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Synopses of Angostura Roem. & Schult. and Conchocarpus J. C. Mikan (Rutaceae)

Authors:

Abstract

Angostura Roem. and Schult. as understood by Engler is defined more narrowly here. The species excluded from Angostura are recognized as species of Conchocarpus J. C. Mikan. Three new species of Angostura (A. alipes Kallunki from Ecuador, A. quinquefolia Kallunki from Para and Maranhao, Brazil, and A. simplex Kallunki from western Amazonas, Brazil, and San Martin, Peru) are described, and three new combinations in this genus are made. In Conchocarpus, 24 new combinations are made, and the following 21 new taxa are described: C. bellus Kallunki, C. cauliflorus Pirani, C. concinnus Kallunki, C. cuneifolius var. confertus Kallunki, C. cyrtanthus Kallunki, C. dasyanthus Kallunki, C. diadematus Pirani, C. fissicalyx Pirani, C. furcatus Kallunki, C. gaudichaudianus subsp. bahiensis Kallunki, C. grandis Kallunki, C. hirsutus Pirani, C. inopinatus Pirani, C. insignis Pirani, C. longipes Kallunki, C. mastigophorus Kallunki, C. modestus Kallunki, C. oppositifolius Kallunki, C. punciatus Kallunki, C. santosii Pirani and Kallunki, and C. sordidus Kallunki. With the exception of C. grandis from Amazonian Brazil, all are native to the coastal forests of eastern Brazil. As a result, seven species of Angostura and 45 of Conchocarpus are recognized. Keys to the taxa of both genera are provided. Lectotypes are designated for Cusparia grandiflora Engl., C. macrocarpa Engl., C. paniculata Engl., C. toxicaria Engl., Calipea odoratissima Lindl., and Lasiostemum silvestre Nees and Mart. and an epitype for Calipea elegans A. St.-Hill. The new combination, Rauia nodosa (Engl.) Kallunki, is made for Cusparia nodosa. The tribe Cusparieae DC. and subtribe Cuspariinae Engl., based on the illegitimate generic name Cusparia Humb., are renamed Galipeeae Kallunki and Galipeinae Kallunki.
Synopses of Angostura Roem. & Schult. and Conchocarpus J. C. Mikan (Rutaceae)
Author(s): Jacquelyn A. Kallunki and José R. Pirani
Source:
Kew Bulletin,
Vol. 53, No. 2 (1998), pp. 257-334
Published by: Springer on behalf of Royal Botanic Gardens, Kew
Stable URL: http://www.jstor.org/stable/4114501
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Synopses of Angostura Roem. & Schult. and
ConchocarpusJ. C. Mikan (Rutaceae)
JACQUELYN A. KALLUNKI' & JosE R. PIRANI2
Summary. Angostura Roem. & Schult. as understood by Engler is defined more narrowly here. The
species excluded from Angostura are recognized as species of ConchocarpusJ. C. Mikan. Three new
species of Angostura (A. alipes Kallunki from Ecuador, A. quinquefolia Kallunki from Pari and
Maranhao, Brazil, and A. simplex
Kallunki from western Amazonas, Brazil, and San Martin, Peru) are
described, and three new combinations in this genus are made. In Conchocarpus,
24 new combinations
are made, and the following 21 new taxa are described: C. bellus Kallunki, C. cauliflorus Pirani, C.
concinnus
Kallunki, C. cuneifolius
var. confertus
Kallunki, C. cyrtanthus
Kallunki, C. dasyanthus
Kallunki, C.
diadematus
Pirani, C.
fissicalyx Pirani, C. furcatus Kallunki, C. gaudichaudianus subsp. bahiensis
Kallunki,
C. grandis Kallunki, C. hirsutus Pirani, C. inopinatus Pirani, C. insignis Pirani, C. longipes Kallunki, C.
mastigophorus
Kallunki, C. modestus
Kallunki, C. oppositifolius
Kallunki, C. punctatus Kallunki, C. santosii
Pirani & Kallunki, and C. sordidus Kallunki. With the exception of C. grandis from Amazonian Brazil, all
are native to the coastal forests of eastern Brazil. As a result, seven species of Angostura and 45 of
Conchocarpus
are recognized. Keys to the taxa of both genera are provided. Lectotypes are designated
for Cusparia grandiflora Engl., C. macrocarpa
Engl., C. paniculata Engl., C. toxicaria Engl., Galipea
odoratissima
Lindl., and Lasiostemum
silvestre Nees & Mart. and an epitype for Galipea elegans
A. St.-Hil.
The new combination, Rauia nodosa (Engl.) Kallunki, is made for Cusparia
nodosa. The tribe Cusparieae
DC. and subtribe Cuspariinae Engl., based on the illegitimate generic name Cusparia Humb., are
renamed Galipeeae
Kallunki and Galipeinae
Kallunki.
INTRODUCTION
Recent collecting in the Atlantic coastal forest of Brazil revealed several new
species of Rutaceae that belong to the neotropical tribe Galipeeae Kallunki trib.
nov. [ - Cusparieae
DC., nom. illegit., Mem. Mus. Hist. Nat. Paris 9: 141 (1822);
type: Galipea trifoliata
Aubl.] and to the subtribe Galipeinae Kallunki subtrib. nov.
[ - Cuspariinae Engl., nom. illegit., Engler & Prantl, Nat. Pflanzenfam. 3(4): 111,
160 (1896)]. These species would be assigned to Angostura Roem. & Schult. ( -
Cusparia Humb.) if the circumscription of the genus proposed by Engler (1874,
1896, 1931) were followed. Since, however, no species described in Angostura
other than its type, A. trifoliata (Willd.) T. S. Elias, has its combination of
characters, Engler's circumscription cannot be followed. Therefore, Angostura
must be defined more narrowly, and the species thus excluded from Angostura
must be accommodated elsewhere in the subtribe. Assessment of the characters
and their variation led us to interpret these excluded species as one polymorphic
genus, ConchocarpusJ. C. Mikan.
Accepted
for publication
October 1997.
1
The New
York Botanical
Garden, Bronx,
NY
10458
- 5126,
U.S.A.
2
Departamento
de BotAnica,
Universidade
de Sio Paulo,
Caixa
Postal
11461,
CEP
05422-970,
Sto Paulo, SP,
Brazil.
257
258 KEW
BULLETIN VOL. 53(2)
Saint-Hilaire (1823, 1824, 1825) cited Cusparia, Angostura, and Conchocarpus
among the synonyms of Galipea
Aubl. and described several new species in Galipea
and two in Ticorea
Aubl. Those he described in Galipea
belong in Conchocarpus,
and
those he described in Ticorea
belong in Galipea.
His erroneous circumscriptions of
these genera were perpetuated by de Candolle (1824),Jussieu (1825), and Bentham
& Hooker (1862). Engler, in his treatments of the Rutaceae in Martius's "Flora
Brasiliensis"
(1874) and in "Die
Natiirlichen
Pflanzenfamilien"
(1896, 1931), defined
these genera somewhat more clearly. He limited Ticorea
to species with five fertile
stamens by transferring to Galipea the two species (with two fertile stamens)
described by Saint-Hilaire. He limited Galipea to species with connate carpels by
excluding from it the apocarpous species, i.e., the species described by Saint-Hilaire
(which, as noted above, belong in Conchocarpus)
and the type of Angostura,
which he
called Cusparia
trifoliata
(Willd.) Engl. He recognized these as one genus, Cusparia,
still confounding Angostura and Conchocarpus.
He also failed to realize that his
Galipea bracteata
(Nees & Mart.) Engl. was congeneric with his C. trifoliata. These
errors are rectified in the present paper.
Elias (1970) showed that Cusparia was only a "provisional name" and that
Angostura was the correct name to use in its place. Subsequently, Albuquerque
(1981) transferred most of the epithets from Cusparia
to Angostura,
not aware that
all species whose epithets he transferred did not belong there. Although Cusparia
was validated by de Candolle (1822), it is an illegitimate later synonym of (and is
based on the same material as) Angostura and, thus, cannot serve as the basis of
tribal or subtribal names.
The Galipeinae
differ from most of the Rutaceae
by tendencies toward zygomorphic
flowers, a tubular corolla, reduction in number of fertile stamens from five to two
with the transformation of stamens into staminodes, and basally appendaged
anthers. Of the 26 genera of the Galipeinae,
10 are monospecific and eight more are
represented by five or fewer species. The genera are defined not by unique
character states but by unique combinations of character states of a limited number
of characters, of which any single state recurs in other genera. Characters vary
independently and similar character states apparently have arisen independently
and through reversals. Delimitation of genera that exhibit this kind of variation in
characters can be difficult because characters must be evaluated in groups, not
singly. Observations of any one character will not permit either accurate assignment
of a species to a genus or a reasonable assessment of relationships among genera.
Use of carpel connation, for example, to group genera will give results different
from those obtained through use of petal connation.
