Tetra Gas Tris

The Genus Tetragastris and the Forests of Eastern Brazil: Studies in Neotropical Burseraceae
III
Author(s): Douglas C. Daly
Source: Kew Bulletin, Vol. 45, No. 1 (1990), pp. 179-194
Published by: Springer on behalf of Royal Botanic Gardens, Kew
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The genus Tetragastrisand the forests of
eastern Brazil
Studies in neotropical Burseraceae III*
DOUGLASC. DALY+
Gaertn., a genus of nine species, is represented by three well-defined species

Summary.Tetragastris
in easternBrazil.All of themare restrictedin distribution,and two are in dangerof extinction.
T. catuaba,lost since it was based on an apparently misplaced collection, is rediscovered and a
neotypeis designated;it figuresin a particularlyconfusingdebatein Brazilianethnobotany.
T. occhionii,the most archaic species in the genus, is endemic to the coastal dune vegetation of
is restrictedto the steepforestedmountainsin andaroundthecityof Rio
Bahia.T. breviacuminata
deJaneiro.
EASTERN BRAZIL
In the context of this paper, eastern Brazil is defined as the coastal regions,
including the Serra do Mar, of the states of Bahia, Espirito Santo, Rio de
Janeiro, and Sao Paulo. I have therefore included parts of both southeastern
and northeastern Brazil as traditionally defined (e.g., Andrade-Lima 1982).
The vegetation of this region is part of the Complexo Mata Atlantica; as
defined in the research program of the Jardim Botanico do Rio de Janeiro
(1988), this includes the vegetation that extended from Rio Grande do Norte
to Rio Grande do Sul along the Atlantic coast in a once-continuous band up
to 150 km wide in places. Although the Complexo Mata Atlantica includes
moist forest, semi-deciduous forest, restinga, cerrado, and other vegetation
types, in the discussion that follows I will focus only on moist forest and
restinga vegetation.
The flora of eastern Brazil is one of the richest in the world. It is an ancient
one, as significant portions of the region were forested since the Tertiary.
This flora is remarkable for its high degree of endemism-even at the generic
level-and for the high number of archaic taxa in a number of plant families
(e.g., Soderstrom & Calder6n 1974; Kubitzki 1975; Mori et al. 1981; Daly
1989).
As early as 1831, Auguste de Saint-Hilaire expressed concern about the
future of the Atlantic forests of Brazil (Fons&ca1985). The populations of the
economically important pau brasil, CaesalpiniaechinataLam., were decimated
already by 1809 (see Mori et al. 1983). For centuries, these forests have been
destroyed or disturbed for such purposes as timber; sugar cane or other
crops; plantations of cacao, oil palm, coffee, and other aborescent plants;
firewood; charcoal; fuel for sugar cane processing or other factories; pasture;
and others. Perhaps only 4% of the original vegetation of the Complexo
Mata Atlantica remains undisturbed (Jardim Botanico do Rio de Janeiro
AcceptedforpublicationMarch1989.
*ContinuedfromBrittonia41:17-27(1989).
+NewYorkBotanicalGarden,Bronx,NY 10458,USA.
179

180 KEW BULLETIN VOL. 45(1)
TABLE1. Distributions of the species of Tetragastris
species distribution
T. altissima(Aublet) Brazil Acre, Amazonas,
Swart Ampt,
Maranhao, Mato
Grosso, Para, Rond6nia
Boliva Pando
Ecuador Napo
French Guiana
Guyana
Peru Madre de Dios
Surinam
Venezuela Bolivar, Delta Amacuro,
Monagas
T. balsamifera(Sw.) Dominican
Oken Republic
Haiti
Puerto Rico
T. breviacuminata
Swart Brazil Rio de Janeiro
T. catuabaCunha Brazil Bahia, Pernambuco
T. hostmannii(Engl.) Brazil Amapai,northern Para"
Kuntze
French Guiana
Guyana
Surinam
Venezuela northern Bolivar
1988); estimates vary, but I have not seen a figure over ten per cent. Only a
small part of the remaining forest is effectively protected, and many of the
reserves that do exist contain only small proportions of primary vegetation
(Mori et al. 1981). It is probable that many of the patches of remaining
primary forest are too small to maintain their integrity as ecosystems, that is,
to sustain their full complement of plant and animal species.
There is an urgent need for taxonomic studies of the plant groups occur-
ring in eastern Brazil. They must analyze overall distributions, pinpoint
centers of endemism, and locate surviving populations. Botanists interested
in the region need to encourage such efforts, to monitor results as they are
published, and to seek congruences in the distribution patterns of endemics.
This information can be used to indicate appropriate localities for reserves
and to argue for their protection. Unfortunately, very little useful work of this
type has been published. A notable exception is that of Lewis (1987), which
provides a great deal of baseline data on the legumes of eastern Bahia. The
work that follows is a modest contribution toward these goals.
TetragastrisGaertn.
The Burseraceae can be studied very profitably in the context of regional
floras in the Neotropics for two reasons. First, Burseraceae
can usually be used
as indicators of primary forest, because few species occur in secondary

