Erythrina (Leguminosae: Papilionoideae):

A Versatile Genus
for Agroforestry Systems
in the Tropics
Ricardo 0. Russo

ABSTRACT. The genus Erythrina is of special interest in the devel-

opment of agroforestry systems because of its adaptability to several
uses (e.g., live posts for fences, shade trees for perennial crops such
as coffee and cacao, forage for livestock, and others). Rapid plant
growth, high biomass production, easy propagation from cuttings,
ability to withstand regular pruning plus subsequent rapid sprouting
and development of strong shoots, are characteristics that make Ely-
thrina an attractive genus to be used in agroforestry. The abundant
nodulation in the rootlets and its potential as a nitrogen fixer, open
interesting possibilities for establishing plantations on low fertility
soils and/or restoring these soils. Some of the most common uses of
Erythrina species are discussed in this review related to specific
agroforestry applications.

Ricardo 0. Russo is affiliated with Yale Universitv. School of Forestw and

Environmental Studies, 370 Prospect Street, New ~ a v & CT , 06511;
The author pratefully appreciates the cooperation and support received from all
the faculty and;taff of ihe'fropical ~~riculiural.~enter for~esearchand Training
(CATIE), Turrialba, Costa Rica, during his stay at the ,Center (1981-1986). Ac-
knowledgment is also extended to the International Development Research Center
(IDRC) from Canada for the financial support received through the Eythrina
Project (1983-1986) and to the personnel of INFORAT at CATIE, Turrialba,
Costa Rica, where the main body of references were obtained. The author also
appreciates the cooperation and suggestions given by Dr. John C. Gordon, Dean
of the Yale School of Forestry and Environmental Studies, during the preparation
of this manuscript.
Journal of Sustainable Agriculture, Vol. l(2) 1990
O 1991 by The Haworth Press, Inc. All rights reserved. 89
90 JOURNAL OF SUSTAINABLE AGRICULTURE

INTRODUCTION
Currently, agroforestry is a rational alternative for land use. It
contributes to food, fiber and wood production through a simple
and generally low-cost technology. This practice combines trees
with crops or pastures in a simultaneous or alternate association,
with the objective of maximizing production per unit area, under
sustainable conditions (Combe and Budowski, 1979).
In a Workshop on Agroforestry held at CATIE, Turrialba, Costa
Rica, March 1979 the participants reached the conclusion that . . .
"When agroforestry systems are utilized, many elements enter into
consideration such as: more efficient use of space, provision of the
basic necessities of the rural populations, improvement of the qual-
ity of life in agricultural communities, avoidance of a reduction in
the productive capacity of the resources involved and a reconcilia-
tion of the short term interests of the farmers with the long term
interests of countries or regions" (Workshop Agro-Forestry Sys-
tems, 1979). Nitrogen fixing trees (NFT) play an important role in
agroforestry, because of their utility and their potential for adding
..
available nitrogen to agroforestry systems . Erythrina spp. are
NFT frequently used in agroforestry systems.
Description of the Genus Etythrina
The genus Erythrina L., belongs taxonomically to the subfamily
Papilionoideae of the Leguminosae. However, in the literature, and
according to botanists that separate the legumes into three families
it also appears as belonging to the Papilionaceae or Fabaceae family
(Krukoff, 1939).
The genus contains over one hundred species (Krukoff and
Barneby, 1974), distributed throughout the tropics and subtropics,
in a wide variety of habitats (Neill, 1984) in both the Old and the
New World. There are 26 species in tropical Africa, five in South-
ern Africa, twelve in Continental Asia and the Pacific Islands, and
two in Australia. In the New World there are 27 species and three
subspecies in Mexico, 25 in Central America and nine in the Carib-
bean (Krukoff, 1982).
Lackey (1981) places Etythrina in the subtribe Erythrininae of
 Research, Reviews, Practices and Technology 91

