Notes on Trattinnickia, including a Synopsis in Eastern Brazil's Atlantic Forest Complex.

Studies in Neotropical Burseraceae IX

Author(s): Douglas C. Daly
Source: Kew Bulletin, Vol. 54, No. 1 (1999), pp. 129-137
Published by: Springer on behalf of Royal Botanic Gardens, Kew
Stable URL: http://www.jstor.org/stable/4111029 .
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 Notes on Trattinnickia, including a synopsis in
eastern Brazil's Atlantic forest complex.
Studies in neotropical BurseraceaeIXI

DOUGLASC. DALY2

Summary. The 13 taxa of Traltinnickiarange from Costa Rica to Eastern Brazil. The genus is rather easily
recognized, but several of the species are difficult to distinguish. The two sections are redefined, one
species is transferred to another section, and two varieties are given specific status. The genus is found to
be represented in Eastern Brazil's Atlantic forest complex by two narrowly endemic species. Trattinnickia
ferruginea Kuhlm., first collected in 1935 in eastern Minas Gerais state, was recently collected for the
second time approx. 110 km north of the type locality. A new species, 7 mensalisDaly, is described here.
It is endemic to a limited area of forest on the subcoastal low tablelands of Espirito Santo state. Both
species may be considered rare and threatened.

INTRODUCTI(ON

As treated here, the neotropical genus Trattinnickia Willd. comprises 13 taxa

ranging from Costa Rica (Puntarenas and Lim6n provinces) through northern
South America (including Trinidad) southeast to the states of Minas Gerais, Espirito
Santo and Bahia in Brazil. The genus usually occurs in moist lowland forests but
reaches 1400 m elevation in Venezuelan Guayana.
Trattinnickia is rather easily distinguished from other Burseraceae by its
sympetalous trimerous flowers with retrorse-appressed trichomes on the corolla,
plus the indehiscent, depressed-ovoid to globose fruits with an oily mesocarp and
bony, rugose endocarp. Most of the taxa have leaflets that are slightly asperous to
scabrous on at least one side.
Examination of leaflet surface characters in the genus using SEM as well as light-
microscope slides of transverse sections has revealed the presence of structures
resembling spiked plates; these appear to be sclereids that arise below the epidermis
and then extrude between epidermal cells. They occur on both surfaces but more
densely on the abaxial surface. More significantly, they occur throughout the genus
but more densely on the leaflets of those taxa with asperous to scabrous leaflets (D.
Daly & Z. Marchal, unpubl. results). These structures were not commented on by
Forman et al. (1989) in their discussion of Burseraceae anatomy.
At both species and varietal rank, Trattinnickia presents taxonomic problems
disproportionate to its small number of taxa. Even compared to other Burseraceae, its

Accepted for publication July 1998.

'Continued from Brittonia 50(4): 517 - 523 (1998).
2New York Botanical Garden, Bronx, NY 10458, U.S.A.

129

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130 KEW BULLETIN VOL. 54(1)

