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Copyright 2005, Board of Trustees, University
Fort Pierce ARC Research Report RL-1974-8 m
DISTRIBUTION, INTRODUCTION AND EVALUATION OF TROPICAL PASTURE SPECIES
1/ OV 8 2 977
Albert E. Kretschmer, Jr.-
Introduction 'A.S. -
..." v. of Florin
The introduction of forage plants is probably as old as cultivated agri-
culture. Man has long recognized the advantages of introducing new species
and genera to improve the agriculture of a local area, but introductions of
species for human consumption received more emphasis than did pasture species
until recently. Until the advent of plant breeding knowledge, genetic improve-
ment came solely from introductions or selections. Plant introduction research
workers and plant breeders of pasture crops use many similar techniques in the
evaluation of new material even though sources of the material are different.
As stated by Williams (280) "Plant introduction aims to remove the genetic
limits to production by providing plants and specific characteristics wither
for direct use or for use in breeding programs." Plant introduction workers
and plant breeders have the same ultimate goal the improvement of an exist-
ing agronomic situation with the introduction work considered as the gross
effort to find new material to include in the more refined and specific
In order to excel at the work of pasture plant introduction, a basic
knowledge is necessary of many agricultural aspects. Knowledge of the areas
of emphasis that will help in performing a better job of plant introduction are:
1. climatic, latitude and altitude effects on plant growth
2. grass and legume anatomy and classification
3. grass and legume agronomic characteristics including growth
habit, general flowering and seeding habits, root type, etc.
4. for legumes, a good knowledge of Rhizobium requirements
5. knowledge of the flora of various countries and of larger areas
with homoclimates (areas of similar agricultural potential with
respect to amounts and distribution of rainfall and temperature
6. basic understanding of plant breeding
7. competitive effects of legumes and grasses for fertilizer,
water and sunlight
8. soil fertility and lime concepts as they affect pasture plant
growth; and in the case of legumes,how they affect the Rhizobium-
9. continuous effort in reading the latest world-wide research
reports, especially those involving the stated objective of the
10. implication of plant effects on livestock such as nutrient contents,
digestibility and intake, and acceptability
11. toxic or sub-toxic compounds in pasture plants that may affect
The author believes that a researcher working in the field of pasture or
forage crop introduction in the underdeveloped areas of the tropics and sub-
tropics has the potential to increase pasture productivity in a localized area
1/ Professor of Agronomy, Institute of Food and Agricultural Sciences,
University of Florida, Agricultural Research Center, P.O. Box 248,
Fort Pierce, Florida.
to a much greater extent than the plant breeder at this particular time in
history. Although a debatable point, the opinion is based on the fact that
pasture improvement in the tropics and sub-tropics is relatively recent. An
excellent review of the poor results of the ryegrass (Lolium perenne) breeding
program even in the temperate zone was done by Raymond (194) in 1969. Further-
more, little is known of the vast grass and legume germ plasm resources in the
tropics. These factors and the recent successes in the experimental and com-
mercial use of new tropical pasture plants in tropical and sub-tropical areas
of the world has far surpassed the use of bred tropical grasses and legumes.
A partial list of areas in the tropical and sub-tropical world where use is
being made of one or more new grass or legume introductionsis shown in Table 1.
Most of the newer legumes come from tropical areas while a few sub-tropical
and temperate Trifolium types (241) have been tried under tropical conditions.
There have been very few instances where plant breeding has led to as
valuable an addition to the tropical and sub-tropical pasture complex as has
plant introduction. One important exception has been the development of the
perennial summer-growing legume, Siratro (Macroptilium atropurpureum) by
Hutton (118) of Australia in 1962. There has been world-wide tropical and
sub-tropical testing and commercial acceptance of this cultivar. However,
there is recent evidence that another type may be more resistant to rust
(Uromyces phaseoli) and web blight (222). Coastal bermudagrass (Cynodon
dactylon), bred and released by Burton (86, 111) in 1943 is an example of the
value of plant breeding in increasing the productivity of permanent pastures
in the lower temperate and sub-tropical areas of the United States.
The ability of the scientist to evaluate the specific need for a new
pasture plant is absolutely necessary (194). There is less reason, for
example, to obtain another high yielding summer-growing grass for an area
that already has 3 or 4 excellent ones, when the limiting factor for year-
round animal production is the lack of feed production during the winter or
dry season. Under these conditions the plant introduction emphasis should be
on finding a more successful winter-growing or dry season-growing grass or
The object of this report is to review various subjects related to
pasture and forage plant introduction. Emphasis is placed on the tropical
and sub-tropical climatic areas and on grasses and legumes for animal rather
than human consumption.
Generally, the tropics are considered to be the areas encompassed by the
tropics of Capricorn and Cancer (230 27' south and north). In this report
the sub-tropical areas refer to a rather ill-defined zone bordering the
tropical zones to the north and south to latitudes of about 30 degrees. This
subject was discussed more thoroughly by Davies (57). Also, there are moun-
tainous areas within the tropics that can be included in the sub-tropical
classification because of lower temperatures.
Table I. Partial list of tropical and sub-tropical areas where new grass
and legume introductions are being used experimentally or
7. Costa Rica
8. Fiji Islands
9. Florida (U.S.)
17. New Guinea
23. Ryukyu Islands
27. Virgin Islands
28. West Africa
7, 11, 23, 54, 69, 80, 149, 241, 259, 262
26, 123, 148, 217
51, 295, 296
101, 136, 137, 141, 142, 143, 144, 157, 189, 268, 281
Centers of Distribution of Grasses and Legumes
Grasses-- Hartley and Williams (90), Hartley (89), Hitchcock (99, 100),
Whitney, Hosaka and Ripperton (275), Whyte (277), Burbidge (27), Booth (16)
and other authors (32, 49, 257) have described various aspects of the origin,
history and development, distribution, and classification of the grasses.
It is believed that grasses emerged as a distinct class of the angiosperm
complex during the late Cretaceous era.
It is believed that the Bovidae (ancestors of our modern cattle) and the
Gramineae could have originated during the same evolutionary era in Eurasia.
This would explain the abundance of excellent pasture species in this largest
center of grass distribution the grasses perhaps having adapted to conditions
of grazing. However, there is no well-founded reason to explain the South
American center of distribution.
The family name Gramineae was first established by Jussieu (27) in 1789.
The International Code of Botanical Numenclature in 1956 decided that Poaceae
should be the preferred family name although the traditional name Gramineae is
permitted and more frequently used (303).
The Gramineae is one of the largest families of plants, containing about
620 genera and 10,000 species (in 50 to 60 tribes)(303). These constitute
about 4.8 and 3.3%, respectively, of the total of the flowering plant com-
munity. Recently (16), in-1964, a tentative new system of classification said
to be more satisfactory for North American grasses listed 6 sub-families and
28 tribes. Hitchcock (100) lists 2 sub-families and 14 tribes, and 168 genera
for the grasses of the United States. There does not appear to be complete
unanimity in sub-family or tribal classification, but the common genera, i.e.,
Panicum, Digitaria, Pennisetum, Paspalum, etc. are better agreed upon by
botanists and agronomists.
Grasses occur on all continents and are exceptionally adaptable to
diverse ecological conditions. Except for Bambuseae (bamboo), all grasses
are herbaceous with relatively little variation in growth form or morphology
compared with other families. In spite of the large number of species of
grasses, it was estimated in 1956 (90) that less than 50 species accounted
for 99% of the planted pasture types. Twenty-five were native in the temperate
region and only 9 in the tropical areas. At present there are more tropical
species in use, although the number of temperate species used is still con-
siderably greater. Consider the potential for finding new types.
The grassland types of plant communities are widely distributed through-
out the world. They occur in the interior of the great continental land
masses, ranging to about 550 N in Asia and North America and southward to 400
in South America. Temperate grasslands can be classified as treeless prairies,
steppes and pampas while in the sub-tropics and tropics, savannah type veg-
etation, with various proportions of trees and shrubs, predominate. Seldom
do the number of grass species in a grassland area comprise a large percentage
of the total flora in these areas. Usually a small number of grass species
predominate the ground cover while a much larger number of non-grass species
provide only a small percentage of the ground cover.
A few grass tribes contain many genera but there are many small tribes
containing only one to several genera. Below are listed three of the larger
tribes, and one smaller one and their general distribution:
1. Andropogoneae predominately sub-tropical and tropical:
found in southeast Asia, India and to lesser extent in Central
Africa in monsoonal type climate: probably spread to the new
world (from South America to southern United States) later in
2. Paniceae largest tribe, found where there are high winter
temperatures and high annual, well-distributed rainfall.
3. Eragrostideae about 50 genera found in arid regions with high
winter temperatures and summer or non-seasonal rainfall.