the tribe Phaseolae (Fabaceae-Faboideae), with the following com-

ment: "The relationship of this genus to the remainder of the Papi-
lionoideae is an absolute mystery. The genus would have long ago
accomodated outside the Phaseoleae had not the foliage suggested
this tribe. The genus certainly deserves separate tribal recognition;
however, such recognition would only compound problemd of gen-
erating other artificial taxonomic groups to accomodate the remain-
der Erythrininae, and in no way would clarify the puzzling status of
Erythrina, already a widely recognized difficulty."
Erythrina is grouped with eight other genera in the subtribe Ery-
thrininae (Strogylodon, Mucuna, Butea, SpatholobuG, Apios,
Cochlianthus, Rhodopis, and Neurudolphia). But the differences
between these genera and Erythrina are large and for that reason,
Raven (1982) proposed that it could be segregated as a monogeneric
tribe.
Erythrina is subdivided into four subgenera: Micropteryx, Ery-
thrina, Chirocalyx, and Erythraster, and further into 26 sections
(Krukoff, 1982). Most species are trees or shrubs, deciduous, with
leaves odd-pinnate, trifoliate, alternate, with glandular stipels on
the petiole (Cook, 1901; Dwyer and D'Arcy, 1980). The stem and
branches are usually armed with stout spines, and sometimes spines
are present also on the rachis. Flowers occur in clusters, in general
located near the apical ends, in lateral inflorescenses, although the
genus exhibits a great diversity in floral structure and inflorescence
orientation (Neill, 1984). Almost all Erythrina species have red-
dish, scarlet, or orange flowers, produce nectars and are adapted to
pollination by avian pollinators, passerines and hummingbirds
(Feinsinger et al., 1979; Neill, 1984). The fruit is a slender two-
valved pod, more or less moniliform, generally dehiscent, contain-
ing several seeds. The seeds contain alkaloids (Folkers and Unna,
1939) and are used medicinally (Duke, 1972).
According to Lewis (1974) the species of this genus have 21 pairs
of chromosomes (2n = 42). Erythrina species have been known as
having curare-like alkaloids (Folkers and Unna, 1939). The genus
Erythrina includes species that are known to bear root nodules (Al-
len and Allen, 1981). These nodules are formed after the infection
of the roots by compatible strains of the soil bacteria Rhizobium,
92 JOURNAL OF SUSTAINABLE AGRICULTURE

and provide Erythrina with the ability to fix atmospheric nitrogen.

Biological nitrogen fixation (BNF), is a highly desirable trait in any
plant as it allows self sufficiency in nitrogen supply. Furthermore,
Erythrina's BNF capability possibly contributes to its rapid growth
and to success in coppicing. Capability for nitrate reduction is an-
other remarkable aspect of Erythrina. It possesses a reduction sys-
tem with a high activity and a low affinity for nitrate reductase, thus
differing significantly from other higher plants (Stewart and Ore-
bamjo, 1979; Orebamjo, Porteous and Stewart, 1982). All of the
Erythrina species studied had in vivo activities of nitrate reductase
in excess of 100 micromole N02hourIg fresh weight, and the en-
zyme was found to use equally well both NADH and NADPH, as
electron donors. The above rates are 10-100 times those reported
for other angiosperms and nitrate reductase from angiosperms that
preferentially use NADH rather than NADPH as the electron donor
(Orebamjo et al., 1982). This unusual nitrate reducing system ap-
pears to be restricted to the genus Erythrina and may be of taxo-
nomic importance. The leaves of various species of Erythrina are
commonly used for fodder. It is a common sight to see people prun-
ing the branches along the living fences mainly in some Central
American countries. In some cases the flowers are eaten by people,
cooked in soups and salads (Martin and Ruberte, 1975).

MAIN SPECIES USED IN AGROFORESTRY

Etythrina poeppigiana (Walpers) O.F. Cook
Its natural distribution ranges from Bolivia to Panama, although
it has proven to be well adapted in other tropical areas where it has
become naturalized (Raven, 1974). Such areas include Central
America, the Caribbean, Africa, and Malaysia, where it was intro-
duced as shade for coffee and cacao (National Academy of Sci-
ences, 1979). The tree is found throughout the lowlands and middle
elevations in the tropical zones with medium to high rainfall (1,500-
3,000 mm/year and more) with a dry season of 0-6 months. Altitude
incosta Rica ranges from sea level to about 1,400 meters (Bor-
chert, 1980); however, in Colombia and Venezuela trees have been
 Research, Reviews, Practices and Technology 93