taxa display remarkable vegetative plasticity while the fruit morphology varies little
among them. It is rarely collected in flower, and two of the species recognized here
are known only from fruiting material. Herbarium collections of sterile branchlets
and especially of juvenile foliage are exceedingly difficult to match with fertile
material. Observations of seedling morphology, which is providing characters useful
at several ranks within the family, are not available for any of the taxa. These
information gaps highlight the value of research on permanent plots, which
facilitate the documentation of variation in individuals and populations at different
developmental stages.
Definition of natural groups in the genus does not come easily. Swart (1942a:
207) divided Trattinnickia into two sections based entirely on leaflet shape. It
should be noted that his section Rhoifoliae is illegitimately named because it
includes the type species of the genus (see Swart 1942b: 420); hereafter it is
referred to as section Trattinnickia. Later (Swart 1942b: 420), he separated the two
sections using a key whose first couplet is unparallel and which seems to divide the
genus into taxa with flowers greater than or equal to 5 mm long versus those with
flowers up to 5 mm long.
The abaxial leaflet surface presents a more qualitative character that can be used
to distinguish two sections in the genus. Guillaumin (1909) was the first to describe
(and illustrate) a character found in the Burseraceae only in some Trattinnickia: the
presence on the leaflet abaxial surface of almost continuous laminar crypts, each
filled with unicellular trichomes curved or hooked toward the centre of the crypt.
These were later described by Metcalfe & Chalk (1957: 343) as "open grooves filled
with small hairs". Guillaumin mistakenly reported the character in T rhoifolia Willd.;
in reality, these crypts occur only in a group of species including 7: burserifolia Mart.,
most of which have relatively smooth leaflets.
Re-examination of this and several other salient characters throughout the genus
revealed inconsistencies that require two new combinations: Trattinnickia lawrancei
Standl. has laminar crypts, while T lawrancei var. boliviana Swart does not; and T
rhoifolia has the calyx glabrous adaxially, while T rhoifolia var. lancifolia Cuatrec. does
not. The two varieties do not resemble their respective typical varieties in other
aspects, nor can they be accomodated as varieties of any other species in the genus,
so they are here given specific status.

Trattinnickia boliviana (Swart) Daly comb. et stat. nov.

Trattinnickia lawrancei Standl. var. boliviana Swart, Acta Bot. Neerl. 1: 249 (1952).
Type: Bolivia. La Paz: Prov. Larecaja, Tuiri (near Mapiri, on left bank of Rio
Mapiri), alt. 490 - 750 ft., 12 - 30 Sept. 1939 (fl), Krukoff 10767 (holotypus U;
isotypi GH, MO, NY).

Trattinnickia lancifolia (Cuatrec.) Daly comb. et stat. nov.

Trattinnickia rhoifolia Willd. var. lancifolia Cuatrec., Webbia 12 (2): 425 - 426 (1957).
Type: Colombia. Amazonas: Vaupes, Rio Apaporis, Raudal de Jirijirimo, 21 Jan.
1952 (fr), Schultes et al. 14952a (holotypus US; isotypi GH, NY).

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TRATTINNICKIA (BURSERACEAE) IN EASTERN BRAZIL 131

As defined here, section Burserifoliae consists of those taxa with trichome-filled

laminar crypts covering all the abaxial surface not occupied by veins or veinlets,
and section Trattinnickia (former sect. Rhoifoliae of Swart) of those lacking laminar
crypts. These are present in T. lawrancei, so it is here transferred to sect.
Burserifoliae. Interestingly, the other taxa published prior to 1942 remain where
Swart placed them.
I have not yet found additional autapomorphies to better define the two sections.
Two of the more salient pairs of character states in the genus - green versus red
flowers, and the calyx adaxial surface pubescent versus glabrous - correlate well
but not perfectly with the sections (see Table 1). In sect. Trattinnickia, all the taxa
have the calyx pubescent adaxially except T glaziovii Swart and T. rhoifolia Willd.,
and all the taxa have green corollas except T aspera (Standl.) Swart, T boliviana
(Swart) Daly and T demerarae Swart, which have red corollas. In sect. Burserifoliae, all
the taxa for which flowers are known have red corollas and the calyx glabrous
adaxially, except for T barbourii Little, which has the calyx pubescent adaxially.
Complete flowers are not known for T barbourii or T lawrancei.

TABLE 1. The sections of Trattinnickia and correlations with two flower characters.

Taxon Calyx within Corolla colour

sect. Trattinnickia
(laminar crypts absent)

7: aspera (Standl.) Swart pubescent red

T boliviana (Swart) Daly pubescent red or green tinged
with red
T' demerarae Swart pubescent red
T glaziovii Swart glabrous usually green (two
reports of brown)
T lancifolia (Cuatrec.) Daly pubescent green
T peruviana Loes. pubescent green
T rhoifolia Willd. glabrous green

sect. Burserifoliae
(laminar crypts present)
T barbourii Little pubescent ?
T burserifolia Mart. glabrous red
T ferruginea Kuhlm. glabrous red
T lawrancei Standl. ? ?
T laxiflora Swart glabrous red
T mensalis Daly glabrous red

N.B.: T multiflora Cuatrec. and T subchoripetala are here considered to be synonyms

of T burserifolia.