Nevertheless, Angostura and Galipea are quite distinct from Conchocarpus
as
illustrated by the following key:
1. Carpels connate and fruit syncarpous before dehiscence; anthers sterile at base
above point of attachment to filament, appendaged at base, connate by their
sterile bases and appendages ............................. Galipea
1. Carpels free and fruit apocarpous even before dehiscence (partially connate in C.
cuneifolius); anthers completely fertile, appendaged or not, but when
appendaged not connate
SYNOPSES
OF ANGOSTURA & CONCHOCARPUS 259
2. Petals connate, adnate to filaments; anthers appendaged at base, exserted
their whole length from corolla tube; trichomes branched ..... Angostura
2. Petals ? coherent, ? adherent to filaments; anthers not appendaged, not
exserted their whole length from corolla tube (except in C. cuneifolius);
trichomes simple ............................... Conchocarpus
In this paper, descriptions of Angostura sensu stricto
and Conchocarpus
are given,
new combinations in both genera are made, and three new species of Angostura
from Amazonian Brazil, Peru, and Ecuador and 21 taxa of Conchocarpus
from Brazil
(all but C. grandis from the Atlantic coastal forests) are described. As a result, we
recognize seven species of Angostura
and 45 of Conchocarpus.
At the end, we indicate
the disposition of those epithets in Angostura
that belong neither in Angostura
sensu
stricto
nor in Conchocarpus
and list as doubtful one species of which we have seen
insufficient material.
All specimens cited were seen by one or both authors unless indicated
otherwise in the case of some type specimens. Herbaria in which specimens are
deposited are indicated by the acronyms found in the eighth edition of "Index
Herbariorum"
(Holmgren et al. 1990). Most measurements given for floral parts
were obtained from rehydrated or pickled flowers. The terminology used to
describe two-dimensional shapes is that of Hickey (1979). The term "partial
inflorescence" (Weberling 1989) refers to the individual elements of the
inflorescence, which in these genera are the variously modified cymose elements
of the compound thyrse. Terms used to describe indumentum are defined at their
first use in each generic description.
ANGOSTURA
Angostura Roem. & Schult.
in Syst. Veg. 4: 188 (1819). Type: A. trifoliata
(Willd.) T. S.
Elias (basionym: Bonplandia trifoliata
Willd.).
Bonplandia Willd. in Mem. Acad. Roy. Sci. Hist. (Berlin) 1802: 26 - 28, pl. 1804
(1802), non Cav. Type: B. trifoliata
Willd.
Cusparia
Humb., Essai Geog. P1. 58 (1807), nom. prov. (see Elias 1970); DC., Mem.
Mus. Hist. Nat. Paris 9: 142, 143 (1822). Type: Cuspariafebrifuga
Humb., nom. prov.
Shrubs or trees to 15 m tall, diffusely branched; wood very hard; bark often pale.
Pubescence
of twigs, vegetative buds, leaves, inflorescences, calyx, and external
surface of corolla of branched trichomes, these either echinoid (i.e., with a convex
or subglobose base and relatively short branches radiating in all directions),
stellate (i.e., with a flat or inconspicuous base and longer or shorter branches
radiating ? horizontally), or a mixture or gradation of such trichomes. Leaves
alternate, usually 3-foliolate (1-foliolate in A. simplex, 5 - 7-foliolate in A.
quinquefolia), petiolate; blade entire, chartaceous. Inflorescence an apparently
terminal thyrse borne near apex of leafy twig and at ? the same level as a vegetative
bud and its subtending leaf, the partial inflorescences cymose, sessile or stalked
(i.e., the secondary axis developed), the higher order axes developed or not. Calyx
of 5, quincuncial sepals + connate at base, persistent, the lobes not overlapping at
260 KEW BULLETIN VOL. 53(2)
anthesis. Corolla
tubular, with spreading or recurved lobes, the innermost in upper
position (at least as seen in A. bracteata), subactinomorphic, white; petals 5, of
imbricate aestivation, connate, the outer surface densely pubescent with echinoid
or stellate trichomes, the inner surface of tube pubescent with simple trichomes,
the inner surface of lobes densely tomentulose (curled, matted trichomes).
Androecium of 2 fertile stamens flanking the innermost petal and 5 staminodes (3
alternate with and 2 opposite to petals); filaments of fertile stamens and
staminodes flattened, connate into a tube, the tube ? equal to corolla tube and
adnate to it for most of its length, sometimes adherent to corolla throat by dense
abaxial pubescence, usually free from corolla to some extent at its apex, the free
part of fertile filaments usually glabrous except sometimes at base, the free part of
staminodes often spathulate, glabrous or minutely pubescent, each bearing a
globose or convex gland at apex; anthers free, + oblong, glabrous, not glandular-
punctate, bearing free, flattened, denticulate or bifid appendages at base, broadly
attached to filament, exserted from corolla tube; pollen grains 5 - 6-short-
colporate, oblate-spheroidal, large, the exine baculate. Disc cupular, shorter than
or equal to ovary, carnose, glabrous, the margin crenate or subentire. Ovary of 5
(6), free carpels united only in the style, apically umbilicate or truncate; stigma
(sub)capitate, exserted from throat at maturity. Fruit of 1 - 5, free mericarps, these
usually beaked apically. Seed 1 per carpel, smooth to slightly rugulose, glabrous, the
testa crustaceous. Embryo curved, the radicle apical, the cotyledons rounded,
conduplicate, plicate, subequal, thin and flexible.
The genus is distributed from Nicaragua to northern Venezuela and south to
northern Bolivia and southern Brazil. Based on their currently known ranges, the
species seem to be allopatric.
The combination of characters that distinguishes Angostura
sensu stricto
from the
other genera of Galipeinae comprises: alternate leaves, stellate or echinoid trichomes,
connate petals, a staminal tube adnate to corolla, 5 staminodes tipped with globose
glands, exserted anthers, free carpels, smooth seeds, and baculate pollen grains. The
branched, stellate or echinoid trichomes are found in no other genus in the
subtribe. Galipea
Aubl., in which some species of Angostura
sensu stricto have been
described, differs, in addition to the characters noted in the preceding key, by simple
trichomes, anthers not exserted from corolla tube, and reticulate pollen grains
(Morton & Kallunki 1993).
KEY TO THE SPECIES OF ANGOSTURA
1. Leaves 1- or 5 - 7-foliolate
2. Leaves 1-foliolate ................... ....*............ A. simplex
2. Leaves 5- 7-foliolate ..............................A. quinquefolia
1. Leaves 3-foliolate
3. Petiolewinged ....................................... A. alipes
3. Petiole not winged
4. Calyx lobes membranous, sparsely stellate-pubescent; fruiting carpels 7 - 8
mm long, glabrous .............................. A. bracteata
SYNOPSES OF ANGOSTURA & CONCHOCARPUS 261
4. Calyx lobes not membranous, densely echinoid-pubescent; fruiting carpels 8
- 15 mm long, densely echinoid-pubescent
5. Secondary axes of inflorescence 5 - 16 mm long; corolla lobes longer
than or equal to corolla tube; fruiting carpels 13 - 15 mm long;
anthers 4.8 - 5 mm long ........................ A. trifoliata
5. Secondary axes of inflorescence to 7 mm long, but almost always much
shorter; corolla lobes shorter than corolla tube; fruiting carpels 8 - 11
mm long; anthers 2.5 - 4 mm long
6. Calyx lobes lanceolate, c. 5 times longer than broad; corolla 47 - 64
mm long ................................. A. longiflora
6. Calyx lobes ovate, c. 1.2 - 1.6 times longer than broad; corolla 23 -
27.5 mm long ............................. A. granulosa
THE SPECIES OF ANGOSTURA
Angostura alipes Kallunki sp. nov. a ceteris congeneribus petiolis late alatis differt.
Typus: Ecuador, Pastaza, Via Auca, V Zak
4018 (holotypus NY, isotypus MO).
Tree 15 m tall; trichomes of twigs, vegetative buds, and leaves minute, tawny,
stellate. Leaves
3-foliolate; petiole 3 - 6.2 cm long, winged, the wings c. 5 mm wide
at apex, cuneate at base; petiolules absent; blade of terminal leaflet oblanceolate,
10.3 - 18.5 x 3 - 4.6 cm, that of lateral leaflet (narrow-) elliptic, 7.7 - 13.4 x 2.9 -
4.6 cm, all acute at base, acuminate at apex, glabrescent, minutely pellucid-
punctate, more obviously so when older, the glands visible on both surfaces but
more so on abaxial one; midvein and secondary veins raised on both surfaces.
Inflorescence
a spiciform thyrse, 16.3 - 30.5 cm long including the peduncle, this 8
- 11 cm long, minutely stellate-pubescent, the partial inflorescences sessile, 1 - 3-
flowered, simple cymes, the secondary axis none; primary bracts or their scars not
seen; pedicels to 0.5 mm long. Calyx dentate, 2 - 3 mm long, minutely stellate-
pubescent without, densely sericeous within, densely fringed at margin by
exsertion of the silky pubescence, persistent in fruit, the lobes broadly rounded, c.
0.2 - 0.5 x 1.2 - 1.4 mm. Corolla
salverform, 45.5 - 57 mm long, minutely stellate-
pubescent without, sericeous on tube within, the tube 38 - 49 x c. 2.5 mm, the
lobes rounded or obtuse, 7.3 - 8 x 2.7 - 3.5 mm, elliptic, spreading to recurved,
flat. Androecium: free part of fertile filament 2.5 - 3 mm long, subglabrous; free
part of staminodes 5.5 - 6.5 mm long, glabrous; anthers narrow-oblong, c. 3 x 0.8
mm, bearing at base a notched flap-like appendage c. 0.25 mm long. Disc 5-
crenate or subentire, c. 1 mm high, ? equal to ovary. Ovary of 5 carpels, truncate
at apex, c. 1 mm high, glabrous; style 40 - 45 mm long, glabrous; stigma
subcapitate. Fruit unknown. Fig. 1G -J.