observed at or slightly above 1,900 meters. This species has long

been known in agroforestry. Early in this century Cook (1901) re-
ported its use as a shade tree in coffee culture in Puerto Rico. It is
known with different common names in different countries such as
"por6 gigante" in Costa Rica, "pito extranjero" in Guatemala,
"clmbulo" or "barbatusco" in Colombia, "bucare" or "ca-
chimbo" in Venezuela, "amasisa" in Peru, and, usually known as
"coral tree" or "immortelle" in English (Cook and Collins, 1903;
Stanley and Steyermark, 1947; Holdridge and Poveda, 1975). Evi-
dence for benefits of Eythrina as opposed to a non-legume tree
Cordia alliodora for shade of cacao was reported in Turrialba,
Costa Rica. Under Erythrina, cacao yields were, on average, 46%
greater than under Cordia during the first four years of the study
(Enriquez,l983) and 23% over a total of seven years of data
(Heuveldop et al., 1988). Litterfall data from this study (Alpizar et
al., 1983; 1986; Heuveldop et al., 1988) indicates that the rates of
cycling for calcium, magnesium and particularly nitrogen through
the cacao litter pathway are greater under Eythrina than under Cor-
dia. Nitrogen fixation (NF) has been little investigated. Escalante,
Herrera and Aranguren (1984) and Lindblad and Russo (1986)
found NF to be around 12-40 kg/ha/year when planted for shade in
coffee or cacao plantations. The tree is also a good source of green
manure; for instance, biomass production from pollarding (lopping
all branches) twice a year provided almost 12 tonsiha/year (Russo
sand Budowski, 1986). Initial experiences at CATIE, Turrialba,
Costa Rica showed promising results for alley cropping (Figure I),
which appears to offer considerable potential as a nitrogen source
for annual crops, since biomass production from Eythrina was
over 8,000 kglha of dry matter contained more than 3% N (Kass,
Russo and Quinlan, 1983; Kass and Barrantes, 1984). E. poeppi-
giana has been also reported in association with grasses (Figure 2)
Cynodon plectostachyus (Bronstein, 1984) and Pennisetum purpu-
mum (Benavides, 1983, 1985; Rodriguez, 1984). In both cases,
grass showed greater production with than without Eythrina. (Fig-
ure 3).
Farmers usually propagate the species through large cuttings, 2.5
94 JOURNAL OF SUSTAINABLE AGRlCULTURE

meters long and 8-12 cm in diameter. Within the first month sprouts
occur (Figure 4) and in just 4-6 months shade the coffee seedlings
(Figure 5, 6). Rooting success varies from 70-90% (Russo, 1984).
When established, total pollarding once or twice a year (Figure 7,
8) is usuqlly practiced and the branches are spread on the ground.
Foliage has been fed to small animals such as rabbits and goats with
good result$, h a v e s contain between 25-30% of crude protein (to-
tal N x 6,25) and are readily eaten by cattle. Digestibility varies
between 50-80% being highest for the bark of young branches.
Swine apparently suffer loss of hair after ingestion.
Because of its fast growth, its capability to produce high amounts
of biomass, its NF capability, its easy propagation by cuttings, its
excellent response to pruning practices, its high content of crude
protein, this tree shows a high potential for various agroforestry
practices beyond use as a coffee or cacao shade tree.

FIGURE 1 . Eythrina poeppigiana large cuttings planted in association with a

maize crop in CATIE, Turrialba, C. Rica.
 Research, Reviews, Practices and Technology 95

FIGURE 2. Pollard Erythrina poeppigiana trees associated with pasture in a farm

of Santa Cruz of Turrialba, Costa Rica (Photos by R. 0.Russo).

Erythrina hsca Loureim (E. glauca Willd)

Another Erythrina species mentioned by Holdridge (1955) and

by Hardy (1961) is E. fissca (best known by its former name E.
glouca). It is reported as originating in S.E. Asia, and it is widely
planted in cocoa farms in Comalcalco, Mexico and Brazil. It also is
used as shade for coffee in some farms in the Central Valley of
Costa Rica (Figure 9). According to Feinsinger et al. (1979) its
.natural reproduction occurs in wet soils such as those in swamps,
badly drained soils or along streams. This also had been observed
previously by Holdridge and Poveda (1975), who reported pure
stands in fresh water swamps in the Pacific lowlands of Costa Rica.
Cadima Zeballos and Alvim (1967) observed that the cacao trees
found around the the E. glauca trees produced more than the cacao
further away from the shade trees. They attributed such differences
to the influence that this species could have on some edaphological
factors related to cacao production. Santana and Cabala-Rosand
(1982) also reported that in a cacao plantation under trees of this
species, higher amounts of nitrogen available to cacao trees were
found. They attributed this to Erythrina litter, which has a high N
content. They also reported that in cacao planted under Erythrina,
 Research, Reviews, Practices and Technology 97

