A monograph of the genus Clitoria (Leguminosae: Glycineae)

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Title:
A monograph of the genus Clitoria (Leguminosae: Glycineae)
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3 v. (xii, 1052 leaves) : ill. ; 28 cm.
Language:
English
Creator:
Fantz, Paul R., 1941-
Publication Date:

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Subjects / Keywords:
Clitoris   ( lcsh )
Genre:
bibliography   ( marcgt )
theses   ( marcgt )
non-fiction   ( marcgt )

Notes

Thesis:
Thesis--University of Florida.
Bibliography:
Includes bibliographical references (leaves 1026-1050).
Statement of Responsibility:
by Paul R. Fantz.
General Note:
Typescript.
General Note:
Vita.

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University of Florida
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All applicable rights reserved by the source institution and holding location.
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aleph - 000011462
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oclc - 03403316
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AA00003919:00001


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A .OGRPi OF 1HE GENUS U1 l 0il i (- GU I,' S p !'


GLYCi i i -


PAUL R. FA%!VZ















A DISS-RTATICG'; PRE~',TED TO THE GRA JATE COUNCIL OF
TFH LUIV'ERSIT 0- F'LORi0,l
iN PARTIAL FULF L;:,T O. T!lE RJIRM1M:L.. FOP TiHE
DEGREE CF G[OC ;- c; PHIi .? !Y






UNIVERSE IT 'Y OF FLORkDF,


1I9771





















COPYRIGHT


By


PAUL R. FANTZ


1977
















ACKNOWLEDGEMENTS


I would like to express my appreciation for the interest, patience,

guidance, and helpful criticisms of Dr. Daniel B. Ward, under whose

supervision this monograph was accomplished. I also wish to thank the

other members of my committee, Dr. Dana G. Griffin III, Dr. Willard

Payne, Dr. Terry Lucansky, Dr. Jack Ewel, and Dr. John Kaufmann, for

their assistance during this study and for reading and providing

helpful editorial evaluations of the manuscript.

I am very grateful to the curators and staff of the herbaria

listed in the chapter on "Systematic Criteria and Taxonomic Notes" for

lending their material, including type-specimens, for use in this study.

Special thanks are extended to Gus Kovalick and the Staff of Hume

Library and Library West for their diligence and extensive aid in

searching for titles and institution of deposit, and obtaining xerox

copies of many older botanical publications; to Dr. Armstrong for her

assistance in providing appropriate geographical indexes, maps, and

atlases during my search for the obscure localities noted by the plant

collectors; and to Dr. Armstrong, Dr. Zimmermann, and Dr. Jones for

providing me with special library privileges to borrow needed references

for extensive periods during this study.

Thanks are extended to Dr. Daniel B. Ward and his staff at the

Herbarium of the Agricultural Experiment Station, for their assistance









in the necessary correspondence and the securing and returning of

borrowed herbaria material used in this study, and for providing the

needed facilities and space for the storage and examination of this

material.

I am indebted to Dr. Robert H. Mohlenbrock of Southern Illinois

University, who made this monograph possible by introducing me to the

world of plants, and whose enthusiasm stimulated me to change careers

and pursue a lifetime study in Vascular Plant Systematics.

I am grateful to Barbara Smerage and her assistant, Linda

Nordstedt, for their time, patience, and cooperation in the typing of

this manuscript.

A special thanks is extended to my wife, Janet, for her patience,

understanding, and the sacrifices she has made during the five years

it took to complete this study and for her assistance in portions of

this study.



















TABLE OF CONTENTS


Page


CHAPTER


ACKNOWLEDGEMENTS .


ABSTRACT . .

INTRODUCTION . .

ECONOMIC IMPORTANCE .

Medicinal Properties .
Leaves .
Seeds .
Flowers .
Additional Notes .
Aphrodisiacs .
Ornamental Properties .
Poisons and Pesticides. .
Dyes and Fibers .
Food for Man .
Fodder and Green Manure .

HISTORY OF CLITORIA .


Origin of the Generic Name . .
General History . . .
Historical Chronology . .
Pre-Linnaean (1650-1753: Polynominal Period).
Post-Linnaean (1753-1824: Descriptive Period)
Post-Linnaean (1825-1858: Revisionary Period)
Post-Linnaean (1859-1932: Floristic Period) .
Post-Linnaean (1932- present: Research Period).

PRIOR RESEARCH ON CLITORIA . .


Anatomical Research .
Chemical Research .
Cytological Research .
Developmental Research. .
Morphological Research. .
Agronomic Research .
Palynological Research. .


. . . i i i


xi


. . 1


. . 2 1


. 79

. 79
. 82
. 84
. 86
. 90
S91
S94


. . .
. . .
. . .
. . .
. . .
. . .
. . .













MORPHOLOGY . .


Habit .
Stems and Bran
Pubescence. .
Leaves .
Leaflets
Stipules
Petiole a
Petiolule
Observati
Inflorescence
Chasmogam
Cleistoga
Chasmocle
Bracts
Pedicels
Bracteole
Observati
Chasmogamous F
Calyx.
Vexillum
Alae .
Carina
Androeciu
Observati
Cleistogamous
Calyx. .
Corolla.
Androeciu
Gynoeciun
Observati
Legume. .
Stipe.
Valves
Observati
Seeds .
Seed Gern
Observati
Summary .
Species [
Character


ches. . .

. . .



and Stipels .
nd Rachis .

ons . .

ous Inflorescences. .
mous Inflorescences .
istogamous Inflorescences .
istogamous Inflorescences
. . .
. . .

ons. ............
s owers. . .

ons . .
lowers . .




Flowers . .
. . .
. . .
. . .
m . .




ons . .


Flowers . .


ion . .
ons . .
m . .
. . .
ons . .
. . .





nation . .
ons . .
. . .


descriptionss
's Occurring


in Relatively


.



. .

. .







. .



. .

.



Few


. .
. .
.

. ., .
. o


.


. .

. .

.
. .

. .
.
Species


DISTRIBUTION OF CLITORIA . . .

SYSTEMATIC CRITERIA AND TAXONOMIC NOTES . .

SYSTEMATIC TREATMENT . . .


95
97
98
100
101
101
102
102
102
103
105
109
109
110
113
113
114
114
118
121
123
124
124
127
133
133
138
138
139
139
140
141
145
148
148
151
151
152
152
153

159

173

180


Page


. 95









Page
Subgenus Bractearia . . ... 193
Section Bractearia . . 198
1. Clitoria arborea. . ... 201
a. var. arborea. . ... 211
b. var. longiramosa. . ... 215
c. var. pseudoamazonica. . .. 216
2. Clitoria fairchildiana. . ... 218
3. Clitoria andrei . .. 230
4. Clitoria juinensis. . .. 234
5. Clitoria moyobambensis . .. 242
6. Clitoria amazonum . ... 247
a. f. amazonum . ... .255
b. f. vulgaris . 256
7. Clitoria nervosa. . ... 258
Section Flexuosa . .... 265
8. Clitoria woytkowskii. . .. 267
9. Clitoria flexuosa . ... .275
a. var. flexuosa . ... .282
b. var. brevibracteola . .... .283
10. Clitoria pozuzoensis. . .284
a. var. pozuzoensis .............. 292
aa. f. pozuzoensis . .... .292
ab. f. subpalmata. . ... 292
b. var. schunkei . ... 293
Section Brachycalyx. ........ ........... .295
11. Clitoria brachystegia . ... 299
12. Clitoria hermannii. . 307
13. Clitoria glaberrima . ... 311
14. Clitoria canescens. . .. 321
15. Clitoria brachycalyx. . .. 326
16. Clitoria dendrina . 332
17. Clitoria froesii. . 344
S. Section Cauliflorae. . . ... 349
18. Clitoria arborescens. . ... 357
19. Clitoria javitensis . 372
a. var. javitensis . .. 398
aa. f. javitensis. . ... 399
ab. f. bracteosubtenda ... 401
b. var. portobellensis . 403
ba. f. portobellensis .. 404
bb. f. pilosa . .. 407
bc. f. truncata . .. 409
c. var. longiloba. . 409
d. var. grandifolia.. ............ .410
e. var. klugii . ... .412
20. Clitoria cavalcantei. . ... 413
21. Clitoria coriacea . ... 418
22. Clitoria tunuhiensis. . ... 425
23. Clitoria sagotii. . ... 429
a. var. sagotii. . 439
b. var. caniculata . ... .442
c. var. sprucei. . ... 444









Page
24. Clitoria kaieteurensis. . ... 446
25. Clitoria pendens. . ... 451
26. Clitoria leptostachya . 457
a. var. leptostachya . 465
b. var. fruticosa . .. 466
27. Clitoria selloi . ... .466
28. Clitoria obidensis. . ... 474
29. Clitoria plumosa. . 480
Subgenus Clitoria . . ... 486
30. Clitoria lasciva. . ... 494
31. Clitoria ternatea . ... .504
a. var. ternatea . ... .536
aa. f. ternatea .. ... .. 536
ab. f. pauciflora. ............ 571
ac. f. fasciculata .. .. 571
ad. f. albiflora .. 574
b. var. angustifolia . 584
c. var. pleniflora . 591
ca. f. pleniflora. .... 596
cb. f. leucopetala . 600
cc. f. subpolyadelpha. .. ... 601
32. Clitoria heterophylla . 602
a. var. heterophylla . .... .609
b. var. pedunculata . .. 610
33. Clitoria biflora . .. 612
34. Clitoria kaessneri. . 619
Subgenus Neurocarpum. . . 628
Section Mexicana . .. .640
35. Clitoria polystachya. . ... 645
a. var. polystachya . .. 654
b. var. pringlei . ... 659
c. var. congesta . .. 659
36. Clitoria monticola. . ... 660
37. Clitoria triflora . ... .665
38. Clitoria mexicana. . ... 671
39. Clitoria humilus. . ... 682
40. Clitoria cordobensis. . ... 686
41. Clitoria fragrans . ... .696
42. Clitoria mariana. . 705
a. var. mariana. . ... 718
aa. f. mariana ... .. 722
ab. f. pedunculata . 743
ac. f. pubescentia .. 746
b. var. orientalis . 749
Section Tanystyloba. . . ... 758
43. Clitoria macrophylla. . .. 762
a. var. macrophylla ............. 774
b. var. sericea. .... .. 777
44. Clitoria javanica . 779
45. Clitoria cordiformis. .... .... 784


viii









Page

46. Clitoria linearis . ... .791
47. Clitoria hanceana . 795
a. var. hanceana . ... 802
b. var. thailanensis . 805
c. var. laureola . 805
d. var. latifolia. . 809
e. var. petiolata. .............. 810
48. Clitoria australis. . ... 811
Section Neurocarpum . .. 820
49. Clitoria stipularis . 826
a. var. stipularis . ... 833
b. var. latifolia . .. 836
50. Clitoria densiflora . ... .837
51. Clitoria irwinii. . .. 846
52. Clitoria laurifolia . ... .852
a. f. laurifolia . ... 863
b. f. glabrior . ... .873
c. f. fasciculata. . ... 879
d. f. parvifolia . ... 879
e. f. petiolata. . ... 880
53. Clitoria guianensis . ... 881
a. var. guianensis . 891
aa. f. uianensis . .. 892
ab. f. macrofructa . .... .903
ac. f. imperfecta. . ... 903
ad. f. unifoliata. . ... 905
b. var. macrocleistogama . .. 905
c. var. chapadensis ............. 906
54. Clitoria epetiolata . 910
a. var. epetiolata . ... .916
b. var. angustissima . 919
c. var. latiuscula . ... 920
55. Clitoria simplicifolia. . 921
56. Clitoria flagellaris. . .. 927
57. Clitoria falcata. . .. 932
a. var. falcata. . ... 943
aa. f. falcata . 943
ab. f. heteromorpha ........... 960
ac. f. stipulacea. . ... 964
ad. f. longirachis . ... .965
b. var. latifolia . .. 966
c. var. aurantiaca .... .. 966
d. var. glabrescens. . .. 971
58. Clitoria nana . ... .972
a. var. nana .. ... ..... 978
b. var. caaguazuensis . .. 979

NUMERICAL LIST OF TAXA . .... ... .. 980

NOMINA ET SYNONYMA CLITORIA COMPLECTENS. . ... 986









Page
PHYLOGENETIC CONSIDERATIONS. . . ... 1007

Evolutionary Trends in Clitoria . .... 1008
The General Plant Structure. . ... 1009
Vegetative Structures. . ... 1013
The Inflorescence Structures . .... 1014
The Flowers . . 1016
The Fruits and Gynoecium . . 1018
Pubescence . . ... 1020
Nonmorphological Data. . ... 1021
Phylogenetic Pathways in Clitoria . .. 1022

BIBLIOGRAPHY . . . 1026

BIOGRAPHICAL SKETCH. . . ... ..... 1051










Abstract of Dissertation Presented to the Graduate Council
of the University of Florida in Partial Fulfillment of the Requirements
for the Degree of Doctor of Philosophy


A MONOGRAPH OF THE GENUS Clitoria (LEGUMINOSAE: GLYCINEAE)


by

Paul R. Fantz

August 1977

Chairman: Daniel B. Ward
Major Department: Botany

This monographic treatment of the genus Clitoria L. (Leguminosae:

Glycineae) is the first since that of George Bentham in 1858. The

genus is treated on a world-wide basis. The study is based upon

examination of approximately 7000 specimens from thirty-three lending

institutions, and from field studies of United States species. Treated

in synonymy are Clitorius Petiv. ex Dill., Clytoria Presl, Macrotrullion

Klotsch ex Schomb., Martia Leandr. Scar., Martiusia Schult., Nauchea

Desc., Neurocarpon Hamilton, Neurocarpum Desv., Neurocarpus Hassk.,

Rhombifolium Rich. ex DC., Rhombolobium Rich. ex H.B.K., Ternatea Tourn.,

Ternatea Tourn. ex Mill., Vexillaria Eaton, and Vexillaria Raf.

Clitoria includes fifty-eight species and fifty-one nontypical

subspecific taxa. Forty-six species are native to the neotropics and

subtropics. One paleotropical species has been introduced and

naturalized in many regions of the neotropics. One species is native

to temperate North America and to tropical Southeast Asia. Eleven

species are native to the paleotropics in Africa and Madagascar, the

Indian subcontinent, Indonesia, and Arnhem Territory of northern

Australia. Two neotropical species have been introduced and naturalized








in portions of Africa and Indonesia. One species is introduced and

naturalized in many Pacific islands.

A history of Clitoria is provided in which the origin of the

generic name is traced to Breyne in 1678. A detailed morphology of the

genus is presented. Cleistogamous flowers, as well as chasmogamous

flowers, are noted to occur in thirteen species. Species of Clitoria

are recorded to have economic uses as medicinal plants, ornamentals,

aphrodisiacs, poisons, pesticides, dyes, fibers, food, fodder, and

green manure. Prior research on species of Clitoria is summarized in

the areas of agronomy, anatomy, chemistry, cytology, developmental

botany, morphology, and palynology.

A systematic treatment is given which recognizes three subgenera

and eight sections. These include sections Bractearia (7 spp.),

Flexuosa (3 spp.), Brachycalyx (7 spp.), and Cauliflorae (12 spp.) of

subgenus Bractearia; subgenus Clitoria (5 spp.); sections Mexicana

(8 spp.), Tanystyloba (6 spp.), and Neurocarpum (10 spp.) of subgenus

Neurocarpum. Each species is described and illustrated. A complete

synonymy is given. A nomenclatural discussion of type collections,

phenology, distribution, subspecific taxa, and citation of examined

specimens is provided under each species. Economic importance and

vernacular names are included for a species when known. Each taxon has

a mapped distribution. Keys are provided to each taxonomic level.

Phylogenetic considerations are given based upon the interpretation

of morphological data presented in this study. Subgenus Bractearia is

regarded as a primitive subgenus with subgenus Neurocarpum as advanced.















INTRODUCTION


The genus Clitoria contains fifty-eight species located mostly

within the tropical belt with a few species distributed within the

temperate zone. The bulk of the genus (ca. eighty percent of the

genera), is neotropical. The genus is absent from Europe, most of

Asia (except the Indian subcontinent), and most of Australia.

Although there is no universal name by which the genus is commonly

known, the most often used vernacular name for the genus is "butterfly

pea." Centrosema, a close morphologically and historically related

genus, bears the same vernacular name or is sometimes distinguished

as the "spurred-butterfly pea." When horticulturalists attempted to

standardize plant names (Kelsey and Dayton, 1942), they adopted

"butterfly pea" for the genus Centrosema and adopted "pigeon wings"

for the genus Clitoria, the latter name rarely used by botanists.

The Papilionaceous genus Clitoria is included by most botanists

within the tribe Phaseoleae, subtribe Glycineae. In the few

traditional tribes, a number of genera are usually included in a

list of exceptions to the typical tribal descriptions. In order to

reduce the number of exceptions, Hutchinson (1964) elevated a number

of subtribes to the tribal level. Thus, he placed Clitoria within the

tribe Glycineae.

Flowering plants of Clitoria are easily determined in the field

by a few unusual characteristics. Papilionaceos, resupinate flowers








which have a bearded, geniculate style identify the genus. Specimens

in fruit become more difficult to recognize due to the variation of

the legume. However, odd-pinnate compound leaves (usually 3-foliate),

persistent stipules, stipels, bracts, and bracteoles, plus a

persistent, five-toothed, infundibular calyx are reliable characters

that, taken collectively indicate the genus Clitoria from other genera.

The genus Centrosema (Glycineae) is often confused with Clitoria.