DISTRIBUTION.
Ecuador (Pastaza); known only from the type locality.
ECUADOR. Pastaza: Via Auca, 115 km S of Coca, PETROCANADA oil well site,
1?15'S, 76?55'W, 13 Feb. 1989 (fl.), Neill et al. 8915 (MO, NY), Carretera de PETRO-
CANADA,
Via Auca, 115 km S de Coca, 4 km S del Rio Tiguino, 1?15'S, 76?55'W,
22
-28 Feb. 1989 (fl.), Zak 4018 (holotype NY, isotype MO).
HABITAT. Tropical moist forest; c. 320 m elevation.
262 KEW
BULLETIN VOL. 53(2)
FIG. 1. Angostura. A, B A. simplex. A apex of branch; B fruit and seed. C - F A. quinquefolia. C apex of branch
with inflorescence; D anther, abaxial, adaxial, and lateral views; E disc and gynoecium with shortened style; F
immature fruit. G -J A. alipes. G apex of branch with inflorescence; H calyx and magnification of stellate hair; J
anther, abaxial and lateral views. A, B from Teixeira
et al. 1213; C - E from Silva & Bahia 2883, F from Rosa 2417;
G -J from Neill et al. 8915. Drawn by Bobbi Angell.
SYNOPSES
OF ANGOSTURA & CONCHOCARPUS 263
This species is easily distinguished from others in the genus by its conspicuously
winged petioles. Its corolla, like those of A. quinquefolia
and A. longiflora,
is longer
than 45 mm, whereas the corolla of the other species (that of A. simplex
is unknown)
is shorter than 27.5 mm. Among these three larger-flowered species, A. alipes is
distinguished by the stellate (vs. echinoid) trichomes on calyx and pedicel, by the
shallowly crenate calyx (the lobes representing 1/7 vs. 1/4 - 3/4 of the length of
the calyx), and by the anthers (3 mm long) within the range of those of A. longiflora
(2.5 - 3.3 mm) but shorter than those of A. quinquefolia
(4.7 - 5 mm).
Angostura bracteata (Nees & Mart.) Kallunki comb. nov.
Sciuris
bracteata
Nees & Mart. in Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat.
Cur. 11: 156, t. 18A & 20 (1823). Type: Brazil, Espirito Santo, Aragatiba, Wied-
Neuwied
s.n. (holotype BR, isotype M).
Ticorea
bracteata
(Nees & Mart.) A. St.-Hil. ex DC., Prodr. 1: 730 (1824).
Galipea
bracteata
(Nees & Mart.) Schult., Mant. 2. Add. I, cl. ii: 195 (248m) (Jan. -
April 1824).
G. bracteata
(Nees & Mart.) Engl. in Mart., Fl. Bras. 12(2): 99 (1874), comb. superfl.
Portenschlagia
trifoliata
Pohl ex Engl. in Mart., Fl. Bras. 12(2): 100 (1874), pro synon.
DISTRIBUTION. Brazil (Bahia, Espirito Santo, Rio de Janeiro, Sao Paulo).
Angostura granulosa (Kallunki) Kallunki
comb. nov.
Galipea granulosa Kallunki in Brittonia 40: 241 (1988). Type: Nicaragua, Zelaya, El
Zapote, Sandino
4720 (holotype NY, isotype MO).
DISTRIBUTION. Nicaragua, Costa Rica.
Angostura longiflora (K. Krause)
Kallunki comb. nov.
Galipea
longiflora
K. Krause in Notizbl. K6nigl. Bot. Gart. Berlin 6: 144 (1914). Type:
Brazil, Acre, Sao Francisco, Rio Acre, Ule 9497 (holotype Bt, photo from F neg
12483 at NY, isotype G, K).
DISTRIBUTION. Brazil (Acre, Mato Grosso), Bolivia.
Angostura quinquefolia Kallunki
sp. nov. a ceteris congeneribus foliis 5 - 7-foliolatis
differt. Typus: Brazil, Part, Marabfa,
M. G. Silva & R. Bahia 2883 (holotypus MG;
isotypus NY).
Shrub or tree 2.3 - 6 m tall; trichomes of twigs, vegetative buds, and leaves
ferrugineous, stellate to echinoid. Leaves 5 - 7-foliolate; petiole 6.3 - 31.8 cm long;
petiolules of terminal blade 0.3 - 1.8 cm long, of lateral blades 0.2 - 1.1 mm long,
slightly swollen at base; blade of terminal leaflet oblanceolate or elliptic, 18 - 32.3 x
5.2 - 10.6 cm, that of lateral leaflets elliptic to narrow-ovate, 9 - 23 x 3 - 9.3 cm, all
acute at base, acuminate (sometimes caudately so) at apex, villous on midvein
adaxially, densely and finely pellucid-punctate, the glands visible on both surfaces;
264 KEW BULLETIN VOL. 53(2)
midvein and secondary veins raised on both surfaces. Inflorescence
a spiciform thyrse
25 - 50.2 cm long including the peduncle, this 7 - 22.8 cm long, stellate- and/or
echinoid-pubescent, the partial inflorescences subsessile, few-flowered, the secondary
axis to 1 mm long, the higher order axes obscure; primary bracts linear, 15 - 19 mm
long at lower nodes to 2 - 4.5 mm long at upper nodes, early deciduous; pedicels 1 -
4 mm long, densely echinoid-pubescent. Calyx dentate, 4 - 5 mm long, densely
echinoid-pubescent without, densely sericeous within, persistent at least in immature
fruit, the lobes triangular, acute, 1 - 1.2 x c. 2 mm. Corolla
salverform, 49 - 56 mm
long, densely echinoid-pubescent (trichomes longer-branched on lobes) without,
densely retrorsely sericeous on tube within, the tube 38 - 44 x 2.5 - 3 mm, the lobes
rounded, 11 - 15 mm long, elliptic or narrow-ovate, spreading, the innermost c. 3
mm wide, canaliculate, the others 4 - 5 mm wide, flat. Androecium: free part of fertile
filaments 6 mm long, glabrous in upper half, tomentulose below; free part of
staminodes 6 mm long, glabrous; anthers narrow-oblong, 4.7 - 5 x 1 - 1.6 mm,
bearing at base a bifid appendage c. 1.1 mm long. Disc 5-crenate, c. 1 mm high, +
equal to ovary. Ovary of 5 (6) carpels, truncate at apex, c. 1 mm high, glabrous,
becoming pubescent soon after anthesis; style c. 47 mm long, glabrous; stigma
capitate. Fruit (very immature) of 1 - 4 mericarps, each 13 - 14 x 6 - 7 mm, sparsely
stellate-pubescent, beaked at least at this young stage. Seed unknown. Fig. IC - F.
DISTRIBUTION. Brazil (Maranhio, Pari).
BRAZIL.
Maranhio: Mun. Santa Luzia, 27 km S of Entroncamento, the
intersection of Hwy. 006 and Hwy. 222, along Hwy. 006, 4?23'S, 46'14'W, 20 March
1983 (bud), Schatz
et al. 946 (NY). Para:
junto a lagoa da Serra dos Carajas, 28 June
1976 (fr), Ribeiro
1423 (US); Sio Domingos do Capim, fazenda Maircia, 40?10'S,
481l0'W, pr6ximo do igarap, Garraf'ao,
18 June 1978 (fr), Rosa 2417 (NY); Marabfi,
km 3 da estrada para a serra dos Carajais,
26 March 1977 (fl.), Silva & Bahia 2883
(holotype MG, isotype NY); Marabai,
Serra dos Carajais,
arredores da serraria, 31
March 1977 (fl.), Silva & Bahia 2960 (MG, NY).
HABITAT.
Tall evergreen forest on terra firme.
This is the only species of Angostura
with 5 - 7-foliolate leaves and, thus, is easily
recognized. The trichomes of its pedicels and calyx are markedly echinoid as are
those of A. trifoliata and A. longiflora. The new species differs, however, from A.
trifoliata by its longer corolla (49 - 56 vs. 21 - 22 mm) and from A. longiflora by its
longer anthers (4.7 - 5 vs. 2.5 - 3.3 mm) and longer fruiting carpels (13 - 14 mm
even when immature vs. 8 - 10 mm). While the trichomes of A. quinquefolia
and A.
trifoliata
are golden to ferrugineous, those of A. longiflora are pale.
Angostura simplex Kallunki
sp. nov. a ceteris congeneribus foliis magnis 1-foliolatis
differt. Typus: Brazil, Amazonas, Mun. Humaitai, L. O. A. Teixeira,
A. J. Fife, K.
McFarland, C. D. A. Mota, J. L. dos Santos, S. P Gomes & B. W Nelson 1213 (holotypus
SPF,
isotypi INPA, n.v., NY and 5 to be distributed to CAS, F, K, MO, U).