FIGURE 4 . Etythrina poeppigiana, a month after planting, sprouting vigorously

in Turrialba, C. Rica.

total N inputs are higher than N removed by a harvest of 1500 kgha

dried cacao beans.
Erythrina bertemana Urban
Its natural distribution ranges from southern Mexico to Costa
Rica in Central America, and probably to South America whether
or not it is the same species as E. mbrinervia H.B.K. (Holdridge
and Poveda, 1975). Its common name is "por6 de cerca" in Costa
Rica, "machete" in Jamaica, "elequeme" in Nicaragua, "gal-
lito," "pernilla de casa" in Panama, "pito" in Colombia, and
"parsu" in Cuna language. The Colombian name "pito" reflects a
children's use of the plant; the corolla of its flower, placed in a
hollow leaf stalk, serves as a whistle (Duke, 1972).
Sanchez and Russo (1985) reported it is the species most fre-
quently (50%) used as live fence posts (Figure 10) in Costa Rica,
according to results of a survey by questionnaries.
98 JOURNAL OF SUSTAINABLE AGRICULTUM

FIGURE 5. Erythrina poeppigiana. 4-6 months after planting, in a new coffee

crop in CATIE, Turrialba, C. Rica (Photos by R. 0.Russo).

Eiythrina berteroana responds to regular pruning with a great

volume of large-leafed shoots in only 3 weeks to 1 month. These
shoots are often used for rabbit feed and are also palatable to live-
stock. Goats are particularly eager to eat these shoots. Biomass pro-
duction of 1 kilometer of live fence of the species varies according
to pruning frequency, as presented in Figure 11.

Other Species of Erythrina

Eythrina costaricensis Micheli (E. steyermarkii Krukofl: This is

another usual live fence species like E. berteroana. Its leaflets are
wider than 10 cm. Its common name is "por6 cimarron" or "poro
de cerca." It is easily propagated by large cuttings or seeds. It is
endemic to Costa Rica, and the species can be usually found in
areas of abundant rainfall (above 1,500 mm). Holdridge and Pov-
eda (1975) consider that E. chiriquensis Krukoff, E. globocalk
FIGURE 6. Etytlrrinn Poeppigiana trees after pollarding, in a coffee crop in Turrialba, C. Rica.
100 JOURNAL OF SUSTAINABLE AGRICULTURE

FIGURE 7. A view of a recently pollard Erythrina poeppigiana tree.

Porsch, and E. steyermarkii Krukoff and Barneby could be syno-

nyms of E. costaricensis.
Eythrina cochleata Standley: This species, also known as poro,
is used in living fences. It is easily distinguished from another Ety-
thrinas because its leaflets are very shiny. Another related species
used as live fence posts in Costa Rica is E. lanceolata Standley
(leaflets smaller than 10 crn width).
Erythrina edulis (Micheli) Triana: This is the only Erythrina spe-
cies with edible seeds. All other species have poisonous seeds, that
are often used in collars or necklaces. Its common name is "balli"
or "basul," "cachafruto" or "poroto." The protein content of the
"balb" seeds varies between 18-21% and it has similar or higher
amounts of most amino-acids compared, to other leguminosae
(Perez et a l . , 1979).
 Research, Reviews, Practices and Technology 101

FIGURE 8. A farmer chopping the Eyrhrina branches, after cutting them down
(Photos by R.O.Russo).