Linnaeus established Clitoria in his "Species Plantarum" in 1753, and

included four species. Two of those species, C. brasiliana and

C. virginiana, were later placed in Clitoria section Centrosema

by de Candolle in 1825. Bentham (1858) segregated Centrosema from

Clitoria and elevated the taxon to the level ot genus. In the keys of

most floristic and revisionary treatments, Clitoria is distinguished

from Centrosema by its "bearded style" or "pubescent style." This

character is often misinterpreted. Clitoria can be distinguished

morphologically from Centrosema by several characters. These

differences are summarized in Table 1.

The last revisionary work on Clitoria was a synopsis published by

Bentham in 1858. This treatment included a total of three sections

with twenty-six species, more than half of which were originally

described by Bentham. Bentham's treatment has been the major

reference relied upon by nearly all authors of floristic works that

included the genus. Bentham's treatment contains a number of

deficiencies, however, and now includes some conflicts with the

International Code of Botanical Nomenclature (Stafleu, 1972).

A number of problems presently exist within the genus that need

resolution, and these can be categorized into six areas. One problem











Table 1. A morphological comparison of the genera Clitoria and
Cent rosema


CHARACTER


Calyx


Vexillum

Alae vs Carina


Style


Ovary


Legume


Leaves


CLITORIA


Infundibular, lobes
broad, shorter than
to nearly equal tube
length (rarely longer)


Spurless


Alae longer than carina

Geniculate, bearded
lengthwise to base


Stipitate (or
subsessile in
Subgenus Clitoria)

Stipitate, ecostate or
with 1 lateral nerve
near midline of valve

Pinnate typically
3 (1) leaflets
(Subgenus Clitoria
commonly 5, 7, 9, or
11 leaflets


CENTROSEIMA


Short-campanulate,
lobes narrow, equal to
or longer than tube
length

Spurred or gibbous

Alae subequal carina

Broadly incurved
(U-shaped), pubescent
only beneath stigma

Subsessile



Subsessile, 2 prominent
nerves laterally, one
near each suture

Pinnate typically
3 leaflets, rarely
5, 7, or 11 leaflets


_~_








is the proliferation of names and their dulineition into specific and

subspecific taxa. Eleven genera must be included in synonymy under

Clitoria. Bentham cited only two, although by tracing back through

his cited references, the genera included in synonymy increased to

six. To date, nearly two-hundred and fifty published binominals (plus

ca. twenty nom. in sched.) are included within the total concept of

the genus Clitoria, senso lato. Of these, one hundred and sixty of

the binominals were published under the naie Clitoria. Bentham

included only sixty of these binominals in his treatment, and since

Bentham's revision, nearly fifty new species have been described,

almost double the total number of species that he recognized. Bentham

did not include subspecific taxa in his synopsis. Twenty-one of the

twenty-three subspecific taxa published were d scribed after Bentham's

treatment.

A second problem is the lack of complete and detailed descriptions

of species, one probable cause of the large abundance of names. Few

of the published specific names were accompanied by detailed

descriptions. Many were published with short diagnostic descriptions

that included a number of characters that are typical of the genus or

subgeneric groups. Many of the descriptions contain relative terms

(i.e. short vs. long, large vs. small) and lacd precise measurement

ranges. Often a character included in one description is lacking in

the description of another species.

Some floristic treatments have included descriptions of their

species, occasionally offering more detail than the original

description. These descriptions can be helpful, but a few have

compounded the problem. An example would be MRicbride's treatment





5


(1943) of the Peruvian species of Clitorid. Many of the specimens

examined by Macbride had been misidentified and did not match the

type collections. This has resulted in descriptions placed under

inaccurate names (e.g., C. aiiazonum and C. nervosa, neither species

occurring in Peru).

A third problem that occurs in conjunction with the first two is

the lack of keys to the species of Clitoria. Bentham did not include

a key in his treatment, although he used some diagnostic phrases that

separated the species into groups. The only keys that exist are those

found within certain floras. Keys which included five or more species

of Clitoria are found in floristic treatments for only a few countries

in South America, a few in Central America, plus one treatment for

Southeast Asia. They often include characters which work well

locally, but not when more species are added from neighboring areas.

A fourth problem is incomplete information, on types. A number of

species had incomplete citations, making type interpretation difficult.

Bentham was partially responsible in that he otten paraphrased data

from herbarium labels. Specimens labeled as "type" in the personal

herbarium of Bentham and in other herbaria which he examined, often

bear a collection number or date which Bentham did not cite.

Occasionally type information has been distorted. For example,

Howard (196/) proposed the name C. fairchildiana as a substitute for

the homonym C. racemosa Benth., correctly citing the Brazilian type

collection of Pohl. Then he distributed speci;,ens (Howard 17052)

from a cultivated tree at the Jennings Estate in Miami, Florida,

United States, as the type tree.









A fifth problem is the number of species that bear illegitimate

names as defined by the International Code of Botanical Nomenclature.

Although Bentham was not bound by a code developed nearly a century

after his revision, he did select names by a process that conflicted

with the Code standards in practice today. Later botanists who

assumed that those names were acceptable under the Code have often

been unaware of these differences and have used names that cannot

stand today.

A sixth problem is lack of information as to the geographic

distribution of the species. Distribution statements have often been

based upon one or a few collections and appear to sometimes have

been wholly hypothetical. The problem has been compounded in some

species which were described originally from cultivars outside the

normal distribution range (e.g., C. arborescens). Distributions in

some species have been distorted when floristic treatments have

mistakenly included a species within their geographical areas or

inaccurately reported the species from other areas.

This author has examined nearly eighty percent of the type

collections for the two hundred and fifty binominals associated with

Clitoria. Many of the types not examined belonged to species described

and placed within the genus Clitoria, but later transferred to other

genera, often Centrosema. In most cases, descriptions of these

species contained diagnostic phrases (i.e. campanulate calyx, spurred

vexillum) sufficient to indicate that the species does not belong to

the genus Clitoria.

Bentham's section Clitorianthes was most poorly understood.

Nearly fifteen percent of the specimens that were examined in this








section were unidentified to species. Foi those specimens with names,

slightly more than fifty percent were misi.lentified. A large portion

of the species in Bentham's section NeurocJrpuYi were also misnamed.

However, many of these specimens were correctly identified to species,

but bore illegitimate synonyms. Relatively few problems were found

within Bentham's section Ternatea.

The purpose of this investigation has been to thoroughly

re-examine available specimens and types, to clarify the nomenclature

and species descriptions, and to prepare Tunctional keys and

illustrations to legitimate taxa.















ECONOMIC IMPORTANCE


With the exception of C. Ternatea, the species within the genus

Clitoria have been of relatively little economic use, except where

they are locally important. Clitoria ternatea is important

economically as a medicinal herb, as an o inamental, as a food and

cover crop, as a dye, as green manure, and as an aphrodisiac.

Clitoria ternatea has been prized as a cultivar at least since the

late seventeenth century, and has been carried by man throughout the

tropics, where it often has escaped and bc,:ome established. As a

result, thi, species now has a pantropical distribution. Attention

has been focused on C. Ternatea as a typical representative of the

genus because it is widespread, well know, and easily accessible for

economic use. Some species within the genus are reported to have some

of the same economic properties as C. tern ,tea, although these species

have not received the same publicity of C. ternatea. Additional

properties have been noted for a few species, for example, the

possible use as powerful insect and vertebrate poisons.

The genus Clitoria has great potential to be an economically

important legume genus. The economic uses of a particular species

properly belong under the treatment of that species. The purpose of

this chapter is to discuss tlhe economic properties known for the genus

and to point the way toward research in the economic botany of

Clitoria as a whole. Clitoria ternatea will be the major








representative of the genus in this discussion. Information obtained

from herbaria is documented with the collector's name and his

collection number enclosed within parentheses, for example, (Hahn

234).


Medicinal Properties

Nearly all the medicinal properties of Clitoria are recorded for

the species C. ternatea as a result of investigations by workers

interested in medicinal Indian, Southeast Asian, and Indonesian plant

species. Remedies and treatments reported are grouped below by

medicinal use for each plant organ used.

Roots

Root or root bark is often reported to be an effective laxative,

but may not always be safe. The degree of cleansing action varies in

the reports. Dalgado (1896), Kirtikar and Basu (1918), and Dutt

(1928) report use of the roots as a laxative. Kirtikar and Basu ibidd)

cited Mr. Mooden Sheriff (1891) who noted that the root bark was a

laxative in some cases. Chopra, Chopra, Honda, and Kapur (1958) noted

that the root bark was used as a laxative. Chopra, Badhwar, and Ghosh

(1949) and Chopra, Chopra, Honda, and Kapur ibidd) reported the roots

as a powerful cathartic similar to Jalap (Exogonium purga,

Convolvulaceae), but that it was not a safe medicine. Dey (1896),

Burkill (1935). Quisumbing (1951) citing Dey, and Gardner and Bennett

(1956) reported the roots as carthartic. Dey reported preparing the

remedy as an alcoholic extract of the roots. Quisumbing ibidd),

citing Rajan (1926), reported the roots as a purgative and narcotic

toxin. Rajan ibidd) indicated symptoms of the narcotic poisoning to








include unconsciousness attended with extreme irritability and a

peculiar loss of memory. Chupra, Badhwar, and Gosh ibidd), citing

O'Shaughnessy (1841), indicated that an alcoholic extract of the root

acts as a brisk purgative in doses of five to ten grains, but produces

gripping anJ tenesmus, and the patient will feel feverish and uneasy.

Quisumbing ibidd), citing Rajan ibidd), Sanyal and Ghose (1934), and

Burkill ibidd), reported the roots as an aperient. Roots were noted

to be a purgative medicine or remedy used in Martinique (Hahn 234).

Roots dre also used in stimulating urination and are useful in

ascites and fevers. Rajan (1926), Dutt (1928), citing the Hindus,

Sanyal and Ghose (1934), Qui umbing (1951), citing the three earlier

works, and Chopra, Chopra, Honda, and Kaputr (1958) reported the roots

as diuretic. Chopra, Chopra, Honda, and l.ipur ibidd) also reported

the root bark used as a diuretic. Kirtikar and Basu (1918), citing

Sanskrit writers, described Lhe root as diuretic, useful in ascites

(an accumul ition of excess fluid in the al.Jomiiial cavity) and fever.

They also cite Mooden Sheriff who noted that an infusion of the root

bark acts as a diuretic.

Quisumiing (1951), citing Sanyal and Ghose (1934) who cited

Mooden Sheriff (1891), and Kirtikar and Basu (1918), who also cited

Mooden Sheriff, reported that Sheriff spole highly of the infusion of

the root bark as a demulcent in cases of irritation of the bladder and

of the urethra.

Roots ire also used as a medicine in promoting menstrual

discharge. Quisumbing (1951) cited Crevost and Petelot (1929) who

reported the roots used as an emmenagogue.








Roots or root juice has been reported to cause nausea and

vomiting and to be useful for lung ailments and removing the phlegm

of chronic bronchitis. Lindley (1938) and Gardner and Bennett (1956)

reported the root as an emetic. Quisumbing (1951), citing Dalgado

(1896) and Crevost and Petelot (1929), indicated that each reported

the roots used as a vomitive. Drury (1873) and Dymock (1885), each

cited by Quisumbing ibidd), quote Mr. Ainslie (1826) who mentions the

use of the root in soup to remove phlegm in chronic bronchitis and to

bring on nausea and vomiting. Kirtikar and Basu (1918), citing Dymock

ibidd), indicated that in the Concan region of India, two tolas1 of

the root juice are given in cold milk to remove the phlegm of chronic

bronchitis through nausea and vomiting. Chopra, Badhwar, and Ghosh

(1949) indicated that opinions differed with regard to the emetic

properties of the root. In the Sudan, the roots are chewed for lung

and throat complaints (Prichard 4).

Dymock (1885), cited by Kirtikar and Basu (1918), indicated that

the juice of the root of a white-flowered form of C. ternatea is blown

up the nostrils as a headache remedy. A similar practice in the Sudan

has other effects. Before a cow is sent to the bull, the root is

chewed and blown up her nostrils to ensure fertility (Pritchard 4).

Roots are reported to be used as an antidote against snake-bite

and scorpion-stings. Chopra, Badhwar, and Gosh (1949) reported the

roots as a possible antidote against snake bites. Chopra, Chopra,

Honda, and Kapur (1958) reported that the roots were used in Indian


1. A tola is an Indian unit of weight equal to one silver rupee,
or 180 grains.








indigenous medicine as a treatment for snuae-bites (Cobras and

Daboias) and for scorpion-stings (genera huthus and Palamnoeus).

However, they found that remedies administered to dogs and rabbits

injected with the snake or scorpion venom in strict conformity with

the directions given in standard Indian medicinal books, had no

preventive, antidotal, or therapeutic effi:ct.

Leaves

The leaves are reported to be used against eruptive conditions

by Chopra, lKadhwar, and Ghosh (1949). Kirtikar and Basu (1918, citing

Watt (1889-96), and each later cited by Quisumbing (1951), reported

that an infusion of the leaves is used for eruptive conditions.

Heyne (1927), later cited by Burkill (1935) and Quisumbing (1951),

reported that the leaves of a white-flowered form of C. ternatea are

used as poultices in Java. Kirtikar and l.su (1918) and Quisumbing

ibidd), each quoting Mr. Taylor,2 stated that the juice of the leaves

mixed with qreen ginger is administered in cases of colliquative

sweating in hectic fever. Quisumbing (ibid) quoted Kirtikar and Basu

ibidd) who quoted Mr. Mukerji (1889-1904) in "Watt's Dictionary" that

juice of the leaves is mixed with common salt and applied all around

the ear for earaches, especially when accompanied with the swelling of

neighboring glands. Quisunbing ibidd) quoted Tavera (1892) who

reported that the leaves are used in the Philippines for swollen

joints. Chopra, Chopra, Honda, and Kapur (1958) reported that the

juice of the leaves is mixed with water to form a jelly which is taken


2. Taylor was quoted by both authorities; however, neither one
documented the reference source of Taylor.








as a cooling medicine for gonorrhea, and used externally for eczema,

prurigo, and impetigo.

Seeds

Seeds are used also as a laxative and considered as an effective,

safer method than the use of roots or root bark. Waring (1868),

Drury (1873), Dey (1896), and Crevost and Petelot (1929), each cited

by Quisumbing (1951), reported that seeds are used as a mild purgative.

Quisumbing ibidd) also cited Kirtikar and Basu (1918) who reported the

seeds as a purgative and an aperient. Chopra, Badhwar, and Ghosh

(1949) indicated that seeds in powdered form have purgative and

aperient properties, and are considered more useful and a safer

medicine than the roots. Quisumbing ibidd) cited Burkill (1935) who

reported that the seeds are used as an apcrient and contained a toxic

alkaloid. Quisumbing ibidd) cited Nadkarni (1927) who reported that

the seeds contained a fixed oil, a bitter acid resin, tannic acid,

glucose, and six percent ash. The testa was brittle and contained

cotyledons full of granular starch. Chopra, Chopra, Honda, and Kapur

(1958) cited Dymock, Warden, and Hooper (1890-93) who reported that

the seeds contained a fixed oil, a bitter resinous principle, and

tannin. (Chemical investigations on the seeds are reported under the

chapter "Prior Research on Clitoria." (Reler to that chapter,

subheading "Chemical Research," for details.)

Burkill (1935) quoted Tavera (1901) lwho reported that the seeds

are used in the Philippines as poultices for swollen joints. In an

earlier edition of his book (1892), Tavera had attributed this

medicinal property to the leaves.








Seeds are supposed to be effective in destroying or expelling

tapeworms. Waring (1868), Drury (1873), Dey (1896), and Crevost and

Petelot (1929), each cited by Quisumbing (1951), reported that the

seeds were used as an antihelminthic.

Flowers

Burkill (1935) cited "Medicinal Book of Malayan Medicine (Gard.

Bull. S.S. 6, 1930, p. 381)" which reported that the floral juice of

a white-flowered form of C. ternatea was used for inflamed eyes.

Additional Notes

Chopra, Chopra, Honda, and Kapur (1958) reported that C. ternatea

is alleged to have anti-dysenteric and anti-tubercular properties.

No plant organ was noted.


Aphrodisiacs


A few species are reported to ensure fertility and are used as

sexual stimulants. This topic is usually not included in the

literature possibly through delicacy complicated by the fanciful

resemblance of the flower to the female sexual apparatus. Yet, in

widely dispersed geographical areas with different species, the native

people use Clitoria plants as aphrodisiacs for man and domestic

animals. It is presumed that the natives follow the ancient

principles of the Doctrine of Signatures, which is based upon belief

that the plant structures which resemble portions of the human body

have been so structured as to advertise their ability to provide

remedies for ailments of those body portions.

The use of the plant in conception was implied by Rumpf (1747)

who noted that the Portuguese name "Fula criqua" was derived from the








flowers, and that Breyne (1678) had appropriately named the plant Flos

clitoridis ternatensibus (this plant now known as Clitoria ternatea).

The Portuguese translation of "Fula criqua" means "to speed or

hurry-up creation." In the Sudan, before a cow is sent to the bull,

the root of C. ternatea is chewed and blown up her nostrils to ensure

fertility (Pritchard 4). Near Anaconda Island on the Rio Napo,

Ecuador, this author attended a wedding presided over by a tribal

witch-doctor. In a discussion with him the night before the wedding,

the flower of C. pozuzoensis (a species that this author was searching

for locally) was described by cupping the hands for the vexillum and the

arching of the thumbs for the keel and wings. The grinning witch-doctor

apparently recognized the plant in question immediately. He accurately

described its white flowers, climbing habit, and vegetative morphology.

He reported through the translator that he gave this plant to couples

that had difficulty bearing children.

In Minas Gerais, Brazil, catuaba (C. guianensis) is put in

cachaca "to make a man out of you" (Williams, Assis, and Moreira

5416),which the collectors noted that the effect was "Potencia"!