TETRAGASTRIS FROM E. BRAZIL (BURSERACEAE III) 181
TABLE1 (contd.)
species distribution
T. mucronata
(Rusby) Swart Trinidad (tentatively) Apure,
Venezuela Aragua, M6rida,
Miranda, Tachira,
Zulia
T. occhionii(Rizzini) Daly Brazil Bahia
T.panamensis(Engl.) Belize
Kuntze Bolivia Beni, La Paz, Pando
Brazil Amapi, Amazonas,
Maranhao, Mato
Grosso, Pardi,Rond6nia,
Roraima
Colombia Antioquia
Costa Rica
Ecuador Napo, Pastaza
French Guiana
Guatemala
Guyana
Honduras
Nicaragua
Peru Junin, Loreto, Madre de
Dios, San Martin
Surinam
Venezuela Apure, Bolivar, Delta
Amacuro, Merida,
Miranda, Zulia
T. sp. nov. Brazil Distrito Federal, Goids,
southern Maranhio,
Mato Grosso
vegetation. Second, this resiniferousfamily usually figures prominently in the

ethnobotany of the regions where it occurs.
The genus Tetragastrisbelongs to the tribe Protieaeand is closely related to
ProtiumBurm.f. The limits of the two genera were redefined recently (Daly
1989). Tetragastrisis found almost exclusively in lowland moist forests in
Central America, northern South America, and parts of the Caribbean. The
distributions of the nine species recognized here are summarized in Table 1,
and taxonomic synonyms are listed in Table 2. The species with the greatest
range is T. panamensis,extending from Belize and Guatemala through much
of northern South America. Over much of its range it is sympatric with
T. altissima;their niche partitioning is not yet understood. In the cerrado
vegetation of central Brazil, they are replaced by an undescribed taxon not
easily distinguished from T. altissima. In part of the lowlands of northern
Venezuela, they are replaced by T. mucronata, which also resembles T. altissi-
ma. Tetragastris
balsamiferais restricted to Hispaniola and Puerto Rico. T. host-
manniiis essentially a Guianan species, although it has also been collected in
Amapai (Brazil), northern Pard (Brazil), and northern Bolivar (Venezuela).

TABLE2. Synonymies of names of Tetragastris
not accepted here
name synonym of
T. balsamifera(Swartz) Oken var.
lancifoliaSwart T. balsamifera
T. osseaGaertn. T. balsamifera
T.panamensis(Engl.) Kuntze var.
grandifoliaSwart T. panamensis
T. panamensis var. hirtellaSwart T. panamensis
T. paraensisCuatrec. T. panamensis
T. phanerosepala Sandw. T. altissima(Aublet) Swart
T. pilosaCuatrec. Protiumpilosum (Cuatrec.) Daly
T. stevensoniiStandl. T. panamensis
T. tomentosa D. Porter (Rose) Engl.
Protiumpittierii
T. trifoliolata(Engl.) Cuatrec. Protiumtrifoliolatum
Engl.
T. unifoliolatum(Engl.) Cuatrec. ProtiumunifoliolatumEngl.
For over four decades, only one species of Tetragastris,

T. breviacuminata,
was
known to occur in eastern Brazil. Recently, however, ProtiumocchioniiRizzini
was transferred to Tetragastris(Daly 1989), and enough evidence has
accumulated from herbarium specimens, field work, and the literature to
confirm the existence of T. catuaba.Characters which can be used to disting-
uish these three species are presented in Table 3. Each of these three species
has a very limited distribution, and each is restricted to a particular vegeta-
tion type. Each is part of a unique flora threatened with wholesale extinction,
in a region where only isolated patches of the original vegetation remain.
Even those areas are being razed.
THE MYSTERYOF catuaba SOLVED-AGAIN
For well over a century, catuabahas been one of the best-known herbal
medicines in Brazil. It was a popular remedy in the mid-nineteenth century,
and by the early part of this century it was sold in the herbal markets of Sao
Paulo city (Hoehne 1920). Today, catuaba'wine' is sold on supermarket
shelves in many regions of the country; grape wine is actually the primary
ingredient, but the others are subject to doubt. Most of the catuaba wines are
made in Saio Paulo state. People in both the cities and the interior of north-
eastern Brazil, including Piaui (Emp6raire 1983), Bahia (G. Pinto, pers.
comm.), Pernambuco (F. G. Serpa, pers. comm.), and Paraiba (L. Xavier
Filho, pers. comm.), still utilize the actual plant as an aphrodisiac or tonic,
and for other purposes. Historically, it has been credited with curing some
rather unusual ailments, including neurasthenia, hypochondria, and nervous
insomnia (Cruz 1979).
The first reference to the meaning of catuabawas made in 1860 by Freire
Alemdo (quoted in Pereira 1978). He indicated that catuabais a compound
word in Tupi meaning 'homem valido', or 'true man'. Pio Correa (1931), on
the other hand, interpreted it as 'good leaf or 'good tree' ('arvore boa'). In
fact, the name catuabaprobably comes from lingua geral, a predominantly
Tupi-Guarani language developed by the Jesuits. It is a polymorphemic