Erythrina americantl:' This species mentioned by Martinez

(1959) is common to the uplands regions of Mexico, but also grows
well at sea level. Its flowers are highly prized for their flavor and
are an additional source of income for the small farmer who usually
sells the flowers in the local market. This author also mentions
other eleven Mexican species of Erythrina: E. breviflora, E. setosa,
E. leptorhiza, E. rubrinervia, E. montanal, E. lanata, E. occiden-
talis, E. flabellifomis, E. foldmanii, E. maicana, and E. herba-
cea, also found in southern United States.
Elythrina lithospenna: This species is mentioned by Urquhart
(1963) as a shade tree for cacao in Western Samoa. It has also been
102 JOURNAL OF SUSTAINABLE AGRICULTURE

FIGURE 9. A view of recently pollard Evthrina ficsca trees shading a coffee

plantation in Tres Rios, Costa Rica.

reported as a shade tree for arabica coffee in India (Achyuta, 1960).

Like E. poeppigiana it is also pruned twice a year. E. lithosperma is
usually planted before cacao seedlings are placed in the field. One
of the advantages reported for this species is that there are some
thornless varieties.
Erythrina umbrosa: This species has been mentioned by Flye
(1945) as a shade tree for coffee plantations in Brazil.

DISCUSSION
The greater potential for Erythrina spp. use lies in the develop-
ment of agroforestry systems. But, all tree species associated with
crops in agroforestry, have advantages and disadvantages. A good
place to start is to recognize or mention cases of successful agro-
forestry systems. A good example of success throughout the years
is E. poeppigiana which is associated with the coffee crop in Costa
 Research, Reviews, Practices and Technology 103

FIGURE 10. An Erythrina berteroana post in a living fence in La Suiza of Tur-

rialba, Costa Rica (Photos by R.O. Russo).

Rica. Another case is the common use of E. berteroana and E.

costaricensis in live fence posts.
Characteristics, Uses, and Potential
of the Genus Erythrina
The following characteristics make this genus particularly attrac-
tive for use in agroforestry: (a) Members are fast growing, and rap-
idly produce considerable amounts of biomass rich in nitrogen; (b)
It is easily propagated by cuttings; (c) It has excellent capabilities to
withstand regular pruning, subsequent rapid sprouting and develop-
FIGURE 11. Production of a live fence of E~ythrinaberferoana pruned every 4, 6, and 8
months in CARE, Turrialba. Costa Rica (Calculated and redrawn from Budowski, Russo
and Mora, 1985)

I
1 I I
4 monlhs 6 months 8 months

frequency of prunlng
 Research,>Reviews, Practices and Technology 105

ment of strong shoots; (d) Its leaves are edible by cattle and rabbits,
and are rich in protein (25-30 % of crude protein content; these
figures are higher than those found in the conventional protein sup-
plements available in the market for cattle, which are usually
around 18%.); (e) The roots are generally associated with the nitro-
gen-fixing bacteria Rhizobium, whose root-nodules have the capa-
bility to fix atmospheric nitrogen, which is a highly desirable trait in
any plant as it allows self sufficiency in nitrogen supply; ( f ) Most of
the species of this genus are adequate to be used in living fences.
The list, of course, could be much longer. I personally inter-
viewed many farmers, between 1982 and 1984, when working for
the CATIEIIDRC-supported Erythrina Project. A high percentage
of farmers agree that "poro is good," which can be interpreted as
evidence that they like the genus because of its beneficial effects.
We should consider whether the genus may have potential to be
used more extensively in agroforestry systems in other countries,
with ecological conditions similar to those where it is currently
used. In Costa Rica, for instance, the use of Erythrina for shading
or nursing other crops is a common agricultural practice in both
coffee and cacao plantations. There is a great deal of evidence
showing its value as a "natural fertilizer" supplier and nutrient cy-
cling helper. The calculated figures show that the return of nitrogen
to the soil and nutrient cycle in coffee, cacao, and also in maize,
can save up to 200 kg Nlha per year.
A considerable research effort in working with this genus has
been done in the Tropical Agricultural Center for Research and
Training (CATIE), Turrialba, Costa Rica through the Evthrina
Project (Centro Agrondmico Tropical de Investigacidn y En-
seiianza, 1986b). This research project supported by the Interna-
tional Development Research Center (IDRC) from Canada, pro-
duced a large amount of research and also compiled a substantial
bibliography on the genus (Centro Agron6mico Tropical de Investi-
gaci6n y Enseiianza, 1986a).
Finally, no doubt agroforestry activities\tend to increase the level
of system complexity. This makes the management of the farm
more difficult. However, if professionals' in forestry and agriculture
aim at the solutions of practical problems, then it is necessary to
deal with the realities that the farmer faces. We cannot sacrifice
106 JOURNAL OF SUSTAINABLE AGRICULTURE

potential benefits for fear of making the problem a more complex

one.