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132 KEW BULLETIN
VOL. 54(1)

Trattinnickia in eastern Brazil

Thanks to recent collections, it is now clear that Trattinnickia is represented in
Brazil's Atlantic forest complex by two narrow endemics. One of them, T ferruginea
Kuhlm., was previously known only from the type collection (Kuhlmann 1936); this
taxon was not mentioned in Swart's (1942b) monograph. The second and only
additional collection was made by graduate students associated with the BHCB
herbarium in 1994, 110 km north of the type locality near Marlieria (the town is
located at 19'43'S, 42?45'W). The other endemic is a new species collected recently
in "tabuleiro" forest on the Reserva Florestal de Linhares in Espirito Santo, in the
course of quantitative forest inventories executed by personnel of the CVRD
herbarium. It is related to the relatively widespread T burserifolia, which does not
reach Eastern Brazil.

Trattinnickia ferruginea Kuhlm., Arq. Inst. Biol. Veg. 3 (1): 45 - 46 (1936). Type:
Brazil. Minas Gerais: silvis primariis ad locum Repreza, proximum ad urbem Vigosa,
16 Nov. 1935 (frt), J. G. Kuhlmann (RB no.) 28.905 (lectotypus [designated here]
RB!; isolectotypus VIC [herb. no. 2283], not seen).
The closely spaced, trichome-filled laminar crypts in the abaxial leaflet surface
place this species in section Burserifoliae Swart. The other taxa in this section have
smooth or at most slightly asperous leaflets, but the adaxial leaflet surface of T.
ferruginea is markedly scabrous. It is also distinguished from the other members of
the section by the presence of flexuous, usually ascending trichomes up to 2 mm
long; on the type collection these form a rust-colored lanate indumentum on the
leaf rachis, leaflet abaxial surface, and infructescence axis, but on the second
collection they are persistent only on the midrib and secondary veins on the abaxial
leaflet surface. Flowers are still unknown, so more collections are needed to
complete the description of the species.
SPECIMENS EXAMINED. BRAZIL.. Minas Gerais: silvis primariis ad locum Repreza,
proximum ad urbem Vi-osa, 16 Nov. 1935 (frt), J. G. Kuhlmann (RB no.) 28.905
(RB, lectotype!; VIC [herb. no. 2283], isolectotype], n.v.); Parque Estadual do Rio
Doce, Marlieria, next to hotel, Sept. 1994 (fr), L. V Costa (BHCB no.) 28650 (BHCB
(n.v.), NY).
DISTRIBUTION AND ECOLOGY. Based on the two collections known to date, this is
a large tree at least 20 m tall, endemic to the isolated moist forests of the Mata
Atlhntica complex in Minas Gerais.
VERNACULAR NAME. "almfacega" (f. G. Kuhlmann2283).

Trattinnickia mensalis Daly sp. nov. Arbor rarior silva humilis Espiritus Sancti incola,
pagina abaxialis folioli cryptis pubescentibus dense instructa, hinc sectioni
Burserifoliae pertinens, sed a Trattinnickia burserifolia his notulis differt: pistillodium
altius parti superae late (nec anguste) conicae leniter (non abrupte) contractum;
filamenta ligulata (nec subulata) in ambobus sexibus; staminodia majora (0.95 - 1.1
mm vs. 0.65 - 0.8 mm) filamenta longiora (0.25 - 0.35 mm vs. 0.05 - 0.15 mm)
fructus ovoideus acuminatus (nec globosus raro leviter obovoideus vel apex breviter

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TRATTINNICKIA IN EASTERNBRAZIL
(BURSERACEAE) 133
acuminatus) major (1.5 - 1.6 x 1 - 1.2 cm vs. 0.8 - 1 cm diam) maturus ut videtur
viridis (nec purpureus). Typus: Brazil. Espirito Santo: Linhares, Reserva Florestal da
CVRD, Est. Mac.-Pele-de-Sapo, ant. 161, km 1,200, 8 March 1978 (Cd fl), J. Spada
65.78 (holotypus CVRD; isotypus NY).