TETRAGASTRIS FROM E BRAZIL (BURSERACEAE III) 183
TABLE 3. Characters separating the species of Tetragastris
occurring in eastern
Brazil
character* T. catuaba T. occhionii T. breviacuminata

distribution Bahia inland Bahia Rio de Janeiro
(sub-coastal)
habitat lowland forest restinga dunes forested slopes
height 15-22 m (0-8) 2-10 m 5-10 (21) m
leaflet
apex retuse to acumen short acumen short and
rounded and broad broad
margin revolute flat flat
flower
length (3-3) 3-5-4-5 2-6-3-1 mm 3-5-4-5 (5-0) mm
(5-0) mm
petals scattered, dense, appressed sparse, spreading
appressed hairs hairs to 2 mm hairs to 0-5 mm
to 2 mm long long long
fruit
colour
(mature) red-tinged green/brown green
size (dry)** 1-4-1-8 x 1-4- 0-8-1-0 x 0-5- 1-3-1-6 x 1-2-
1-5 cm 0-7 cm 1-3 cm
vesture glabrous brown-pubescent glabrous
cotyledons plano-convex U-shaped J-shaped
* measurements taken from dry specimens
**measurements
applyto dry fruitswith only one loculedeveloping(andthereforenot lobed)
descriptive term with two components. Katu unquestionably means 'good'.

The bound form for 'leaf' is -rdba;for 'tree' it is -iba;and for 'person' it is abd.
Though neither corresponds perfectly, it is most likely that 'good leaf is the
true meaning for catuaba.One of the secondary names given by Pio Correa is
catuiba,which would give the 'good tree' interpretation.*
Unfortunately, no one seems to be sure just what the plant is. Depending
on which source is consulted, catuabais a plant of the Erythroxylaceae, Bigno-
niaceae,Sapotaceae,Euphorbiaceae, Myrtaceae,Meliaceae,Apocynaceae, or Burser-
aceae.The issue has been complicated by two primary factors. First, aphrodi-
siacs, like snake bite remedies, always enjoy a certain mystique; consequent-
ly, they suffer from taxonomic ambiguity. Second, the diaspora of northeast-
ern Brazilians fleeing repeated catastrophic droughts over the centuries has
resulted in pulses of the distinct northeastern culture being spread through
other parts of the country. If indeed catuabaoriginally indicated a single
*I am indebted to W. Balie for the etymological analysis.

species, the displaced Northeasterners would seek substitutes, which they

might call by the same name, since folk terms for useful plants are often
synonymous with the products or materials obtained from them.
This dilemma has attracted the attention of many of Brazil's greatest
botanists over the years, among them Adolpho Ducke. His last paper (Ducke
1966), published posthumously, addressed the problem of identifying catuaba;
it is hoped that the frustration of this undertaking did not contribute to his
demise. The confusion surrounding its identity dramatically illustrates what
happens when publications on economic plants fail to cite evidence or refer-
ences, and when identifications are asserted without citing vouchers. The
result is an example of the Law of Monstrous Durability of Error (Gillett
1980): essentially, once mistakes are in the literature, it is very difficult to get
them out. For example, in the dictionary of useful Brazilian plants compiled
by Cruz (1979), he dutifully reiterated errors in the older literature stating
that catuabais Erythroxylum catuabaMart., and that there is a second species of
Erythroxylum found throughout northeastern Brazil and Amazonia which is so
common it forms dense forests. In fact E. catuabais a nomennudum,and none of
the various species identified as catuabashows that distribution, nor does any
of them form dense forests.
The great nineteenth century Brazilian botanist and physician Francisco
Freire Alemao Cisneiros, referred to in the literature as Freire Alem5ao,spent
two years in the northeastern state of Ceara, where he reportedly collected
20000 plants (Mello Leitao 1937). He clearly regarded catuabaas a species of
Erythroxylum, and waxed poetic about its properties:
'The wood of catuaba,by its sturdiness, serves as a staff for the brave; the
infirm fortify themselves with its bark. I mean to say that catuabais a
well-known aphrodisiac. In order to recognize the foundation for the belief
in its efficacy, we have the means of direct experience; it seems to me that
such an estimable plant is already unnecessary in this country.. . . One
makes a comparison... with the celebrated coca-a plant of the same
genus-of the libidinous Peruvian .... It is not, however, only for its
aphrodisiacal properties that one recommends the very bitter catuaba.It is
an appreciable tonic possessed, as I have just surmised, of curious prop-
erties; to an extent it could make up for the lack of guarand.It is appropri-
ate for those who are . .. feverish, or weak in the stomach. I would dare to
utilize it, in the form of a slow cordial, for malignant nervous fevers. And
turning to the virilizing force of catuaba, . . . it operates slowly, reconstitut-
ing, promoting life in those organs which have lost it through weak-
ness. ... This is surely one of the treasures of the materiamedicaof Cear~a,
and studying it will furnish beautiful pages in the history of our pharma-
cology.' (my translation of a quote in Pereira 1978).
Ducke spent the last years of his life in Fortaleza, Ceara, and he wrote
extensively about the Northeastern flora (Ducke 1959). In 1958, a collector
he sent to the Serra do Araripe in Ceardibrought back specimens of the plant
known locally as catuaba:Eythroxylumvaccinifolium Mart. (T. L. Guedes528,
NY!). In addition to Ceara, it occurs in Bahia, the Distrito Federal, Minas
Gerais, Rio deJaneiro, Sao Paulo, and Santa Catarina. Monteiro Silva (Silva
1951) also identified catuabaas an Erythroxylum,although he used the nomen
nudumE. catuaba(see below), and he presented rather detailed instructions