Tree 5 - 6 m tall; trichomes of twigs, vegetative buds, and leaves minute, tawny,
echinoid. Leaves 1-foliolate; petiole 3.5 - 6.5 cm long, glabrescent; blade narrow-
obovate, elliptic, or lanceolate, acute at base, subcaudately acuminate at apex, 27 -
SYNOPSES OF ANGOSTURA & CONCHOCARPUS 265
40 x 9.2 - 13.2 cm, glabrescent, minutely pellucid-punctate, the glands visible on the
abaxial surface, the midvein and secondary veins raised on both surfaces. Inflorescence
a narrow thyrse 35.5 - 39 cm long including the peduncle, this 13 - 23 cm long,
minutely echinoid- and stellate-pubescent, the partial inflorescences stalked, few-
flowered, the secondary axis 2 - 3 mm long in bud to 1 - 2.5 (4 - 5) mm long in
fruit, the higher order axes obscure; primary bracts c. 4.5 mm long, deciduous;
pedicels 3.5 - 6 mm long in fruit, minutely echinoid-pubescent. Calyx
lobed, 3.5 - 4
(7) mm long in fruit, minutely echinoid-pubescent without, densely sericeous or
villous within, persistent in fruit, the lobes narrow-ovate or -oblong, acute, 3 - 4 (6 -
6.2) x 1 - 1.2 (2.5 - 2.9) mm. Corolla,
androecium,
disc not known. Ovary
of 5 carpels,
densely minutely echinoid-pubescent (glabrous) after anthesis. Fruit of 2 - 5
mericarps, each 12 - 14 (20) x 8 (11.5) mm, sparsely minutely echinoid-pubescent
and glabrescent (glabrous), transversely ridged, pale glandular tuberculate, beaked
(or not) apically, the blunt beak when present c. 1.5 - 2 mm long. Seed
subrugulose
dorsally, 7 - 7.5 x 5 - 5.5 mm, oblong or suborbicular, brown. Fig. 1A, B.
DISTRIBUTION. Brazil (Amazonas), Peru (San Martin).
BRAZIL. Amazonas: Mun. Humaiti, estrada Humaiti-Jacarecanga, km 64, a 1 km
ao N, 7045'S, 62032'W, 18 June 1982 (fr.), Teixeira et al. 1213 (holotype SPF, isotypes
CAS, F, K, MO, NY, U).
PERU. San Martin: Prov. Mariscal Caceres, Dtto. Tocache Nuevo, Quebrada de
Cascarilla, NE de Puerto Pizana, 30 July 1973 (bud, fr.), Schunke V 6569 (MO).
HABITAT. Tall forest on terra firme.
This is the only species of Angostura
with unifoliolate leaves. In the absence of
flowering specimens, an assessment of its relationships is impossible.
The type collection (Teixeira
et al. 1213 from Brazil) bears fruits, which I assume
are of mature size because the coat of the seed within is well developed and the
embryo almost fills the seed. The other specimen (Schunke
V 6569 from Peru) bears
one inflorescence with very immature buds, another older inflorescence from which
the corollas have dropped, leaving calyces and slightly enlarged carpels, and one
unattached mericarp, which may not be of mature size because the seeds within are
quite small. Even at this early stage, however, the mericarp of this latter specimen is
larger than that of the type (12 - 14 x 8 vs. 20 x 11.5 mm long). The specimen from
Peru differs further from the type in its considerably larger calyces (7 vs. 3.5 - 4 mm
long) and its carpels that are glabrous (vs. echinoid-pubescent) after anthesis. Given
that the vegetative parts of these two specimens are virtually identical, I am
assuming for the moment that these differences are merely intraspecific.
Angostura trifoliata (Wilid.) T S. Elias in Taxon 19: 575 (1970).
Bonplandia
trifoliata
Willd. in Mem. Acad. Roy. Sci. Hist. Berlin 1802: 27, fig. (1804).
Type: Venezuela, golfe Sta. F6, pros Cumanit, Bonpland 1048 (holotype P, n.v.,
photo from F neg 36819 at NY, isotype P - 2 sheets).
Cuspariafebrifuga
Humb., Essai G6og. P1.
58 (1807), nom. prov.
A. cuspare
Roem. & Schult., Syst. Veg. 4:188 (1819).
Galipea
cusparia
A. St.-Hil. ex DC., Prodr. 1: 731 (1824).
266 KEW
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VOL. 53(2)
B. angustura Spreng., Syst. Veg. 1: 780 (1825).
G. officinalisJ. Hancock in Trans. Med. Bot. Soc. London 1: 25 (1829), illegitimate
substitute for G. cusparia.
Sciuris
officinalis
(J. Hancock) Oken, Allg. Naturgesch. 3(2): 1268 (1841).
C. trifoliata
(Willd.) Engl. in Mart., Fl. Bras. 12(2): 113 (1874).
Portenschlagia
trifoliata
Pohl ex Engl. in Mart., Fl. Bras. 12(2): 100 (1874), pro synon.
G.
febrifuga
(Humb.) Baillon, Traite Bot. Med. Phan. 2: 851 (1884).
B. angostura
Rich. ex B. D. Jacks. in Index Kew. 1: 322 (1893).
C. officinalis (J. Hancock) Engl. in Engl. & Prantl, Nat. Pflanzenfam. 3, Abt. 4: 167
(1896), nom. prov.
C. angostura
(Rich. ex B. D. Jacks.) A. Lyons, P1.
Nam., ed. 2, 153 (1907).
DISTRIBUTION.
Venezuela.
CONCHOCARPUS
ConchocarpusJ. C. Mikan, Del. Fl. Faun. Bras. t. 2 (1820, see Stearn 1956). Type: C.
macrophyllusJ. C. Mikan.
Diglottis
Nees & Mart. in Nova Acta Phys.-Med.
Acad. Caes. Leop.-Carol. Nat. Cur. 11:
151, t. 19, fig. F, t. 25 (1823). Type: D. obovata Nees & Mart. [= C. obovatus (Nees &
Mart.) Kallunki & Pirani].
Lasiostemum Nees & Mart. in Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat.
Cur. 11: 152, t. 19, fig. G, t. 26 (1823). Type: L. silvestre Nees & Mart. [= C. silvestris
(Nees & Mart.) Kallunki & Pirani].
Obentonia
Vell., Fl. Flumin. [text] 1: 18 (1829), [icones] 1: t. 46 (1831). Type: O.
castrata Vell. [= C. macrophyllusJ. C. Mikan].
Dangervilla Vell., Fl. Flumin. [text] 1: 27 (1829), [icones] 1: t. 66 (1831). Type: D.
spinosa
Vell. [= C. longifolius
(A. St.-Hil.) Kallunki & Pirani].
Rossenia Vell., Fl. Flumin. [text] 1: 31 (1829), [icones] 1: t. 77 (1831). Type: R.
pentaphylla Vell. [= C. heterophyllus (A. St.-Hil.) Kallunki & Pirani].
Shrubs or small trees to 7 m tall, usually single-stemmed or with few, orthotropic,
relatively thick branches bearing leaves (often congested) at apex of branch, less often
diffusely branched and leaved. Pubescence
of vegetative parts when present of simple,
sometimes mealy trichomes, usually inconspicuous and deciduous (brown-hirsute
and
persistent in C. hirsutus). Leaves alternate (opposite in only C. oppositifolius), usually 1-
foliolate, sometimes 3- 7-foliolate, rarely apparently simple (i.e., lacking a pulvinus),
usually petiolate, in C. cuneifolius 1-foliolate and sessile (i.e., non-petiolate) but
petiolulate; petiole usually canaliculate or flattened adaxially,
rarely subterete, usually
swollen, rarely not, often geniculate at apex; blade entire, chartaceous to coriaceous.