BIBLIOGRAPHY
Achyuta, R.Y.R. 1969. Shade tree for arabica coffee: Erythrina lithospenna.
Indian Coffee 24(12):500-505.
Allen, O.N. and A.K. Allen. 1981. Leguminosae, a source book of characteris-
tics, uses, and nodulation. Madison, Wisconsin, USA, University of Wiscon-
sin Press. 212 p.
Alpizar, L.A., H.W. Fassbender and J. Heuveldop. 1983. Estudio de sistemas
agroforestales en el Experiment0 Central del CATIE, Turrialba. 111. Produc-
ci6n de residuos vegetales. CATIE, Turrialba, Costa Rica. 13 p. (mimeo).
Alpizar, L., H.W. Fassbender, J. Heuveldop, H. Folster and G. Enriquez. 1986.
Modelling agroforestry systems cacao (Theobroma cacao), with laurel (Cordia
alliodora), and poro (Eythrina poeppigiana) in Costa Rica. I. Inventory of
organic matter and nutrients. Agroforestry Systems 4(3):175-190.
Benavides, J.E. 1983. Investigacidn en arbales forrajeros. In Curso Corto Inten-
sivo sobre Tbcnicas Agroforestales Tradicionales. CATIE, Turrialba, Costa
Rica, 8-18 Nov., 1983. 27 p.
Benavides, J.E. 1985. Produccidn y calidad nutritiva del forraje de pasto king-
grass (Penniseturn putpureurn x P. typhoides) y por6 (Erythrina poeppigiana)
sembrados en asociaci6n. CATIE, Turrialba, Costa Rica. 37 p. (mimeo).
Borchert, R. 1980. Phenology and ecophysiology of tropical trees: Etythrina
poeppigiana O.F. Cook. Ecology 61(5):1065-1074.
Bronstein, G.E. 1984. Producci6n comparadade una pastura de Cynodon plectos-
tachyus asociada con Arboles de Cordia alliodora, asociada con Irboles de
Erythn'na poeppigiana y sin &boles. M.Sc. Thesis UCR-CATIE, Turrialba,
Costa Rica. 109 p.
Budowski, G . , R.O. Russo and E. Mora. 1985. Productividad de una cerca viva
de Erythrina betiemana Urban en Turrialba, Costa Rica. Turrialba 35(1):83-
86.
Cadima Zeballos, A. and P. de T. Alvim. 1967. Influencia del Brbol de sombra
Erythrina glauca sobre algunos factores edafoldgicos relacionados con la pro-
ducci6n del cacaotero. Turrialba 17(3):330-336.
Centro Agronbmico Tropical de Investigaci6n y Ensefianza. 1986a Bibliography
on the genus Eythrina. CATIE, Renewable Natural Resources Department,
Turrialba, Costa Rica. 182 p.
Centro AgronBmico Tropical de Investigacidn y Ensefianza. 1986b. Final project
technical report: Erythrina spp. (phae 1 : 3-p-82-0015). CATIE, Turrialba,
Costa Rica. 182 p.
Cook, O.F. 1901. Shade in coffee culture. U.S. Department of Agriculture, Divi-
sion of Botany, Washington, D.C. 79 p.
 Research, Reviews, Practices and Technology 107