4. Oryzeae seven genera, tropical and warm temperate regions,
The distribution of the first three tribes listed above appear to have
developed as a result of climatic factors while Oryzeae developed as a result
of specific soil characteristics with climate playing a minor role.
Among the smaller, less widely distributed tribes, Hartley (27) states
that their distribution may reflect historical patterns more than climate
and soil variations since they may not have reached all areas of the world
where the climate is suitable for development. It is likely that there are
many species within these smaller tribes that could be successfully utilized
for pastures in areas of homoclimates.
Although the distribution of genera is difficult to assess, such a genus
as Poa is known to thrive in areas of high latitudes or high altitudes. It
is absent or rare in the tropics except in mountainous areas. On the other
hand, moisture-loving species such as Glyceria fluitans and Phalaris arundin-
aceae are widely distributed. Man and animal have helped in the distribution
of species of Bromus, Cenchrus, Cynodon, Digitaria, Echinochloa, and Paspalum
among others. It is interesting to note that in several instances (including
Phalaris tuberosa), introduced and successful cultivated pasture grasses have
not been cultivated in their place of origin.
The greatest concentration (Eurasian) of cultivated grasses came from the
north central Mideterranean region where more than half originated (90). All
of the remaining came from a zone including all but northern Europe and parts
of Africa and Turkey. In eastern Canada and northeastern United States, 6 or
7 of the grasses may be indigenous even though their centers of varietal poly-
morphism are probably in Eurasia (examples are Agrostis, Festuca, Phalaris,
A smaller concentration of the grasses are found in east Africa (Kenya and
Uganda) and possibly extend into southern Rhodesia. Examples are buffelgrass
(Cenchrus ciliaris), paragrass (Brachdaria mutica), molassesgrass (Melinus
minutiflora), guineagrass (Panicum maximum), Rhodesgrass (Chloris gayana),
kikuyugrass (Pennisetum clandestinum), elephantgrass (P. purpureum), and
Bermudagrass (Cynodon dactylon). Except for buffelgrass and Bermudagrass
none are indigenous in the Eurasian region.
There are minor concentrations in South America represented by carpet-
grass (Axonopus affinis), rescuegrass (Bromus catharticus), dallisgrass
(Paspalum dilatatum), bahiagrass (P. notatu), and Sorghum almum.
In 1956 it was noted (90) that only three grasses, those indigenous to
the western United States (rescuegrass, Bromus marginatus), the Sudan
(sudangrass, Sorghum sudanense) and the Transvala in south Africa (pangola,
Digitaria decumbens), were not indigenous to one or more of the three main
regions discussed above.
With the exception of rescuegrass, no grass indigenous to the large land
mass of the United States is used to any great extent elsewhere in the world.
More recently, a cultivar or cultivars of alemangrass ("Pasto Aleman")
(Echinochloa polystachya HBK. Hitchc.), which is indigenous or at least
naturalized in the southern United States (100, p. 711), but not cultivated,
is being successfully used in high rainfall areas of central and tropical
south America; and is thought by many to be superior to paragrass. In the
great grassland areas of the world few, if any of the dominant species are
of importance as cultivated grasses. On the other hand, paragrass and pangola,
which come from very restricted areas of natural distribution, have been
used commercially world-wide in the tropics and sub-tropics where local
climate and soil conditions are suitable. Undoubtedly there are many other
localized species that would prove to be of great economic importance to
pasture development and improvement if introduced to areas where specific
underproduction periods occur because of the presently used species.
If one is to obtain grass introductions for use in a particular area,
knowledge of the grass flora and descriptions of grasses of similar homo-
climates would appear to be of extreme usefulness in selecting species or
genera that may be adaptable. This assumption may not always be true
because of other environmental factors. In any event, it would be pre-
sumptuous to assume that the use of world maps, and a complete knowledge
of the flora alone would permit an individual to introduce an untested grass
or legume from some isolated area with a similar homoclimate and have a
better species than those that are locally being used. As a matter of fact,
Semple (209, p. 152) in 1970 pointed out that introduced grasses can be
successful in climatic conditions that are widely different from those in
which they originated.
There are several excellent published accounts on the flora and descriptive
accounts of grasses in the literature. Work with United States' grasses by
Hitchcock (100), grasses of Central America (98), and grasses in the West
Indies by the same author (99) cover temperate, sub-tropical, and tropical
grasses of the Western Hemisphere. Work by Martinez-Crovetto (158) in Argen-
tina, Van Donselaar (255) in Surinam and French Guiana, Lucas de Febres (151)
in Venezuala and Leithead and Yarlatt and Shiffet (146) in the southern United
States give additional information and references to native species in the sub-
tropical and tropical areas of the Western Hemisphere. These works sometimes
indicate when a grass has been introduced and if it is naturalized as of the
date of publication. For example, guineagrass was considered naturalized in
the West Indies by 1930.
Bolivian native grasses were reported by Braun (19) and classified
according to areas of occurrence; between the jungle and pampas, natural
pastures, and grasses found in low areas or areas where periodic flooding
occurs. Information on some of the introduced, cultivated grasses is also
reported for tropical areas in general by Hubbell (113), Canoniero (43) and
The grasses in Haiti as reported by Barker and Dardeau (10) include 11
tribes with 22 genera found in the Paniceae, including 7 species of Setaria,
4 of Pennisetum, 5 of Cenchrus, 6 of Digitaria, 1 Erichloa, 1 Axonopus, 32 of
Paspalum,3 of Echinochloa (not including E. polystachya) and 33 of Panicum.
Some of these were naturalized. Whether a grass is native or naturalized
should have little effect on its possible use as an introduced type, since a
species that has become naturalized may have different agronomic character-
istics from those of the original introduction. It would be expected, for
example, that various strains or cultivars of Rhodesgrass could be found in
an area where it had been introduced 100 or more years ago because of its
cross-pollination characteristics (15).
Probably the most comprehensive work on grasses of South Africa is that
of Chippindall (46) who described 173 genera in 25 tribes. Gupta and Nanda
(84) described the natural grassland types in the western Himalayas based on
altitude, into alpine, temperate, sub-tropical and tropical types. Kernick
(132), for near east and north Africa, discussed the ecology of rangelands
where climatic differences are pronounced. In addition, Hepper (93, 94) for
the northern Nigerian highland, Tuley (247) for eastern Nigeria, Pratt,
Greenway and Gwynne (187) for east Africa, Sacco (204) for the northern, arid
zone of Somaliland with its xerophyllic climate, and Verboom (263) for Zambia,
described native grasses in smaller areas in Africa. Descriptions of the grass
flora of west tropical Africa by Hutchinson and Dalziel (299) and east tropical
Africa by Clayton (303) and by Clayton, Phillips and Renvoize (304) are very
In Table 2 are listed selected references dealing with native or cultivated
grass flora not listed in Table 1.
Table 2. Selected list of references
dealing with native or cultivated
Country or Area References
1. Worldwide 172, 277, 209
2. Latin America 113
3. Canary Islands 232, 233, 234
4. India 53
5. Australia 65
6. United States 257
7. Philippines 192
8. Puerto Rico 266
There are numerous descriptive reports dealing with specific grass
species or genera that are invaluable for plant introduction researchers.
Besides describing the grass, many times valuable references are included
regarding climatic and soil adaptation, history, ability to combine with
legumes and nutritive value. In Table 3 are a selected list of references
dealing with specific grasses.
The problem of increasing the quality of a temperate species has been
discussed by Raymond (194). Increasing the quality of an existing tropical
grass pasture by replacing it with another grass may be more difficult.
Probably the overwhelming factor involved here is the dry matter (or net
energy) intake by the animals rather than the digestibility; and intake, un-
fortunately, is expensive and difficult to assess. However, introduction of
grasses can result in markedly increased yearly animal production if the
grasses "fill-in" the feed shortage periods. The greatest impact on a yearly
grazing management system will occur where the cultivated species which dies
or grows very slowly during a period of stress (i.e. cool or dry seasons) is
replaced or supplemented by a grass that grows and remains green during this
period. The quality of the new grass in these instances is relatively un-
important since the choice essentially is between no grass and green grass.
Cattlemen understand this point very well and for this reason many cattlemen
in some of the wet-dry American tropical areas are using African star bermuda-
grass ("Estrell& Africana") (Cynodon plectostachyus probably incorrect
scientific name (86) for C. nlemfuensis Vanderyst or C. aethiopicus Clayton et
Harlan). African star bermuda which remains green four or more months without
rainfall has begun to replace jaraguagrass (Hyparrehenia rufa), which is not
drought tolerant. It is doubtful that there is much difference in digestibility
or intake between the two at equal stages of growth during the growing season;
but there would be a considerable difference between intakes during the dry
season when comparing dead stubble or lack of grass (jaragua) with green,
growing grass (Bermuda).