Inflorescence
a variously modified thyrse, usually terminal or extra-axillary, rarely
axillary, borne usually on leafy stems, rarely (in C. cauliflorus
and C. ramiflorus)
on
leafless stems, the partial inflorescences usually alternately attached along a + elongate
rachis, rarely (in C. guyanensis and C. cuneifolius var. confertus) closely proximate at
apex of long peduncle, cymose, sessile or stalked (i.e., the secondary axis developed),
usually dichasial at first node (and rarely also at second nodes) and shortly
SYNOPSES OF ANGOSTURA & CONCHOCARPUS 267
monochasial at higher nodes, sometimes the monochasial branchlets developing as
scaly short-shoots or as secund racemiform branchlets, sometimes the secondary and
higher order axes often so reduced that the flowers are borne in dense cymules and
the axes are indiscernible. Calyx
of (4) 5 sepals, these quincuncial, usually connate at
base and the lobes not overlapping at anthesis, sometimes free or shortly connate and
overlapping at anthesis, coriaceous, deciduous (sometimes circumscissile) or
persistent. Corolla
? tubular, with spreading or recurved lobes, actinomorphic or
zygomorphic, usually subactinomorphic due to unequal divergence of lobes and
presence of only two fertile stamens, usually creamy white, rarely pink; petals (4) 5, of
imbricate aestivation, usually coherent by pubescence (at least at a point above the
base) forming a tube, rarely coherent only at base and connate at a point at top of
tube, connate at base of tube and coherent above, or free from each other but
adherent to the filaments and still forming a tubular corolla, the outer surface usually
densely sinuate-strigose (? flexuous, appressed, relatively long trichomes) but
glabrous at very base, rarely glabrous on lobes, the inner surface usually tomentulose
(curled, matted trichomes) especially at throat but glabrous at very base, rarely
glabrous on lobes. Androecium
usually of 2 fertile stamens always flanking the
innermost petal and 3 (or in C. macrophyllus
5) staminodes, sometimes of 3 fertile
stamens (of which 2 usually flank the innermost petal and 1 is adjacent) and 2
staminodes or of 5 fertile stamens; filaments usually free from each other (connate
only in C. odoratissimus),
usually free from the corolla at very base but adherent to it at
the throat by the intertwining of their abaxial tomentulose pubescence with that of
the corolla, infrequently also adnate to the corolla either at base or at a point near top
of tube; filaments of the fertile stamens flattened, adaxially glabrous at base and
usually villous (? flexuous, spreading trichomes) at apex, rarely completely glabrous,
not exserted from the corolla tube; the staminodes flattened and usually adaxially
glabrous at base for the length of the corolla tube, usually ? villous at throat, the
exserted apical portion attenuate, subterete or flattened, glabrous or pubescent;
anthers usually free, connate in only C. cuneifolius,
C. nicaraguensis,
and C. ucayalinus,
ovate, oblong, or lanceolate, glabrous or appressed-pubescent, sometimes glandular-
punctate especially on connective, lacking basal appendages (the bases of the thecae
in C. guyanensis sterile), sometimes bearing an apiculum or flattened apical
appendage, narrowly
or less often broadly attached to the filament, not exserted from
the corolla tube; pollen grains 3 - 6-colporate or pantoporate, euprolate, subprolate,
or spheroidal, medium to large, the exine perforate, foveolate, reticulate, or very
densely baculate. Disc cupular, shorter than or equal to the ovary, carnose, glabrous,
the margin entire to crenate, sometimes inflexed and concealing the ovary. Ovary
of
5, free carpels usually united only in the single style (styles free in C. longifolius)
or
rarely (only in C. cuneifolius)
connate axially and basally, apically umbilicate or tapered
into the style, glabrous or pubescent; the stigma enlarged or not, often 5-sulcate and
sometimes finally 5-lobed, usually included in the corolla throat at anthesis. Fruit of 1
- 5 mericarps, these free or rarely (in C. cuneifolius)
connate at base, usually rounded,
sometimes beaked or carinate apically. Seed
1 per carpel, smooth, subrugulose, or
tuberculate, glabrous, the testa papery, leathery, or crustaceous. Embryo usually
curved, less often straight, the radicle adaxial, that of curved embryos apical, that of
straight embryos either lateral or basal, the cotyledons of curved embryos rounded,
268 KEW
BULLETIN
VOL. 53(2)
conduplicate, plicate or not, sub- or unequal, relatively thin and flexible, those of
straight embryos + plano-convex, subequal, thick and stiff.
This genus of 45 species ranges from Nicaragua to northern Bolivia and
southeastern Brazil.
As can be seen from the description, Conchocarpus
is quite variable in most
characters and is composed of the species that do not fit into the other more easily
definable genera. Though variable, Conchocarpus
is delimited by a combination of
character states of the same characters used in the other genera. That combination
comprises: an arborescent or shrubby habit; alternate leaves; a relatively small calyx;
a + tubular corolla formed by coherent or connate petals that are in turn adherent
or adnate to the staminal filaments; flattened staminal filaments; completely fertile,
basally unappendaged anthers; a cupular disc; and free carpels. The few exceptions
are: C. oppositifolius with opposite leaves, C. concinnus and C. cyrtanthus with ? free
calyx lobes, C. guyanensis
with thecae bases that are sterile but still retain the form of
thecae and not of appendages, and C. cuneifolius
with basally and axially connate
carpels. Although these species exhibit these atypical character states, most other
character states place them in Conchocarpus.
KEY TO THE SPECIES OF CONCHOCARPUS
In the following key and descriptions, the term "scaly short-shoot" refers to
branches of the inflorescences of which the nodes (often marked in specimens by
the pedicel scars) and their bracteoles (scales) are contiguous, while "secund
racemiform branch" refers to branches of which the nodes are separated by
minimal but discernible internodes. The presence or absence of an apical
appendage can be determined most easily by observation of the adaxial surface of
the anther. Characters of the ovary often can be observed in the undeveloped
carpels that remain in place after the fall of the corolla and stamens or that persist
at the base of a mature fruiting carpel. Be aware, however, that in some species
(e.g., C. insignis), the ovary is glabrous in flower but becomes pubescent at some
time after the corolla drops. The geographic ranges of the species that occur
outside of eastern Brazil are given in the key in order to make its use more efficient.
1. Leaves 3 - 7-foliolate
2. Inflorescence borne in an axillary position on leafless stems, a short-
pedunculate dichasium with a central flower and eventually two secund
racemiform branches; leaves 3-foliolate [Rio Negro, Brazil, and adjacent
Venezuela] .................................... C. ramiflorus
2. Inflorescence borne on leafy branches, a long-pedunculate, narrow thyrse;
leaves 3 - 7-foliolate
3. Fertile stamens 2 or 3, the anthers narrow-ovate; ovary pubescent in flower;
cotyledons conduplicate, plicate, thin (Fig. 6J); leaves 3 - 7-foliolate
4. Partial inflorescences sessile, congested (axes not visible), the
inflorescence perennating, finally elongate and whip-like; corolla 3.8 -
6.2 mm long; fertile stamens 2; ovary tapered into style; fruit sparsely
appressed-pubescent, glabrescent ............. C. mastigophorus
SYNOPSES OF ANGOSTURA & CONCHOCARPUS 269
4. Partial inflorescences stalked, lax (axes apparent), the inflorescence not
perennating, deciduous after one flowering season; corolla 9 - 20 mm
long; fertile stamens 3; ovary umbilicate; mericarp densely rufous-
hirsutulous (unknown in C. silvestris)
5. Corolla bright yellow; all leaves 3-foliolate ............ C. silvestris
5. Corolla creamy white; most leaves 5-foliolate ...... C. heterophyllus
3. Fertile stamens 5, the anthers linear; ovary glabrous in flower; cotyledons
conduplicate, not plicate, thick; leaves 5 - 7-foliolate
6. Corolla 10 - 15 mm long; anthers 4 - 6.7 mm long; cotyledons very
unequal, the inner one less than one-quarter as long as the outer one
[western Venezuela; Acre, Brazil, and adjacent Peru and Bolivia] ....
...................................... C. transitionalis
6. Corolla 15 - 25 mm long; anthers 7.5 - 14.2 mm long; cotyledons usually
equal, rarely the inner one 1.25 times longer than the outer one
7. Mericarp 18 - 25 x 15 - 16 mm [upland cloud forests of western
Venezuela at 1100 - 2100 m] .................... C. larensis
7. Mericarp 10 - 13 x 10 - 11 mm [lowland forests of Panama, the
Guianas, Amazonian Brazil and Peru] ............ C. toxicarius
1. Leaves simple or 1-foliolate
8. Leaves opposite ................................ C. oppositifolius
8. Leaves alternate
9. Inflorescence borne at base of trunk .................. C. cauliflorus
9. Inflorescence borne on leafy branches
10. Foliaceous (green), suborbicular to narrowly elliptic bracts present at base
of partial inflorescences (Figs. 2A, 6A), or the branchlets of the partial
inflorescences developed as scaly short-shoots (Figs. 2C, 6A, 8H)
11. Leaf-blades oblanceolate, long-cuneate at base, 8 - 52 times longer than
the petiole; anthers connate at anthesis; carpels connate laterally at
base and, before dehiscence, axially also, usually 4 or 5 maturing
12. Partial inflorescences dispersed along rachis, the bracts usually 0.8 -
3.4 x 0.4 - 1.4 cm; corolla 6.9 - 12.2 mm long ..............
............................ C. cuneifolius var. cuneifolius
12. Partial inflorescences clustered near apex of peduncle, the bracts
usually 1.5 - 8.5 x 1.3 - 5 cm; corolla 12.5 - 14 mm long .......
............................ C. cuneifolius var. confertus
11. Leaf-blades elliptic, oblong, or lanceolate (rarely oblanceolate), acute,
obtuse, or rounded at base, 1.4 - 10 times longer than petiole;
anthers free; carpels free, usually only 1 maturing
13. Ovary glabrous; corolla 11 - 24 mm long, pink, white, or lilac,
zygomorphic -......................... C. macrophyllus
13. Ovary pubescent; corolla 4.5 - 16 mm long, white, actinomorphic
14. Ovary tapered into style, the style strigulose like ovary (Figs. 9G,
13K); corolla 10 - 16 mm long; anthers 2, not glandular-
punctate, usually with an apical appendage; foliaceous bracts
subtending partial inflorescences 17- 40 mm long, deciduous
...
................................. C. adenantherus
270 KEW
BULLETIN
VOL. 53(2)
14. Ovary umbilicate, the style glabrous (Figs. 2E, 8K); corolla 4.5 -
8.3 mm long; anthers 3, glandular-punctate, lacking an apical
appendage; foliaceous bracts when present 10 - 26 mm long
15. Leaf-blades 1.5 - 2.3 times longer than petiole; corolla 8 - 8.3
mm long; bracts subtending partial inflorescences
foliaceous, 10 - 26 mm long, deciduous ........ C. bellus
15. Leaf-blades 2.7 - 6 times longer than petiole; corolla 4.8 - 5.8
mm long; bracts subtending partial inflorescences not
foliaceous, 1.5 - 3 mm long, deciduous .... C. diadematus
10. Foliaceous bracts not present, the branchlets of the partial inflorescences
not developed as scaly short-shoots
16. Inflorescence a terminal, long-pedunculate, corymbiform thyrse
bearing 3 or 4 closely proximate partial inflorescences near apex [C.
America; northern S America; Amazonian Brazil and Peru] ......