Ornamental Properties


Most of the species of the genus Clitoria have potential for

becoming economically important as ornamentals because of their large

and beautifully-colored, showy flowers and the various habits they

possess. Frequently prized are those species which display climbing

habits, used commonly for garden trellises, and less frequently on

other vegetation. Also much esteemed are those species which possess








the "azure blue" flowers and those of "delicate shades of pink."

Other species are noted for their foliage. A few examples will be

cited here.

The oldest and most widely cultivated ornamental is C. ternatea.

This species is probably indigenous to Eastern Africa with subsequent

expansion into central and western Africa and the Indian subcontinent.

Man has transported this species throughout the tropical belt where it

often has escaped and become established as a wild plant. Today it is

commonly cultivated and is found wild in many Pacific Islands, in

portions of Australia, Indonesia, Africa, and the Americas, and in the

Antilles. Breyne (1678) reported the species as a cultivar, taken

from the Mollucan island of Ternate and planted in European gardens,

where Curtis (1812) reported that it needed protection from the

climate, and that it would not set seed. Because of its brilliant

blue flowers of large size which bloom nearly year round, it is a

popular cultivar in tropical regions, growing rarely as a small shrub,

but most commonly as a perennial climber. The plant grows rapidly,

forming many intertwined stems, and when placed upon open ground,

develops tangled mats. It can be trained through cutting to form

low-growing shrubs. The white-flowered and pale blue-flowered strains

are not as popular as the azure blue-flowered strains.

Clitoria ternatea is also important because of a double-flowered

variety. These plants produce large, actinomorphic, free, banner-like

petals of deep azure to bright purple colors, and rarely white flowers.

This variety was first reported by Commelin (1701). Today it is

commonly cultivated and is naturalized in portions of the Caribbean,

Sri Lanka (=Ceylon), and Indonesia, with isolated populations

elsewhere in the neotropics.








Another early prized ornamental was C. heterophylla which was

reported by Curtis (1820). This species is endemic to the islands of

Madagascar and Mauritius. It is valued for its climbing habit, blue

flowers, and unusual pinnate leaves of two distinct shapes found upon

a single plant.

The most commonly cultivated, erect plant species has been

C. laurifolia (synonym: C. cajanifolia). It is native to the

neotropics and introduced into central Africa (naturalized in Zaire)

and Southeast Asia-Indonesia. In the latter areas, it is cultivated

as a shrub with several plants often planted closely together to

form a hedge.

Two woody lianas from the neotropics are popular in protected

gardens: C. arborescens of northern South America has large leaves,

silky beneath, many racemes of numerous purple flowers, and it is a

tall climber; C. javitensis of South America and Panama is valued for

its climbing habit and numerous lilac to delicate pink shaded flowers.

A less popular liana species, C. lasciva, a native of Madagascar, is

valued for its luxurient foliage and blue flowers.

The tall erect shrub of C. amazonum is desirable in protected

gardens because of its erect habit with upper branches climbing, and

its huge, showy, purplish flowers which are numerous in each of many

inflorescences.

Clitoria fairchildiana (synonym: C. racemosa Benth.) is a

popular tree species. It is endemic to the deltas of the Amazon River

and the other large rivers to the southeast. This tree is planted

extensively along beach roads in Rio de Janeiro, and valued for its

elongated racemes of numerous purple flowers, and for the luxurient








foliage borne on branches which arch and droop to the ground, forming

a canopy.


Poisons and Pesticides


Several species are known to have toxic properties. It has

already been pointed out (Rajan, 1926) that the roots of C. ternatea

taken as a purgative can produce narcotic poisoning in man. Chopra,

Badhwar, and Kapur (1949) included C. ternatea in a list of important

plants poisonous to man and livestock.

Gardner and Bennett (1956) reported that C. arborescens was used

as a fish poison, the seeds containing alkaloids with weak curariform

action. The seeds of a related species from Colombia, C. arborea,

were reported as alkaloid positive (Schultes 24120). In Amazonas,

Brazil, C. froesii was reported as being considered the most toxic

plant in the region of Rio Icana, and was planted by the Indians and

found in fair quantities (Froes 12441/185).

The genus is not noted for its pesticidal properties, although

some species contain the insecticide ingredient rotenone. In Surinam,

the stems and fleshy roots of C. falcata have a strong odor of

rotenone (Archer 2848). A cultivated population of C. laurifolia in

Sao Paulo, Brazil, was noted not to be attacked by fungus or insects

(Norris 278). In Thailand, tuber juices of C. macrophylla are sprayed

on vegetables to kill green flies and root juice is used to kill worms

in the backs of buffaloes (Collins 1441).








Dyes and Fibers


Clitoria ternatea is the only species of the genus that is

reported to be used as a dye plant. Rumpf (1747) and Uphof (1968)

reported that in the Moluccan island Ambon, the flowers were boiled

with rice to give the rice a blue tinge. Burkill (1935) reported

identical use in Malaya. Burkill also noted that the juice of the

leaves is used on some occasions in the Dutch Indies to color food

green. He also reported that the flowers were used as a dye to give a

temporary color to white cloth. The dye was used in the Rhio

Archipelago (=Riouw Archipelogo, a portion of the Malaya Archipelago)

and as a reagent for detecting acid and alkaline solutions, as does

litmus paper.

Liana species are used locally by the natives to make ropes.

Uphof (1968) reported that C. lasciva stems are the source of a fiber

used by natives of Madagascar for making ropes. Clitoria sagotii

was noted as a bush rope by Jenman 4930, in Guyana.


Food for Man


Uphof (1968) reported that the pods of C. ternatea are consumed

as boiled vegetables.


Fodder and Green Manure


A number of references report that C. ternatea is a good forage

crop and is edible by cattle, sheep, and goats. Burkill (1935), for

example, reported that sheep and goats will eat the foliage. In

Venezuela, the species is used as a forage crop (Zambrano 105).

Several experimental studies have reported on the establishment of


L








these species in pastures, and other studies indicate its nutritional

value. These studies will be detailed in the later chapter "Prior

Research on Clitoria," under the section "Agroromical Research."

Burkill (1935) reported that C. ternatea has been used as a green

manure, but that it climbs too much, although it was fairly successful

in some respects, on land that was to be fallow for a short time.

Burkill (1935) reported that C. laurifolia was only fair as a green

manure and that better plants existed for the purpose. Uphof (1968)

reported that this species was used as a jreen manure in warm

countries. Holland and Joachin (1933) found that C. laurifolia was

well adapted as a hedge plant to control terraces on steep slopes in

tea plantations in Ceylon (=Sri Lanka). tBrkill (1935) noted that

the erect stems made this species well suited ,:s a hedge plant on

contoured terraces.















HISTORY OF CLITORIA


The history of Clitoria begins nearly a century before its

officially recognized origin in 1753. Although nomenclaturely

irrelevant, the pre-Linnaean names published and the accompanied

data presented provides better understanding of the genus and its

close relationships with the genus Centrosema.


Origin of the Generic Name


The debatable origin of the generic name has been a very

distasteful subject to many botanists in the past. The most commonly

supposed origin of the name is from the Latin "clitoris," an

anatomical term by which it is indicated that the flower bears a

resemblance to the human female sexual apparatus. This unscientific

notion of comparing the flower with the female reproductive organs, a

practice noted by some as one of questionable taste or immoral

behavior, has been disturbing to a number of botanists. Solutions to

this problem have been various. First, ignore the problem by making

no reference to the subject. Second, circumvent the issue by

concluding that the origin of the name is lost in history, and today

has no significant meaning. Third, gently touch upon the problem by

indicating that the origin is from the flower's fantasized resemblance

to "a female organ," or to "a female structure," or to "an organ."

Fourth, circumvent and attack the problem by indicating that the


I








origin is from the Greek "cloisto: to enclose' which refers to the

manner in which the organs of fructification ate enclosed within the

keel. Or one can use as the origin, Clitor, a town in Arcadia

(Stearn's "Botanical Latin," 1966, has Arcddia = Arkadhia,

Peloponnisos, S. Greece, a European country side outside the

distribution of the genus). Fifth, a direct attack upon the problem

by indicating the immoral practice and unscientific method of

selection of the generic name, which should be selected to reflect

some botanical structure, to reflect a geographic origin, or to honor

a distinguished person by naming the genus after him. Hence the genera

established as substitutes, Ternatea Tourn. for Clitorius Petiver, and

Nauchea Desc. and Vexillaria Eaton substituted for Clitoria L.

To this author, it seems clear that tHe o0 igin of the name can be

traced to Breyne (1678) who published Flos clitoridis Ternatensibus

for an exotic cultivar from the Moluccan island of Ternate. He

indicated the vernacular name to be "Bokyni cotele de Principisse

Clitoris." Rumpf (1747) expanded upon this by indicating the Ternatice

names to be "Saja Cotele" and "Bokyma Cotele" Irom which Breyne erected

Flos clitoridis (flowers clitoris-like) based upon clitoris principisse

(or origin from clitoris), and from which the Portuguese name, Fula

criqua (to speed or hasten breeding or creation) were derived and

appropriately named. The Portuguese and Ternatice names were probably

based upon the ancient "Doctrine of Signatures," a belief that plant

structures which resembled human structures were created to furnish

remedies for the ailments of those body structures. A number of

Clitoria species are used as aphrodisiacs, both for man and for

animals, in widely separated geographical -reas and cultures. Breyne








evidently recognized a resemblance in the flower to the female anatomy

as others hkid, because ne iin:orporated the con rpt that the flowers

were clitoris-like in his polynominal name


G neral Histor,,


Breyne (1678), a German botanist, was the first to describe and

to illustrate d representative of the genu, Clitoria. In "Exoticarum

Plantarumn Cnturia Priiia" he described a plant, from the Indian

subcontinent and transferred to Europe for cultivation, as Flos

clitoridis ler!natensibus. SBieyne describe I th- showy papilionaceous

flowers as inverted, with the vexillum ventrali/ located, a unique

characteristic that segregates Clitoria fmin n mt other legumes.

However, in tho excellent illustration (pi te I, drawn by Stech1),

the flowers ar r not resipinale (possibly .:, arl ist's or engraver's

error). The v,,xillum appear ;d dorsally a, in typical papilionaceous

flowers. 'Breyne's description included hi,,it, calyx, bracteoles

(Breyne used the term "leafl ts"), and fruit (i.ot illustrated), major

characters sedl by later authors in segreg tini Clitoria species. The

illustration clearly shows tie odd-pinnat( y c poundd leaves and

persisting bra:ts and sLipuls (the stipul *s ai e drawn too large for

the species), additional characters used .irly in the determination of

species. Breyne's represent.ition is easily recognized as Clitoria

ternatea L.. the type species for the genu .reyne reported the plant

from the Moluccan island of Ternate, and ited that it bore the name


1. Hunt Iot. Cat. 1:37.1 attribute d ,winc, to Andreas Stech and
engraved by Issac Saal. Plate bears initi Is "AS" at left bottom,
"IS" at rigit iottoii.








of "Bokyni iotele de Principisse Clitoris.' the source for Breyne's

polynominal name. Breyne reported a coll, tioi by Sevenhuysen in

1667 from Domiingo (=Doiiinican Republic) a: related to the Ternate

specimen. If verified as a dlitoria, it would d represent an earlier

record of tie genus.

Breyne also described and illustrated (plate 32) a second

representative as Plantt Legiminosa Brasiliana, Phaseoli facie, flore

purpureo maAiro. This specimen provided Ire earliest record of the


species known as Clitoria braisiliana L., 1

becoming the type species of the genus Cen

Plukenat (1691) in his 'Phytographia'

Foenum1 grae. u!iin phaseoloides ;'irginiana fle

the earliest record of Lhe species known o

later corre tly transfer' red LO the genus (

Petiver (1704) in his Appendix to hi.

Siccis Petiverianus" published two polynoh

data. His number 54 was Cli :orius Marianu,

record of this polynominal has been found

number 55 w.s Clitorius Marinnus trifolia(

the earliest record of the species known n.

was the first to use Clitorii or a form oi

"generic" nine.

Tourneforc (1706) in hi, article "Sui

queques Nouveaux Genres de Plantes" descri


iter transferred to and

Lrost ,a.

described and illustrated

'e aiiplo caeruleo. This is

Cl toria virginiana L.,

ntr semla.

"Ca' logus Plantarum Hortis

nal both lacking additional

trifoliatus viridis. No

iteo after this date. His

I sibtus glaucus. This is

Cl toria Iiariana L. Petiver

the basic root word as the



ie di L'etablissement de

,ed the genus Ternatea. He


provided an illustration of the flower wil cal/x, individual petals,

fruit, and :,eels. Thre.t meniiers of the g,us 'are included. His

first species was Ternaea flore sim plici, caeijleo based upon Breyne's








specimen. Breyne's specimen became the type for the genus and the

source of the generic name, which Tournefort indicated was named after

the Moluccan island of Ternate. Tournefort's second species was

Ternatea flore pleno, caeruleo based upon a double-flowered specimen

described by Commelin in 1697. His third species was Ternatea flore

simplici albido, a white-flowered specimen recognized for the first

time. All three species are now Clitoria ternatea L.

Dillenius (1732) in his "Hortus Elthamensis" described and

illustrated Clitorius trifolius, flore minore caeruleo. He segregated

this specimen from those of Breyne, Petiver and Tournefort, and

concluded that it represented a new member of the genus Clitorius as

defined by Petiver. His specimen is Clitoria virginiana L., although

no reference was made to Plukenet (1691), the earliest describer of

this species.

Linnaeus (1737b)in his "Hortus Cliffortianus" became the first to

use the generic name Clitoria. He recognized two species. His first

species was Clitoria foliis pinnatis, based upon specimens described

by Breyne (1678), Rheede (1688), Plukenet (1691), Rivinius (1691)

Commelin (1697), Tournefort (1706), Dillenius (1719), and Burman

(1737). Breyne's Flos clitoridis ternatensibus was the source of the

generic name. Neither Petiver (1704) nor Dillenius (1737), both of

whom had used Clitorius, was cited. The diagnosis was based upon

Phaseolus foliis pinnatis Riv. The distribution was given as

Malabaria (=S.W. India), Zeylona (=Ceylon, now named Sri Lanka), and

Ternate. Linnaeus recognized three varieties. His typical variety had

simple blue flowers (=Tournefort's first member). His variety alpha

was white flowered (Tournefort's third member). His variety beta had








double blue flowers (Tournefort's second member). In addition,

Linnaeus gave a brief Latin description of variety beta. His second

species was Clitoria foliis ternatis based upon Breyne's Brazilian

specimen and Plumier (1693).

Linnaeus (1748) in his "Hortus Upsaliensis" recognized the same

two species, but used binominal names. His first species Clitoria

foliis pinnatis became Clitoria ternatea. His species Clitoria foliis

ternatis became Clitoria brasiliana.

Nomenclaturally, the official history of the genus began with

Linnaeus (1753) in his "Species Plantarum." Linnaeus established the

genus Clitoria (p. 753) with four species: C. ternatea and C.

brasiliana from his earlier works (1737band 1748), with the addition

of C. virginiana (based upon Dillenius, 1737, and Gronovius, 1739) and

C. mariana (based upon Petiver, 1704). His brief Latin diagnosis for

each species was based upon leaflet number and calyx shape. India,

Brazil, Virginia, and northern America were given as the ranges of

his four species.

The International Code of Botanical Nomenclature (Stafleu, 1972)

officially recognizes that those generic names which first appear in

first edition (1753) of Linnaeus's "Species Plantarum" (e.g. Clitoria)

are associated with the first subsequent description given under those

names in the fifth edition (1754) of Linnaeus's "Genera Plantarum."

Clitoria is described under genus number 796 (p. 334) of Linnaeus's

"Genera Plantarum" as:


CAL. Perianthium monophyllum, erectum, tubulatum,
quinquedentatum, persistens. COR. Vexillum maximum
rectum, emarginatum, margin undulatum, patens. Alae
oblongae, rectae, obtusae, vexillo breviores. Carina








alis breviores. STAM. Filamenta diadelpha (simplex
& novemsidum). Antherae simplices. PIST. Germen
oblongum. Stylus adsendens. Stigma obtusum. PER.
Legumen longissimum, lineare, compressum, uniloculare.
SEM. plura, reinformia.


This diagnosis represents the first validly published description

of the genus Clitoria L.

Miller (1754) in his "Gardner's Dictionary" provided the first

encyclopedic reference work on the genus. He adapted Tournefort's

treatment (1706) and recognized the genus Ternatea. Thus, three of the

four species he recognized followed Tournefort's treatment, not

Linnaeus's (1737) who treated these as three varieties of one species.

A fourth species, Ternatea Americana perennis flore caerulea from

Jamaica, was included. In a later edition of his work, Miller (1759)

adopted Linnaeus's generic name Clitoria and his four species. The

three species of Tournefort included in his earlier edition were

included as C. ternatea in the later edition. The treatment of the

Jamaican species is unclear, as no reference to this specimen was

made. It may have been included under C. brasiliana or C. virginiana,

or treated under another genus, but it could not have been included

in C. mariana, a species which does not occur in Jamaica.

Miller described the flower as papilionaceous, with the vexillum

longer .than and hiding the keel and wings, the fruit opening by the

sutures, and the seeds kidney-shaped. The first description of the

stamens and stigma were given. He referred to the flower as a

butterfly type.