Canopy or emergent tree, reproductive size 10 - 22 m high x 20 - 60 cm diam.

Trunk cylindric. Bark rough. Leaves (2) 3 - 5-jugate, 17 - 30 cm; petiole 4.8 - 9.8 cm x
2 - 3.1 mm, canaliculate, margin slightly involute; interjuga shorter than the petiole,
(0.8) 1.6 - 2.8 cm; petiole and rachis provided with sparse twisted glandular hairs less
than 0.05 mm and sparse appressed hairs less than 0.05 mm, rarely also with dense to
sparse ascending hairs to 0.7 mm (R. P Belem & Magalhaes 826); petiolules narrowly
canaliculate, relatively long and slender, lateral ones 0.4 - 1.1 cm, terminal one 0.9 -
2.4 cm, usually shorter than the interjuga, lateral pulvinuli relatively inconspicuous
or sometimes apparently absent; leaflets coriaceous, drying beige below and light
grey-brown above, leaflets usually lanceolate, less often ovate or oblong-lanceolate to
subelliptic, base subequal, truncate to slightly cordate, apex gradually and broadly
acuminate (acute to abruptly acuminate on R. P. Belem & Magalhaes 826); venation
brochidodromous, on abaxial surface the midrib prominent, secondary veins
prominulous, and higher-order veins prominulous to flat, on adaxial surface the
midrib prominulous but sunk in a groove, secondary veins slightly impressed to
prominulous, and higher-order veins prominulous to flat, abaxial surface slightly
asperous, all the abaxial surface covered by laminar crypts (where not occupied by
veins or veinlets), these densely filled with introrse hairs to 0.1 mm long, adaxial
surface smooth, without raised glands, glabrous. Inflorescence (sub)terminal,
exceeding the petiole when axillary, to 12 cm x 3 - 5 mm (to 30 cm x 6.6 mm on
fruiting specimens), secondary axes well developed, to 7.5 cm long, higher-order
axes congested, axes provided with sparse to dense glandular hairs to 0.05 mm and
sparse to dense erect hairs to 0.1 mm (these scattered on infructescences); bracts on
axes and subtending cymules ovate, to 4 mm, bracteoles subtending flowers
lanceolate to subulate, to 1.5 mm; pedicel terete, those on staminate inflorescences
1.5 - 2 x 0.7 - 0.8 mm, those on pistillate ones 2.3 - 5 x 0.7 - 0.9 mm. Flowers
(measurements from rehydrated material): calyx abaxial surface densely provided
with glandular hairs and erect hairs to 0.1 mm, margin ciliate, adaxial surface
glabrous; corolla wine-colored, tubular, fleshy, the inflexed apicula relatively
conspicuous, approx. 0.25 mm long, abaxial surface sparsely to densely provided
with glandular hairs and densely so with appressed-retrorse hairs to 0.05 mm, margin
papillate and ciliate with erect hairs to 0.1 mm, apiculum with sparse introrse hairs to
0.2 mm; staminate flowers 3.5 - 4.2 mm overall; calyx cupular, 1.5 - 1.8 x 2 - 2.2 mm
overall, often level with apex of the ovariodisk, lobes rounded depressed-deltate to
rounded deltate, 0.4 - 0.5 mm, sometimes irregularly split and the lobes appearing
1.1 - 1.2 mm long (the split portion not ciliate), the apex acute to acuminate; corolla
2.9 - 3.5 mm overall, the lobes rounded-deltate, 1.1 - 1.2 mm; stamens partially
adnate to the base of the ovariodisk, the episepalous series inserted higher and so
the apex higher than the epipetalous series, the former level with the ovariodisk
apex, free portion of stamens 0.9 - 1.1 mm, filaments broadly strap-shaped, fleshy,
anthers ovate-oblong (anthers of epipetalous stamens sometimes more ovate and