. ...................................... C. guyanensis
16. Inflorescence a terminal or lateral, long- or short-pedunculate, loose or
dense, + elongate thyrse, bearing partial inflorescences along rachis
17. Calyx in flower of free or shortly connate, usually overlapping and
rounded or obtuse sepals (Figs. 4B, 11B), persistent in fruit;
corolla lobes usually glabrous abaxially (partially pubescent in C.
insignis)
18. Twigs, petioles, and abaxial surface of leaves hirsute with erect,
brownish trichomes ...................... C. hirsutus
18. Twigs, petioles, and abaxial surface of leaves + pubescent with
short, appressed, pale trichomes or glabrescent
19. Inflorescence, fruits, and calyx densely pubescent with
yellowish-grey trichomes (underlying surface not visible) -
.......................................... * C.
insignis
19. Inflorescence and fruits sparsely pubescent and calyx glabrous
or pubescent (but underlying surface visible)
20. Leaves usually wide-oblong or -elliptic, broadly rounded at
base (and often at apex); corolla tube formed by
coherent petals .................... C. concinnus
20. Leaves oblanceolate, usually cuneate, sometimes acute at
base; corolla tube formed by connate petals
21. Inflorescence apparently axillary, often >1 per branch;
internodes of twig obscured by densely clustered leaves
22. Inflorescence axis 1 - 1.5 cm long; corolla 10.4 - 18
mm long; mericarp 18 - 19 x 11 mm [Rio de
Janeiro, Santa Catarina, Minas Gerais] ..........
S..... C. gaudichaudianus subsp. gaudichaudianus
22. Inflorescence axis 5.7 - 8 cm long; corolla c. 6.8 mm
long; mericarp 14 x 7 mm [Bahia] .............
............... C. gaudichaudianus subsp. bahiensis
21. Inflorescence terminal, only 1 per branch; internodes of
twig evident, not obscured by leaves
SYNOPSES OF ANGOSTURA & CONCHOCARPUS 271
23. Anthers free, broadly attached to filament, without an
apical appendage; petals connate into a tube .....
.............0......................... C. cyrtanthus
23. Anthers connate, narrowly attached to filament, with or
without a blunt, puberulent apical appendage; petals
coherent into a tube
24. Anther (including sterile apiculum) 4.5 - 6.8 mm
long, the apiculum 0.3 - 0.9 mm long; mericarp
14 - 20 x 11 - 15 mm, densely appressed-pubescent
[Central America; western Colombia and
Ecuador] ................. C. nicaraguensis
24. Anther (including sterile apiculum when present)
2.5 - 3.4 mm long, the apiculum usually absent,
rarely present and then 0.2 - 0.6 mm long; fruit
11 - 16 x 8 - 13 mm, glabrous or sparsely
appressed-pubescent [northern and Amazonian
South America] .............. C. ucayalinus
17. Calyx in flower cupular, with non-overlapping, acute or obtuse lobes
(Figs. 10H, 13J), rarely entire, deciduous or persistent in fruit;
corolla lobes usually densely pubescent abaxially (sparsely so and
glabrescent in C. grandiflorus)
25. Ovary glabrous
26. Inflorescence appearing axillary, to 1.5 cm long; style c. 20 mm
long, sparsely hirsutulous (at least near apex); filament c. 3.5
times longer than anther ............. C. grandiflorus
26. Inflorescence terminal, 3 - 30 cm long; style to 8 mm long,
glabrous; filament shorter than anther to c. 1.2 times longer
27. Peduncle more than twice as long as the rachis of the
inflorescence; mericarp oblong in long-section; cotyledons
thick, conduplicate, inner much smaller than outer
[Guyana, Amazonian Brazil] ........... C. fanshawei
27. Peduncle equal to or shorter than the rachis of the
inflorescence; mericarp subcircular in long-section;
cotyledons thick, plano-convex, + equal
28. Calyx entire; pedicel 3 - 6 mm long in flower, 8 - 12 mm
long in fruit; corolla c. 16 mm long; mericarp 20 - 25 x
17 - 20 mm [Amazonian Brazil; perhaps Venezuela] ?
....................... 0.......*............ C. grandis
28. Calyx dentate; pedicel 0 - 1.5 mm long in flower, 3.5 - 4
mm long in fruit; corolla 7 - 10 mm long; mericarp c.
10 x 9 mm [French Guiana; upper Amazonian Brazil
and adjacent Bolivia] ............. C. acuminatus
25. Ovary pubescent
29. Ovary umbilicate, the style glabrous (Fig. 8E, K); fertile
stamens 3
272 KEW BULLETIN
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30. Internodes of twig obscured by the densely clustered leaves;
leaf-blade oblanceolate, long-cuneate at base, 44.5 - 64 cm
long, 14 - 45 times longer than petiole ... C. dasyanthus
30. Internodes of twig not obscured; leaf-blade elliptic or
oblong, rounded to acute at base, 8.5 - 36.5 cm long, 2.5
- 6 times longer than petiole
31. Young twigs and vegetative buds densely brownish-
sericeous or -hirsute; lower surface of leaf with
appressed trichomes, tardily glabrescent; corolla 10
mm long, yellow; mericarp visibly hirsutulous ......
.............................. C. marginatus
31. Young twigs and vegetative buds minutely appressed-
pubescent; lower surface of leaf glabrous; corolla 4.5 -
5.8 mm long, creamy white; mericarp inconspicuously
sparsely puberulent .............. C. diadematus
29. Ovary tapered (or in C. longipes abruptly narrowed) into the
style(s), the style pubescent (Figs. 9G, 13K); fertile stamens
usually 2, rarely 5 or 3
32. Leaf-blade short-petiolate (i.e., usually >10 times longer than
petiole), widest above the middle, usually >20 cm long
33. Leaf-blade >80 times longer than petiole, the petiole 0-
0.7 cm long; styles 5, divergent (i.e., each carpel
bearing a separate style); peduncle 2 - 18.5 cm long,
the inflorescence usually 6 - 42 cm long, loosely
flowered, the internodes obvious and secondary axes
developed [northern S America, eastern Brazil] ....
. . . .
.........* ......... ... .. . C. longifolius
33. Leaf-blade 8 - 40 times longer than petiole, the petiole
0.9 - 4 cm long; style 1 (i.e., styles of all carpels
coalesced); peduncle to 1 cm long, the inflorescence 1
- 7.5 cm long, densely flowered, the internodes
obscured and secondary axes not developed
34. Anther appendage two times longer than the thecae
(Fig. 9F); inflorescence 6.5 - 7.5 cm long, borne
near apex of branch; calyx at anthesis slit to near its
base on one side; filaments free ..... C. fissicalyx
34. Anther appendage absent; inflorescence to 3.5 cm
long, appearing axillary along stem; calyx at anthesis
usually not slit to its base; filaments connate (usually
for full length, rarely only in basal 2 mm) .......
..................................... C. odoratissimus
32. Leaf-blade long-petiolate (i.e., <10 times longer than
petiole), usually widest at or below (rarely above) the
middle, 5- 36 cm long
35. Anther with apical appendage (Fig. 3G)
36. Flowers pedicellate ............. C. adenantherus
SYNOPSES OF ANGOSTURA & CONCHOCARPUS 273
36. Flowers sessile
37. Corolla 14 - 15 mm long; mericarp 18 - 25 mm
long; inflorescence 4.5 - 17.5 cm long ........
..... .
....... ...... *....*....... C. macrocarpus
37. Corolla 8- 10 mm long; mericarp 12- 16 mm long;
inflorescence 1 - 5.5 cm long ..... C. obovatus
35. Anther without apical appendage
38. Leaves densely clustered at apex of twig, obscuring the
internodes ................... C. inopinatus
38. Leaves not densely clustered, the internodes of twig not
obscured
39. Partial inflorescences (at least the lower ones with
open flowers) stalked (i.e., the secondary axis
developed) (Figs. 10F, 15A, F), higher order axes
also developed (at least in older inflorescences)
40. Flower buds ovoid (Fig. 10G); blade of most leaves
1.1 - 5 times longer than petiole; branches of
older partial inflorescences not racemiform and
secund
41. Leaf-blades narrow-elliptic, 1.8 - 5 times longer
than petiole; calyx densely mealy-pubescent,
the lobes wider than long; carpels tapered
into the style ............ C. paniculatus
41. Leaf-blades narrow-obovate or oblanceolate, 1.1
- 2 times longer than petiole; calyx glabrous
(or essentially so), the lobes longer than
wide; carpels abruptly narrowed into the
style .................... C. longipes
40. Flower buds cylindrical (Fig. 17B); blade of most
leaves 5 - 13 times longer than the petiole;
branches of older partial inflorescences
racemiform and secund (Figs. 15F, 17G)
42. Anthers glandular-punctate (Fig. 15K); ovary
with ascending trichomes; flower buds 2.7 -
4.3 mm wide; corolla 16 - 20 mm long; part
of staminode free from corolla hirsutulous
................0........... C. punctatus
42. Anthers not glandular-punctate; ovary with
appressed trichomes; flower buds 1.7 - 2.4
mm wide; corolla 14 - 16 mm long; part of
staminode free from corolla glabrous .....
...................... .