Buchoz (1775) in his "Histoire Universelle du Regne V6egtal ou

Nouveau Dictionnaire Physique et Economique de toutes les Plantes qui

Croissent sur la Surface due Globe" became the first to describe the








genus Clitoria in greater detail. Clitoria was described as: calyx

tubular, 5-toothed, persistent; corolla papilionaceous; vexillum very

large, straight, notched, with undulating margins; wings oblong,

straight, obtuse, much shorter than standard; carina much shorter than

wings, round underneath, furrowed; stamens diadelphous, anthers simple;

ovary oblong, style ascending, stigma obtuse; legume very long, linear,

flat, bivalved, awl-tipped; seeds numerous, reniform. Buchoz followed

the Linnaeai treatment and included five species, C. ternatea L.,

C. brasiliana L., C. virqiniana L., C. mariana L., and C. galactia

Crantz. Each species ,was described with notes on origin, culture, and

economic importance.

Lamarcl. (1786) in his "incyclopedie tiethodique, Botanique"

described seven species, of which Clitorid heterophylla (East Indies)

and Clitori, falcata (Santo Domingo) were newly described.

Description, lacked measurements, but included habit, stem, leaf

shape and pubescence, leaflet number, peduicle location and relative

size as compared to petiole size, calyx, flower size and color, and

fruits. All sizes were relative and compared to similar structures of

other species. These characters, along with the addition of bracteoles

and stipules, were to become the major characters utilized by later

authors in describing and segregating Clitoria species. Lamarck's

style of describing Clitoria species established a trend that continued

through Bentham's revision of the genus in 185' and beyond. Poiret

(1811), in supplement to Lamarck's work, described three new species

and brought the total number to thirteen.

Desvaux (1813) in his journal article "Precis de Caracteres de

plusieures ienres, de la Famille des Legui. ineuses" described a new








genus Neurocarpum, with a tubular calyx of five, equal teeth and a

stalked fruit characterized by a longitudinal nerve on each side. He
2
indicated Aublet's Crotalaria gajanensis (published by Aublet as

Crotalaria guianensis) and a new species, N. ellipticum Desv., were

placed in the genus. In a succeeding article a year later, Desvaux

(1814) described the new species N. ellipticum and included

Crotalaria guyanensis Aubl. and Crotalaria longifolia Lam. as

synonyms of a species called Neurocarpum janense Desv.

Eaton (1817) in his "Manual of Botany for the Northern States"

established the genus Vexillaria as a substitute for Clitoria because

J. E. Smith severely censured the name Clitoria (reference of the name

reflecting the female anatomy) in Rees' "Cyclopaedia" (1807).3 Rickett

and Stafleu (1959), in an expanded footnote regarding Vexillaria Benth.

(=Centrosema), quoted Smith as:


CLITORIA, in Botany, (from XAElw, claudo, include),
expressing the manner in which the essential organs
of fructification are enclosed or shut up in the
keel and wings of the corolla. Whatever may have
been in the thoughts of Petiver, by whom the name
was first introduced into botany; or of the
illustrious naturalists, by whom it has since been
continued, reformed, or sanctioned, we cannot refrain
from entering our decided protest against every
attempt to associate it directly with an anatomical
term, to which, though derived from the same Greek
theme, it has in fact only a very remote,


2. Desvaux erred in his interpretation of Aublet's species.
Desvaux's type (P-59!) does not agree with Aublet's type (BM!).
Therefore, Aublet's name is not synonymized with Desvaux's name. A
complete discussion can be found under the type treatment of subgenus
Neurocarpum, p.
3. This author has not been able to obtain a copy of Eaton's
publication, nor Smith's article. However, a copy of the eight edition
of Eaton's Manual (1840) has been obtained. Clitoria was not mentioned.
Vexillaria Eaton contained three species, V. virginiana, V. mariana,
and V. plumieri, each treated as a Clitoria species by almost all other
botanists.








fictitious analogy. It is greatly to be lamented,
that a fondness for these gross allusions should
ever have been indulged by any, who, in all other
respects, have deserved highly of natural science,
and whose splendid talents should have rendered
them far superior to such grovelling ideas. By
this conduct they may have done all in their power
to pollute a study, which is, perhaps, more than
all others, suited to the loviest part of the
human race, and which, without concealing any
essential part of the sexual system, may easily be
so conducted, as not to excite an unpleasant
sensation in the most delicate female mind. We
do not mean to exempt from the full severity of
this censure our great master, Linnaeus, himself;
for when, not only the purity of moral feeling,
but also the common decorum of polished life, is
infringed, the nullis in verba of the poet will, we
trust, be uniformly our principle and our practice).


Vexillaria Eaton does not appear in Willis' Dictionary, Index Kewensis,

Gray Card Index, nor other standardized index references, nor in

articles associated with Clitoria. Vexillaria Eaton apparently was

not accepted by other botanists of the time, and was thus easily

overlooked by the compliers of the major index references. Vexillaria

Eaton is a probable synonym of the genus Clitoria.

Rafinesque (1818) in a two-line note in a journal article remarked

"Clitoria mariana must form a particular genus Vexillaria." Thus

Vexillaria Raf. was established and recorded by the major standardized

index references. Other botanists did not agree with Rafinesque and

neither. Vexillaria mariana Raf. was used nor were other species of

Clitoria transferred to the genus Vexillaria Raf.

Leandro do Sacramento (1821) in a journal article described a new

species under a new genus Martia, named in honor of the German botanist

Carl F. P. Martius. The description of Martia physodes was detailed

with an illustration provided. The generic characters which indicated

a distinct genus were the lack of a corolla, small stamens with








rudimentary filaments, a style in contact with its own anthers, and a

legume with a protruding lateral nerve. The description was based

upon a cleistogamous flowering collection of Clitoria falcata from

Brazil.

Schultes (1822) in his "Mantissa in Volumen Primum Systematis

Vegetabiliuii" published the species of Leandro do Sacramento under the

name Martiusia physalodes. Perhaps Schultes changed the generic name

because of the homonym Martia Sprengel (1819) which is synonymous with

a non-legume genus.

Humboldt, Bonpland, and Kunth (1824) in their "Nova Genera et

Species Plaiitarum" described four new species of Neurocarpum. A new

genus was cited in synonymy as "Rhombolobium Rich. mss.," probably

based upon a Paris specimen of Richard (P- 2!) whose label bears the

generic description of Rhombolobium, and less likely based upon a

Geneva specimen of Richard (G-545!) whose label bears the data

"Rhombolobiumj Richard, ijenre nouveau qui doit etre place a coti du

Clitoria. 11 y a un angle que [handwriting illegible for this

term] du milieu du Legume du chaque coti." Clitoria L. and Ternatea

Tourn. were treated as distinct genera by Humboldt, Bonpland, and

Kunth.

A. P. de Candolle (1825) published the first major revision of

Clitoria in his "Prodromus Systematis Naturalis Regni Vegetabilis."

Almost published simultaneously was his 'Ti iijires sur la Famille des

Legumineuses" which gave explanations of his reasons underlying the

classification adopted in his "Prodromus." A. P. de Candolle

characterized the genus Clitoria as: caly-, of five teeth with a pair

of bracteoles at its base; the corolla and stamens inserted at the








calyx base at a point visible to the exterior; a large round standard;

a style more or less dilated at the summit; a resupinate flower; and

a fruit of two valves with seeds separated by a cellular partition.

The genus Ternatea Tourn. was placed in synonymy. A. P. de Candolle

recognized four sections within the genus, segregated by the calyx

shape, bracteoles, presence or lack of a vexillum spur, and a leaflet

number.

A. P. de Candolle called section I "Ternatea," based upon Kunth's

treatment of Ternatea in 1824. This section was characterized by a

tubular calyx, spurless vexillum, and five or seven leaflets. Two

species were included, Clitoria heterophylla Lam. and Clitoria

ternatea L. A. P. de Candolle named section II "Euclitoria," based

upon his statement that this section included the true clitorias.

Section II was characterized by a tubular calyx, spurless vexillum,

and three leaflets. Six species were included: Clitoria mariana L.,

Clitoria mexicana Link, Clitoria angustifolia H.B.K., Clitoria

formosa H.B.K., Clitoria glycinoides DC., and Clitoria poitaei DC.,

the latter two newly described species. It should be noted that the

type species for the genus, C. ternatea L., is not in this section,

and thus the section cannot contain the "true" clitorias. Section III

was named "Centrosema" by de Candolle, based upon the spurred vexillum,

a unique characteristic in the legumes. Section III was characterized

by a campanulate calyx of five divisions, a spurred vexillum,

longitudinally striated bracteoles, and three leaflets. Three species

were included: Clitorid virginiana L., Clitoria brasiliana L., and

Clitoria plumieri Turp. Section IV was named "Glycinopsis" by

de Candolle, based upon the resemblance to the genus Glycine.








Section IV was characterized by a campanul.te calyx of five teeth,

corolla and stamens inserted at the calyx uase, style dilated at

summit, longitudinally striated bracteoles, and three leaflets. One

new species, Clitoria berteriana DC., was placed here. A. P.

de Candolle also noted five additional species that may possibly

belong in the genus Clitoria, bringing the total number of species

included to seventeen. The five species that were briefly noted, and

not placed in any of the four distinct sections, were: Clitoria

coccinea Schrad., Clitoria speciosa Cav., Clitoria laurifolia Poir.,

Clitoria vicioides Nees. & Mart., and Clitoria arborescens Ait.

For each species placed in a section, de Candolle gave a brief

Latin description in his "Prodromus." He included the general

distribution of the species dnd cited collections he had seen for

the new species.

A. P. de Candolle treated the genus Nieurocarpum Desv. as distinct

from Clitoria L. in his "Prodromus" following the tradition of other

botanists of his time. He characterized the genus Neurocarpum as:

calyx tubular with five, subequal, acuminate teeth bearing two

bracteoles at its base; vexillum large, subrotund; stamens diadelphous,

legume stipitate, compressed, subtetragonous, valves bearing a

prominent mnidial longitudinal nerve; and with seeds in cellular

partitions. Seven species were included: Neurocarpum simplicifolium

Kunth, NeurLcarpum angustifolium Kunth, Neurocarpum guianensis Desv.,

Neurocarpum ellipticum iesv., Neurocarpum javitense H.B.K., Neurocarpum

macrophyllum H.B.K., and Neurocarpum falcatum (Lam.) DC. A. P.

de Candolle included a new genus, Rhombifolium Rich., in synonymy

based upon a specimen iii Richard's herbari im.








A. P. de Candolle treated the genus Martiusia Schult. as also

distinct from Clitoria and Neurocarpum. He characterized the genus

Martiusia as calyx slightly bilabiate with five teeth, lowermost

longer; corolla lacking; stamens four with two sterile and two fertile,

subciliate anthers; filaments much shorter than the ovary; legume

stipitate, compressed, subtetragonous, bearing a medial longitudinal

nerve. A. P. de Candolle noted that the calyx and legume were those

of Neurocarpum, and that the genus may not be distinct from Neurocarpum.

He included only one species, Martiusia physalodes Schult. The genus

Martia Leandr.-Sacr. was treated in synonymy.

Descourtilz (1826) in his journal article "Sur les Nauchees,

Genre Nouveau dans la Famille des Legumineuses" criticized the use of

the name Clitoria and its reference to an organ from which the name

was derived. He felt that the botanical language should consecrate

the memory of those illustrious persons of importance by adopting

names which recognized their contributions. He proposed the

substitution of the name Nauchea for Clitoria. The genus Nauchea was

named in honor of M. fe docteur Nauche.4 Twelve species of Clitoria

were transferred to Nauchea and a new species was described. Each

species included a description and a distribution. Descourtilz gave

each species a common name, based upon the specific epithet (i.e.

Nauchea multiflora was "Nauchee a fleurs nombreuses"). Other botanists

did not adopt the generic name of Nauchea.

Desvaux (1826) in his article "Observations sur la Famille des

Legumineuses" proposed a revisionary change in the genus Neurocarpum.


4. A prominent French physician, Jacques-Louis Nauches (1776-
1843), who pioneered work in electrical shock therapy.








Two species were placed near those described in 1813. They were

Neurocarpum barbatum Nees, newly describe(, and Neurocarpum laurifolium

Desv., a transfer of Poiret's species (thus should have been N.

laurifolium (Poir.) Desv.). Three species were recognized as different.

They were Neurocarpum rubiginosum Desv., a transfer of Jussieu's

species published by Persoon (thus should nave been N. rubiginosum

(Juss. ex Pers.) Desv.), Neurocarpum glycinoides N.,5 a transfer of

de Candolle's species (thus should have been N. glycinoides (DC.)

Nees), and Nleurocarpum villosum N., newly Jescribed. Desvaux proposed

the name Pilanthum for this group based upon Puiret's name used for

the type of C. gycinoides DC. (=N. glyciilides Nees). The genus

Pilanthus Puir. ex End]. (Genera Plantarum p. 1289, 1841), presently

synonymized with the genus Centrosema, may be based upon Poiteau's

name Pilantlhos, although Endlicher's article has not been seen. The

specimen referred to by Desvaux and also b/ de Candolle (Pilanthum

cited under C. glycinoides in his Prodromun 2:234, 1825), bears the

name Pilanthos tetragonus Poiret. Two new species of Clitoria were

also described, C. laurifolia Nees, a homonym for C. laurifolia

Poiret (1811), and C. sinuata Nees.

Don (1832) in his "A General History of the Dichlamydeous Plants"

followed de Candolle's treatment of Clito ria, Neurocarpum, and Martiusia

with minor revisions. Of the five dubious species listed by

de Candolle under the uenus Clitoria, Don laced two in the section

Centrosema (C. speciosa Cav. and C. arborescens Ait.), one in the

section Glycinopsis (C. coccinea Schrad.), and left one in a dubious


5. Desvaux's use of N. was an abbreviation for Nees.








list (C. vi(ioides Nees & Mart.). The filth species was transferred

to the genus Neurocarpuii (C. laurifolia Poir.) along with C.

glycinoides DC. from section Euclitoria. lie also described two new

species, Clitoria raceimosa and Clitoria alba, and placed them both in


the section Euclitoria.

and Martiusia with the

Bentham (1837) in

de Candolle's treatment

There were three major

segregated the section

to the level of genus,

botanists have adopted

century later, botanist

and Centrosema (DC.) Be

Glycinopsis by transfer


No changes were iimde in the genera Neurocarpum

exception of the transfers already noted.

a journal article on legume genera followed

of these genera with some revisionary changes.

changes in the genus Clitoria. First, Bentham

Centrosema from Clitoria and elevated the taxon

using de Candolle's name Centrosema. Succeeding

this revision, and presently, more than a

s continue to reco!;gize both genera, Clitoria L.

rith. Second, Benthdm eliminated the section

ring Clitoriana berteriana DC. to a


non-clitoritn genus, Periandra, and placed Clitoria coccinea Schrad.

in a dubious status. Succeeding botanists adopted this revision as

the section Glycinopsis disappeared from literature after this date.

Third, Bentham recognized de Candolle's sections Ternatea and

Neurocarpum, and added to them a new section, Bractearia Mart.ex

Benth. Bracteria was described as fruticose species with trifoliate

leaves and bracteoles equal to or longer than the calyx. To this new

section, Bentham transferred Clitoria racemiosa Don and Clitoria

poitaei DC. He also described three new species and placed them in

this section, Clitoria amazonum, Clitoria acuminata, and Clitoria

arborea.








Benthain's treatment of section Ternatea followed de Candolle and

Don with the addition of a new species, Clitoria lasciva Boj. ex Benth.

Section Euclitoria underwent minor revision. Clitoria formosa H.B.K.

and Clitoria angustifolia H.U.K. were transferred to the genus

Centrosema. Bentham added a newly described species, Clitoria

pedunculata Boj. ex Benth., to the two remaining species, Clitoria

mariana L. and Clitoria mexicana Link. Bentham also added notes on a

number of miscellaneous species published as Clitoria, but which had

been transferred to other genera.

Benthain's treatment of the genus Neurocarpum followed de Candolle

and Don with two important changes. First, Bentham placed the genera

Martia Leandr.-Sacr. and Martiusia Schult. in synonymy. Second,

Bentham remarked on the close relationship between Neurocarpum and

Clitoria by stating that Neurocarpum is a Clitoria except that it has

a different legume (i.e. Neurocarpum has a costate legume; Clitoria

has an ecostate legume). Eleven species ware recognized and placed in

two unnamed, artificial groups. Group 1 iid suberect stems whereas

group 2 had procumbent or climbing stems. Five species were newly

described and several species were placed in dubious status.

Bentham (1839), in a publication on British Guianan plants

collected by Schomburgk, substituted the nime Dendrocyamus for his

section Bractearia in the genus Clitoria. He reasoned that the name

Bracteria was inappropriate since it had been given to a genus of the

Rubiaceae and had been used as a section of the genus Chaetogastra

(Melastomaceae).

Bentham (1858) published a second revision of the genus Clitoria

in a Journal of the Linnean Society article. This was the last revision








of the genus until this work over a century later, and was adopted by

all succeeding botanists in their treatment of the genus. The major

change from his earlier revision (1937) was the uniting of the genus

Neurocarpum with Clitoria. Bentham reasoned on p. 35:


In a systematic point of view I had formerly
endeavoured to render Clitoria more natural, by the
elimination of DeCandolle's section Centrosema; and
I now find it necessary for the same purpose to unite
it with the Neurocarpum of Desvaux, hitherto universally
adopted by other botanists, myself included. This
entails the giving up, as a generic character, one which,
in Leguminosae, is usually considered as absolute, the
raised longitudinal nerve or wing along the center of
each valve of the pod. It is the same peculiarity which
has induced the separation of Tetragonolobus from Lotus
among European plants. But in the division of Lotus, as
well as in that of Clitoria, this purely technical
character is unaccompanied by any other differences, and
I now have instances in Clitoria where it is inconsistent
in one and the same species, and even on the same
specimen.