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134 KEW BULLETIN VOL. 54(1)

slightly shorter), the apex obtuse to slightly emarginate, 0.8 - 0.9 mm; ovariodisk 1.2
- 1.5 mm high overall, the basal (disk) portion 1 - 1.2 mm diam, gradually
contracted into a conical distal portion that is minutely 3-lobulate at apex, glabrous;
pistillate flowers 4.5 - 4.7 mm long overall; calyx 1.5 - 1.8 x 2.4 - 2.6 mm overall,
exceeding the disk, more regularly divided, the lobes depressed-deltate, 0.5 - 0.7
mm; corolla more narrowly tubular, 4 - 4.2 mm, the lobes rounded-triangular, 1.1 -
1.3 mm; staminodes inserted on margin of the undulate disk, 0.95 - 1.1 mm, the
epipetalous series inserted on higher portions of the disk, the filaments strap-shaped,
thin, anthers (ovate-) oblong, the apex obtuse to truncate, reaching partway up the
pistil, 0.7 - 0.75 and 0.75 - 0.8 mm (those of the epipetalous staminodes often
slightly shorter and sometimes more ovate); disk high and thin, 0.35 - 0.5 x 0.1 mm;
pistil narrowly ovoid, 2.1 - 2.3 x 0.85 - 1 mm overall, glabrous, the base sometimes
slightly substipitate, the style approx. 0.15 mm, not easily distinguished from the
ovary, the stigmatal area 0.25 - 0.3 mm, the three capitate stigmas not easily
distinguished. In fruit the pedicel 2.5 - 7.2 x 1.4 - 3 mm; fruit reportedly maturing
green (Folli 503), ovoid, 1.4 - 1.6 x 1 - 1.2 cm when dry, the base obtuse to acute, the
apex acuminate; pyrene bony, rugose. Cotyledons and seedlings unknown. Fig. 1.
The specific epithet refers to the low tablelands where the species occurs.
SPECIMENS EXAMINED. BRAZIL. Bahia: Camacfi-Canavieiras Highway, 30 km W of
Canavieiras, 13 May 1965 (fr), R. P Belem & Magalhdes 826 (NY). Espirito Santo:
Linhares, Reserva Florestal da CVRD, Est. Flamengo, ant. X-1, km 6.161, left side,
20 Jan. 1984 (9 fl & immat. fr), D. A. Folli 490 (NY), 26 June 1984 (fr), D. A. Folli
503 (NY).
DISTRIBUTION AND ECOLOGY. see below.
VERNACULAR NAMES. "amescla tapina"; "amescla".
NOTES. Trattinnickia mensalis resembles 7' burserifolia, from which it is most easily
distinguished by the fruit, which in 7' mensalis is ovoid, 1.5 - 1.6 x 1 - 1.2 cm, sharply
acuminate, and reportedly matures green, while the latter's fruit is globose or rarely
slightly obovate and sometimes short-acuminate, 0.8 - 1 cm diam., and matures red
(- purple). The new species is further distinguished by the following: adaxial leaflet
surface drying light grey-brown (vs. brown or greenish-brown); staminate flowers
usually larger (3.5 - 4.2 mm long rehydrated, vs. 2.3 - 3.6 mm); the pistillode slightly
longer (1.2 - 1.5 mm vs. 1 - 1.15 mm) and gradually (not abruptly) contracted into a
conical (not narrowly conical) upper portion; the filaments strap-shaped (not
subulate) in both sexes; the pistillate flowers usually longer (4.5 - 4.7 mm vs. 3.5 - 4.5
mm); the staminodes longer (0.95 - 1.1 mm vs. 0.65 - 0.8 mm), with longer filaments
(0.25 - 0.35 mm vs. 0.05 - 0.15 mm); and the disk usually narrower (0.1 mm thick vs.
0.1 - 0.2 mm) and not thickened at the base.

