, Heidelberg. New York, and Tokyo. 

Monocotyledon Evolution: Characters and Phylogenetic Estimation. 

In M.K. Hecht, B. Wallace, and G.T. Prance, editors. Evolutionary 

Biology, 16:255-395. 

Dahlgren, R.M.T., and F.N. Rasmussen 

1983. 

Daumann, E. 

1970. Das Blutennektanum der Monocotyledonen unter besonderer 

Berucksichtigung seiner systematischen und phylogenetischen 

Bedeutung. Feddes Repertorium Specierwn Novarwn Regni Vegetabilk, 80:463-590. 

Davidse, G.. editor 

1975. The Bases of Angiosperm Phylogeny. Annals of the Missouri 

Botanical Garden, 62515-834. 

Davis, G.L. 

1966. Systematic Embryology of the Angwsperm. New York. 

Davis, P.H., and V.H. Heywood 

1963. Principles of Angiosperm Tpronomy. hdon. 

1955. The Evolution of the Plants and the Taxonomy of the Monocotyledons. Sbornik Narodniho Musea v Praze, 11B(6):1-143. 

DeyZ M. 

Dressler, RL. 

1981. The Orchids, Natural History and Classijkztwn. Cambridge, 

Massachusetts. 

1960. Classification and Phylogeny in the Orchidaceae. Annals of the 

Missouri Botanical Garden, 47:25-68. 

Eames, A. 

1961. Morphology of the Angiosperm. New Yolk. 

Ehdorfer, F., and R. Dahlgm. editors 

1983. New Evidence of Relationships and Modem Systems of Classification of the Angiosperms. Nordic Journal cfBotany, 3:3-155. 

1980. Phylogenetic Patterns and the E~olutW~ry Process. New York. 

1960. Lea Vegetaux vasculaires. In M. Chadefaud et L. Emberger, Traite

de Botanique Systematique. Paris. 

Engler, A. 

Dressler, R.L., and C.H. Dodson 

Eldredge, N., and J. Cracraft 

Emberger, L 

1881-1988. Botanirche Jahrbucher fur Systematik, Pflanzengeschichte 

1900-1953. DPP Pflanzenreich. 107 volumes. hipzig and Berlin. 

1926. Angiospermae: Kurze Erlauterung der Bluten- und 

Fortpflanmgsvehaltnisse. In A. Engler and K. Prantl, Die 

naturlickn Pflanzenfamilien. 2nd edition, volume 148. Leipdg. 

1924. Syllabus der Pflanzenfamilien. 9th and loth editions. Berlin. 

1887-1889. Die natuLlichen Pflanzenfamilien. 1st edition, part 2. Lcipzig. 

1930-1940. Die naturlichen Pjlanzenfm’lien. 2nd edition, volumes 

und Pjlanzengeographu. Leipzig. 

Engler, A., and E. Gilg 

Engler, A,. and E. Prantl 

14d-150. Ldpdg. 

Erdtman. G. 

1952 Pollen Morphology and Plant Tpu~omy: Angiosperm. Stockholm. 

Esau, K. 

1977. Anatomy ofseed Plants. 2nd edition. New York. 

71 
72 SMITHSOMAN CONTRIBUTIONS TO BOTANY 

Faegri, K., and L. van der Pi$ 

1978. The Principles of Pollination Ecology. 3rd edition. Word and New 

York. 

Fedorov, A.A., editor 

1969. Chromosome Nwnbers of Flowering Plants. Leningrad. 

Gibbs, R.D. 

1974. Chemotaxonomy of Ftowering Plants. 4 volumes. Montreal and 

London. 

Goldberg, A. 

1986. Classification, Evolution, and Phylogeny of the Families of 

Dicotyledons. Smithsonian Contributions to Botany, 58: 314 pages 

1981, 1984. Index to Plant Chromosome Numbers for 1975-1981. 2 

Goldblatt, P., editor 

volumes. St. Louis. 

Good, R. 

1974a. Features of Evolution in the Flowering Plants. New York. 

1974b. The Geography of the Flowering Plants. 4th edition. London. 

Hallier, H. 

1912. L'Origine et le systeme phyletique des angiospermes. Archives 

Neerlandaise des Sciences Exacter et Naturelles, series 3B. 

1 : 146-234. 

Harbome, J.B., and B.L Turner 

Hegnauer, R. 

1984. Plant Chemosystematics. London and New York. 

1962-1986. Chemotaxonomie der Pflanzen. 7 volumes. Base1 and 

Stuttgan. 

Heywood, V.H., D.M. Moore, I.B.K. Richardson, and W.T. Stearn, editors 

Hooker, W.J. and J.D. Hooker 

Huber, H. 

1978. 

1837-1982. Hooker's Icones Plantarum. Volumes 1-38. London. 

1969. Die Samenmerkmale und Verwandtschaftsvarhisse der Liliifloren. Mitteilungen der botanischen Staatssammlung Munchen, 

Flowering Plants of the World. New York. 

8~219-538. 

Hutchinson, J. 