Table 3. Selected list of references dealing with specific grass
genera or species.
Grass Name Reference
1. Pennisetum clandestinum (Kikuyu) 105, 129, 160, 282
2. Cenchrus ciliaris (Buffel) 106, 116
3. Panicum maximum var. trichoglume (Green panic) 115
4. Setaria sphacelata (Setaria) 85
5. Panicum maximum (Guinea) 81
6. Panicum coloratum var. makarikariense (Makarikari) 149
7. Chloris gayana (Rhodes) 15, 42
8. Pennisetum purpureum (Elephant) 80
9. Paspalum sp. 102, 109, 240
10. Cynodon sp. (Bermuda) 30, 59, 60. 86, 87,
88, 111, 198
11. Digitaria decumbens (Pangola) 101, 107, 173, 178,
238, 267, 270
JP. Leersia sp. 190
13. Hemarthria altissima (Limpo) 281
14. Brachiaria mutica (Para) 126
15. Melinis minutiflora (Molasses) 108
Legumes-- Two excellent reviews on the origin and history of legume development
from an ecological viewpoint were written by Andrews (3) in 1914 and by Tutin
(2'"4) in 1958. Other historical aspects of legumes were reported by Burkart
(30) in Argentina, Whyte (278), Semple (209) and in the United States (257).
The family Leguminosae is one of the largest of flowering plants, with an
estimated 700 genera and 14,000 species. It is believed that legumes had their
beginnings in upper Cretaceous times with development beginning from large trees
growing under wet tropical conditions. Later development resulted in the appear-
ance of shrubs and woody climbers; and more recently, emergence of perennial
and then annual herbs. From a climatic view the development began under wet
tropical conditions then in the dry tropics, and later in the sub-tropics and
temperate climatic zones. Only about 200 genera and 4,000 species are con-
sidered to be temperate legumes. Of these probably less than 40 genera and
200 species have been used in commercial agriculture. Strong evidence is
provided that the subfamilies, Mimosoideae and Caesalpinioideae of the family
Leguminosae contain species that are older than many of those in the subfamily
Papilionoideae. This is interesting since our more frequently used legumes in
temperate and sub-tropical areas of the world have been of more recent eco-
logical development, coming from predominately temperate tribes of the Papilion-
oideae. Examples are species of Trifolium, Medicago and Lotus. This sub-
family is characteristic of cooler rather than tropical regions, and a few
rare species extend both northward and southward to the limits of dicotyledenous
growth in the very cold regions. It is also of interest that with the ex-
ception of two or three, the commercial tropical legumes are from tropical
tribes of the subfamily Papilionoideae. The tropical tribes of Papilionoideae
are believed to be ecologically older than the temperate tribes.
When working with legumes it is necessary to have an understanding of the
symbiosis between host plant and Rhizobium. An excellent historical review of
Rhizobia was published by Norris (174) in 1956, and information concerning
Rhizobia and sub-tropical legumes was reported in 1964 (175). Abstracts
of papers concerning nitrogen fixation from 1948-1967 were published by
Brockwell (20). No attempt will be made to discuss this aspect of legumes
Probably the main advantage that legumes have compared to grasses is
their ability to fix nitrogen. Although up to 500 kilograms of nitrogen per
hectare per year (56) can be fixed by legumes, a more realistic value would
be between 100 and 300 kilos under grazing (56, 142). A second and possibly
equally important advantage of legumes is that their quality does not decrease
with time as rapidly as that of grasses, especially tropical grasses. Further-
more, animal intake of most tropical legumes is higher than that of maturing
tropical grasses with the same digestibilities (163).
It is doubtful that pasture legumes could or should be used under all
conditions. Their use in the southern United States has decreased recently
mainly because of the increased use of nitrogen fertilizer under intensive
pasture production and because of the alfalfa weevil that has caused a large
reduction in the alfalfa acreage. In Florida, the sub-tropics and tropics of
Australia and in the remainder of a large part of the sub-tropical and tropical
world there has been increased interest in and commercial use of tropical
legumes. This is particularly true as a result of the large increases in
fertilizer nitrogen costs in 1973-74. Most of these areas are under extensive
pasture management systems where the use of nitrogen is too costly. Whether
the interest and work with these legumes is always justified is not known;
but according to Whyte (276) the thought of finding a "white clover" for the
tropics seems remote. From another viewpoint, there are many commercial
pasture areas in the world that are not using nitrogen and do not have a
legume component in the pasture. If a legume can be found that will maintain
itself in a grass pasture there is no doubt that the quality of the pasture
will be improved.
Generally, centers of tropical legume distribution in the Western Hemi-
sphere are to be found in Central and South America, with Mexico being one
of the main centers. Other large centers are in tropical Africa and Asia.
Smaller zones in Africa are the source of tropical, i.e., (Glycine wightii),
and sub-tropical types of legumes, i.e., (Trifolium semipilosum, and
Lotononis bainesii). The largest center of temperate legumes is the Mediterr-
anean area where many Trifolium and Medicago species are believed to have
developed. The ecology of this area is discussed by Rossiter (199). A
list of the origin of the principal cultivated legumes is presented by Burkart
(30 p. 68). He also describes the legumes used for human consumption, green
manures and forages, medicinal legumes and those used for lumber and other
industrial purposes. Towe (245) discussed the forages available for Argen-
There are excellent reviews and discussions of legumes used for human
consumption, i.e. Pigeon pea, Cajan cajanus (73, 74, 77, 135, 180), cc.i',
Vigna sinensis (181, 183), and general, (8, 156, 195, 218). This subje?-.
will not be discussed here.
Rather than describe the flora of various areas, a list of the number of
species reported for selected genera is presented in Table 4. Additional
information on the Galegeae, and Mimosoideae and Caesalpinioideae of the north
central States (of The U.S.A.) are reported by Welsh (269) and Isley (128).
Part of the South American Flora (mainly Peru) was presented by McBride (152).
Many of the naturalized species may have different agronomic characteristics
than those for plants in their original habitats. The number of species
listed represent only a small part of the indigenous and introduced flora.
There have been many more species included in the introduction programs of
almost all the major agricultural countries. Further identification could
be made according to species but this would require a number of tables. As
an example, a partial list of the Desmodium species reported in the literature
are presented in Table 5. Desmodium species are indigenous to all land masses
except New Zealand and the western United States and Europe. One large center
of distribution in the Western Hemisphere is in Mexico. Among others, the
species that are being or have been used commercially as pasture or cover
crops in several countries are: D. intortum, D. uncinatum, D. sandiwicense,
D. canum, D. barbatum, D. gyroides, D. tortuosum, and D. ovalifolium. Similar
lists can be made for other genera that would be of help to plant introduction
workers or plant breeders.
Climate, Latitude and Altitude
Climate as it affects cattle in tropical countries has been amply des-
cribed by Burgas (28). Both Davies (55) and Ebersohn (65) discuss homo-
climates in general and with respect to plant introduction for arid or semi-
arid zouns of Austrllia while Semple (209) discussed the subject more broadly.
Forage productivity in difficult environments was discussed in the book,
Australian Grasslands (258) in 1960; and there are several reports concerning
climate of various countries, i.e. Australia (52) and Costa Rica (207). The
effects of macro- and micro-climates on pasture production was presented by
Prescott (188) in 1956. Temperatures have an important role in the adaptation
of pasture species. This is especially true for the extremes (coldest and
hottest periods). Rainfall has an additional modifying influence.
Latitude and altitude effects are related to a pasture plant's adaptability.
There is a general belief, for example, that plants adapted to high altitudes
in the tropical zone (lower latitudes) can be used or introduced successfully
to higher latitudes and lower altitudes. Therefore, introduction of high
altitude-low latitude pasture species into sub-tropical areas is believed to
be a better method of finding adapted species than attempting to introduce
temperate species into the sub-tropics. An excellent discussion of this point
was presented by Atkinson (9). Although there have been successful intro-
ductions of tropical grasses into the sub-tropics, the effect of altitude is
not always apparent. Kikuyugrass (P. clandestinium) is indigenous of the high
mountainous areas in tropical Africa and has been successfully used in sub-
tropical Australia and Hawaii at lower altitudes but most of its use has been
limited to mountainous areas within the tropics (more longitudinal movement
than the low latitude-high altitude to higher latitude-lower altitude movement).
Pangola (Digitaria decumbens) comes from a relatively low altitude and high
latitude, an area considered tropical; yet the northern and southern range of
Kikuyu and pangola growth are similar. Pangola, however, is able to grow in
the true tropical areas, while Kikuyu will not. On the other hand, successful
white clover introduction from temperate zones into sub-tropical zones has
been as great, latitudinally speaking. Tropical legumes from high altitude
areas (Glycine wightii, Desmodium intortum, Lotononis bainesii) generally
perform better during cooler periods in the sub-tropics than do low altitude
tropical legumes (Centrosema pubescens, Phaseolus sp.). In any case, the
effect of latitudinal movements because of daylength influences can have a
pronounced effect on flowering responses in a number of genera. The differ-
ences in number of daylight hours at different latitudes is shown in Table 6.