for preparation of tinctures and decoctions. Ducke (1959) noted that another
plant from the Serra do Araripe (T. L. Guedes639, NY!), known as catuaba
cipd,proved to be the apocynaceous Secondatiafloribunda A.DC. This species is
known from Ceara, Espirito Santo, Bahia, and Minas Gerais.
The late T. Plowman, specialist in the Erythroxylaceae, visited the Serra do
Araripe in the early 1980s, and he noted that the bark had been stripped
from all the adult individuals of Erythroxylum vaccinifolium.
In an ethnobotanical study of an area in the interior of the northeastern
state of Piaui, Emp6raire (1983) found that the plant locally called catuaba
brancais a Myrtaceae,while catuabapreta,or 'black catuaba',is an Erythroxylum.
(No collections are cited, nor is there any indication of where the specimens
are deposited, but she did note that herbarium material was identified by D.
de Andrade Lima and M. Ataide at the Instituto de Pesquisas Agronomicas
de Recife). The bark of the latter is smoked, burned ('brfile'), or consumed as
an infusion or a syrup. It is used as an aphrodisiac and a hallucinogen. The
infusion is also used to treat stomach aches. It is interesting to note that, like
Monteiro Silva (Silva 1951), she recorded that the wood of that catuabais
used to make handles for tools.
In 1983, M. J. Balick found that the Apinaj6 Indians of Goias state were
using a plant with the Portuguese name catuabaas a male aphrodisiac. Using
the bark, they drank a decoction or an alcohol infusion. Fertile vouchers of
this plant (M. J. Balick et al. 1613, NY!) were identified by T. Plowman as
Erythroxylum subracemosum Turcz., a species that extends down into southern
Mato Grosso, over to Rondonia, and up to the southern parts of Maranhdo
and Par-i.
Frederico Carlos Hoehne (1920) was also interested by catuaba.However,
the material he collected in the markets of SaioPaulo and Rio de Janeiro was
identified as Anemopaegma mirandum(Cham.) A.DC. [=A. arvense (Vell.)
Stellf.] (Bignoniaceae),although he did not indicate the origin of the samples or
how the determination was made. In a later work (Hoehne 1939), he noted
that the catuabawas made into decoctions and tinctures, and even used in the
form of a fumigant, as a 'stimulant for the internal organs'. In the herbarium
at NY, there is a collection of A. mirandum(M. G. Silva & A. Pinheiro4420)
made in 1979 near the junction of the Cuiabai-Santarem and Cuiabai-Porto
Velho highways in Mato Grosso. The label states that the common name
among the Parecis Indians is catuaba,that a tincture of the ground root is
used as an aphrodisiac, and that they had bundles of the roots for sale.
Pio Correa (1931), the great authority on Brazilian economic botany,
provided several possible identities for catuaba,although he agreed with
Hoehne that catuabaverdadeira,or 'true catuaba',is Anemopaegma mirandum.
Under catuaba,he recorded Anemopaegma glaucum Mart. ex DC. as an anti-
syphilitic, while he attributed tonic, stimulant, and aphrodisiacal properties
to PhyllanthusnobilisMuell. Arg. (=Margaritaria nobilis L.f). He identified
catuabadomatoas Ilex conocarpa Reiss., a species found in Minas Gerais and the
Distrito Federal, which purportedly has been used as a tonic and a diuretic,
and for stomach problems. He listed all the above uses-except as a diure-
tic-for A. mirandum.
Both Hoehne (1939) and Ducke (1966) wrote that Trichilia(Meliaceae)had
been identified with catuabain the literature, although I have not been able to
find where that idea originated. Hamet (1936), working with bark identified
as catuabaand provided by a Professor Goris, concluded from an anatomical

TABLE 4. Distributions of taxa known as catuaba*
taxon family distribution
Anemopaegma arvense Bignoniaceae Distrito Federal, Goias,

(including A. mirandum) Minas Gerais, Mato Grosso
do Sul, Sdo Paulo, Paraguay
A. glaucum western Bahia, Distrito
Federal, Goias, Minas
Gerais, southern Mato
Grosso
Erythroxylum Erythroxylaceae Goiats,southern Maranhio,
subracemosum southern Mato Grosso,
southern Para, Rond6nia
E. vaccinifolium western Bahia, Ceara,
Distrito Federal, Minas
Gerais, Rio deJaneiro,
Santa Catarina, Sdo Paulo
Ilex conocarpa Aquifoliaceae Distrito Federal, Minas
Gerais
Micropholissp. Sapotaceae Maranhio (and elsewhere?)
Secondatiafloribunda Apocynaceae western Bahia, Ceara,
Espirito Santo, Minas
Gerais
catuaba
Tetragastris Burseraceae eastern Bahia, Pernambuco
*Only those taxa are included which have been mentioned in the literature or noted on herbar-
ium specimens.
analysis of the material that it belonged to a plant in the Meliaceae,a family