1969. Evolution and Phytogeny of Flowering Plants. London and New 

York. 

1973. The Families of Flowering Plants. 3rd edition. Oxford, England. 

Jackson, B.D. 

1928. A Glossary of Botanic Terms. 4th edition. London and New York. 

Kanis, A. 

1981. An Introduction to the System of Classification Used in the Flora 

of Australia. In Flora of Australia, 1:77-111. Canberra. 

Kevin, P.G., and H.G. Baker 

1983. Insects as Flower Visitors and Pollinators. Annual Review of 

Entomology, 28:407-453. 

Syseme et phylogelie des Monocotyl6eones. Notulae Systematicae, 

Kimura, Y. 

1956. 

15: 137-159. 

Kubitzki. K.. editor 

1977. Flowering Plants, Evolution and Classification of Higher Categories. 

In Plant Systematics and Evolution, supplementum 1. Vienna and 

New York. 

Lawrence, G.H.M. 

1951. Taronomy of Vascular Plants. New Yo&. 

Le Maout, E., and J. Decaisne 

1873. A General System of Botany. Landon, Boston. and New York. 

LemQ, A. 

1929-1951. Dictwnaire descriptif et synonym'que des genres de plantep 

phanerogames. 9 volumes. Brest, France. 

The Vegetable Kingdom. 3rd edition. London. 

Triuridaceae. Flora Neotropica, monograph 40. 

Lindley, J. 

1853. 

Maas, P.J.M., and T Rubsamen 

1986. 

Maas, P.J.M., H. Maas-van de Kamer, J. van Benthem, H.C.M. Snelders, and 

T Rubsamen 

1986. Burmanniaceae. Flora Neotropica, monograph 42. 

1950. An Introduction to the Embryology of the Angiosperms. New York. 

1946. Comparative Internal Morphology of Seeds. American Midland 

Naturalist, 36513-660. 

Maheshwari, P. 

Martin, A.C. 

Martius, C.F.P. 

1840-1906. Flora Brasiliensis. 15 volumes (in 40 parts). Munich, Vienna, 

Meeuse, A.D.J. 

and Leipzig. 

1970. The Descent of the Flowering Plants. Acta Botanica Neerlandica, 

19~61-72, 133-140. 

Melchior, H.. editor 

1964. Engler's Syllabus der Pflanzenfamilien. 1wI edition, volume 2. 

Berlin. 

Metcalfe, C.R., editor 

1960-1982. Anatomy of the Monocotyledons. 7 volumes. Oxford, 

England. 

Moore, H.E., Jr. 

1973. The Major Groups of Palms and Their Distribution. Gentes 

Herbarum, 11(2):27-141. 

Moore, H.E., Jr., and N.W. Uhl 

1982. Major Trends of Evolution in Palms. The Botanical Review, 

48~1-69. 

Muller. J. 

1981. Fossil Pollen Records of Extant Angiosperms. The Botanical 

Review, 47:l-142. 

Omduff, R., and R.J. Moore, editors 

1967-77. Index to Plant Chromosome Numbers for 1965-1974. 5 

volumes. Utrecht. 

Proctor, M., and P. Ye0 

1973. The Pollination of Flowers. London. 

Me, A.A. 

1952 Compendium van de Terminologie, Nomenclatuw en Systematiek 

der hadplanten. 3rd edition. Utrecht 

Radfoni, A.E., W.C. Dickism. J.R. Massey, and C.R. Bell 

Vascular Plant Systematics. New York, Evanston, San Francisco, 

and London. 

1974. 

Real, L, editor 

Rendle, A.B. 

1983. Potlination Biology. Orlando. 

1930. The Classjfiawn of Flowering Plants. 2nd edition, volume 1. 

Cambridge, England. 

Rouleau, E. 

1981. Guide to the Generic Names Appearing in the Index Kewensir and 

Its F$teen Supplements. Pars N: Conspectus Systematis. Cowansville, Quebec, Canada. 

Royal Botanic Gardens, Kew 

1974-1988. Kew Record of Taronomic Lifenature Relating to Vascular 

Plants for 1971-1988. 13 volumes. London. 

Smith, A.C. 

1971. An Appraisal of the Orders and Families of Primitive Extant 

Angiosperms. Journal of the Indian Botanical Society, M: 

215-226. 

Smith, LB., and E.S. Ayensu 

1976. A Revisiar of American Velloziaceae. Sm'thsonian Contributions 

to Botany, 30: 172 pages. 

Smith. LB., and R.J. Downs 

Sneath, P.H., and R.R. Sdtal 

1974-1979. 

1973. Numericd Tmonomy: The Principles and Practice of Numerical 

Classjfiatwn. San Francisco. 

Bromeliaceae. Flora Neotropica, monograph 14, parts 1-3. 
NUMBER 71 13 

Scderstrom, TR., K.W. Hilu, C.S. Campbell, and M.E. Barkworth, editors 

Soo, C.R. de 

1987. 

1961. 

Grms Systematics and Evolution. Washington and London. 