Table 6. Effects of latitude on the approximate range and maximum
number of daylight hours difference from the shortest to
longest day (28).
North or South Maximum range Total number of
of Equator of daylight Hours difference
Degrees Hours: minutes Hours: minutes
0 12 hours continuous 0
10 11:25 to 12:35 1:10
20 10:48 to 13:12 2:24
30 10:04 to 13:56 3:52
40 9:08 to 14:52 5:44
Many introductions of the legumes Stylosanthes guyanensis, Centrosema
pubescens and other genera obtained from lower latitudes (tropics) fail to
flower sufficiently early in the fall in Florida and elsewhere in the sub-
tropics. Conseqently seed production is not always assured because of frosts.
Within the tropical zones the effect of latitude is less defined and probably
relatively unimportant compared with local rainfall patterns and altitude.
Generally, there is little chance of the adaptation of tropical grasses out-
side the sub-tropics. Tropical legumes that produce seeds early enough
before the first killing frost may be adapted as self-regenerating annuals
in temperate areas under certain conditions; but there appears to be less
hope of obtaining a tropical legume that would perenniate in the temperate
areas without large-scale breeding efforts. Annuals such as Lablab purpureus
(L.) Sweet. (Dolichos lablab) could provide excellent quality summer feed in
the temperate areas. Farmer use would depend on adequate seed supplies from
tropical areas, however. Temperate grasses and legumes do not perform satis-
factorily in the tropics except at high-altitudes. Kretschmer (140, 143)
discussed altitude, latitude and daylength effects on tropical legume growth
and presented a diagram of the effects of these factors plus rainfall on
adaptability of white clover and tropical legumes within the 300 N and S
latitude zone. Jones (131) discussed the effect of frost damage on the
establishment of tropical grasses and legumes in the sub-tropics of Aus-
tralia, and Coaldrake and Russell (48) and Tothill (246) reported on the
effect of fire on establishment and persistence of legumes in native grass
After a researcher has decided that an introduced species may be of more
value than any of those locally available, the task of defining the scope of
the introduction program must be considered. As mentioned previously, the
introduction of grasses would probably be considered of prime importance if
additional dry matter were needed during some period of pasture stress (cold,
dry, or flooded conditions). In the sub-tropics where temperatures of 0 C
or less occur occasionally during the year and where tropical grasses are
the basis for the pasture program, it is doubtful that a perennial grass
will be found that can overcome the "winter stress" period. West (270) has
shown that at least some tropical grasses have a mechanism that prevent rapid
growth when temperatures of 100 C are reached. Some temperate grasses grow
well during winters in sub-tropical areas but plants generally will not
successfully survive the summer season. An alternative to the use of per-
ennial species in solving the "winter stress" problem is to utilize annual
winter grasses (Lolium sp., Avena sp., etc.) or temperate legumes (Trifolium
sp., Medicago sp., etc.j. Many times the difficulty with this approach is
the necessity of seeding the pasture every year because of the poor self-
regenerating abilities of most of the legumes and particularly the grasses.
If the local problem requiring the introduction of plants is one of
quality in the tropics or sub-tropics then the search should be made for a legume
that is adaptable. A general rule of thumb in deciding whether to try annual
or perennial legumes is the following. Perennial legumes should be emphasized
in the introduction program unless extremely low winter temperatures or
extremely dry periods of 7 to 9 months duration occur in the area. If a
perennial legume perenniates under the local conditions, and produces seeds
consistently, then quality and quantity generally would be greater than
that obtained from a self-regenerating annual legume. In areas of high,
well-distributed rainfall in the hot tropics annual legumes should not be
tested except under unusual circumstances. It will probably be a difficult
task to find a perennial legume that can compete with a cultivated grass when
soil fertility is high; or at least the management aspects of maintaining a
satisfactory population will be more difficult.
It has been the author's opinion that after the decision of what broad
type or types of plants may be desired, and after initial screening of various
representatives of the species, one or several genera should be selected for
further testing. Then, every effort should be made to obtain seeds of as many
species within the genus (or as many cultivars of a species) as possible.
This approach is much better than comparing only a few of the species or
cultivars. It will prevent retesting additional introductions of the genus
or species each year and will shorten the overall evaluation period. The
procedure was used very successfully in the search for a rapid-growing winter
clover to be used as a compliment to white clover in south Florida. From the
initial testing of many Trifolium alexandrinum (berseem clover) cultivars in
1961 (136), a description and commercial use of this high yielding winter
legume began in 1964 (137, 139).
Another approach can be used at the beginning of an introduction program
to find a suitable tropical legume. Tropical legumes that have been used
successfully in commercial plantings can be tried first to determine which
are adapted. It is believed that this method will simplify the initial
program and will result in finding a cultivar that would be adaptable. Because
of the more readily available commercial seed supplies, rapid commercial use
could be made of those that are adaptable. Glycine wightii (G. javanica) (221),
Centrosema pubescens, Stylosanthes guyanensis, Desmodium intortum, D. uncinatum,
Macroptilium atropurpureum (Siratro), Pueraria phaseoloides (Kudzu tropical),
and Leucaena luecocephala (L. glauca) are some of the perennial legumes that
are used commercially. Aeschynomene americana and Stylosanthes humilis are
two successful annual legumes. Kretschmer in 1971 (143) listed these and
others with their respective agronomic characteristics.
The introduction program must be accompanied by good record keeping. Such
items as country of origin, date of introduction, date of planting, and
accession number are essential to maintain continuity of the program and to
prevent duplication. Requests for seeds of introductions should include
requests for specific origins especially from second-country seed supplies.
An excellent method of maintaining a filing system for introductions is
to place all the available data on cards. Various sizes from 3 by 5 to 5 by
8 inches can be used depending on the amount of data desired. Smaller cards
would have only the basic information concerning the accession. Larger cards
have room for additional initial testing information such as, plant population,
seedling vigor, leafiness, seed production, winter or drought hardiness,
flowering date, and disease and insect resistance. Other characteristics
such as perenniality, root type, type of planting material used (seeds or
vegetative) can be included. On the back of the cards a tabular form can be
printed with which notations of height, spread, yield, etc. can be plotted
against differential fertilizer rates or time.
These cards can be conveniently placed in loose-leaf notebooks in
numerical order of accession; or separate books can be used for types of
plants, i.e., perennial legumes, annual legumes, tropical legumes, perennial
grasses and annual grasses. The author has filed accession cards in4 separate
books, one each for temperate or tropical legumes or grasses. Also separate
books are used for cards of introductions that have been discarded. These
cards can be used for taking field notes if necessary but there is a less
difficult complimentary system to keep field records for researchers dealing
with hundreds of introductions. In addition to the card catalogue, two lists
are maintained on typing paper one each for grasses and legumes (or 4 lists
to conform with the 4 books described above). A very brief description in-
cluding scientific name, origin and other country's accession numbers is
included in these lists. The lists greatly aid in searching the files for
the accession numbers of a particular genus or species. If particular
emphasis is being placed on the introduction of Centrosema species, for
example, an additional list on typing paper can be made to include only
For data collection, in addition to a plot outline, a list of the planted
introductions for a particular test should be typed to conform with the plot
outline. Columns, to the right of the names and brief descriptions, can be
used to record information on flowering dates, growth habit, insect damage,
etc. This system can be used for single plant plots, rows, or for replicated
or non-replicated small plots. For example, if 10 tropical legumes of
several genera and species were being tested as single plant plots with two
replications, a plot layout as shown below could be drawn and numbered
according to accession numbers as follows:
plot Dianrsm of'
Single Tropical Legume Introductions
Seeded June 10, 1974
I, I t
210 211 8 27
6 212 211 25
10 9 210 40
27 25 9 10
40 8 6 212
The list of appropriate columns should be made as follows:
Evaluation of Single Tropical Legume Introductions, Seeded
Origin and No.
" Australia (CPI 32742)*
ex Costa Rica
Ltus Costa Rica
i Australia ("Tinaroo")**
June 10, 1974
July 27 Aug. 15
atropurpureum Australia ("Siratro")**
muconoides Brazil (IRI 372)
8 Vigna repens Florida (local)
Australian accession number but seeds originally from Costa Rica.
** Common commercial names.
The list would continue in the order of observations for replicate B, i.e.,
6, 9, 210, 211, 8, 27, etc. and plant names could be omitted. This system is
particularly helpful when dealing with large numbers of introductions by saving
time in the recording of field data. Data can also be recorded on the plot
diagram sheet. Later, for replicated tests, the information can be transferred
to statistical sheets.