closely related to the Burseraceaein the order Sapindales. However, inter-
pretation of the rather archaic French anatomical terminology shows that he
may have been describing resin canals, in which case the material could well
have been from a burseraceous plant (D. W. Stevenson, pers. comm.).
The occurrence of catuabain Amazonia has been a subject of some debate.
Mello Moraes (1881) wrote that catuabais a plant of Par-aand Maranhio,
although he did not identify it. Hoehne (1920) noted that it was J. G. Kuhl-
mann, then a curator at the Jardim Botinico do Rio de Janeiro, who iden-
nobilis(=Margaritaria nobilis). Ducke (1966) doubted
tified it as Phyllanthus
this because, so far as he knew, no plant with the name catuabawas known to
the native population of Amazonia. Also, there is no reference to catuabain
van den Berg's (1982) ethnobotanical survey of the famous Vir-o-Peso mar-
ket in Belem, Para. On the island of Sao Luis, Maranhao, where the original
vegetation (now almost completely destroyed) contained elements of both
the Amazonian and Northeastern floras, Ducke and R. L. Fr6es collected a
Micropholis(Sapotaceae)identified by local people as catuaba(Ducke 1966).
Ducke noted that in Amazonia proper the role of catuabais filled by Ptychopeta-
lumuncinatum and P. olacoides(Olacaceae),both known as muirapuama.

Ducke had access to most of the evidence presented above, and he reached
the reasonable conclusion that the name catuabacorresponds to different
species in different regions, even if the uses are the same. He believed that the
true catuabais Anemopaegma mirandum,but that where it does not occur it is
replaced by other taxa: Erythroxylum vaccinifoliumin Pernambuco and part of
Ceard; Secondatiafloribunda elsewhere in Ceara; Micropholisin Maranhi~o;and
Trichilia(Meliaceae)in Bahia (see discussion below). Indeed, the species of
Anemopaegma referred to catuabaare confined to Paraguay, central Brazil, part
of Sio Paulo state, and western Bahia. Table 4 presents the distributions of
the species that have been identified as catuaba.
The identity of catuabain eastern Bahia has been a mystery in itself. Most
of the confusion stems from a doctoral thesis written by A. J. Silva in Salva-
dor in 1904 (cited by Ducke 1966; also by Cunha 1939). According to Ducke
(1966), Silva generated a nomennudum,Erythroxylum catuaba,and the descrip-
tion of the plant is a composite of three distinct species. Cunha (1939)
averred that the description corresponded more to Anemopaegma than to
Erythroxylum. There is, however, an illustration of the plant in Silva's thesis.
By all reports, it shows a plant with alternate, compound leaves and a
capsular fruit, leading Ducke (1966) to conclude that it was the Trichiliahe
had seen referred to in the literature as catuaba.Narciso Soares da Cunha
(Cunha 1939), an expert on Bahian medicinal plants and a founding member
of the Sociedade Baiana de Historia Natural, was intrigued by this problem,
and he organized an expedition to Mata de SdioJoao in search of the catuaba
of Bahia. The participants included the venerable Padre Camille Torrend
and Dr Alexandre Leal Costa (G. Pinto, pers. comm.). They collected her-
barium material of the tree indicated by local people as catuaba.After study-
ing it himself and consulting with J. G. Kuhlmann, Cunha published it in
Tribuna Farmaceuitica (a journal issued by the Universidade Federal do
Parani) as a new species of Tetragastris,T. catuabaCunha. The publication
includes a photograph of the tree and of a herbarium specimen.
An exhaustive search, including correspondence with a participant in
Cunha's expedition (G. Pinto, pers. comm.), has failed to turn up the type
collection. However, parts of Cunha's thorough description leave no doubt
that the plant he collected was indeed a Burseraceae, and specifically a Tetra-
gastris, with the following characters: schizogenous resin canals present;
flower diplostemonous; petals connate ? their length, with a thickened apicu-
lum; filaments broadly laminar and very short; disk 8-10-sulcate; ovary
4-5-locular; ovules 2 per locule; style short; stigmas capitate; fruit a drupe
with 4-5 distinct pyrenes.
It is probable that the plant identified by Ducke (1966) from other litera-
ture and from the illustration in J. B. Silva's thesis as Trichiliais actually the
Tetragastriscatuabaof Cunha. In an illustration, this species could be mistaken
for a Trichilia,because it has alternate, compound leaves with no lateral
pulvinuli, and its dehiscent drupe (so the unit of dispersal is a pyrene, and
not a seed) could be mistaken for a capsule. Ducke (1966) wrote that Cunha
admitted there were indeed trichilias known as catuabain Bahia, but I found
no such statement in Cunha's (1939) published work.
The photographs and other parts of the description-notably the long
petiole and the broadly elliptic to obovate leaflets with a rounded or retuse
apex-have made it possible to match Cunha's species with several existing
collections of Tetragastrismade in eastern Bahia. These include one made

near Ilheus (R. L. Fr6es 1090G) which has catuabaindicated as the common
name. It is interesting to note that one collection from Bahia (T. S. dosSantos
1102) has the common name cocodantarecorded on the label, and the plant
from Pernambuco (F. Paiva 1688) was called cajueirinho.
The question remains as to why the same name, catuaba,has been applied
to taxonomically unrelated plants. If this occurred because they share certain
attributes or activity signatures (uses), then it is a case of a special purpose
classification in the folk taxonomy (cf. Balke & Daly 1989). This can also
occur when unrelated taxa look alike, but most of the catuabasdo not resem-
ble each other at all. However, some specimens of Anemopaegma glaucum,as
well as others determined by A. Gentry as hybrids between A. glaucumand
A. arvense(Irwinet al. 9724; Maguireet al. 56202) have leaves that are similar
in overall appearance to some leaves of Tetragastris catuababecause they are
ternate and have coriaceous, obovate leaflets with revolute margins.
The neotype of T. catuabais selected here and is followed by a description
of the species based on the material collected to date:
Tetragastris catuaba Cunhain Tribuna Farmacefitica 7(3): 45-52 (1939).