Bentham continued to recognize three sections, although he changed

their names. Section Ternatea remained unchanged, whereas Neurocarpum

was substituted for Euclitoria, and Clitorianthese was substituted for

Bractearia (=Dendrocyamus in 1839) used in Bentham's earlier revision

(1837). No comment was made on the change in section names, nor was

there any reference to the earlier sectional names. However, a

comparison of the two revisions by Bentham indicated that the sectional

diagnosis agreed (although the 1837 description was expanded upon in

1858), the species placed in each section agreed, the order in

which each section was treated agreed, and the number given by Bentham

for each section agreed.

Section Ternatea was characterized as: stems herbaceous or

rarely fruticose, prostrate, twining or climbing; leaflets 5-9, rarely








3; legume valve flat or slightly convex, non-costate, seeds subreniform,

compressed, smooth. Six species were recognized, all known previously.

They included Clitoria lasciva Boj. ex Benth., Clitoria ternatea L.,

Clitoria pilosula Wall., Clitoria heterophylla Lam., Clitoria biflora

Dalz., and Clitoria pedunculata Boj. ex. Benth.

Section Neurocarpum was characterized as herb, twining or

prostrate or short erect; leaflets 1 or 3; legume valve convex, medial

longitudinal costa or rarely ecostate; seeds globose, ovoid, or thick

subreniform, with a viscid coat. This section contained eleven species

placed into three groups based upon the stem habit. The groups were

unnamed and distinguished by one to three "*" symbols. The first group

with twining stems included: Clitoria macrophylla Wall., Clitoria

mariana L., and Clitoria glycinoides DC. The second group with

prostrate stems included two newly described species: Clitoria

flagellaris and Clitoria rufescens. Group three with stems arising

from a lignose rhizome, erect to ascending, included: Clitoria

cajanifolia Presl, Clitoria simplicifolia (Kunth) Benth., Clitoria

guianensis (Aubl.) Benth., Clitoria densiflora (Benth.) Benth., and

Clitoria stipularis (Mart.) Benth., plus a newly described species,

Clitoria nana.

Section Clitorianthes was characterized as erect shrub or tall

climbing; leaflets 3; legume valve flat or slightly convex, coriaceous;

seeds in matured state unknown. This section contained nine species

divided into two unnamed groups based upon the bracteoles. The first

group with narrow bracteoles, or bracteoles much shorter than the calyx,

included Clitoria polystachya Benth., Clitoria brachystegia Benth.,

Clitoria arborescens Ait., Clitoria javitensis (H.B.K.) Benth., plus








two newly described species, Clitoria sellui and Clitoria leptostachya.

The second group with ovate, coriaceous bracteoles subequal to the

calyx, included: Clitoria aiiazonum Mart. ex Bunth., Clitoria racemosa

Benth., and Clitoria hoffmanseggii Benth.

The treatment of each species included a Latin diagnosis, the

distribution as then known, the citation ol one to a few collections,

and synonymy/. Characters used in the diagnosis of the species

included: stem habit; leaf shape and pubescence, and leaflet number;

peduncle size, location, and number of flowers borne, calyx;

bracteoles; legume curvature and costa; ard sometimes stipules or

vexillum pubescence. -enthaii occasionally gave additional descriptive

data with a minimum quantity of measurements.

After lentham's revision, the next century of publications on

Clitoria was minly of a floristic nature, not revisionary. Isolated

descriptions of newly described species placed in the genus also

occurred. Two exceptions to the lack of revision are worth noting.

First, Baker (1879), in a floristic treatment of legumes in "Hooker's

Flora of British India," elevated Bentham's sections to the level of

subgenus. Subgenus Ternatea was characterized as having flat legumes

without a costa. Clitoria mariana L. and Clitoria macrophylla Wall.

were transferred here from Bentham's section Neurocarpum. Subgenus

Neurocarpum was characterized as having a turgid, costate legume.

Bentham's section Clitorianthes was not treated, probably because the

species within this section do not occur in the Paleotropics. Second,

Kuntze (1891) transferred all Clitoria species to the genus Ternatea,

a revision not adopted by later botanists. Occasionally, floras would

include Clitoria and Martiusia as separate genera, the latter genus








containing the species of Neurocarpur (e.jg., Small, 1933; Britton,

1924).

In the earlier parr. of Lhe twentieth century, research in various

areas began on Clitoria (e.g., agronomy, anatomy, chemistry, cytology,

development). The results of these studies are enumerated in the

chapter "Prior Research on Clitoria."

These selected articles represented a partial, highlighted history

of the genus Clitoria. The important portions of the nomenclatural

history-of the genus, above Ihe level of species, are summarized in

Figures 1 ard 2. For both figures, genera were expressed in capital

letters and enclosed in a "box" for better visualization. Genera

included in synonymy were enclosed in parentheses. For the few

polynominals included, ihich are important in tracing the origin of

some generit- names, the key terms are used and underlined, followed

by "..." which represented the missing, nonessential terms. Sections

are numbered and subgenera are preceded by the abbreviation "Subg."


Historical Chronol gy


A more complete historical chronology is presented below. Each

article is presented in a chronological order, by year, with a brief

synopsis of the publication, and nomenclatiral notes. Those articles

marked (*) are expanded upon in the "General History" section

previously discussed, and the reader is referred to its coverage.

Those articles marked (R) are expanded upon in the next chapter, "Prior

Research on Clitoria," and the reader is referred to that chapter for

details.

















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Pre-Linnean (1650-1753: _Po/lnominal Peri;l)

1667: Sevenhuysen was reported by Bieyne (1678) to have collected

a specimen Irom Santo Domingo that was related to the Moluccan specimen

Flos clitoridis ternatensibus. If this specimen is verified as a

Clitoria, this would be the earliest known record of the genus.

1678: Breyne (*) described and illustrated two specimens that are

the framework upon which Linnaeus established two of the original four

species of Clitoria. His Table 31 described Flos clitoridis

ternatensibus (=Clitoria ternatea L.) froii the island of Ternate,

Moluccan Islands. His Table 32 described Planta leguminosae

Brasiliana, phaseoli fcie, flore purpureo maxirno (= Clitoria

brasiliana L.).

1688: Rheede tot Draakestein described Schonga cuspi from Malabar

(=S.W. Indid) which Linrnaeus later cited in synonymy for Clitoria

ternatea. From its description and a photograph of Schongi cuspi as

illustrated in his Table 38, one can easily conclude that this plant

is not a Clitoria specimen.

1691: Plukenet published Foenum graecum phaseoloides Virginianus

flore amnplo caeruleo which is the origin of the third original Linnaean

species, Cl itoria viriniana L.

1693: Plumier described Phaseolus am;ple flore peltato, siliquis

nigris et anqulosis (=Clitoria plumieri Turp.) based upon a specimen

from Santo Domingo.

1696: In his catalogue of Jamaican plants, Sloane included

Phaseolus minor lactescens flore purpureo (=Clitoria galactia Crantz).

1698: Dr. David Kreig brought a specimen to Great Britain that

was collected in Maryland. [his specimen is now deposited in the Royal








Botanic Garden Herbarium at Edinburgh (E-127, Herb. Dubois!). This

specimen represents the oldest known representative of the genus

Clitoria (=C. mariana L.).

1700: Plukenet published Phaseolus Indicus, coeruleus,

Glycyrrhizae foliis alatis, flore amplo Cliterio (=Clitoria ternatea

L.) and Phaseolus flore vexillo, silique rectis teretibus (= Clitoria

brasiliana L.).

1701: Commelin described and illustrated in his Table 24,

Phaseolus Indicus, glycyrrhizae foliis, flore amplo, caeruleo, pleno.

This was the first known example of the unique double flower of

Clitoria ternatea L.

1703: Dr. Edward Bulkley sent a specimen to Great Britain that

was collected from Fort St. George in the East Indies. This specimen

is now deposited in the Royal Botanic Garden Herbarium at Edinburgh

(E-69, Herb. Dubois!). This specimen represents the oldest known

specimen of the type species of the genus Clitoria (=C. ternatea L.).

1704: Petiver (*) published two polynominals in an appendix, the

first to use "Clitorius" as the leading key term of the polynominal.

His number 54 was Clitorius Marianus trifoliatus viridis which

disappeared from Clitoria literature. His number 55 was Clitorius

Marianus trifoliatus subtus ajaucus which was the origin of the fourth

original Linnaean species, Clitoria mariana L.

1706: Tournefort (*) established the genus Ternatea based upon

a specimen from Ternate, a Moluccan Island. Three elements were

published which later served as the framework for the three varieties

of Clitoria ternatea L. These were Ternatea flore simplici, caeruleo








(agreed with Breyne, 1678); Ternatea flore pleno, caeruleo (agreed

with Commelin, 1701); and Ternatea flore simnplici albido.

1709: Petiver published Plukenet's rime (1691) and illustrated

a specimen in his Table 104, Figure 19.

1732: Dillenius (*) described and illustrated Clitorius trifolius

flore minor caeruleo (=Clitoria virginiana L.) which he concluded was

segregated from, yet related to the specimens of Breyne (1678) and

Petiver (1704). No reference was made to Plukenet (1691).

1737: Burmann reported Flos clitorius flore coeruleo (=Clitoria

ternatea L.) from Ceylon (=Sri Lanka presently) citing Breyne (1678)

and Plukenet (1700).

1737: Linnaeus (*) established the genus Clitoria and recognized

two species. The first species was Clitoria foliis pinnatis (=Clitoria

ternatea L.) based upon Breyne (1678), Rheede tot Draakestein (1688),

Plukenet (1700), Commelin (1701), Tournefort (1706), and Burmann (1737).

Three "varieties" were recognized using Tournefort's segregation. The

second species was Clitoria foliis ternatis (=Clitoria brasiliana L.)

based upon Breyne (1678) and Phaseolus flore vexillo amplitismo,

filiquis rectis teretibus published by Plumier in "Spec. 8."6

1739: Gronovius published Clitoria foliis ternatis, calycibus

oblongus, from Virginia (=U.S.A.), citing Petiver (1704) number 55 in

synonymy. He provided the first record of a specimen citation for

Clitoria, Clayton 108 (=Clitoria mariana L.). He also reported

Clitoria foliis ternatis, calycibus campanulatis, based upon Dillenius

(1732), and Clayton 112 (-Clitoria virginiana L.).


6. This author was unable to establish the publication title and
therefore was unable to obtain a copy of Plumier's publication.








1740: Royen reported three species from Batavia (=Java). The

first species was Clitoria foliis pinnatis of Linnaeus (1737). The

second species was Clitoria foliis pinnatis, caule decumbente (=non-

Clitoria). The third species was Clitoria foliis ternatis of Linnaeus

(1737).

1747: Rumpf (Rumphius) described "Bongu Biru" from the island of

Amboin, a Moluccan Island, as Flos coeruleus (=Clitoria ternatea L.).

Vernacular names in several languages were noted.

1748: Linnaeus (*) established binominals for his two species

published in 1737. The two species became Clitoria ternatea and

Clitoria brasiliana, respectively.

Post-Linnaeain (1753-1824: Descriptive Period

1753: Linnaeus (*) officially established the genus Clitoria,

and recognized four species, Clitoria ternatea and Clitoria brasiliana

based upon his prior works (1737; 1748), Clitoria virginiana based upon

Dillenius and Gronovius, and Clitoria mariana based upon specimen

number 55, Petiver (1704). India, Brazil, Virginia, and northern

America were given as the ranges of the four species.

1754: Linnaeus (*) published a description of the genus

Clitoria. This description is the first description published that is

officially recognized by present botanists through the rules established

in the International Code of Botanical Nomenclature (Stafleu, 1972).

1754: Miller (*) in his fourth edition of "Gardner's Dictionary"

followed Tournefort (1706) in the treatment of the genus Ternatea. Thus

the genus Ternatea Tourn. ex Miller became officially established. In

the seventh edition (1759) Miller adopted the Linnaean treatment. This








was the first encyclopedia type of reference work on the genus.

Descriptions and horticultural notes were provided.

1756: Browne reported two species from Jamaica with Linnaean

polynominals cited in synonymy. They were Clitoria major scandens,

foliis subrotundo-ovatis, floribus qeminatis (=Clitoria virginiana L.)

and Clitoria minor scandens, foliis subvillosus oblongo-ovatis,

floribus ge!inatis (=Clitoria ternatea). His Table 32 bore the title

"Clitoria galactia"; however, the plant was described under the genus

Galactia in the text, with Sloane's Phaseolus minor lactescens cited

in synonymy. From the illustration and description, this plant is not

a Clitoria.

1758: Linnaeus in his tenth edition of "Systema Naturae" cited

two additional species in addition to those established in 1753.

Neither were numbered as the other species were numbered. Clitoria

lactescens disappeared from Clitoria literature after this date, except

for Richter (1840), who cited the species in synonymy for Clitoria

galactia L. The second species, Clitoria zoophthalmum, also disappeared

from the literature until cited by Richter as synonymous with the

non-Clitorian species, Dolichos urens L.

1762: Gronovius published the same two polynominals in the second

edition of his Virginia Flora as he did in 1739. A third non-Clitorian

species published in 1739 (Clitoria foliis pinnatis, caule decumbente)

was now transferred to the genus Cracca.

1763: Linnaeus described a new species, Clitoria galactia, based

upon Sloane's specimen cited by Browne (1756). No reference was made

to Clitoria lactescens.








1766: Crantz followed Linnaeus in his treatment of C1itoria.

He listed tne five species in Linnaeus "Species Plantarum" (1763), but

altered the spelling on two species, published as C. ternatensium and

C. galactea.

1775: Aublet described a new species, Crotalaria guianensis, from

French Guiana. He was the first to use bracteole and stipule

characteristics, and the first to note the prominent costa on the fruit,

a characteristic that was important in the segregation of genera and

section's by later botanists. Aublet had cJllected several plants from

French Guiana, which included two species similar in gross appearance

that seemingly matched the illustration provided in his Plate 305.

Thus, from the misidentification, the namL Crotalaria guianensis Aubl.

would become associated with two distinct species (i.e. Clitoria

guianensis iAubl.) Benth. and Clitoria laurifolia Poir.) and provide

confusion ii later literature and specimen identification.

1775: Buchoz (*) followed the Linnaedn treatment and provided

the first detailed generic description of Clitoria, which included

perianth, stamens, pistil, legume, and seeds. Each species was

described aid notes on origin, culture, and economic properties were

included.

1775: Forskal (or Forsskal) described a new species, Lathyrus

spectabilis (=Clitoria ternatea L.) from Egypt. He was the first to

indicate the villous style typical of the genus.

1786: Lamarck (*) described two new species of Clitoria, C.

heterophylla and C. falcata, along with the basic five Linnaean species.

He also described Crotalaria longifolia citing Aublet's Crotalaria

guianensis in synonymy. Thus, Lamarck's riime is superfluous for

Aublet's name.








1786: Scopoli described Clitoria micrantha and segregated Royen's

C. brasiliana from the species and called it C!itoria quadralupensis.

Both species plus C. galactic L. were described with detailed

descriptions and illustrated.

1788: Swartz described ?Clitoria multiflora with a two-line

Latin diagnosis based upon a plant from Santo Domingo.

1788: Walter's Carolina flora recognized two species, C. mariana

L. and C. virginiana L., segregated on leaflet number and calyx type.

1789: Browne republished his treatment of Clitoria originally

published in 1756, still not adopting the Linnaean treatment. His

Table 32, Figure 2 continued to bear the totle "Clitoria galactia"

while described under the genus Galactia.

1791: Gaertner, in his publication on fruits and seeds of various

plants, described and illustrated those of C. ternatea L.

1792: Richard described Clitoria capitata with characteristics

that are too vague for placement of the species without the specimen

in hand. No type was given.

1796: Salisbury published Clitoria spectabilis which was based

upon Lathyrus spectabilis Forssk. and C. ternatea L. Salisbury's name

was superfluous for the Linnaean species.

1797: Raeuschel published Clitoria aquiliupensis, a nomen nudum,

in his list of Clitoria species.

1798: Roth described Clitoria amoena with a very detailed Latin

description (clearly marking it as a Centrosema member). It was

published in Roemer's "Archiv fur die Botanisk" and republished by

Roth in his "Catalecta Botanica" in 1800.

1802: Cavanielles described a new species, Clitoria speciosa.








1804: Desfontaine in his "Tableau dt 1 'ecole de Botanique" listed

Clitoria glbella and Clitoria galactia glbella as Clitoria glabre.

Both C. glalella Desf. and C. glabre Desf. were without any description

or discussion and thus, nomen nudum.

1806: Salisbury described and illustrated Clitoria calcarigera.

Three varieties were described based upon leaflet shape. Variety alpha

was the typical representative. Variety beta was based upon Dillenius

(1732). Variety gamma was based upon Clitoria virginiana L. Thus

Salisbury's name was superfluous for the Linnaean name, C. virginiana.

1807: Persoon included ten species of Clitoria in his treatment

of the genus. Each species had a Latin diagnosis with habitat. Two

species fro., Domingo were newly described. Clitoria plumieri Turp. was

based upon a specimen sent to him by Turpin bearing a Latin diagnosis

under the nime. The species was named for Plumier who described the

species in 1693. The second new species w,.s Clitoria rubiginosa Juss.

ex Pers. often cited as C. rubiginosa Pers.

1811: Poiret in a supplement to Lamarck's "Encyclopedie

Methodique Botanique" added seven species to those listed by Lamarck,

the last one, C. capitata Rich., lacking a number. Newly described

were Clitoria polyphylla and Clitoria laurifolia from Porto Rico and

Clitoria bracteata of unknown origin. The other three species included

the new pair described in Persoon plus Swartz' species. Clitoria

amoena Roth was synonymized with C. brasiliana. Clitoria galactia was

reported as the type for a new genus Galactia Brown which was adopted

by Michaux.