Cook, O.F. and G.N. Collins. 1903. Economic plants of Porto Rico. In Contribu-
tions from the US. National Herbarium 8(2):57-269.
Duke, J.A. 1972. Isthmian etnobotanical dictionary. Fulton, Maryland, J.A.
Duke. 96 p.
Dwyer, J. and W.G. D'Arcy. 1980. Eythrina. In Flora of Panama. Part V. An-
nals of the Missouri Botanical Garden 67(3):686-697.
Enriquez, G. 1983. Breve reslimen de 10s resultados del Experiment0 Central de
plantas perennes de La Montafia, CATIE, Turrialba, Costa Rica. Agroforestry
Short Course 1983. CATIE, Turrialba, Costa Rica. 13 p. (mimeo).
Escalante, G., R. Herrera and J. Aranguren. 1984. Fijaci6n de nitr6geno en hr-
boles de sombra (Erythrina poeppigiana) en cacaotales del norte de Venezuela.
Pesquisa Agropecuaria Brasileira 19(suppl.):223-230.
Fassbender, H.W., L. Alpizar, J. Heuveldop, H. Folster and G. Enriquez. 1986.
Modelling agroforestry systems cacao (Theobroma cacao), with laurel (Cordia
alliodora), and poro (Etythrina poeppigiana) in Costa Rica. 111. Cycles of
organic matter and nutrients. Agroforestry Systems 6(1):49-62.
Feinsinger, P., Y.B. Linhart, L A . Swarm and J.A. Wolfe. 1979. Aspects of the
pollination biology of three Etythrina species on Trinidad and Tobago. Annals
of the Missouri Botanical Graden 66(3):451-471.
Folkers, K. and K. Unna. 1939. Etythrina alkaloids. V. Comparative curare-like
potencies of species of the genus Eryfhrina. J. Am. Pharm. Assoc. 28:1019-
1028.
Flye, W.D. 1945. Erylhrina umbrosa e a sombra "inbmortal" para e cafeiro.
Boletin Superintendencia dos Servicios do Cafe (Brazil) 20:878-879.
Glover, N. and J. Beer. 1984. Spatial and temporal fluctuations of litterfall in
agroforestry associations Coffea ambica var. Caturra-Eythrina poeppigiana
and C. arabica var. Caturra-E. poeppigiano-Cordia alliodora. CATIE, Tur-
rialba, Costa Rica. 43 p. (mimeograph).
Glover, N. and J. Beer. 1986. Nutrient cycling in two traditional Central Ameri-
can agroforestry systems. Agroforestry Systems 4(2):77-88.
Hardy, F. 1961. Manual de cacao. Turrialba, Costa Rica, IICA. p. 28.
Heuveldop, I., H.W. Fassbender, L. Alpizar, G. Enriquez and H. Folster. 1986.
Modelling agroforestry systems cacao (Theobroma cacao), with laurel (Cordia
alliodora), and poro (Elythrina poeppigiana) in Costa Rica. 11. Cacao and
wood production, litter and decomposition. Agroforestry Systems 6(1):37-48.
Holdridge, L.R. 1955. La agricultura y la dasonomia en Costa Rica: Competencia
o coexistencia. Comunicaciones en Turrialba no. 56. Setiembre 1955.
Holdridge, L.R. 1967. Arboles de sombra para el cacao. In Manual del Curso de
Cacao. IICA, Turrialba, Costa Rica. pp. 113-117.
Holdridge, L.R. and L. Poveda, 1975. Arboles de Costa Rica. v.1. San Jost,
Costa Rica, Centro Cientifico Tropical. 546 p.
Kass, D.C.L., R.O. Russo and M.M. Quinlan, 1983. Leguminous trees as nitro-
gen source for annual crops. Agronomy Abstracts 1983. p. 45.
Kass, D.C.L. and A. Banantes. 1984. Leguminous trees as nitrogen source for
annual crops. CATIE, Turrialba, Costa Rica. 31 p. (mimeo).
108 JOURNAL OF SUSTAINABLE AGRICULTURE

Krukoff, B.A. 1939. The American species of Etythrina. Brittonia 3:205-239.