Type: Brazil. BAHIA: EstaCdo Experimental de Belmonte/CEPLAC,
16 Sept. 1970 (fl), T. S. dos Santos1102 [neotype (designated here) CEPEC;
isoneotypes K, NY, UEC].
Canopy tree, reproductive height 15-22 m x 25-35 cm (DBH); bark grey,
rough, thick, shed in large irregular plates; resin cream-coloured, sticky.
Branchletsrobust, 3-7 mm diam. near apex, with sparse, appressed hairs to
0-1 mm long, soon glabrescent. Leaves1-3-jugate, 16-6-34-5 cm long; petiole
semi-terete, longer than interjuga, 5-5-10-2 cm x 1-5-3-5 mm, glabrous; in-
terjuga relatively short, 2-2-6-4 cm long; petiolules semi-terete, 1-
canaliculate, laterals 0-2-1-4 cm long, terminal 1-5-4 cm long, pulvinulus
inconspicuous. Leafletscoriaceous, glabrous, waxy and glossy, broadly ellip-
tic, less often slightly ovate (laterals) or slightly obovate (terminal); laterals
5-4-14 x 2-6-6-8 cm, base subequal and acute to slightly obtuse or rarely
truncate, narrowly attenuate; terminal 6-4-13 x cm, base acute and
rounded and 3"6-6-2
retuse or
narrowly attenuate; apex slightly rarely abruptly and
broadly short-acuminate with rounded acumen; margin entire, revolute;
abaxial surface with all veins prominent, adaxial surface with midvein and
secondary veins prominulous, rest of veins prominulous to flat. Inflorescences
paniculate, congested near apex, slender, 6-15 cm x 1-2-3 mm, exceeding
petiole of subtending leaves (or subtended by fleshy, lanceolate bracts 2-
4-6 mm long); secondary axes poorly developed, to 3 cm long; axes with
scattered, appressed hairs to 0-15 mm long or glabrescent; pedicel terete,
0-4-2-2 x 0-4-0-8 mm. Flowers(only staminate ones known) 5-merous, 4-2-
5-7 mm long; calyx 1-1-45 x 2-1-2-5 mm overall, lobes depressed-deltate,
rounded, apiculate; corolla green, fleshy, tubular, 3-7-4-7 mm long, abaxially
with scattered, appressed hairs to 0-15 mm long, lobes narrowly triangular,
1-3-1-7 mm long, apex acute, with conspicuous, inflexed apiculum to 0-7 mm
long, margins short-papillate; stamens 10, isomorphic, mm long,
filaments strap-shaped, mm long, anthers elliptic 1"5-1"95
to ovate in dor-
siventral view, mm0"3-0"45
long, base entire; ovariodisk rounded-conical, glab-
1-1"5
mm high, mm wide at base. Fruitsslightly pendent on
rous,
terete 1"1-1"5
pedicel 0"9-1"3
x 1-5-2 mm; fruits maturing green tinged with red,
2"5-3

smooth, glabrous, glossy, not stipitate, very broadly and obliquely ovoid,
x 1-3 cm when dry (to 2 cm diam. when more than one pyrene), apex
1"5-2
rounded, base subtruncate; columella and interior of valves red; pseudaril
white, pulpy, covering all of pyrene except for a subapical, laterally '8'-
shaped area accounting for 4 of ventral surface and corresponding to funicu-
lar scar; pyrene somewhat bony, 1-2-1-3 x 0-9-1 cm, in dorsiventral view
broadly ovate with acute apex and truncate base, in lateral view slightly
obliquely broad-lanceolate, glabrous on removal of pseudaril; site of funicle I
distance to apex of pyrene: seed 0-9 x 0-6 mm, testa thin and smooth,
cotyledons entire and piano-convex.
This species is represented by a total of six collections (three of them
sterile) from only four localities (see Map 1). Two of these localities are
probably no longer forested (G. Pinto, pers. comm.; F. G. Serpa, pers.
comm.). The forests of the other two localities are not adequately protected:
one is a private ranch, and the other is an experimental station of Brazil's
cacao research center (CEPEC).
Tetragastriscatuabais a medium-sized to large tree, clearly rare and res-
tricted to the lowland moist forests of eastern Bahia, as well as to one of the
isolated patches of moist forest in the northeastern highlands. Ecologically,
these are two of the most threatened regions in the world (Mori et al. 1983;
Andrade-Lima 1982; Mayo & Fevereiro 1982). Much of the forest destruc-
tion in eastern Bahia is due to charcoal production and timber extraction; in
relation to the latter, lumber companies that have exhausted the legal forest
resources in neighboring Espirito Santo state are now very active in southern
Bahia (Mori et al. 1983; pers. obs.).
In most of the northeastern Brazilian states (with the exceptions of Maran-
hao and Piaui), isolated patches of moist forest occur (or occurred) on the
brejos*.These are granitic plateaus of up to 1000 m elevation, found 30-
90 km from the coast. Humid microclimates occur at higher elevations on the
southern and eastern slopes, which are exposed to coastal winds. In
Cearai they have been called serrasfrescas. Floristically, the forests of the
brejoscontain(ed) elements of the Serra do Mar (Ducke 1959) as well as
Amazonia (e.g. Andrade-Lima 1982; pers. obs. in herbaria). The prevailing
view is that the brejoforests are remnants of forests which were continuous
with or effectively (in the sense of dispersal) connected with the Amazonian
and especially with the eastern coastal forests at one or more times during the
Tertiary (e.g. Andrade-Lima 1982).
The brejoshave been under tremendous pressure for centuries, as they are
the primary source of timber and firewood where they occur. It is not known
how many of them have any primary forest cover at all now, and few (e.g.
Mayo & Fevereiro 1982) have been studied in detail. Already by 1959,
Ducke wrote of the brejoforests in the conditional or in the past tense, and he
lamented the rudimentary knowledge we have of them:
'They are humid and they are or were covered by rain forest, the floristic
composition of which we possess only some notions. ... In the past, there
were opulent evergreen forests, today reduced to secondary forest with rare
*Jacques Huber worked in the Serra de Baturite in Ceara in 1897, and Ducke (1959) credited
Huber with being the first botanist to study the flora of the brejos,although he wrote that Freire
Alemio made collections (deposited at the Museu Nacional) in the same Serra around 1860.