There is a helpful field technique which can be used when evaluating
initial flowering dates or other data when the positive observance precludes
further observations. If an evaluation for initial flowering includes 100
introductions of legumes with three replicates, for example, considerable time
is spent observing the 300 plots if done weekly. A colored plastic tape or
other easily seen marker can be attached to plants that began flowering during
the first week of observations (and the data properly recorded). The same
technique is used in succeeding weeks. The advantage of marking plants in
this manner is that they do not have to be examined in succeeding weeks. As
time passes, there will be less and less plants to observe. There have been
several comprehensive reports concerning plant introduction or methods used
in evaluating newly introduced plants. In Table 7 are presented the selected
references numbers with appropriate comments concerning the topics covered.
Field Plot Design, Weed Control, Germination and Plant Spacings
Plot Designs--For initial and further evaluation of pasture species it is the
author's belief that three or more replicates should be used wherever possible.
Although this requires more labor and space the advantages of being able to
discard a large number of introductions the first or second year with a good
degree of certainty over-shadows the disadvantages. Soil nutrient and moisture
variations and missing plants that die from known or unknown causes do not
result in the loss of one-year's data.
Weed Control--Weed growth will probably become a problem. Several things can
be done to combat weeds. If the area is free of tree stumps soil incorporation
of 35 to 50 gallons per acre of "Vorlex" (methyl mustard oil--CH3-N=C=S), or
other soil sterilant such as methyl bromide will kill vegetative material as
well as most of the weed seeds. Planting should not be made and the soil
should not be disturbed for about 3 weeks after application of Vorlex. At
the Agricultural Research Center, Fort Pierce, the following procedure was
successful in preventing excessive weed growth for from 4 to 6 months after
treatment. The area was deep-plowed, leveled with a disk, and the Vorlex was
applied into the soil about 4 inches deep with a special tractor drawn appli-
cator. The area was then packed with a heavy roller to delay the escape of
the gas produced. Application should be made when the soil is moist. This
is an expensive but useful method. The material should be kept away from
the skin and eyes because of its irritating and toxic characteristics.
Another system, if hand labor is expensive, or unavailable, is to use
paraquat at recommended rates. After the pasture seeds have germinated,
small tin cans or other containers are placed over the plants and the area
sprayed. A 3 gallon hand sprayer can be used effectively. Spraying should be
..... done only after there are numerous weeds evident, but before a majority of
weeds-reach-the height of about 2 centimeters. After plants are established,
the plants need not be covered for additional sprayings if care is used and
there is little or no wind. If this procedure is used, the soil should not be
disturbed again, or a large number of weed seeds will be exposed to the surface
and another germination will occur. Treatment about every two months would
prevent a large weed population from becoming established. Paraquat will
not kill well-established weeds or cultivated grasses or legumes; and because
it is a contact rather than a translocated herbicide, small quantities drifting
onto established introductions will not kill them.
"Treflan", "Eptam" and probably others may also be successfully used as
preemergence herbicides when incorporated into the soil for legume introdutions;
but application rates are not sufficiently known for the many soil types en-
For vegetatively planted grasses 2 kg/ha of simazin or 2,4--D can be
applied about 2-3 days after planting.. (aes--oon-as-weed germination commences).
The rate to apply depends on soil type.
Tractor-drawn disks or harrows can also be used successfully if plant
spacings, etc. are fitted to the type of equipment to be used.
Where labor is available, normal hand cultural methods can be used if
care is taken not to damage the plants. For mechanical or hand cultivation,
early cultivation is much more desirable and efficient than late cultivation
when weeds are difficult to kill.
Germination and Plant Establishment--With vegetative material, direct planting
into the field is desirable, with irrigation and fertilization as necessary to
With temperate legume seeds or grasses, if sufficient seeds are available,
several seeds can be planted in one spot (if single-spaced plants are used)
and later plants can be thinned to a desired number; for row or small plot
plantings the seeds can also be seeded directly if the germination percentage
If there are insufficient seeds (as usually occurs for initial intro-
ductions) then a different procedure will have to be used. Seeds can be
planted in soil in the greenhouse or in a seedbed outside and then plants can
be transplanted as needed. A problem may arise for this or the direct field
seeding method,however. Hardseededness for tropical legumes (37) or poor
germination percentage for tropical grasses plus the added danger of soil borne
diseases, insects, or inclement weather (if planted outside) may reduce or
eliminate the necessary number of plants needed. In any event, all of the
seed of a cultivar or species should not be planted because of the difficulty
of replacement should insufficient germination occur.
If only a few seeds of each introduction are available, germination in
petri dishes is a satisfactory method for establishment. Seeds are not lost
with this method, and if hardseededness prevents germination, seeds can be
scarified. In this method, one or two pieces of filter paper (or other
appropriate paper) are placed in numbered petri dishes and the seeds kept
moist with water. For grasses a 0.2% potassium nitrate solution should be
used initially to enhance germination. As seeds germinate they can be
placed in soil in the greenhouse or in pressed peat pots (commercially
available in various sizes 2 by 2 inch peat pots are satisfactory for most
species). Later they are transplanted to the field. Little damage is done
to the root system with this method. The top edges of peat pots should be
removed at least to the soil level before placing them into the soil. The
exposed edges of peat pots cause dessication of the soil and root zone of the
plant through capillary action.
Hardseededness of certain tropical legume seeds can be overcome by
individual scratching of each seed with a sharp instrument, the use of fine
sandpaper for larger quantities, or by moistening seeds with concentrated or
half-strength sulfuric acid for about 15 to 30 minutes. Soaking of seeds
of Stylosanthes humilis or Macroptilium atropurpureum (Siratro) in hot water
(700C) for 30 minutes or placing them in the oven (700C) for the same length
of time has proved to be a satisfactory means of increasing the percentage of
readily germinatable seeds. These treatments may reduce germination of certain
legume seeds so laboratory tests with a few seeds of a particular species
should be tried before all seeds are treated.
Plant Spacings--With single plant plots the seeds of species being used should
be pure so that each plant represents the type. The problem arising from the
use of non-pure seeds is obvious; the replicates may represent different
rather than a single type. Single plant plots offer the advantage that the
plant can be more easily observed and notes on rate of growth, type of root
system, flowering date, and other measurements are more precise. Number of
seeds produced per plant and number of seeds that germinate also can be
estimated more readily. In contrast, rows or small plots of plants make it
more difficult to observe single plants, by permitting a number of plants to
obscure the individual plant and its growth. The plants should be spaced to
give the shortest possible distance between plants without allowing the plants
to grow into each other. With several stoloniferous types of grasses and
tropical legumes, plant spacing of even 2 meters in the row and between rows
may not be sufficient to completely evaluate growth rate. However, in these
instances a compromise usually must be made and plants should be cut back as
necessary (or the growing tips sprayed with paraquat) during the growing
season. With Centrosema or other similar trailing species, posts can be
placed by each plant, and the plants can be trained to grow up the posts
and tied with string when needed. This is satisfactory for obtaining
flowering information, insect or disease resistance or for breeding purposes,
and requires less space. If introductions with various growth habits are to
be planted, and the general growth habit is known in advance, the bunch or
upright types can be planted in separate blocks, at shorter intervals than
the stoloniferous types.
Rows of convenient length (2 to 7 or more meters) have been used ex-
tensively for testing introductions and breeding lines. These have an ad-
vantage compared to single plant plots because the death of one or more plants
does not necessarily deter from the data to be recorded. A shortage of seeds
makes this type of plot more difficult or impossible to use; and if direct
seeding is not used, the transplanting problem with respect to time and labor
becomes a function of the number of plants per plot multiplied by the time
and labor necessary to use the single plant plot technique. A modification
of the row system, if transplanting is necessary, would be to limit each row
to three plants spaced closely together (15 to 60 centimeters).
Small plots (i.e. 1 by 3 to 3 by 5 meters) can be most useful when working
with grasses or grass-legume mixtures; but generally this technique should be
used in the later evaluation stages of work or with known cultivars in the
initial introduction phases. A better idea of growth under simulated grazing
(clipping the plots periodically) can be attained with this method. Also,
disease and insect problems that could occur under field conditions are more
likely to be simulated in this type of planting as compared with the single
row or single plant method; and competition with weeds can be more effectively
For any field design it is helpful to use stakes in appropriate places to
mark the plants or plots. It is even more helpful if these stakes are marked
with the easily read appropriate accession numbers. This reduces the chances
of making mistakes when collecting field data since the stake number can be
checked with the number on the introduction or plot diagram sheet when data
is being recorded.
Any of the three methods can be used in the initial testing phases in
commercial pastures if the owner is cooperative. A group of replicated plots
can be established at the Research Center and a similar set in a commercial
pasture so as to shorten the initial evaluation period. Using any of the
three types of plots and working with legumes, seeds can be planted in com-
mercial field at the same time that a productive grass of the area is planted
so as to be able to evaluate the competition effects.