FIc. 1. Trattinnickia mensalis Daly. A flowering branchlet with staminate inflorescence; B areole of veinlet,
showing laminar crypt filled with introrse trichomes; C staminate flower; D ovariodisk with some stamens
removed, and longisection of staminate flower; E dorsal and ventral views of stamen; F leaf and pistillate
inflorescence; G pistillate flower, and same with portion of calyx and corolla removed; H transverse section of
ovary, and longisection of pistil; J ventral and dorsal view of staminode; K fruit, and infructescence. A - E from
Spada 65.78, F -J from Folli 490, K from Folli 503. Drawn by Bobbi Angell.

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TRATTINNICKIA (BURSERACEAE) IN EASTERN BRAZIL 135

C,

3 cm<
2 mm

B NE
Imm

I1 '

m
/

3 cm 1
" '

cmI

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136 KEW BULLETIN VOL. 54(1)
Restricted to the Reserva Florestal de Linhares in Espirito Santo state, with a
disjunct population further north in Bahia beyond the Rio Doce. The single
collection from Bahia (R. P Belem & Magalhdes 826) differs from the Espirito Santo
material in three ways: the petiole and rachis provided with dense to sparse
ascending hairs to 0.7 mm (vs. absent or caducous); the leaflet apex acute to
abruptly acuminate (vs. gradually acuminate); and the fruit of the same shape but
larger (1.5 - 1.6 x 1.2 cm vs. 1.4 - 1.5 x 1 cm).
The differences in the Bahian population may be explained by its location across
the Rio Doce, which clearly has functioned as a barrier to gene flow for a number of
taxa. Rainfall in the lower Rio Doce valley is lower, more seasonal, and less
consistent than it is farther north or south, and the differences may have been
accentuated in the geological past (see Voeks 1987). This partial climatic and
physical barrier has resulted in varying degrees of divergence, including the
evolution of varietal and species pairs (Mori et al. 1981; and Daly 1992 (here the Rio
Doce misnamed as the Rio Claro)). Still, I maintain the Bahian material in T
mensalis unless additional collections are found in Bahia that present additional and
consistent differences. This is assuming the species is not already extinct there.
In Espirito Santo, T mensalis apparently occurs in two forest types that are part
of the "mata de tabuleiro" vegetation complex in that region of low flat tablelands
28 - 90 m above sea level. The climate, with approx. 1,100 - 1,400 mm annual
precipitation and a dry season from May to September, supports "somewhat semi-
deciduous" forests. One of the forest types, called mata alta, has been studied by
personnel from the CVRD herbarium (e.g., Peixoto & Silva 1997, Peixoto et al.
1995). It has a discontinuous canopy approx. 24 m high, and is found on red-
yellow dystrophic podzols. An approx. one-hectare inventory included two
individuals of 7: mensalis, 16 m high x 10.6 cm diam. and 12 m high x 8 cm diam.
(and listed as 7' burserifolia); they were considered "trees of the present" sensu
Oldeman (1974) of the intermediate stratum.
The new species also occurs an emergent or canopy species in a rarer
vegetation type known as mussununga or floresta baixa. This forest type is found on
sandy ridges and is characterized by a more open and lower canopy and more
sclerophyllous species.
Like the majority of the taxa of Burseraceae that occur in Brazil's Atlantic forest
complex (see Daly 1990, 1992), both 7T mensalis and 7T ferruginea are narrow
endemics and should be considered threatened, even though both are known to
occur in conservation units.

ACKNOWLEDGEMENTS

I thank Jiulio Lombardi of the BHCB herbarium for background information on

their collectors and excellent collections, Dra. Ariane Peixoto for supplemental
information about the tabuleiro forests, Rupert Barneby for assistance with the Latin
diagnosis, Bobbi Angell for the fine illustration, and an anonymous reviewer for
corrections and constructive comments.

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TRATTINNICKIA IN EASTERNBRAZIL
(BURSERACEAE) 137
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