*T, breviacuminata
* T, ocohionil 1
* T, catuaba
I. ..
30
MAP1. Distributionof Tetragastris

in easternBrazil.
remains of the primary flora surviving in places of difficult access for man
and for fire.' (my translation).
Although Andrade-Lima (1982) was of the opinion that we now have a good
idea of the plant species occurring in the brejos,there is not much in the
literature to indicate that is so.
It is possible that further botanical exploration planned for the near future
will reveal other localities for T. catuabain Bahia, Espirito Santo, and/or the
brejos.This species is no longer so mysterious, but certainly endangered.

Other specimens examined. Brazil. BAHIA: Mun. Belmonte, EstaCao Ex-
perimental Greg6rio Bondar (EGREB), km 47 of Itapebi-Belmonte road,
7 km E of BarrolAndia,c. 16?14'S, 39000'W, 12 Jan. 1985 (fr), Daly et al. 4062
(CEPLAC; K; MO; NY; US); Ilheus, 16 March 1943 (sterile), Fr6es1090G
(A); Mun. Una, Maruim, border of fazendas Maruim and Dois de Julho,
33 km SW of Olivenga on road to Buerarema, 26 April 1981 (sterile), Mori
et al. 13742 (CEPEC, NY), Mori et al. 13764 (CEPEC). PERNAMBUCO:Amar-
aji, Engenho Sete Rancho, Mata Tres Bacias, 6 Sept. 1963 (fl), F. Paiva 1688
(Herb. Sdrgio Tavares no. 1335) (US).
Tetragastris occhionii (Rizzini) Daly in Brittonia 41: 23 (1989). Type: Bra-

zil. BAHIA: Salvador, restinganear seashore, 5 Dec. 1973 (fl), Duartes.n.
(holotype RB 170553).
ProtiumocchioniiRizzini in Leandra 4-5 (ano 3-4): 15 (1974).
Tetragastris occhioniiwas only recently transferredto the genus from Protium
(Daly 1989). Two characters make this species somewhat anomalous in
Tetragastris.Although the cotyledons are entire, they are distally uncinately
curved much like those of Crepidospermum Hook.f. Also, this species has an
articulation in the ovariodisk-a conical structure derived from the disk and
the pistillode-while in the rest of the genus it is a homogeneous structure.
This less specialized condition in the ovariodisk may indicate T. occhioniias
the most archaic species in the genus (Daly 1989). This conclusion is consis-
tent with that of others who have perceived a remarkable concentration of
archaic taxa in the forests of eastern Brazil (e.g. Cowan 1981a; Kubitzki
1975; Soderstrom & Calder6n 1974).*
The label information for two collections of T. occhionii(Harley22152, Mori
et al. 11415) contain the first reports of latex (more probably a milky resin) in
the genus. However, at least one other collection (L. A. M. Silva 1846)
indicated the sap was colorless.
Twelve collections of T. occhioniiare known over an aerial distance of
c. 300 km along the Bahian coast (Fig. 1). This species forms part of the
unique restingaflora of the Mata Atlantica complex. The restingas,evolved on
the eastern coastal plain of Brazil, range from scrub vegetation on dunes to
relatively tall forest, but they always occur in the lowlands near the coast, on
sandy soils with a perched water table. Tetragastris occhioniiis a small tree or
less often a shrub, and it is found in lower forests, forest edges, and dune
vegetation. This extremely fragile habitat is being destroyed at a rapid pace
due to felling of trees for firewood, timber, and charcoal; clearing of land for
coconut plantations; construction of roads and vacation homes; the expan-
sion of coastal towns and cities; camping; and other activities (Hueck, 1972;
Lacerda et al. 1984).
*There are indications that, at least in Bahia, some of the endemic taxa represent derived
members of archaic groups. Harleyodendron Cowan is a derived member of the Swartzieae,now
regarded as an archaic tribe in the Papilionoideae
(Cowan 1981a). ArapatiellaRizzini & Mattos is
a derived member of the archaic tribe Caesalpinieae(Polhill & Vidal 1981). Among the more
archaic groups within PiperL., subgenus Ottoniashows several derived characters (R. Callejas,
pers. comm.). On the other hand, Barbrodriguesia Cowan is the least specialized of a natural
group of genera in the derived tribe Detarieae (Cowan 1981b), and I have found that Tetragastris
occhionii(Rizzini) Daly is a less specialized member of the relatively derived genus Tetragastris
(Daly 1989).