It should always be kept in mind that many species of certain genera, i.e.
Crotalaria (278), Indigofera (122) and other species (210, 211, 278) are toxic
to cattle. These should not be evaluated except under closely controlled
After initial screening, there are two or three additional steps in the
evaluation technique that generally should be taken prior to the "release" of
a cultivar for commercial use. Small-plot clipping experiments are valuable
for comparisons between established cultivars and the new cultivar. Clipping
interval tests (82, 125) can be used to simulate the effects of frequent or
continuous grazing. However, there is no substitute for the animal since
trampling and selective grazing cannot easily be simulated by clipping tests.
The effect of the animal on the plant can be determined by a technique where
a large number of cattle are turned into a pasture comprised of a number of
replicated plots of different species. The object is to graze intensively
for a short period so all plots are defoliated and palatability effects will
be reduced. The quicker the grazing time, however, the more closely this
technique and clipping approach. With the proper research facilities new
grass varieties (and legumes) can be tested for their effect on animal gains
by grazing trials (25, 44, 81). This is another subject that will not be
discussed here, but is one that requires a large input of capital to be an
effective research tool.
Many analyses have been made for crude protein, fiber, fat, carbohydrates,
etc. in an effort to evaluate the nutritive value of grasses, legumes and their
mixtures. With grass testing, after initial evaluations, it may be desirable
to compare the trends of crude protein and other chemical constituents of new
versus the commercial grasses. Once this trend is established then little is
gained by additional analyses (145). It is generally accepted that at a given
cutting interval (or grazing interval), the more nitrogen applied to a tropical
grass the greater will be the crude protein content, while the crude fiber and
dry matter percentages will be less. At a fixed level of nitrogen the longer
the interval after application, the less the protein content and the greater
the fiber and dry matter contents will be. Also, crude protein yields (dry
matter yield times crude protein percent) of grasses are almost invariably
more positively related to dry matter yields than to crude protein contents;
i.e. when yields increase then crude protein yields also increase. Furthermore,
there is general agreement that above a certain minimum protein content, in-
creasing nitrogen rates increase the pounds of beef produced per area but
seldom increase the pounds of beef per animal when compared to a lower, but
adequate, nitrogen rate at a very low ("free-choice") stocking rate that
permits maximum grazing selection.
The value of "in vitro" or "in vivo" digestibility analyses of tropical
grasses and legumes is limited compared to their value in evaluating temperate
species where a good positive relationship exists between dry matter intake and
"in vitro" (and "in vivo") digestibility. For tropical species the relation-
ship between "in vitro" and "in vivo" digestibilities is sometimes good; but
may not have any relationship with dry matter intake. Excellent reports by
Minson (165, 163, 290, 291) and Minson and Haydock (164) indicate the problems
involved with "in vitro" digestibility and other laboratory methods for
assessing the quality of pasture species. The conclusion was that there is
no simple or even a complicated laboratory method to estimate dry matter in-
take by cattle or sheep.
Some legumes such as Coronilla varia (29), Lespedeza cuneata (62), and
Desmodium sp. (120, 200) contain tannin, a compound that may reduce intake if
found in sufficiently high concentrations. It would be necessary to know the
tannin contents of these types of legumes so that this information can be
utilized in selecting the best cultivar.
Other Factors that May Affect Initial Evaluation
Rhizobium--In most instances, "cow-pea" type bacteria can be used to inoculate
the tropical pasture legume species. Important exceptions are in the genus
Lotononis (22), and certain cultivars of Stylosanthes hamata, and Leucaena
leucocephala. Some temperate Trifolium species, i.e. T. semipilosum are not
effectively inoculated with commercial white clover bacteria. Dark green,
vigorous growing plants indicate effective nodulation. Roots should be
examined for nodules on those plants that appear to be non-vigorous and light
green to yellow in color.
Lime and fertilizer--The optimum amounts of lime, phosphorus, potassium, and
a mixture of trace elements (including molybdenum) should be applied and in-
corporated into the soil prior to planting legume introductions. For grass,
in addition, sufficient nitrogen should be used to promote vigorous growth.
The object is to assure adequate soil nutrients so they do not limit plant
Insects and diseases--Normally, above-ground insects should be controlled
during the initial stages of growth until the plants become established.
After that, notes should be made concerning insect types and preferences.
Nematodes and below-ground insects are difficult to control unless a soil
sterilant is used prior to planting. The site-for-the introduction garden
should be chosencaefy-s~f o-that-hee problems are minimized or a soil
sterilant-should be used if a potential problem is suspected.
Soil borne diseases such as Rhizoctonia sp. can kill germinating and
-...-.. -young seedlings. Every effort should be made to prevent this from occurring
in order to obtain the plant population desired. After establishment,
diseases attacking plants should be identified and the severity of damage
recorded. A few of the references concerning diseases are: 2, 67, 68, 222,
272 and 292.
Other pests--Damage to youin and old plants from rats and rabbits can affect
the evaluation of growth rates, seed production, etc. Differential "grazing"
by these animals can result in false evaluations, i.e. the more heavily grazed
introductions may be rated as less vigorous than the non-grazed ones, even
though they are faster-growing. A small-meshed wire fence (with holes about
3 centimeters in diameter), a meter or more high was effective at the
Agricultural Research Center, Fort Pierce in keeping rabbits out; but rats
and mice had to be poisoned.
Additional References Concerning Specific Genera or Species
In Table 8 is presented a list of various references that will be useful
in obtaining initial information concerning some of the important legumes
being used throughout the world. The list does not include many legumes that
are being used because of the impossibility of reviewing all of the publications.
Alfalfa is not included because of the numerous books and scientific publica-
tions that are readily available in most libraries.
1. Ahlgren, Gilbert H. and R. F. Fuelleman. 1950. Ladino clover. Advances
in Agronomy II. Academic Press. New York. pp. 207-232.
2. Aldrick, Susan J. 1971. Bacterial wilt (Pseudomas solanacearum) of Stylos-
anthes humilis in the Northern Territory. Trop. Grass. 5:23-26.
3. Andrews, E. C. 1914. The development and distribution of the natural
order Leguminosae. Jour. Roy. Soc. N.S.W. (Australia) 48:333-407.
4. Anonymous. 1939. Bibliography on white clover (Trifolium repens). Imp.
Bur. Past. Forage Crops. Mimeo Pub. 5. (Great BritainT. pp. 30.
5. Anonymous. 1939. Bibliography on.red clover (Trifolium pratense) Imp.
.Bur. Past. Forage Crops. ,Mimeo Pub. 4. Aberystwyth. pp. 60.
6. 1' 1960. Persian clover a legume for the south. U.S.D.A.
Leaf. 484. .
7. 1965. Pasture legumes and grasses. Bank of New South Wales
S(Australia). pp. 76.
8.' 8 :.. USDA, ARS, USAID. :-1968. Regional pulse:improvement project.
Prog. Rpt. No.6. pp. 232.; .. .:
9. Adkinson, W. T. 1970. High altitude-low latitude forage plants from Mexico
Sand Latin America. Proc. 11th Int. Grass. Cong. pp. 181-184.
10. Barker, Henry D. and William~S. Dardeau. 1930. Flore D'Haiti. Serv.
STech. Dep. Agr. Port-Au-Prince. pp. 456.
11. Barnard, C. 1969. Australian herbage plant species. :C.S.I.R.O. Div.
Plant Ind. Canberra. pp. 154.
12. Barnes, D. L.: '1968. Dryland pastures. Rhodesia.Agr. Jour. 65:1-8.
13. Bogdan, A. V. 1964. Cultivated varieties of tropical and subtropical
herbage plants in Kenya. East African Agr. For. Jour. 30 (4):
14. 1966: 'Plant introduction, selection, breeding, and multi-
plication. Tropical pastures.. Faber and Faber, London. pp. 75-88.
15. 1969. Rhodes grass. Herb. Abstr. 39:1-13.
16. Booth, W. Edwin. 1964. Agristology. Edwards Brothers. Ann Arbor,
Michigan. pp. 222.
17. Boultwood, J. N. 1964. Horse marmalade and Kuru vine. Rhodesia Agr.
Jour. Bul; 2258. pp. 4.
18. Boyd, Frederick T.' and V. G. Perry. 1969. The effect of sting nematodes
on establishment, yield and growth of forage grasses on Florida
sandy soil. Soil Crop Sci. Soc. Florida Proc. 29:288-300.
19. Braun, Otto. Pastos del tr6pico humedo de Bolivia. Min. Agr. Bol. Exp. 31.
20. Brockwell, J. .1969. I.B.P. (section PP-nitrogen fixation) Bibliograph-
ical report. CSIRO Div. P1. Ind. Tech. Pap. 28. pp. 167.