1812: Curtis illustrated C. ternatea in color in his "Botanical

Magazine" in plate 1540.








1813: Balbis listed three species ot Clitoria in his "Catalogus

Stirpium." Clitoria broussonetii, named for the French botanist who

sent the specimen to Balbis under a name loti coerulei, was newly

described in a footnote. The footnote contained a Latin diagnosis and

placement of the species near C. galactia.

1813: Desvaux (*) described the genus Neurocarpum and indicated

that it included Aublet's Crotalaria gajanensis ( published as

C. guianensis by Aublet) and a new species, N. ellipticum, which lacked

a description and is a nomen nudum.

1814: Desvaux (*) described Neurocarpum ellipticum and included

Crotalaria guyanensis Aubl. (Aublet published as: guianensis) and

Crotalaria longifolia Lam. as synonyms of Neurocarpum janensis Desv.

1814: Roxburgh listed five species of Clitoria cultivated in

Calcutta. Clitoria erecta, a new species, lacked a description and

thus was a nomen nudum.

1814: Brown described a new species, Clitoria arborescens, in

Aiton's "Hortus Kewensis," based upon a cultivar from Trinidad. This

species is usually cited as C. arborescens Ait.

1817: Eaton (*) substituted the name Vexillaria for Clitoria

because of the criticism regarding the alleged resemblance of the

Clitoria flower to the female reproductive anatomy. Eaton's

superfluous name was not adopted by botanists.

1818: Edwards illustrated and described in detail Clitoria

plumieri Turp., with notes presented on the history of the species.

1818: Rafinesque (*) established the genus Vexillaria Raf. by

his published note "Clitoria mariana must form a particular genus








Vexillaria." Neither the generic name nor Vy. mariana (L.) Raf. were

adopted by botanists.

1820: Curtis published a color plate and notes on Clitoria

heterophylla Lam. (Plate 2111).

1821: Schrader described Clitoria coccinea, and noted it near,

yet distinct from Clitoria falcata Lam.

1821: Nees included a note that Clitoria coccinea Schrad. was

near his species, but uncertain where Clitoria falcata Lam. belonged.

This was the source of Clitoria falcata Nees, a name associated with

C. coccinea Schrad. in synonymy in later literature, but the name was

not validly published. Hence, C. falcata Nees is an illegitimate name.

1821: Trdttinnick cited fourteen species in his discussion of

the genus Clitoria, and described and illustrated two cultivars,

C. ternatea and C. brasiliana.

1821: Leandro do Sacramento (*) described a new genus Martia

based upon a cleistogamous specimen from Brazil. The genus was named

in honor of the German botanist Carl Martius. One species, Martia

physodes, was placed in the genus. Martia is a homonym of Martia

Spreng. (1818).

1822: Schultes (*) published Leandro do Sacramento's species

under the name Martiusia physalodes. The change in generic name is

presumed to be due to the recognition of Martia as a homonym. The

change in the spelling of "physodes" is unclear.

1824: Kunth described in detail and illustrated two new species

of Neurocarpum, N. simplicifolium (plate LIX) and N. angustifolium

(plate LX). Both species were from Brazil.








1824: Humboldt, Bonpland and Kunth (k) described four species,

two of which were published three months errlicr by Kunth. The other

two were newly described in detail. They were N. javitense from

Javitam (=Yavita, Amazonas, Venezuela) and N. iacrophyllum from

Novo-GranatJ (=Columbia). A new genus Rhoinbolobium Rich. based upon

Richard's manuscript was cited in synonymy. The genus Ternatea Tourn.

was recognized as separate and included T. vul(aris, a new name for the

Linnaean C. ternatea. Under a third genus, Clitoria, two new species,

C. angustifolia and C. forlmosa, were described from Brazil.

1824: Nees and Martius described two new species from Brazil,

Clitoria vi:ioides and Clitoria angustifol ia. A Latin diagnosis was

given for Clitoria tomentosa which was observed as differing from

C. rubiginosa Pers.

1824: Schrank described Clitoria gladiata, a new species from

Brazil.

Post-Linnaeon (1825-1853: Revisionary Period)

1825: Hamilton miscited the genus Neurocdrpum Desv. as

"Neurocarpon." He split Desvaux's species (1814) such that N.

ellipticum was based upon a specimen in Dcsvaux Herb. and N. guianensis

included Crotalaria guianensis Aubl. and (rotalaria longifolia Lam.

He noted the relationship to Clitoria laurifolia Poir. and C.

rubiginosa Pers., both bearing the prominent lateral nerve on the

legume.

1825: Velloso described and illustrated four new species,

Clitoria flumensis (plate 123), Clitoria brasiliana (plate 129),

Cl-itoria gewna (plate 130) and Clitoria insulana (plate 131).








1825: A. P. de Candolle (*) published in mid November the first

major revision of the genus Clitoria in his "Prodromous." The genus

was divided into four sections. Section I (Ternatea Kunth) included

two species. Section II (Euclitoria) included six species, C.

glycinoides and C. poitaei newly described. Section III (Centrosema)

included three species. Section IV (Glycinopsis) included only the

newly described C. berteriana. In discussion of C. glycinoides

de Candolle referred to Poiteau's Pilanthus. This was an earlier

record of Pilanthus Poit. ex Endl. (Genera Plantarum 1289, 1841). The

genera Martiusia (one species) and Neurocarpum (seven species) were

treated as separate genera. This publication is the source for

N. falcatum DC. which was an error in citation by later botanists.

Lamarck's species was transferred by de Candolle from Clitoria and

should have been cited as N. falcatum (Lam.) DC.

1826: A. P. de Candolle (*) published his "Memoires sur la

Famille des Legumineuses" nearly simultaneously (early February) with

his "Prodromous," in which he explained his reasons for the classifi-

cation used in his "Prodromous." This was the source cited for the

origin of C. berteriana; however, the "Prodromous" predated this work

by almost three months.

1826: Desvaux (*) proposed a new group within Neurocarpum that

would bear Poiteau's name Pilanthum. This section included two

transfers cited as N. glycinoides N. (C. glycinoides DC.) and

N. rubiginosum Desv. (C. rubiginosa Pers.) plus newly described

N. villosum N. Two species were placed near Desvaux' species described

in 1813, N. laurifolium Desv. (C. laurifolia Poir.) and N. barbatum N.,

newly described. Two species of Clitoria were newly described,

C. laurifolia Nees, a homonym of Poiret (1811), and C. sinuata Nees.








1826: Descourtilz (*) substituted the superfluous name Nauchea

for Clitoria and listed thirteen species, one newly described as

N. pudica.

1827: Tussax described and illustrated Turpin's C. plumieri from

the West Indies.

1828: Wallich listed four species in his catalogue. Three were

new species which lacked descriptions, but had specimens cited for

each. They included Clitoria macrophylla Wall. (no. 5345), Clitoria

acumina-ta Grah. (no. 5346) and Clitoria pilosula Wall. (no. 5347), each

a nomen nudum.

1829: Descourtilz described the genus Nauchea, and illustrated

and described three species, N. pudica, N. virginiana and N. rubiginosa,

each with medicinal notes.

1830: Sweet listed twelve species in his catalogue with Clitoria

occidentalis listed as a substitute for the non-Linnaean virginianaa

B.R. 1047." The species is not described and the citation is unclear.

1830: Guillenim and Perrottel in their "flora Senegambiae

Tentamen" describe a new species, Clitoria micrantha, a homonym of

Scopoli (1786).

1832: Curtis illustrated C. arborescens. This illustration was

a composite of two species, C. arborescens and C. javitensis which

resulted from additions made to an unpublished drawing of 1822.

1832: Presl described and illustrated Neurocarpum cajanifolium.

1832: Geel described and illustrated the "Plumier Clitoria."

1832: Zuccarini described Martia mexicana in detail.

1832: Don (*) adopted de Candolle's trealinent of Clitoria,

Martiusia and Neurocarpum. He described Clitoiia racemosa and Clitoria








alba and placed them in Sect. Euclitoria. Four of the five dubious

species listed by de Candolle were placed. Clitoria broussonetii was

transferred to the genus Cologania. Clitoria phyrne Juss., C. mariana

ioc. and Sesse ined., C. anustifolia Nees and Mart. and C. rubiginosa

Nees and Mart. were synonymized with Galactia species.

1835: Pritzel listed Nauchea clypeata Desc. based upon "Desc.

Ant. 8,591," a publication not obtained by this author.

1837: Hooker described and illustrated Clitoria virdiflora Bouton

ex Hook'.

1837: Bojer described Clitoria lasciva and reported Clitoria

virdiflora Bouton mss. Both Hooker and Bojer cite "Bouton mss. in hb.

nostr." Stafleu's "Taxonomic Literature" (no. 537) indicated Hooker

published sometime within October 7 to November 8. The date of Bojer's

publication was unknown except for the year, although if it preceded

Hooker, then C. virdiflora Boj. was a nomen nudum. Bojer made no

reference to Hooker.

1837: Schlechtendal reported Clitoria mariana L. from Mexico.

This was the source of Clitoria mariana Schlecht. cited by other

botanists. Schlechtendal had misidentified his specimens, which are

Clitoria mexicana Link.

1837: Bentham (*) published a revision of de Candolle's treatment

of Clitoria, Neurocarpum and Martiusia. Two sections of Clitoria were

eliminated. Section Centrosema was elevated to a genus level. Section

Glycinopsis was eliminated through the transfer of its only species to

another genus. To the remaining two de Candolle sections, Bentham

added the section Bractearia. Martiusia and Martia were synonymized

with Neurocarpum. Bentham noted the close relationship of Clitoria and








Neurocarpum segregated only by the prominent costa on the legume. New

species included Clitoria amazonum Mart. ex Beiith., Clitoria acuminata

Benth., Clitoria arborea Benth., Clitoria raceinosa Benth., Clitoria

pedunculata Boj. ex Benth., Neurocarpum longifolium Mart. ex Benth.,

Neurocarpum frigidulum Mart. ex Benth., Neurocarpum rufescens Benth.,

Neurocarpum densiflorum Benth., and Neurocarpum bracteatum Mart. ex

Benth.

1838: Schlechtendal described Clitoria schiedeana.

1838: Lindley reported that C. ternatea roots were emetic.

1839: Bentham's 1837 article was republished with the title

revised.

1839: Bentham reported on Schomburgk's Biitish Guiana collections

and substituted the name Dendrocyamus for his section Bractearia. He

reasoned that the name Bractearia was preoccupied by a genus in the

Rubiaceae and in a section of Chaetogastra, and thus inappropriate for

Clitoria.

1839: Bentham described Clitoria polystachya in his enumeration

of Hartweg collections.

1840: Bentham described Neurocarpum flagellare and synonymized

N. frigidulum with N. longifolium in Hooker's "Journal of Botany."

1840: Steudel published a list of Clitoria species and species

transferred. The following binominals, each a nomen nudum, appeared for

the first time: C. brasiliana Arrab., C. !Irahdmi Steud. (C. acuminata

Grah.), C. insulana Arrab., C. micrantha Smith, C. phryne Commers. and

C. pudica Steud. (Nauchea pudica Desc.). The latter name was a

transferred name and legitimate. Clitoria grahami was an apparent








substitute name for the homonym C. acuminata Grah. which to this date

was still undescribed.

1841: Steudel's second volume published two new binominals for

Neurocarpum, each a nomen nudum. They included Martia brasiliensis

Zuccar. (N. ellipticum Desv.) and N. mexicanum Steud. (Martia mexicana

Zuccar.), both transferred to Neurocarpum, the latter described

previously, the former undescribed but placed in synonymy.

1841: Hooker and Arnott described Neurocarpum multiflorum in their

enumeration of the plants collected on the Beechey's voyage.

1841: Brunbury described Neurocarpum restipinatum from Brazil.

1842: Hasskarl described two new species from Java, Clitoria

oblonga (C. vir~jniana L. var elliptica DC.) arid Neurocarpus retusus.

1842: Bertoloni described and illustrated Clitoria alabamensis.

1843: Martius and Galeotti described Cliloria multiflora and

Clitoria grandiflora from Mexico.

1844: Bentham described Clitoria brachystegia from Ecuador in his

enumeration of plants collected on the voyage of the Sulphur.

1844: Hasskarl published Neurocarpus, an orthographic variant of

Neurocarpum, and Neurocar us retusus.

1844: Paxton illustrated in color and described Clitoria fulgens

(Centrosema coccinea).

1845: Voigt transferred Clitoria erecta Roxb. to Neurocarpum. He

listed six species of Clitoria cultivated in Calcutta.

1845: Blanco reported one species, C. ternatea, from the

Philippines.

1845: Tenore described Clitoria tristis.








1847: Burnett illustrated and described a new variety, C.

ternatea var. major, based upon a cultivar raised from seeds sent from

New South Wales. He also illustrated and discussed Paxton's species.

1848: Hasskarl provided a detailed description of his species,

Neurocarpus retusus, published in 1844.

1348: Schomburgk published three new species and a new genus

based upon a manuscript of Klotzsch. Each was a nomen nudum. The

genus was Macrotrullion K. with two species, M. spendens K1. and

M. elegans K1. The third species was Neurocarpum speciosum Kl.

1850: Dalzell described Clitoria biflora, a new species, from

India.

1851: Miquel described Clitoria amoena, a new species from the

lower Marrowyne River.

1852: Bentham described two species in a footnote that were

published previously as a nomen nudum. They were Clitoria macrophylla

Wall. (published 1828) and Clitoria grahani Steud. (published in 1852;

synonym C. acuminata Wall. published in 1828.)

1853: Duchass and Walpers describe Neurocarpum argentum, a new

species from Panama.

1854: Beurling published Clitoria portobcllensis, a new species

from Panama.

1856: Miquel described a new species front Java, Clitoria javanica.

1858: Bentham (*) published his second revision of the genus

Clitoria which included Ternatea, Neurocarpum, Martia and Martiusia in

synonymy. The major change was the synonyjiizing of the latter three

genera with Clitoria. Bentham noted that the character of the legume

costa broke down. Another change was the riame Clitoriathus used for








section three instead of Bentham's earlier names Bractearia (1837) or

Dendrocyamus (1839). New species described were C. flagellaris,

C. rufescens, C. nana, C. selloi, C. leptostachya; C. hoffmanseggii

(superfluous name for C. arborea) and C. stipularis. The species

transferred from Neurocarpum included C. simplicifolia, C. guianensis

and C. densiflora.

Post-Linnaean (1859-1932): Floristic Period)

1862: Bentham in Martius and Eichler's "Flora Brasiliensis"

listed *fifteen species with descriptions, synoinymy, and a key to the

species. This was the first record of a key to Clitoria species and

used characters of: habit; leaflet shape, pubescence and number;

legume costa; peduncle length and number of flowers borne; and

bracteole shape and length relative to calyx or pedicel. New

binominals included C. yuyanensis (orthographic variant of C. guianensis)

and C. cajanlifolia (transfer of species from N(urocarpum). A new

variety described was C. glycinoides var. juraritiaca.

1864: Griesbach reported four species of Clitoria in his flora

of the British West Indies.

1864: Bentham described a new species in his "Flora Australiensis,"

Clitoria australis.

1866: Griesbach described a new species in his "Catalogus

Plantarum Cubensium," Clitoria glomerata.

1871: Oliver reported one species from tropical West Africal in

his flora.

1875: Micheli described a new species from Brazil, Clitoria

pedunculata. This name was a homonym for l[ojet ex Benth. (1837).

Micheli placed his species in Section Clil riarthes.








1878: Lowis provided some observations on the Indian C. ternatea.

1878: Hance reported on rare Chinese plants and listed Clitoria

macrophylla Wall. for a specimen in his herbarium collected by Sampson.

This was the source for C. macrophylla Hance cited by later botanists.

1878: In a second Philippine flora, Blanco continued to report

one species of Clitoria as he had in 1845.

1878: Vatke described Clitoria zanziharensis, a new species from

East Africa.

1879: Baker (*) reported five species of Clitoria in his flora of

British India. He made a revisionary change that elevated Ternatea

and Neurocarpum to the subgenus level, the first botanist to do so.

Section Clitorianthes was not reported, presumably because the species

within this group did not occur in India. A new variety was reported,

C. ternatea var. Oilosula (C. pilosula Wall.).

1882: Sagot reported five species of Clitoria in his French

Guiana flora. A new variety, C. javitensis var. gjabra, was described.

1887: Watson described two new species from Mexico. Clitoria

triflora was placed near C. quianensis in the section Neurocarpum.

Clitoria sericea was described from a fruiting specimen. Watson did not

place this species in any section.

1887: Hemsley described Clitoria Hanceana based upon a specimen

in Hance's herbarium collected by Sampson. He noted that this was

equivalent to C. macrophylla Hance (cf 1878), but not C. macrophylla

Wall.

1887: Sesse and Mocino described two new species from Mexico.

Clitoria racemosa (homonym of Don, 1832, and Bentham, 1837) was first

reported in synonymy by Don (1832) and probably by de Candolle (1825)








as a synonym of C. plumieri based upon a iinusL.ript of Mocino and Sesse

cited as "fl. Mex. ic. ined." The second species Clitoria uirginiana,

was probably an orthographic error (cf. 1 93) because Gronovius'

"Flora Virginiana" was cited (also typographic error with "U" for "V"

and corrected by Sesse and Mocino in the second edition of their

publication.

1889: Deflers reported one species from Yemen.

1893: Sesse and Mocino published their second edition of "Plantae

Novae Hispaniae" and cited C. virginiana. This corrected the

orthographic error of their first edition (cf. 1887).

1891: Kuntze (*) transferred twenty-nine species to the genus

Ternatea because Clitoria was a homonym of Clitorius Dillenius

(1732).