* C. santosii
39. Partial inflorescences sessile on main axis (i.e., the
secondary axis not developed) (Figs. 10A, 15B,
17H), higher order axes also not developed
(except in older inflorescences of C.
furcatus)
274 KEW BULLETIN
VOL. 53(2)
43. Inflorescence 1 - 3.5 cm long and the partial inflo-
rescences usually reduced to a single flower and
the pedicel 1.5 - 4 mm long .... C. pentandrus
43. Inflorescence 5.5 - 37.5 cm long or the partial
inflorescences of few to several flowers or the
flowers sessile
44. Fertile stamens 5; ovary with ascending
trichomes; flowers sessile, 1 or a few at each
node; leaf-blade length/width ratio 1.7 - 3.1
............................................ C. ovatus
44. Fertile stamens 2 (3); ovary with appressed
trichomes; flowers sessile or pedicellate, few
to several at each node; leaf-blade
length/width ratio 2.4 - 4.8
45. Inflorescence rachis persistently
clothed with a
dense, dull yellowish-brown pubescence;
abaxial surface of leaf sparsely but
persistently mealy-pubescent - ? C. sordidus
45. Inflorescence rachis glabrous or +
glabrescent; abaxial surface of leaf glabrous
46. Calyx circumscissile (Figs. 2F, 8K); corolla
in bud ovoid, often with obvious
retrorsely spreading trichomes at base, at
anthesis 7 - 9.7 mm long ... C. elegans
46. Calyx persistent or the lobes sometimes
breaking off singly; corolla in bud +
cylindrical, with appressed trichomes at
base, at anthesis 3.5 - 7.5 mm long
47. Mature partial inflorescence a sessile
pair of racemiform, secund branches
(Fig. 10B); pedicel and calyx densely
mealy-pubescent; corolla 7.2 - 7.5
mm long ........... C. furcatus
47. Mature partial inflorescence a cluster of
few to several flowers (Fig. 15B);
pedicel and calyx glabrous or
sparsely pubescent; corolla 3.5 - 6.3
mm long
48. Leaf-blade 3 - 10 times longer than
petiole; inflorescence 3 - 25 cm
long; mericarp transversely ridged
............. C. fontanesianus
48. Leaf-blade 10- 13 times longer than
petiole; inflorescence 2 - 4.5 cm
long; mericarp smooth.......
...................... C. modestus
SYNOPSES
OF ANGOSTURA
& CONCHOCARPUS 275
THE SPECIES OF CONCHOCARPUS
Conchocarpus acuminatus (Pilg.) Kallunki & Pirani comb. nov.
Cusparia
acuminata Pilg. in Verh. Bot. Vereins Prov. Brandenburg 47: 154 (1905).
Type: Brazil, Amazonas, Juru~a
Miry, Rio Jurua, Ule 5805 (holotype Bt, photo
from F neg 12499 at NY; isotypes G, HBG, K).
Angostura
acuminata (Pilg.) Albuq. in Acta Amazon. 11: 850 (1981).
DISTRIBUTION. French Guiana, Brazil (Amazonas, Acre), Bolivia (Beni).
Conchocarpus adenantherus (Rizzini) Kallunki & Pirani comb. nov.
Angostura
adenanthera Rizzini in Revista Brasil. Biol. 50(4): 1085 (1990). Type: Brazil,
Espirito Santo, Reserva de Sooretama, D. Sucre 5515 (holotype RB; isotypes RB-2
sheets, SPF).
DISTRIBUTION. Brazil (Pernambuco, Bahia, Espirito Santo).
Conchocarpus bellus Kallunki sp. nov. a ceteris congeneribus (i.e., C. dasycarpo
et C.
diademato)
staminibus fertilibus tribus, antherae connectivo glanduloso-punctato, et
ovario umbilicato e carpellis strigulosis styloque glabro constanti provisis,
inflorescentia bracteas foliaceas conspicuas ferenti et foliorum lamina quam
petiolum 1.5 - 2.3-ies longiore differt. Typus: Brazil, Espirito Santo, Mun. Rio
Bananal,J. A. Kallunki,
J. R. Pirani &J. B. Fernandes 711 (holotypus SPF, isotypi NY-
2 sheets and 1 to be distributed to MO).
Treelet 1 - 2 m tall, unbranched or with up to 6 orthotropic branches, bearing
congested leaves near apex of stem, the internodes obscured. Leaves 1-foliolate;
petiole 9.2 - 15.5 cm long; blade usually elliptic, sometimes oblanceolate, rounded
at very base, obtuse or (bluntly) acute at apex, 17 - 33 x 5.1 - 9.9 cm, densely
pellucid-punctate, the glands visible on both surfaces; midvein slightly convex
adaxially, a fimbrial vein present. Inflorescence
an erect, spiciform thyrse, extra-
axillary, more than one borne on a stem, 14- 18 cm long including the peduncle,
this 4.7 - 8.5 cm long, the axes puberulent, glabrescent, the partial inflorescences
sessile cymules, with age two apparently monochasial branchlets developing as
short-shoots to 7 mm long; primary bracts foliaceous, 10 - 26 x 2 - 4 mm,
deciduous; pedicels 3 - 5 mm long. Calyx dentate, c. 2.2 mm long, puberulent
without, glabrous within, green (in vivo), circumscissile, the lobes not overlapping
at anthesis, broadly triangular, obtuse, c. 0.6 x 1.5 mm. Corolla actinomorphic,
sinuate-strigose on tube and lobes without, tomentulose within, white; petals 8 - 8.3
x c. 1.9 mm, oblanceolate, coherent into a tube c. 4 mm long, the lobes acute, 4-
4.3 mm long, spreading to recurved, flat. Androecium of 3 fertile stamens and 2
staminodes; filaments adherent to the corolla, those of fertile stamens c. 3.4 mm
long, villous at apex adaxially, the staminodes c. 5.6 mm long, villous in upper half
adaxially, subterete at apex; anthers oblong, c. 2.2 x 1.1 mm, hirsutulous mainly
adaxially, conspicuously glandular-punctate on connective, unappendaged at base,
narrowly attached to the filament. Disc c. 1 mm high, equal to the ovary. Ovary
of 5
276 KEW BULLETIN VOL. 53(2)
carpels free except in the style, c. 0.9 mm high, umbilicate, densely strigulose; style
c. 1.5 mm long (including stigma), glabrous; stigma not exserted from the throat at
anthesis. Fruit of 1 mericarp, 7.5 - 10 x 5.5 - 7 mm, puberulent, transversely ridged,
rounded apically. Seed unknown. Fig. 2.
DISTRIBUTION. Brazil (Espirito Santo); known only by the type collection.
BRAZIL. Espirito Santo: Rio Bananal, Fazenda Sao Jorge, 18 km from church in
Bananal on paved road from Bananal to Novo Brasil, behind house, c. 19012' -
19013'S, 40025' - 40026'W, 21 April 1995 (fl.), Kallunki et al. 711 (holotype SPF,
isotypes MO, NY).
HABITAT. Disturbed forest remnant.
This new species is most similar to C. dasyanthus
and C. diadematus;
they are the
only three species in the genus that have the combination of a fimbrial vein on the
leaf blade, a circumscissile calyx, three fertile stamens with glandular-punctate
anthers, and an umbilicate ovary with pubescent carpels and glabrous style. The
new species is quite distinct, however, in its conspicuously bracteate inflorescences
and its leaves borne on relatively long, subterete petioles. In the new species, the
blade is 1.5 - 2.3 times longer than the petiole, while in C. dasyanthus it is 14 - 44
times longer and in C. diadematus it is 2.7 - 6 times longer. It is distinguished further
from C. diadematus
by its larger flowers, of which, for example, the petals are 8 - 8.3
(vs. 4.5 - 5.8) mm long, and by its leaves which are clustered at the apex of the stem,
obscuring the internodes, rather than distributed along the stem such that
internodes are obvious.
Conchocarpus cauliflorus Pirani sp. nov. ab omnibus congeneribus inflorescentiis
ad basem caulis congestis optime distincta. Typus: Brazil, Epirito Santo, Rio
Bananal, J. A. Kallunki,
J. R. Pirani &J. B. Fernandes 690 (holotypus SPF;
isotypus NY
and 1 to be distributed to MO).
Treelet 0.6 - 1.6 m tall, to 1.5 cm dbh, unbranched or with 1 - 3 ascending
branches, bearing congested leaves near apex of stem, the internodes obscured.
Leaves 1-foliolate; petiole 5.5 - 16 (20) cm long; blade narrow-elliptic to
oblanceolate, cuneate or acute at base, acute and sometimes shortly apiculate at
apex, (15) 19 - 32 (40) x (2.5) 4.8 - 10 cm, conspicuously pellucid-punctate, the
glands visible or not on both surfaces; midvein raised adaxially, a fimbrial vein
absent. Inflorescence
a short, racemiform thyrse, borne in clusters near base of
trunk, 2.5 - 11 cm long including the peduncle, this very short or absent, the axes
densely pubescent, glabrescent, the partial inflorescences sessile, 1 - 3-flowered;
primary bracts 1.8 - 2.5 mm long, deciduous; pedicels c. 3 mm in flower, to 8 mm
in fruit. Calyx lobed, 2 - 2.6 mm long, appressed-pubescent without, glabrous
within, green to dark red (in vivo), persistent in fruit, the lobes not overlapping at
anthesis, triangular, acute, 1.2 mm x 0.8 - 1.3 mm wide. Corolla
actinomorphic,
densely sinuate-strigose without, sparsely tomentulose within, creamy white at
anthesis, pink on lobes in bud; petals 10 - 13 x 2 - 2.4 mm, narrow-oblong,
coherent forming a tube c. 4 - 5.5 mm long, the lobes acute, 6 - 9 mm long,
spreading, flat but with a slightly raised midvein. Androecium
usually of 2 fertile
SYNOPSES OF ANGOSTURA & CONCHOCARPUS 277
FIG. 2. Conchocarpus
bellus. A apex of stem; B part of inflorescence; C older partial inflorescence showing short -
shoots; D flower; E gynoecium and disc with part of calyx removed; F calyx with circumscissile abscission; G
stamen, adaxial and abaxial views; H staminode; J fruit. All from Kallunki et al. 711. Drawn by Bobbi Angell.