21. Bronckers, F. et:B. de Keyser. 1966. Contribution A 1' etude poly-
nologique des Papilionaceae Phaseoleae Phaseolinae. I.
SObservations sur:quatre especes de Dolichos L. et sur ;Lablab niger
Medic.. Bul.'du J ardin Bot. de L'etat. 36:57-63.
-22. Bryan,W. W. 1961. Lotononis bainesii Baker, a legume for sub-tropical
.;,* pastures. 'Australian Jour.,Exp. Agr. and Anim. Huab. 1:4-10.
23. '1963. "A search for tropical pasture legumes a progress
report. Jour. Australian Inst. Agr. Sci. 29:149-153.
24. .1969. Desmodium intortum and Desmodium uncinatum.
Herb. Abstr. 39:103-191. .
S 25. and T. R. Evans. 1971. A comparison of beef production from
nitrogen fertilized pangola grassand from a pangola-grass legume
:pasture. Trop. Grass. .5:89-98.
26. Buller, R. E., et. al. 1970.. Performance.of tropical legumes in the up-
land savannah of central Brazil. Proc. 11th Int. Grass. Cong.
; .pp. 143-146.
27. Burbidge,:N. T. 1964. Grass systematics. Grasses and'grasslands.
C.S.I.R.O. Canberra. MacMillan. New York. pp. 13-28.
. 28. Burgas, Juan L. 1969. El clima en. la production del ganado tropical.
S Ganaderia Tropical. Tomo 1. Libraria "El Ateneo".- Buenos Aires.
29. Burns,:R E., P. R.'Henson, and D. G. Cnmmins. 1967. Tanninmcontent of
crown vetch (Coronilla varia L.) herbage. Agron. Jour. 59:284-285.
30. Burkart, Arturo. 1952. Las leguminosas Argentinas silvestres and cultiv-
ades. Acme Agency. Buenos Aires, Argentina.. pp..569.
31. Burth R. L., :et al. 1970. Assessing the agronomic potential of the genus
Stylosanthes in Australia. Proc. 11th Int. Grass. Cong. pp. 219-223.
32. Burton, Glenn W. 1951.' The adaptability:and breeding of suitable grasses
for the southeastern States. Advances in Agronomy III. -Academic
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202. and Ukio Urata. 1966. Some agronomic observations in
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203. Sacco, Tommaso. 1964. Contribute allo studio della flora pabulare di
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208. Seanay, Robert R. and Paul R. Henson. 1970. Birdsfoot trefoil. Advances
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210. Shone, D. K. 1966. Poisonousplants of-Rhodesia (Part I). Rhodesian
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211. 1967. Poisonous plants of Rhodesia (Part II). Rhodesian
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212. Schubert, Bernice G. 1943. Desmodium. Field Mus. Nat. Hist. 13:413-4:
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216 --sillar, D. I. 1967. Effect of shade on growth of Townsville lucerne
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217. Skerman, P. J. 1970. Stylosanthes mucronata Willd., an important natural
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219. Smith, W. K. and.H. J. Gorz. 1965. Sweetclover improvement. Advances in
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220. Smith, C. A. 1962. Tropical graas/legume pastures in northern Rhodesia.
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221. Smith, C. Earle, Jr. Recent nomenclatural changes for cultivated plants.
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233. 1971. First records and new combinations in the vascular
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238. Thomas, 0. A. and McLaren, L. E. 1971. Some studies on the digestibility
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252. 1959. The legumes of Texas. Univ. Texas Press. Austin
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254. Tutin, T. G. 1958. Classification of the legumes. Nutrition of the
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255. Van Donselaar, J. 1969. Observations on savanna vegetation-types in the
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256. Van Rensburg, H. J. 1967. Pasture legumes and grasses in Zambia.
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257. Various authors. 1948. Grass yearbook of agriculture. U.S.D.A.
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258. 1960. Forage provision in difficult environments.
Proc. 8th Int. Grass. Cong. pp. 154-169.
259. 1970. Australian grasslands. Australian Nat. Univ.
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260. Various authors. 1970. Selection and testing of temperate species and
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262. '. 1971. The ability of pastures in the coastal lowlands
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264. 1966. The grassland communities of Barotseland. Trop.
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265. 1968. Grassland successions and associations in Pahang,
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266.: Vicente-Chandler, Jose, et al.. 1967. El manejo intensive de forrajeras
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267. Villamizar R., Fernando and Jaime Lotero C. 1967. Respuesta del past
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268. Wallace, A. T. 1957. Hairy indigo, a summer legume for Florida. Florida
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269. Welsh, Stanley L. 1960. Legumes of the north-central states: Galegeae.
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270. West, S. H. 1970. Biochemical mechanism of photosyntheses and growth
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272. .Whiteside, J. 0. 1959. Diseases of legume crops in Southern Rhodesia.
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275. Whitney, L. D., E. Y. Hosaka, and J. C. Ripperton. 1939. Grasses of the
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276. Whyte, R. 0. 1962. The myth of tropical grasslands. Trop. Agr. Trin.
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278. ___, G. Nilsson-Leissner and H. C. Trumble. 1953. Legumes in
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279. Wilbur, Robert L. 1963. The leguminous plants of North Carolina. North
'Carolina Agr. Exp. Sta. Tech. Bul. 151. pp. 294.
280. Williams, R. J. 1964. Plant introduction. Some concepts and methods in
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281. Wilms, H. J., J. W. Carmichael and S. C. Schank. 1970. Cytological and
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282. Wilson, G. P. M. 1970. Method and practicability of Kikuyu grass seed
production. Proc. llth Int. Grass. Cong. pp. 312-315.
283. ___and G. J. Murtagh. 1963. Lablab new forage crop
for the north coast. New South Wales Dept. of Agr., Div. of PL..._--
284. Wilson, J. R., K. P. Haydock and M. F. Robins. 1970. The development
in time of stress effects in two species of Glycine differing in
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285. Wright, C. B. 1964. Lotononis in the Wallum. Queensland Agr. Jour.
(Feb.) pp. 93-96.
286. Wutch, J. G., E. M. Hutton and A. J. Pritchard. 1968. Inheritance of
flowering time, yield and stolon development in Glycine javanica.
Australian Jour. Exp. Agr. Anim. Husb. 8:316-322.
287. 1968. The effects of
photoperiod and temperature on flowering in Glycine javanica.
Australian Jour. Exp. Agr. Anim. Husb. 8:543-547.
288. Younge, O. R., D. L. Plucknett and Peter P. Rotar. 1964. Culture and
yield performance of Desmodium intortum and D. canum in Hawaii.
Hawaii Agr. Exp. Sta. Tech. Bul. 59. PP. 22.
289. Zandstra, Ilse I. and William F. Grant. 1967. The biosystematics of the
genus Lotus (Leguminosae) in Canada. I. Cytotaxonomy. Canadian
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290. Minson, J. D. 1972. The digestibility and voluntary intake by sheep
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292. Sonoda, R. M. 1974. Diseases of Stylosanthes spp. Fort Pierce ARC
Res. Rpt. RL-1974-4.
293. Neal, Rodney H. 1972. Some pasture grasses in Belize. Bul. 1, Min. Agr.
294. Fretes, Ruben, Ricardo Samudio, and Cahrles Gay. 1970. Las praderas
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295. Paladines, Osvaldo. 1974. Potential for increasing beef production in
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296. Lotero, Jaime C., Chaverra G., Hernan, and Crowder. Loy V. 19. Gramineas
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pecuario Manual No. 10.
297. Various authors. 1971. Peru veterinary institute for tropical and high
altitude research rpt.
298. 1971. Peru veterinary institute for tropical and high
altitude research rpt. No. 21.
299. Hutchinson, J. and J. M. Dalziel. 1958. Flora of west tropical Africa.
Vol. 1 (2). pp. 828. H.M.S.O. P. 0. Box 569, London, S.E.1.
300. Liberato, Maria Candida. 1972. Flora de S. Tome e Principe Papilionaceae.
pp. 90. Min. do Ultramar. Lisboa, Portugal.
301. D'orey, Jose, and Maria Candida Liberato. 1971. Flora da Guine Portuguesa
Papilionaceae. pp. 172. Min. do Ultramar. Lisboa, Portugal.
302. Exell, A. W. e A. Fernandes. 1966. Conspectus florae Angolensis.
Leguminosae. Vol. II. pp. 407. Junta de Investigacoes do
Ultramar. Lisboa, Portugal.
303. Clayton, W. D. 1970. Flora of east tropical Africa. Gramineae.(Pt. 1).
pp. 176. H.M.S.O. P. O. Box 569, London, S.E.1.