1894: Taubert in Engler and Prantl's "Die Naturlichen Pflanzen

Familien" adopted Bentham's treatment. They indicated about thirty

species in the genus were found in the tropical zone, but their species

were reported by sections and totaled to only twenty-four. They placed

ca. six species in Section Ternatea (Asia and Africa), ca. fifteen

species in Section Neurocarpum (Asia, Australia, and America), and ca.

three species in Section Clitorianthes (tropical America).

1894: Nairne reported one species of Clitoria in his Western

India flora.

1894: Robinson described the flowers of C. sericea (cf. Watson,

1887) and noted that the style was similar to Centrosema, yet had other

flower characters of Clitoria.

1894: Sesse and Mocino's "Flora Mexicana' followed their 1893

treatment of Mexican plants.








1894: Siebert and Voss reported Clitoria coelestis as a horti-

cultural synonym of C. ternatea. This new binomial is a nomen nudum.

1897: Britton and Brown reported one species of Clitoria in their

flora of the northern United States.

1897: Duss reported two species from Martinique that occur

natively plus one cultivated species. A new variety, C. glycinoides

var. ecostata, was a noiien nudum.

1897: Micheli described a new species from East Africa, Clitoria

tangani-censis.

1899: Rose reported nine species of Clitoria in Mexico with six

additional species as doubtful and excluded. Two new species were

Clitoria humilis and Clitoria subsessilis. Rose indicated that Clitoria

sericea may be a Cracca and that Clitoria grandiflora Mart. & Gal.

and Clitoria schiedeana suggested Centrosema.

1900: Bailey reported two Australian species of Clitoria in his

Queensland flora.

1901: Huber described a new species from Brazil, Clitoria cearensis

and noted its affinities with C. glycinoides, C. guyanensis and

C. cajanifolia.

1903: Merill reported vernacular names of C. ternatea in his

Philippine Dictionary.

1904: Chodat and Hassler described a new variety from Brazil,

Clitoria cajanifolia var. latifolia.

1907: Harms reported the first occurrence of cleistogamy in

Clitoria in a detailed article. Cleistogaious flowers in three species

were described and illustrated. They were C. glycinoides, C.

guianensis crnd C. cajanifolia. Harms discussed the history of Martia








physodes, a species described based upon a cleistogamous flowered

specimen from Brazil (cf. 1821).

1908: Mattei described a new species from northern Africa,

Clitoria albiflora. He noted its affinities to C. ternatea and C.

tanganicensis.

1909: Bailey listed the same two species of Clitoria in his

Queensland catalogue that he reported in his flora (1900).

1909: Gibbs reported one species of Clitoria in his Fiji flora.

1909: Herzog described a new species from Bolivia, Clitoria

nervosa. He noted its affinities to C. hoffmansegii and C. racemosa

Benth.

1909: Huber described a new species from Brazil, Clitoria

obidensis.

1910: Hassler described a new variety of Clitoria from Paraguay,

C. densiflora Benth. var. mucronata.

1911: Dunn added C. mariana to the list of species found in

China.

1911: Hayata reported one species of Clitoria from Formosa.

1912: Merrill reported one species of Clitoria from Manila in

his flora.

1912: Compton (R) described the seedling morphology and vascular

structure of C. ternatea. This was the first non-floristic, non-

nomenclatural, non-horticultural research undertaken on the genus

Clitoria.

1913: Harms described a new species of Clitoria from the Congo,

Clitoria kaessneri.








1914: Baevecke reported one species of Clitoria from the Eastern

United States in his flora.

1915: Gagnepain described a new species from Laos, Clitoria

linearis, and placed it near C. hanceana.

1916: LeCompte reported six species of Clitoria from Indo-China.

A key, descriptions, distributions, and citations were included.

Clitoria linearis was illustrated. A new variety was described,

C. hanceana Hemsl. var. laureola.

1917: Ewart reported one species of Clitoria in his Australian

flora on the Northern Territory.

1918: Britton reported one species of Clitoria from Bermuda in

his flora.

1918: Kirtikar and Basu reported the Indian vernacular names and

medicinal properties of C. ternatea.

1918: Pittier described and illustrated the keel and wings of a

new Venezuelan species, Clitoria dendrina.

1919: Brandegee described a new species from Baja California,

Clitoria monticola, and segregated it from C. mariana.

1920: Britton reported one species of Clitoria from the Bahamas

in his flora.

1920: Fawcett reported two species of Clitoria from Jamaica in

his flora. They were segregated by leaflet number and flower color.

1921: Harms described a new species from Brazil, Clitoria

brachycalyx and reported its affinities with C. brachystegia.

1922: Ducke described a new species from Brazil, Clitoria

snethlageae, and noted its affinities with C. leptostachya.








1922: Ridley reported two species of Clitoria from the Malayan

Peninsula in his flora and noted that C. cajanifolia was introduced

into Indonesia on Brazilian cattle which had the viscid seeds in their

hair.

1922: Standley reported six species of Clitorian trees and shrubs

from Mexico in his flora. A key and brief diagnosis were included.

1923: Merrill reported one species of Clitoria from the

Philippines in his flora.

1924: Britton reported three species from Puerto Rico and the

Virgin Islands in the survey conducted by the New York Academy of

Science. One species was reported under the genus Clitoria. Two

species from the Neurocarpum section were reported under the genus

Martiusia. Even through the original species N. ellipticum Desv. was

listed in synonymy, Britton did not use the older generic name

Neurocarpum. The two transferred species were Martiusia rubiginosa

(Juss.) Britton and Martiusia laurifolia (Poir.) Britton. This was the

first example of a botanist diverting from Bentham's treatment of 1858.

This trend of recognizing two genera, Clitoria and Martiusia, was

followed by a few American botanists.

1925: Fragoso and Ciferri (R) described several ascomycete

species from branches and legumes of "siccis Clitoriae triandrae"

collected in the Dominican Republic. This was the source for two

names, Clitoria triandra and Clitoria riandra.

1925: De Wild described a new species from British East Africa,

Clitoria merrnsi.

1926: Small described a new species from Florida, Clitoria

fragrans andl noted its affinities with C. mIaridna.








1928: Standley reported four species of Clitoria from the Panama

Canal Zone in his flora. A key and habitats were included.

1929: Baker reported four species of Clitoria in his treatment

of Tropical Africa legumes. A key and diagnosis were included. He

noted that C. albiflora and C. mearnsi were equivalent to C. ternatea.

A new variety that had been nom. in sched. was described as Clitoria

ternatea var. angustifolia Hochst. ex Bak. Baker published Clitoria

tanganyicensis Micheli as an orthographic correction for C.

tanganicensis licheli (1897).

1929: Hutchinson & Dalziel transferred C. racemosa Don to the

genus Vigna.

1929: Standley transferred C. sericea Wats. to the genus

Tephrosia.

1930: Barker and Darbeau reported four species of Clitoria from

Haiti which included C. plumieri Tuss., a nomen nudum.

1930: Ducke described a new species from Brazil, Clitoria

grandifolia, and noted its affinities to C. arborescens.

1930: Macbride described a new species from Peru, Clitoria

pozuzoensis. He noted that some of the characters that Bentham used to

distinguish the species of Section Clitorianthes broke down since

Macbride's new species keyed to C. javitensis from which it was totally

different.

1931: Gleason reported a species of Clitoria collected on the

Tyler-Duida expedition to Colombia.

1931: Malme described a new species from Brazil, Clitoria

chapadensis and noted its affinities with C. guyanensis. He also

described a new variety, Clitoria qlycinoides var. megapotamica.








1931: Sandwith reported his observations on the type of

nomenclatural problems of C. javitensis.

1931: Standley reported two species of Clitoria from the

Lancetilla Valley of Honduras.

1931: Williams reported four species of C1itoria from Trinidad

and Tobago. A key is included.

Post-Linnaeanr (1932-present: Research Period)

1932: Tschechow and Kartaschowa (R) reported the first chromosome

count for the qenus with 2n=16 for C. terrntea. The chromosomes were

illustrated in a camera lucida drawing. Relationships of C. ternatea

with other species in tne Phdseoleae tribe were discussed.

1932: Rylberg reported one species of Clitoria in his flora of

the Central North Ameri.an plains and prairies.

1933: Small reported oie species of :litoria and two species of

Martiusia it the Southestern United States. His floristic study

adopted Britton's treatment (1924). Small's species (1926) was

transferred as Martiusii frayrans Small.

1933: Holland and Joachin (R) reported their results on soil

erosion experiments with C. (ajanifolia in Ceylon (=Sri Lanka).

1935: Burkill reported the economic uses of two species of

Clitoria of thte Malayan Peninsula.

1936: Brown reported one species of :litoria in his flora of

Southeastern Polynesia.

1936: Standley reported two species of Clitoria from British

Honduras. Either a key nor descriptions were included.

1936: Stahl included three species or Clitoria from Puerto Rico.

1937: Standley reported four species of Clitoria from Costa Rica.








1937: Stehle reported on the colonizationi of some plants in

Martinique ond included Clitoria rubiginos.i Juss. var. ecostata (Urb.)

Stehle, a numen nudum.

1939: Amshoff reported eight species of Clitoria in Pulle's

Suriname flora. A key was included. Amshoff adopted Bentham's

classification (1858).

1940: Standley described a new species from Panama, Clitoria

velutina.

1941: Burkart described and illustrated j new species from

Argentina, Clitoria cordobensis. He discussed cleistogamous flowers

in the species and its affinities with C. Iariana, C. guyanensis,

C. multiflora ind C. rubiginosa.

1943: Kol, Machado and Meirelles (R) reported their results of

utilizing C ternatea ihy for sheep digestion.

1943: Ma,.bride reported eight species of Clitoria from Peru. A

key, descriptions, and citations were included. However, this treatment

was of poor quality because of the numerous misidentified specimens that

Macbride examined and the difficulty within the group Clitorianthes.

Macbride reported variability in several species, and along with his

additional notes, indicated his awareness of the problem. Some species

reported do not occur in Peru. Other species not reported do occur in

Peru. Peru has several endemics not recogiiized by Macbride.

1943: Mocoso reported four species of Cl itoria from the Dominican

Republic in his flora. Clitoria polyphylla was included in synonymy

with a Barbieria species.

1943: Pittier described a new specie, from Venezuela, Clitoria

glaberrima. He noted its affinities with C. arborescens and C. dendrina.








The description included "Pedicelli 4-5 cm' which was a typographical

error for "mm."

1944: Pittier published a second description of C. dendrina

(cf. 1918) ind illustrated the perianth.

1945: Hassell (R) reported his results of experiments to

establish C. ternatea in pastures of Queensland, Australia.

1946: Standley reported five species of Clitoria in his flora of

Guatemala. A key was included.

1946: Strang reported ihe ornamental properties of C. racemosa

Benth. in Brazil.

1948: Robyns reported one species ot Clitoria in his flora of

Albert National Park in Africa.

1948: Stehle and Quentin reported t,,j species of Clitoria from

Guadeloupe in their flora. They included rnbiinosa var. genuina

(illegitimate name for the typical variety) and C. rubiginosa var.

ecostata (Ub.) Stehle, a noiren nudum (cf. 1937).

1949: Burkart described and illustrated a new species from

Argentina, Clitoria epetiolata. He noted its affinities and compared

the species to C. guiarrensis. The variety C. epetiolata var.

latiuscula w as described.

1949: Chopra, Nath, Badhwar, and Ghosh reported the economic uses

and problems of C. ternatea in their "Indijenous Drugs of India."

1949: Henderson reported C. laurifolia as an established Malayan

wildflower introduced from Brazil early in the nineteenth century.

1949: Johnston reported C. portobellonsis from San Jose Island,

Gulf of Panama, in his flora and discussed its confusion with Clitoria

arborescens, the name the Panama plants were given by botanists.








1949: Merrill in his "Index Rafines(eue" reported that C. parviflora

Raf. was C. ternatea.

1950: Metcalfe and Chalke (R) included the genus Clitoria in their

descriptive anatomy of the Papilionaceae. Leaf, axis and wood

characteristics were included. The species studied were not reported.

1950: Velez and Overbech included C. terratea as a desirable

cultivar in their book.

1951: Krapovickas and Frapovickas (R) reported a chromosome count

of 2n=24 for C. cordoboensis and discussed its relationship to C.

ternatea.

1951: Leon reported three species of- Clitoria from Cuba in his

flora. Clitonria glomerita was included in synonymy with a Galactia

species.

1951: Quisumbing reported the medicinal properties of C. ternatea

in the Philippines.

1951: Rau (R) described endosperm development in C. ternatea.

1952: Andrews reported one species of Clitoria from the Sudan in

his flora.

1952: Burkart reported five species of Clitoria from Argentina.

A key was included.

1952: Lemee reported six species of Clitoria in his French Guiana

flora.

1952: Schery described a new species, Clitoria coriacea, from

Brazil.

1953: Frahm-Leliveld (R) reported a chroniosome count of 2n=16 for

C. ternatea.








1954: Berhaut reported two species or Clitoria from Senegal in

his flora. Within his key he published C. ternatea var. alba.

1954: Wilcez reported three species of Clitoria in his Belgium

Congo flora. A key and descriptions were included. He recognized

Baker's variety (1929) of C. ternatea and included C. tanganicensis

and C. mearnsi as synonyms.

1955: Sirdeshmukh (R) reported a double flower occurring within

C. ternatea.

1956: Bunting and Lea (R) reported their observations on the

introduction of three strains of C. ternatea in Sudan pastures.

1956: Fidalgo (R) reported his anatomical study of the Brazilian

tree ornamental, C. rac.emosa Benth. (=C. rairchildiana). This was the

first and only anatomictAl study of a Clitotia species, except for the

survey of MNtcilfe and Chalk (1950). TwerLy plates of illustrated

anatomical structures were presented.

1956: Gardner and Bennett included C. ternatea as a poisonous

plant in their book "Toxic Plants of Western Australia" and reported

the curariform properties of the seeds of U. atborescens.

1957: Cowan described a new species from Venezuela, Clitoria

cerifera, and noted its affinities to C. a:borescens. The species was

illustrated in his Figure 65.

1958: Chopra, Chopra, Honda, and Kapur included C. ternatea in

their book "Indigenous Drugs of India" and reported the species' use and

effects as: a cathartic, ; remedy for snake bites and scorpion sting,

an anti-dysentaric drug, a diuretic drug, j laxative, a drug for gonor,

and a skin jelly for eczema, prurigo and iipetigo.








1958: Santapau reported one species of Clitoria in his flora of

Purandhar, India.

1959: Rizzini described a new speciL, from Paraguay, Clitoria

burkartii.

1959: Tiwari and Gupta (R) reported the first chemical study of a

Clitoria species and reported a new chemical "dparajitin" extracted

from the leaves of C. ternatea.

1960: Sinha (R) identified a sterol from the yellow oil of the

seeds of C. ternatea. In a second article, Sinha (R) reported a

chemical and its properties extracted froii the leaves of C. mariana.

Although unnamed by Sinha, the author of the present study found the

properties LO agree with those of "aparajitin.

1961: Hundley reported two species uf Cl itoria from Burma and

their vernacular names.

1961: Sen (R) reported and illustrated binucleate pollen mother

cells in C. ternatea.

1961: Sen and Krishnan (R) reported their genetic experimental

results with the double flowers of C. territea and concluded that the

double flowers originated through a dominant gene mutation.

1961: Saroja (R) reported a chromosome count of n=8 for the

double flowered C. ternitea. He also reported his results on the

fertility of its pollen grains with those grains of the papilionaceous

flower type. Saroja concluded that the mutant gene was pleiotropic.

1962: Lind and Tallantire reported one Clitoria species from

Uganda.

1962: White reported one species of Clitoria from Northern

Rhodesia (-'ambia).








1963: Backer and Bakhuizen von Brink reported four species of

Clitoria from Java within a detailed descriptive key. They synonymized

C. javanica with C. rubiginoa.

1963: Maheshwari reported one species of Clitoria from Delhi.

1963. Rizzini reported twenty-three species of Clitoria from

Brazil. His treatment included keys, descriptions, morphological notes,

extensive citations, and illustrations of leaflets and some germination

diagrams, lie reported that for the few species studied, seed

germination type was an excellent sectional character. He found that

Sect. Ternatea and Sect. Clitorianthes was of the epigean type and that

Sect. Neurocarpumi was of the hypogean type. Newly described were

Clitoria rubiginosa f. longifolia, C. rubiginosa f. latifolia, C.

guyanensis i. chapadensis (Milme) Rizz., L. amizonum f. rotundifolia

and C. raceimosa f. obovatifolia. Despite 1 feu; nomenclatural errors,

Rizzini's floristic treatment was the best of any seen by the author of

the present study.

1965: Angley reported three species of Clitoria from Parana,

Brazil without naming the species.

1965: Gooding, Loveless,and Proctor reported one species of

Clitoria from Barbados.

1965: Liogier transferred two species from Martiusia in Britton

and Wilson's flora (1924) to Clitoria, which corresponded to Bentham's

treatment.

1966: Chavan reported two species of Clitoria from Gujarat, India.

1966: Hutchinson and Dalziel reported two species of Clitoria

from West Tropical Africa.








1966: Lourtig discussed the specimens within the herbarium of

Johan Burman and reported thit Flos Clito) ius flore coeruleo Burm was

Clitoria ternratea (cf. 1737).

1966: Padmanabhai (R) reported the tirst tissue culture studies

on a Clitorian species. His results on the excised embryonal axes of

C. ternatea were discussed.

1967: Boulos reported one species of Clitoria in his weed flora

of Aswan, Egypt.

1967: D'Arcy reported one species ot Clitoria from the Virgin

Islands.