Krukoff, B.A. 1979. Notes on the species of Erythrina. 111. Annals of the Mis-
souri Botanical Garden 66(3):422-445.
Krukoff, B.A. 1982. Notes on the species of Erythrina. XVIII. In Erythrina Sym-
posium IV. Allertonia 3(1):121-154.
Lackey, J.A. 1981. Phaseolaceae. In Polhill, R.M. and,P. Raven (eds.) Advances
in legume systematics. Royal Botanic Gradens, Kew. pp. 301-327.
Lewis, W.H. 1974. Chromosomes and phylogeny of Etythrina (Fabaceae).
Lloydia 37:460-464.
Lindblad, P. and R.O. Russo. 1986. cH,-reduction by Etythrina poeppigiana in
a Costa Rican coffee plantation. Agroforestry Systems 4:33-37.
Martin, F.W. and R.M. Ruberte. 1975. Edible leaves of the tropics. Mayaguez
Institute of Tropical Agriculture, A.R.S., U.S.D.A., Mayaguez, Puerto Rico.
pp. 59-60.
Martinez, M. 1959. Plantas 5tiles de la flora mexicana. Botas, Mexico, D.F.
p. 159.
Neill, D.A. 1984. Experimental studies on species relationships in Etythrina (Le-
guminosae: Papilionoideae). Ph.D. Dissertation, Washington University, St.
Louis, Missouri. 226 p.
Orebamjo, T.O., G. Porteous and G.R. Stewart. 1982. Nitrate reductase in the
genus Etythrina. In Erythnna Symposium IV. Allertonia 3(1):11-18.
PCrez, G . , C. de Martinez and E. Diaz. 1979. Evaluacibn de la calidad de la
proteina de la Etythrina edulis (ba15). Archivos Latinoamericanos de Nutri-
cibn 29(2):193-207.
Raven, P.H. 1974. Etythrina (Fabaceae): Achievements and opportunities.
Lloydia 37:321-331.
Raven, P.H. 1982. Erythrina (Fabaceae: Faboideae): Introduction to Symposium
IV. Allertonia 3(1):1-6.
Record, S.J. and R.W. Hess. 1947. Timbers of the new world. New Haven, Yale
University Press. 640 p.
Rodriguez, R.A. 1984. Produccibn de biomasa de porb gigante (Erythrina poep-
pigiana (Walpers) O.F. Cook y King grass (Pennisetum purpureum x P.
typhoides) intercalados en funcibn de la siembra y frecuencia de poda del porb.
MAC. Thesis UCR-CATIE, Turrialba, Costa Rica. 96 p.
Russo, R.O. 1984. Studies on Etyrhrina poeppigiana (Walpers) O.F. Cook, a
versatile tree in Costa Rican farms. Ph.D. Dissertation Southeastern Univ.,
New Orleans, LA. 145 p.
Russo, R.O. and G . Budowski. 1986. Effect of pollarding frequency on biomass
of Erythrina poeppigiona as a coffee shade tree. Agroforestry Systems
4(2):145-162.
Sanchez, G.A. and R.O. Russo. 1985. Biomass production of Erythrina spp.
(Papilionaceae) in the humid tropics. Paper presented to the Conference "Re-
search in Forestry for Energy." Copenhague, Denmark, October 28 - Novem-
ber 11, 1985.
 Research, Reviews, Practices and Technology I09

Santana, M.B.M. and P. Cabala-Rosand. 1982. Dynamics o f nitrogen in a shaded

cacao plantation. Plant and Soil 67:271-281.
Stanley, P.C. and J.A. Steyermark. 1947. Flora o f Guatemala. Fieldiana: Botany,
Chicago Natural History Museum 24(5): 258-260.
Stewart, G.R. and T.O. Orebamjo. 1979. Some unusual characteristics o f nitrate
reduction in Elythrina senegale& D.C. New Phytologist 83:311-319.
Workshop Agro-Forestry Systems in Latin America, Proceedings. March 26-30,
1979. Edited by G. de las Salas. CATIE, Tu~~ialba, Costa Rica. p. 211.

RECEIVED: 12/14/89
REVISED: 1/23/90
ACCEPTED: 1/26/90

for laculfy/prolessionals wilh journal subscription recomrnendalion

..
authorily lor their inslilulionallibrary.
Ifyou have read a reprint or photocopy of this article, would
you like to make sure that your library also subscribes lo
this journal? If you have the aulhority lo recommend sub-
scriptions lo your library, we will send you a tree sample
copy lor review with your librarian. Just fill oul the form below-and make
sure that you type or wrlte out clearly both the name of the lournel and
your own name and address.
'
( ) Yes, please send me a complimentary sample copy of this journal:

(please write in complete journal title here-do not leave blank)

I will show lhis journal to our instilulional or agency library lor a possible
subscriplion.
The name of my instilutional/agencylibrary is:

INSTITUTION:
ADDRESS:
CITY: STATE. ZIP:
Retbrn lo Sample Copy Department. The Haworln Press. Inc.,
10 Alce Street. B~nghamton.NY 13904-1580