304. ___ S. M. Phillips and Renvoize. 1974. Gramineae. (Pt. 2).
pp. 450. H.M.S.O. P. 0. Box 569, London, S.E.1.
305. Verdcourt, B. 1970-71. Studies of the Leguminosae Papilioneideae for
the flora of tropical east Africa. Kew Bul. 24,25 (Pt. I-V).
J; : ;' i ,^ *- ;::
-- u *B oTiottA-I
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i H 0 r-4 *V400r 0 0 m N
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Table 4. (cont'd.)
Country or Area
31. East Africa
32. Sao Tome &
33. Portugese Gi
Sg d 1 2
0 o a e a a m ao
OH O- k d +> l3
M O t 4 0 H wO O 30 0 S3 ri
S0 0 0 0 a
r4U U U U O Id 1 O O 0
3 3 51*- 2 2 8 1 57 -
1 1 2 58 1 7 3 1 2 3 -
2 1 1 20 4 -26 2 5 1 3 -10 1
2 5 1 2 r 14 26 1 3 34 2 5 2 28 3
Approximate Approximate Approxim
Latitude Altitude Annual R
Country or Area References Degrees m mm
158 \ 25.5-280
255 \ 3-50N
10 \ 18-20N
I _~ __ _
Co try or Area
N4. Carolina (U.s.)
Sao Tome & Principe Is.
Only named species counted. In some instances local or introduced types were unidentified except for genera.
/Short dry season (1-3 months). 3/ Various types of rainfall pattern. 4/ Moderate dry season (3-6 months).
/Severe dry season (greater than 6 months). 6/ Evenly distributed. 7/ Only species not reported in other
ambia reference. 8/ Species introduced. 9/ Personal communication from Ing. Mario Arana, M.A.G., Guatemala;
om an old list of Guatemalan flora. Including closely related genera.
___ __References __
barbatum 10,19,30,158,202,212,214,255,263,299,302 2/
Table 5. List of Desmodium Species from Various Parts of
adscendens 10,30,124,212,299,300,301,302 38.
affine 10,30,212,223,214,263 2/ 39.
africns^ 263 10.
ang.sta.-e 2/ 41.
ang's- tS-PcF *- 223, _/ 42.
asperun 223 43.
axillare 10,212,214,215,223 4.
the World with Appropriate Referencesl/
glabellum 279 301,302
gyroides 124, 197,202 2/
linearifolium 299,301 2/
Table 5. (cont'd.)
sa idwicense 124,202
sco piurus 10,27,124,202,212,223,299 2/
ser ceum 223
sess tifolium 127,252,279
seti arum 214,247,299,302
spiral e 263,265 2/
stolz 1 214
strob lacium 223
schwei furthii 299
triquetrum 202 301
Table 5. (cont'd.)
143. umbellatum 47
144. uncinatum 30,47,124,197,2b2,212,265 2/
145. vargasianum 212
146. varians 124,202
147. velutinum 124,202,214,235,247,263,299,300,301,302
148. venosum 30
149. viridiflorum 124,127,213,252,279
153. Zonatum 47
l/ No attempt was made to alter species names even though there may be some duplication since, for example,
D. tortuosum has also beed named D. spirale (214).
2/ Personal communication frop Ing. Mario Arena, M.A.G., Guatemala with a copy of an old list of Flora of
Guatemala; all were listed as Meibomia rather than Desmodium.
Table 7. A Selected List of Research Work Dealing with Plant Introduction
41. Stylosanthes humilis
35. -- "
37. StylPsaEth6s humilis
72. Glycine wightii
clipping tests after initial evaluation
20 locations from sea level to 4,000 meters
high altitude-low latitude plants for subtropical
screening for subtropics
primary evaluation in subtropics
semi-arid screening and introduction
dairy animals, effects of legumes and grasses on
nutritive value of legumes and grasses in subtropics
legumes and grasses
general discussion of introduction possibilities
discussion of aims of plant breeding; applicable to
laboratory method and discussion of mechanism that
could be used for cold tolerance evaluation
agronomicc potential assessed
methods of screening plants
II It 11 II
high rainfall tropics-animal
collection and evaluation in
evaluation on various
initial evaluation of 11 introductions and coliectio
methods of obtaining data on initial flowering dates
of 92 types
replicated grass clipping evaluations for subtropics
type of Rhizobia needed for various tropical legumes
methodology for hard-seededness evaluation
morphological and agronomic characterization of 127
Table 7. (cont'd.)
241. African Trifolium sp.
45. ~r-L go truncatula
50. Medicago sativa
244. African Trifolium sp.
i79. Digitaria sp.
147. Legume-grass mixtures
evaluation of low latitude-high altitude plants in
low altitude suibtropics
methodology of ev',l.i.u::on and reporting variability
Of 206 intr du.ac.~to! in temperate area
evaluation of 4, c~u; ~;
evaluation of 40 Ic. .-s using yield and height data
evaluation of flowering and growth of 13 species using
selection for drought resistance
selection of grasses for nematode resistance
small plot technique for initial evaluation of mixture
Table 8. Selected List of References Concerning Specific Legumes
Scientific Name References Subject Matter
Alysicarpus vaginalis 294 Description and use in Florida
Arachis sp. 289 History, description and use in Florida
Canavalia sp. 191 Botanical classification
i" Tr- 205 Revision of genus
Cassia fasciculata 74 Management and yields
1 -- 73 Response to fertilizers and plant populations
Centrosema pubescens 230 Beef production
n "1 117 Flowering and pollination
Desmanthus sp. 249,250,253 Species in Texas and Mexico and chromosome numbers
Desmodium sp. 121 Breeding
i- Tr- 133 Contribution to yield and quality in mixtures with pangola
120,200 Crude protein and tannin contents
202 Seed weight differences among species
271 Effect of temperature on growth of 2 species
Desmodium canum 288 Culture and yields
Desmodium discolor 17 General use in Rhodesia
Dqsmodium intortum 24 General review
125 Defoliation and moisture stress
201 Agronomic characterization
17 General use in Rhodesia
i" 288 Culture and yields
Desmodium uncinatum 24 General revieww
----. 117 Flowering and pollination
Dolichos sp. 21,153,154,155 Chromosome comparisons
Table 8. (cont'd.)
MacrOtyloma uniform Vevdc. 229
(Doiichos uniflorus = D. biflorus)
Lablab purpureus (L.) Sweet 283
; -" 184
it "I 96
Glycine wightii 72
it. "t 284
It It 66
Indigofera hirsuta 268
Indigofera spicata 117
Lespedeza sp. 91
Lespedeza stipulacea 73
Leucaena leucocephala 79,176
Lotononis bainesii 22,285
-n- --r 34
Lotus sp. 104
Lotus corniculatus 208
Lotus uliginosus 112
Medicago tornata 185
Description and use in Australia
Use in subtropics of Australia
Description of Rongai cultivar
Effect of temperature on flowering
Characterization of various introductions in Australia
Effect of salt stress
Inheritance of flowering time and yield
Effect of photoperiod and temperature on flowering and growth
Yields of 38 varieties in 3 Australian environments
Effect of temperature on seedling germination
Description and use in Florida
Flowering and pollination
Breeding and toxicity
Response to fertilizers and plant populations
Production and processing
General description for use in Australian subtropics
Breeding system and chromosome numbers
Taxonomy of Hawaiian species and value as pasture legumes
Chemataxonomy of 9 species
Morphological and cytological characterization of numerous typ
General description and use in Oregon, U.S.A.
Cultivar, Tornafield, described
Table 8. (cont'd.)
Macroptilium atropurpureum (DC).
Macroptilium lathyroides (L.)
Testing and growth of 200 types
General review covering all phases
Revision of U.S. species
Description and use in Florida
Description and use in Florida
Flowering and pollination
Taxonomic studies using numerical procedures
Revision of genus
Chromosome numbers and morphology
General in Africa and Madagascar
Description of seeds and revision
Descriptions of species from Central America and Mexico
Photoperiod effects on flowering and growth of 4 species
Review and botanical description
Flowering and pollination
Effect of cutting on growth of 3 ecotypes
Hardseededness and seed dormancy
Daylength and temperature effects
Description and use in Florida
History in Australia
Flowering and natural distribution in Australia
Controlled environmental studies
Effect of latitude and time of sowing or flowering
Effect of shade on growth
General habitat and description
Botanical review of cultivated species
Effect of phhtoperiod anp night temperatures on flowering and
growth or 3j A rican species
Table 8. (cont'd.)
----- I- -I-
General agronomic evaluation of African species
Description and use in Florida
Use in Yugoslavia
Description and use in California, U.S.A.
Description and use in Alabama, U.S.A.
Complete review of Ladino clover
General description and use
Description and use in Alabama, U.S.A.
Discussion of various tropical legumes (and grasses)
Effect of soil temperature on nodulation
Inheritance of flowering