1967: Howard reported his observatiios on the woody Clitoria

species and substituted the iame Clitoria iairchildiana for the homonym

C. racemosa Benth.

1967: Mullick, Pra:kash and Chatterji (R) reported their results

on seed germination of C. ternatea under v.irious ecological and

physiological conditions.

1967: Mullick and Chatterji reported their results of the effects

of sodium cyanide on seed germination of C. ternatea.

1967: Mitra and Datta reported a chromosome count of n=8 for

C. ternatea.

1968: Baum discussed binominals used by Linnaeus in his "Systema

Naturae, ed. 10" and reported C. lactescens was C. galactia and that

C. zoophthalmum was Dolicho> urens.

1968: Gupta and Lil (R) reported their isolation of a hexacosanol,

a sitosterol and an anthoxanthin glucoside froii the seeds of C. ternatea.

1968: Lakshmann and Padmanabhan (R) reported their results of the

antibiotic .scuchitine on th. growth of C. ternatea in vitro.








1968: Radford reported one species of Clitoria from the Carolinas.

1968: Uphof reported the economic iI~portance of three Clitoria

species in his dictionary.

1970: Datta and Sjha (R) reported tl, floral vasculature of

C. ternatea and concluded that it represented an advance stage in the

Phaseolae.

1970: Katiyar, Ranjhan and Shukla (R) reported their experimental

results on the yield and nutritive of C. tcrnatea for sheep.

1970: Oakes (R) reported his results on C. ternatea grown for

livestock consumption.

1970: Correll and Johnston reported one native species and one

cultivated species of Clitoria in Texas.

1970: Verdcourt reported one species of Clitoria from Tropical

East Africa and described a new variety C. rubiginosa var. glabrescens.

He proposed that the following species of the "rubiginosa-mariana"

complex be reduced to subspecific levels under the two species

C. mariana and C. rubiqinosa based upon stipule shape. His complex

included C. mariana L., C. grahami Steud. ex Benth., C. javanica Miq.,

C. australis Benth., C. frag'ans Small, C. mexicana Link and

C. rubiyinol d Pers.

1971: Cameron reported C. ternatea reacted favorably to a small

extent in becoming established where trifluralin was used to control the

black pigweed Trianthemna portulascastrum.

1971: D'Orey and iberato reported one species of Clitoria from

Portugese Guinea.

1971: Gillett, Pohill and Verdcourt reported three species of

Clitoria from Tropical fast Africa. Clitoria ternatea was concluded









to be a polymorphic species that included C. zanzibarensis, C. mearnsii,

C. tanganicensis and C. ternatea var. angustifolia, and no variety was

recognized. Refer to the African members Jf C. ternatea in this study.

1972: Adams reported two species of Clitoria from Jamaica.

1972: Liberato reported one species of Clitoria from S. Tome and

Principe.

1973: Aiyar, Narayanan, Seshadri and Vydeeswaran (R) reported

three glycosides of kaempferol extracted from the leaves of C. ternatea.

1974: Croat reported six species of Clitoria in Panama, with the

new combination C. guianensis var. subsessilis. Croat correctly

provided the first explanation of the plants known by the name C.

arborescens as represented by two species, a liana C. javitensis and

the tree, C. glaberrima.

1975: Fantz sent a manuscript on Clitoria for inclusion within

the Leguminosae in the Flora of Panama series, which was accepted and

approved by the editor. Six species were reported and described. They

represented the same six species identified by Croat, but with some

changes in the species names to correct nomenciatural errors to conform

with the Code. Croat's variety was not recognized as his segregating

characters broke down. The publication will appear when the entire

family has been treated.

1976: Fantz presented a preliminary paper comparing diagnostic

characters that segregate Clitoria from Centrosema. This was in

response to some individual suggestions and herbaria proposals that the

two genera should be recombined, a proposal he does not agree with as

the genera are distinct entities.















PRIOR RESEARCH ON CLITORIA


Most of the prior research on Clitoria that is non-floristic and

non-nomenclatural occurred after 1950. The amount of research on

Clitoria has been relatively small given the size of the genus. Most

of the work has involved the species Clitoria ternatea. This species

is commonly cultivated in gardens and often escapes, becoming

naturalized, giving it the largest distribution (pantropical and

subtropical) in the genus. Because of its easy accessibility,

C. ternatea was usually selected as the representative of the genus

Clitoria when research problems included several legume genera.


Anatomical Research


The few anatomical studies on Clitoria have resulted from surveys

including a number of genera. Only Fidalgo (1956) provided a detailed

anatomical study of a species within the genus.

Metcalfe and Chalk (1950) included Clitoria as one of the genera

in their descriptive anatomy of the Papilionaceae. No reference was

made as to which Clitoria species were used in their study. They often

described an anatomical characteristic followed by a list of genera

that included species with this characteristic. The following

anatomical characteristics were noted for Clitoria: Leaf: Non-glandular,

hooked hairs, with short basal cells, and a larger bent, terminal cell

(p. 505); lower surface papillose or subpappilose (p. 506); epidermis









often including a proportion of mucilaginous cells (p. 506); rod-shaped

crystals (styloids) in the palisade tissue (p. 511). Axis: Young stem

with cork arising in the middle of the cortex (p. 513); tanniniferous

cells, colored brown in dried material, and sometimes containing

proteins, mucilage, and other materials as well, often occurring in

groups situated in the pith and phloem or in the primary cortex

(p. 515). Wood: Parenchyma very abundant, in broad, moderately

regular bands 4 cells or more wide in some species (p. 521); rays

exclusively uniseriate or with only occasional biseriate rays in some

species (p. ;24); rays up to 1 mm in height in some species (p. 524);

sheath cells present in some species (p. 525); fibres forming small

islands on the cross-section owing to the abundance of wood and ray

parenchyma (p. 525).

Datta and Saha (1971) included Clitori a ternatea as one

representative member in their study of the floral vasculature of the

tribe Phaseoieae. Their findings include tiat the common bundles for

perianth, stamen and disc remained undividd.l for a short distance; that

the disc traces are feebler, branching immediately after origin in the

swellings at the bases of the stamens for a short distance; that the

common bundles for the perianth and stamens are elongated; that the

style is hollow having one dorsal and two marginal traces, with the

upper part solid having a single trace; that the vascular trace

continues almost up to the stigma tip. A camera lucida drawing of a

transaction of the floral axis is illustrated in Figure 1 and a graphic

diagram of the floral vasculature is illustrated in Figure 17.

Fidalgo (1956) did ,in anatomical study of Clitoria fairchildiana

(published under the synonym (litoria racemiisa), a tree species









cultivated in parts of Brazil as an ornamental, and infrequently

cultivated elsewhere. He provided 43 page, of his techniques and

descriptive anatomical results on the roots, steis, leaves, flowers,

fruits, and seeds. He included 20 plates (52 figures) of drawings that

illustrate the anatomical structures. He includedd that there is a

great occurrence of calcium oxalate, monoclinal, principally in the

cortical parenchyma; small particles of starch were found in woody

parenchymous cells, in great abundance in the pith; tannins occur

frequently, not from internal differentiation of cell, but from

secretary pockets or canals; mucilaginous canals commonly not in

parenchymatous pith rays; external epidermis provided with uniseriate

protective hairs (not peduncle) with 1 or 2 basal cells, partially or

totally caducous with developing organ; secretary hairs occur in hidden

parts of peduncle, with a single basal cell and a terminal swelling;

development of 20 structures of stem emerges between layers of apical

collenchyma, strata of suberized cells occupy various positions;

petioles invaginated from bark by activity of cambium and isolated

woody elements forming a secondary internal woody ring of bark;

pulvinus and rachis express an interesting evolution of cortical weave

which permits movement of the leaves; leaves present numerous

adaptations to xerophytic conditions, with great cavities in the lower

epidermis, obliterated by long protective hairs over large, scanty

stomata; fruit with two patterns of sclerenchymous layers orientated

in different directions; seed coat formed of numerous weave patterns

in the external epidermis, not containing toxic proteins, presenting

great advantages for their use through cattle alimentary tracts.









Chemical Research


Chemical research has been conducted by Indian investigators on

the seeds and leaves of Clitoria species that occur natively in India

and are reported by the Indians to be of medicinal value.

Tiwari and Gupta (1959) reported a new chemical they named

"aparajitin" that was obtained from an alcoholic extract of the dried

leaves of C. ternatea. The chemical name was derived from the plant's

Indian vernacular name of "aparajita." Based on experimental results,

aparajitin (C26H5002) was determined to be a o-lactone of 2-methyl-4-

hydroxy-n-pentacosanoic acid with the chemical structure:

CH

CH3(CH2) 18-CH2CCHCH2 CH2HCO

0----

The properties of aparajitin were described as insoluble in water,

cold alcohol, chloroform, ether, and petroleum ether; sparingly soluble

in acetone; soluble in hot alcohol, benzene, ethyl acetate and

pyridine; did not decolorize bromide water or alkaline potassium

permangate solution and gave no acetyl derivative; did not reduce

Fehling's solution nor respond to any keto group test; laevo-rotatory

with M.W. of 391 and m.p. of 92-93'; treatment with hydroiodic acid

and hydrobromic acid produced 2-methyl-4-iodo-n-pentacosanoic acid

(C26H51021) and methyl-4-bromo-pentacosanoic acid (C26H5102Br)

respectively; esterification produced methyl acid, 2-methyl-4-hydroxy-

n-pentacosanoic acid, which formed the acetyl derivative of 2-methyl-

4-hydroxymethyl-n-pentacosanoate; oxidation produced arachidic acid

and B-methylglutanic acid.









Sinha (1960) identified the yellow fixed oil (yield 18.78%)

obtained from the seeds of C. ternatea as a y-sitosterol based upon

the preparation of acetate and benzoic derivatives and comparing their

comparative compositions, melting points and rotations with those of

known sitosterols. The sterol's properties were described as:

obtained as colorless shiny plates; molecular formula of C20H500;

negative test for N,P,S, and halogens; positive Salkowski reaction

and digitonin test; coloration assumed purple to blue, then to green

in Liebermann Burchard reaction; purple coloration in Steinkle

Kehlenberg reaction, which turned cobalt blue on exposure to light.

Sinha (1960) described a o-lactone compound from alcoholic extract

of the dried leaves of Clitoria mariana with a molecular formula of

C26H5002. Front the nearly identical properties described and

treatments of the new extract, the chemical is "aparajitin," although

Sinha has no reference to the work of Tiwari and Gupta.

Gupta and Lal (1968) reported the isolation of hexacosanol,

B-sitosterol and an anthoxanthin glucoside from the seeds of

C. ternatea. On acid hydrolysis, the anthoxanthin glucoside yielded

quercetin and glucose. Chromatography of the amino acid composition

enabled the identification of the essential amino acids lysine, valine,

methionine, phenylalanine and isoleucine in the protein hydrolysate,

along with aspartic acid, serine, glycine, alanine, glutamic acid,

tryosine, proline, argisine and histidine.

Kulshrestha, Kumar and Khare (1968) detected cersulfate positive

compounds by thin layer chromatography in the butanol-soluble fraction

of the seeds of C. ternatea. Four of the six crystalline compounds

were identified as adenosine, kaempferol-3-rhamnoglucoside,









p-hydroxy-cinnamic acid and ethyl-alpha-D-galactopyranoside. The

remaining two compounds were a polypeptide and a phenolglycoside.

Aiyar, Narayanan, Seshadri, and Vydeeswaran (1973) reported three

glycosides of kaempferol from the leaves of C. ternatea. The ethyl

acetate soluble part of the ethanol extract gave three compounds on

chromatography identified as follows: Compound A (m.p. 198-200) as

kaempferol-3-monoglucoside; Compound B (m.p. 215-70) as kaempferol-3-

0-rhamnosyl (1-6) glucoside; Compound C (m.p. 188-90) as kaempferol-

3-rhamnosyl (1-6) galactoside. Light petroleum ether and ether

extracts on chromatography yielded waxy matter, chlorophyll and

B-sitosterol.


Cytological Research


Very little research has been done on the chromosomes of Clitoria.

From the few counts reported, the species appear to fall into two

distinct groups which correlate with the morphological characters which

indicate two subgeneric lines for the species involved. The subgenus

Clitoria has x=8, and is based on one species. The subgenus

Neurocarpum has x=12, and is based on four species. The chromosome

counts reported for Clitoria species are tabulated in Table 2.

Tschechow and Kartaschowa (1932) included C. ternatea as one of

the species in their cytological survey of the tribe Phaseoleae. They

provided the first chromosome count for the genus (2n=16). A camera

lucida drawing of the chromosomes was provided. They remarked that

C. ternatea differs uniquely from the other species in the tribe both

by the number of x=8 (versus x=ll, 12, 19, 20 in other species, the

latter pair of counts pertaining to other species in the same subtribe)














Table 2. Reported chromosome numbers in Clitoria species.



Subgenus CLITORIA


Species

ternatea

ternatea

ternatea

ternatea

ternatea

ternatea

ternatea (double)


n 2n date

16 1932

8 1938

16 1953

16 1957

16 1962

8 1967

8 1961


reporter

Tschechow & Kartaschowa

Senn

Frahm-Leliveld

Frahm-Leliveld

Shibata

Mitra & Datta

Saroja


Subgenus NEUROCARPUM


Species n ?n date

cordobensis 24 1951

falcata* 24 1960

laurifolia 24 1957

macrophylla 12 1971


*Chromosome number reported under the


reporter

Krapovickas & Krapovickas

Frahm-Leliveld

Frahm-Leliveld

Larsen


ynonym C. rubiginosa








and by the two distinct chromosome sizes, four long chromosomes and

four short ones. Even among the two groups, these eight chromosomes

are not equal in size, although there appears to be a 3:1 size

relationship between the long and short chromosomes.

Krapovickas and Krapovickas (1951) provided a count (2n=24) and

an illustration of the chromosomes of Clitoria cordobensis. They

concluded, after comparison to C. ternatea, that there are either two

distinct groups of species with x=8 and x=12 respectively, or that

C. cordobensis was a triploid.

Sen and Krishnan (1961b) reported that binucleate pollen mother

cells occur at a very low frequency in C. ternatea. One of the two

nuclei was always smaller. Meiosis was not observed in these cells

beyond early diakinesis. An illustration of a binucleated cell with

both nuclei at pachytene was presented.


Developmental Research


Developmental research has been undertaken almost entirely upon

C. ternatea, primarily by Indian researchers. Studies have been

conducted on endosperm, embryo growth, seed germination, and seedling

growth.

Rau (1951) described the endosperm development in C. ternatea

along with four other selected papilionaceous species. The endosperm

is of the nuclear type, with the free endosperm nuclei arranged along

the sides of the inner layer of the inner integument. As the seed

cavity at the chalazal end widens in later stages, the massive nucellus

projects into it. The antipodal end of the embryo sac is wedged into

this nucellar mass. Free endosperm nuclei lying in dense cytoplasm








gather here. Cytokinesis of endosperm nuclei occurs when the embryo is

in an advanced stage of development. The cell formation is restricted

to just one or two layers around the embryo; the remaining part retains

free nuclei.

Padmanabhan (1966) reported growing excised embryonal axes in

vitro on Nitsch's basal medium supplemented with vitamins. He

concluded that the course of axis growth was remarkably similar to

natural germination events, namely; (1) growth of root (day 2-3),

(2) elongation of hypocotyl (day 4), and (3) activation of shoot bud

(after day 20). Elongation was not affected by removal of the apical

bud. From these results the observation that upper hypocotyl curves,

he concluded that the embryonal organs are capable of independent

development when isolated, but follow the same developmental pattern

as in the entire embryo.

Lakshmann and Padmanabhan (1968) reported the effect of the

antibiotic ascochitine on in vitro growth of Clitoria ternatea. They

concluded that, in general, the toxin exerted its effects on the

embryos grown in vitro at much lower concentrations than is required

to cause perceptible effects on the germinating seeds. Ascochitine

produced total inhibition of root growth, presumably because root

proteins were easily denatured. A low percentage of embryos did

produce callusing of the root apex and the hypocotyl. The shoot

meristem was not affected by the toxin, presumably because the shoot

meristem converts a large quantity of the toxin to dihydroascochitine,

rendering this toxin benign.

Mullick and Chatterji (1967a) reported on seed germination

of C. ternatea. Imbibition and germination behavior was








studied under various eco-physiological conditions in the laboratory,

conditions similar to those found in the wild. They concluded: light

had no effect on either behavior; seeds germinated mostly at a

temperature of 35' 20C; better imbibition and germination behavior

were obtained when seeds were subjected to physical and chemical

treatments of impaction with sand, impaction without sand, hot water,

concentrated sulphuric acid, absolute alcohol and normal potassium

hydroxide. Seed imbibition and germination behavior were about fifteen

per cent under laboratory conditions of light and temperature.

Mullick and Chatterji (1967b) reported the effects of sodium

cyanide on seed germination of C. ternatea. They concluded that seeds

soaked for 24 hours in 100 ppm of aqueous sodium cyanide solution

accelerated imbibition, germination, and promoted early growth of the

seedlings. Increased concentrations produced inhibitory effects

leading to the suppression of normal seed germination and growth.

Light and temperature did not influence the chemical effect on the

seeds.

A. P. de Candolle (1825) described seedling development based upon

three species of Clitoria and illustrated various stages in Plate IX

(Figures 33-35). Two of these species are now placed in the genus

Centrosema. The illustrations show plants with 3-foliate leaves, but

the leaflet shape tends to indicate de Candolle's Figure 33 as

C. ternatea by elimination. Clitoria ternatea typically is 5- and

7-foliate; however, Rizzini's study (1963) clarifies the apparent

discrepancy in leaflet number. With the data representing a mixture

of two genera, de Candolle's treatment becomes a non-reliable source

of seedling development information in Clitoria.




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