Present Aspects of Evolutionary History of Telomophyta Andes 

Universkatis Scientiarwn Budapestinensis: Sectio Biologica, 

4~167-178. 

Spome, K.R. 

1975. The Morphology of Angiosperms: The Structure and Evolution of 

Flowering Plants. New York. 

Stafleu, F.A., and R.S. Cowan 

19761988. Taxonomic Literature. 2nd edition, 7 volumes. Utrecht and 

The Hague. 

Stapf, 0. 

1929-1931. Indezlondinensis to IlluFtrations of Flowering Plants, Ferns 

and Fern Allies. 6 volumes. Oxford. England. (Additional 2-volume 

supplement by W.C. Worsdell issued in 1941.) 

Flowering Plants, Evolution above the Species Level. Cambridge, 

Massachusetts. 

Paleobotany and the Evolution of Plants. Cambridge, England and 

New York. 

Botanical Classifications: A Comprison of Eight Systems of 

Angiosperm Classification. Hamden, Connecticut. 

1969. Flowering Plants: Origin and Dispersal. Edinburgh, Scotland. 

1980. Outline of the Classification of Flowering Plants. The Botanical 

Review, 46:225-359. 

Stebbins, G.L. 

1974. 

Stewart, W.N. 

1983. 

Swift, L.H. 

1974. 

Takhtajan. A. 

1986. Floristic Regions of the World. Berkeley and LAX Angeles. 

1987. 

1972-1986. Index Bibliographique sur la morphologie des pollens 

d'hgiospennes. In Travau de la Section Scientifqw et Technique 

de l'lnstitui Francais de Pondichery, India, volumes 12 @arts 1 and 

2). 13.18. and 22. 

A Phylogenetic Classification of the Angiosperms. In M.K. Hecht, 

W.C. Steere, and B. Wallace. Evolutionary Biology, 9:35-106. New 

York and London. 

Proposed New Realignments in the Angiosperms. Nordic Journal 

of Botany, 3:85-117. 

Califomia. 

[System Magnoliophytorwn.] Leningrad. [In Russian.] 

Thanikaimcni, G. 

Thome, R.F. 

1976. 

1983. 

Uhl, N,W., and I. Dransfield 

1987. Genera Palmarum. Lawrence, Kansas. 

Vester, H. 

1940. Die Areale und Arealtypen der Angiaspermen-Familien. Leipzig. 

Wagner, P. 

1977. Vessel Trpes of the Monocotyledons: A Survey. Botanisko Notiser, 

130~383-402. 

Wettstein, R 

1935. Handbuch der systematischen Botanik 4th edition. Leipzig and 

Vienna. 

Willis, J.C. 

1973. A Dictionary of the Flowering Plants and Ferns. 8th edition. 

Cambridge, England. 

Phytochemistry and Angiasperm Phylogeny. New York. 

Young, D.A.. and D.S. Seigler, editors 

1981. 
Agavaceae, 42 

Alismataceae, 11 

Alismatales, 11 

Amaryllidaceae, 50 

Aponogetonaceae, 17 

Araceae, 36 

Arales, 36 

Arecaceae. 29 

Arecales, 29 

Bromeliaceae, 50 

Bromeliales, 50 

Burmanniaceae, 66 

Butomaceae, 11 

Cannaceae, 61 

Centrolepidaceae, 23 

Commelinaceae, 53 

Commelinales, 53 

Corsiaceae, 68 

Cyclanthaceae, 34 

Cyclanthales, 34 

Cyperaceae, 26 

Cyperales, 26 

Dioscoreaceae, 47 

Eriocaulaceae, 55 

Flagellariaceae, 26 

Haemodoraceae, 64 

Index to Orders and Families 

Hydrocharitaceae, 11 

Iridaceae, 66 

Iridales, 64 

Juncaceae, 23 

Juncales, 23 

Juncaginaceae, 14 

Juncaginales, 14 

Lemnaceae, 39 

Lilaeaceae, 14 

Liliaceae, 40 

Liliales, 39 

Lowiaceae, 59 

Marantaceae, 61 

Mayacaceae, 55 

Musaceae, 59 

Najadaceae, 20 

Kajadales, 17 

Orchidaceae, 68 

Orchidales, 66 

Pandanaceae, 9 

Pandanales, 9 

Philesiaceae, 42 

Philydraceae, 64 

Poaceae, 29 

Poales, 27 

Pontederiaceae, 47 

Posidoniaceae, 20 

Potamogetonaceae, 17 

Rapateaceae, 55 

Restionaceae, 26 

Scheuchzeriaceae, 14 

Smilacaceae, 40 

Sparganiaceae, 20 

Stemonaceae, 45 

Strelitziaceae, 59 

Taccaceae, 47 

Thumiaceae, 23 

Trilliaceae, 39 

Triuridaceae, 14 

Triuridales, 14 

Typhaceae, 20 

Typhales, 20 

Velloziaceae, 50 

Xanthorrhoeaceae, 42 

Xyridaceae, 55 

Zanichelliaceae, 17 

Zingiberaceae, 61 

Zingiberales, 57 

Zosteraceae. 17 

74