Other specimens examined. Brazil. BAHIA: Marafi, coastal forest 4 km S of
Marau, 11 Oct. 1968 (fl), J. Almeida & T. S. Santos 138 (CEPEC, NY);
Salvador, Lagoa Abaet6, restinga, 25 Jan. 1965 (fl), R. P. Belim & J. M.
Mendes296 (IAN, US); R. P. Belim &J. M. Mendes310 (UB); Mun. Una,
Ilheus-Una road, 27 km S of Olivenga, coastal forest, 2 Dec. 1981 (fl), M.
Carvalho& G. Lewis 861 (K); 65 km NE of Itabuna at mouth of Rio de
Contas on N bank opposite Itacard, high restinga forest, 14'16'S, 39000'W,
30Jan. 1977 (fr), Harley 18423 (CEPEC); coastal zone near Marafi, forest
margin, 14"10'S, 39000'W, 16 May 1980, Harley 22152 (CEPEC, NY);
Olivenqa, restinga, 1 Dec 1970 (fl), C. E. Mello Filho 3585 (CEPEC); Mun.
Maraui,km 8 of road from Ponta do Muta (Porto de Campinhos) to Maraui,
6 Feb. 1979 (fr), Moriet al. 11415 (CEPEC, NY); Una-Olivenqa road, coas-
tal forest, 28 Oct. 1971 (fl), R. S. Pinheiro1661 (CEPEC); Maraui, coastal
forest, 17 Nov. 1971 (fl), T. S. Santos2174 (CEPEC); Mun. Ilheus, Fazenda
Guanabara (next to Fazenda Barra do Manguinho), on side road 4 km W of
km 10 of Ilheus-Olivenga road, 7 March 1985 (fr), L. A. M. Silva et al. 1846
(CEPEC, NY).
Tetragastris breviacuminata Swartin Rev. Tray. Bot. Neerl. 39: 206. 1942.
Type: Brazil. RIODEJANEIRO: near Rio de Janeiro in mountainous forest, Oct.
1832 (fl), Riedel 1070 (holotype LE; isotypes BR, GH, K, L, M, NY, RB, S,
US,W).
Although Tetragastris breviacuminata
is represented by an appreciable num-
ber of collections, these are from few collecting localities, and the species is
apparently restricted to the immediate vicinity of the city of Rio de Janeiro.
It occurs as an understory or canopy tree in primary or occasionally old
secondary moist forests on lower elevations of steep mountain slopes,
although it has been found on ridgetops (Daly et al. 4073). Some of these
localities are fortunately protected, such as the Parque Nacional de Itatiaia
[D. Constantino (RB no.) 21035] and the Horto Florestal of the Jardim Botani-
co do Rio de Janeiro. Since even the nearby Serra dos Orgaos is not well
collected (P. Carauta, pers. comm.), it is possible that the young and active
staff of RB will encounter T. breviacuminata elsewhere in the state where the
mountain slopes are still forested.
Representative specimens examined. Brazil. RIODEJANEIRO: city of Rio de
Janeiro, Vista Chinesa, 13 Oct. 1935 (buds, fr), D. Constantino(RB no.) 21035
(RB, NY); Jardim Botanico do Rio de Janeiro, Horto Florestal, 18Jan. 1985
(fl), Daly et al. 4072 (RB, NY), 18Jan. 1985 (fr), Daly et al. 4073 (NY, RB);
Sumare [Corcovado], 4 March 1932 (fl),J. G. Kuhlmann(RB no.) 62351 (NY,
RB); Estrada da Mesa do Imperador, 8Jan. 1945 (fr), Occhioni140 (RB);
Mata do Rumo, 12 Dec. 1971 (immature fr), Sucre8093 (RB no. 166037) (NY,
RB).
ACKNOWLEDGEMENTS
For all their help, especially in the rather tedious historical aspects of this
investigation, thanks are due to W. Balke, M. J. Balick, H. P. Bautista, J.
Brennan, R. Callejas P., E. Elisabetsky, A. H. Gentry, H. C. Lima, J. T.
Motta, B. W. and S. Nelson, G. C. P. Pinto, T. Plowman, I. A. Rodrigues, K.
F. Rodrigues, F. G. Serpa, L. A. Mattos Silva, P. Silva, D. W. Stevenson,
and L. Xavier Filho.

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The Genus Tetragastris and the Forests of Eastern Brazil: Studies in Neotropical Burseraceae

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Gaertn., a genus of nine species, is represented by three well-defined species

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Tetragastris catuaba Cunhain Tribuna Farmacefitica 7(3): 45-52 (1939).

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MAP1. Distributionof Tetragastris

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Tetragastris occhionii (Rizzini) Daly in Brittonia 41: 23 (1989). Type: Bra-

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