278 KEW BULLETIN
VOL. 53(2)
stamens and 3 staminodes or seldom of 1 fertile stamen and 4 staminodes;
filaments adherent to the corolla, those of fertile stamens c. 3 - 3.9 mm long,
sparsely villous at apex adaxially, the staminodes 9 - 11 mm long, sparsely villous at
throat adaxially, subterete and hirsutulous in apical half; anthers oblong, 2.3 - 2.9
x c. 1 mm, glabrous, yellow, eglandular on connective, unappendaged at base,
narrowly attached to the filament, bearing a flattened, obtuse, puberulent, apical
appendage 1 mm long exserted from the throat at anthesis. Disc 1.5 - 1.6 mm high,
taller than the ovary. Ovary of 5 carpels free except in the style, c. 1 mm long,
tapered into the style, densely short-sericeous; style 1.2 - 1.4 mm long (including
stigma), densely short-sericeous; stigma not exserted from the throat at anthesis.
Fruit of 1 or 2 free mericarps, each 12 - 15 x 8 - 10 mm, sparsely appressed-
pubescent, transversely ridged, ventrally apiculate. Seeds not known. Seedling
phanerocotylar, the cotyledons orbicular, cordate at base, rounded at apex. Fig. 3.
DISTRIBUTION. Brazil (Espirito Santo); known only from the type locality.
BRAZIL. Espirito Santo: Rio Bananal, forest behind Fazenda Lagoa Nova,
reached by dirt road turning left (S) at 24 km from bridge at W edge of Linhares on
road to Bananal, c. 19017'S, 40012'W, 20 Feb. 1994 (fl., fr.), Kallunki et al. 590 (NY,
SPF), 19 April 1995 (fl.), Kallunki et al. 690 (holotype SPF, isotypes MO, NY- 2
sheets), 5 Dec. 1994 (fl.), Pirani et al. 3455 (NY, SPF).
HABITAT. Disturbed forest.
Conchocarpus cauliflorus is similar to C. obovatus and C. macrocarpus
in the structure
of the spiciform inflorescence, i.e., the mature partial inflorescences are sessile
cymules of I - 3 flowers, and in the presence of a flattened apical appendage on the
anther. It is easily distinguished from these and all other congeners because its
inflorescences are borne in clusters near the base of the trunk, rather than on leafy
branches.
Conchocarpus concinnus Kallunki sp. nov. ab omnibus congeneribus foliis late
oblongis vel ellipticis basi apiceque rotundatis differt. Typus: Brazil, Bahia,
Olivenga, J. R. Pirani &J. A. Kallunki
2743 (holotypus SPF;
isotypi CEPEC, NY, and 6
to be distributed to CAS, K, MBM, MO, RB, U).
Shrub or treelet 0.4 - 3 m tall, to 2 cm dbh, unbranched, bearing leaves near
apex of stem, the internodes evident. Leaves
1-foliolate; petiole 3.7 - 17 cm long, as
small as 0.7 cm long on the small leaves noted below; blade usually wide-oblong or
wide-elliptic, sometimes oblong, rarely ovate, rounded to (sub)cordate at base,
rounded or obtuse and usually retuse at apex, 13 - 19.5 x 7 - 12.5 cm, rarely (on
sprouts from cut stems) as small as 4.4 cm long and 2.4 cm wide, not pellucid-
punctate, the glands visible as minute warts on adaxial surface of older leaves and
as brownish dots on abaxial surface; midvein plane or subcanaliculate adaxially, a
fimbrial vein absent. Inflorescence
an erect, cylindrical thyrse, terminal, becoming
lateral and leaf-opposed after development of the bud in the axil of the opposed
leaf, sometimes more than one borne on the stem, 3.2 - 15 cm long including the
peduncle, this 2 - 7.2 cm long, the axes hirsutulous, the partial inflorescences
stalked, dichasial at first and second nodes, briefly monochasial at higher nodes,
SYNOPSES OF ANGOSTURA & CONCHOCARPUS 279
FIG. 3. Conchocarpus
cauliflorus. A habit, showing cauline inflorescences; B apex of stem; C old inflorescences at
base of stem; D inflorescence; E flower, lateral and frontal views; F opened corolla showing androecium; G stamen,
abaxial and adaxial views; H disc and gynoecium with part of calyx removed and gynoecium with longitudinal
section of disc; J fruit. A, C from Kallunki et al. 590; B, D -J from Kallunki et al. 690. Drawn by Bobbi Angell.
280 KEW BULLETIN VOL.
53(2)
the secondary axis 2 - 4 mm long; primary bracts very early deciduous (only scars
seen); pedicels 1.5 - 2.3 mm long in flower, to 4 mm in fruit. Calyx
of free sepals,
these overlapping at anthesis, sparsely appressed-pubescent without especially near
margins, appressed-pubescent within, ciliate, creamy white with pale green apex
(in vivo), persistent in fruit, the outer ones orbicular, 1.3 - 1.5 x 1.5 - 1.7 mm, the
inner ones oblong, rounded, 1.2 - 1.5 x 1.1 mm. Corolla
subactinomorphic, with
innermost petal in upper position, densely sericeous on tube and glabrous on
lobes without, tomentulose at throat and glabrous on lobes within, creamy white;
petals 5.4 - 6.5 x 1.2 - 1.5 mm, narrow-oblong, coherent into a tube c. 2 mm long,
the lobes rounded, 3.4 - 4.5 mm long, spreading, the innermost folded
longitudinally, cucullate at apex, the others flat. Androecium
of 2 fertile stamens and
3 staminodes; filaments adherent to the corolla, those of fertile stamens 2 - 2.2 mm
long, villous at apex adaxially, the staminodes 3.5 - 4.6 mm long, villous adaxially,
subterete and glabrous at apex; anthers narrow-ovate or ovate, 1.5 - 1.7 x 0.7 - 0.9
mm, glabrous, eglandular on connective, unappendaged at base, broadly attached
to the filament; pollen 3-colporate, subprolate, reticulate. Disc 0.5 - 0.6 mm high,
equal to the ovary. Ovary
of 5 carpels free except in the style, 0.5 - 0.6 mm high,
umbilicate, glabrous in flower but soon sparsely appressed-pubescent; style 1.1 -
1.6 mm long (including stigma), glabrous; stigma not exserted from the throat at
anthesis. Fruit of 1 - 5 free mericarps, each 10 - 11 x 8 - 9 mm, sparsely appressed-
pubescent, transversely ridged, rounded apically. Seed
subrugulose, glabrous, c. 7
mm x 5 mm, the testa papery. Embryo straight, with adaxial radicle, the cotyledons
plano-convex. Seedling
phanerocotylar, the expanded cotyledons ovate, cordate at
base, obtuse at apex. Fig. 4.
DISTRIBUTION.
Brazil (Bahia); known only from the vicinity of the type locality.
BRAZIL. Bahia: Mun. Ilheus, 5 - 6 km SW de Olivenga, na estrada que liga
Olivenga ao povoado do Maruim, 29 July 1993 (fl., fr.), Jardim et al. 254 (CEPEC,
NY, SPF), 6 km from OlivenCa on road to Serra das Trempes, 14'45'S, 39002'W, 30
March 1995 (fl., fr.), Kallunki et al. 686 (CEPEC, NY), OlivenCa, estrada que liga a
Estad?o Hidromineral de Olivenca ao Povoado de Vila Brasil, 5 km ao SW de
Olivenoa, 8 Feb. 1982 (fl.), Mattos Silva et al. 1480 (CEPEC, NY, SPF) & 26 April
1983 (fl.), Mattos Silva et al. 1690 (CEPEC), antiga estrada que liga Oliven4a ao
Povoado de Vila Brasil, 8 km a SW de Olivenca, 14 Jan. 1985 (fl.), Mattos Silva et al.
1822 (CEPEC, NY, SPF), a 6 km a partir da Igreja de Olivenga na estrada para
Maruim, approx. 14'57'S, 39'00'W, 6 Feb. 1993 (fl., fr.), Pirani & Kallunki 2743
(holotype SPF; isotypes CAS, CEPEC, K, MBM, MO, NY, RB, U), estrada vicinal de
Olivenga a Buerarema, 6 km de Olivenga, 12 Feb. 1994 (fl.), Pirani et al. 2943 (NY,
SPF), road from Olivenga to Serra das Trempes, 6 km from Olivenca, 39003'W,
14'59'S, 1 Feb. 1992, Thomas
et al. 9015 (CEPEC, NY).
HABITAT. Disturbed restinga forest.
This species is one of five in the genus with anthers broadly attached to the
filament, almost free sepals that still overlap at anthesis, and, where known, plano-
convex cotyledons. This species is most easily recognized by its distinctive leaves,
which are usually oblong or elliptic and rounded at both ends. In the other four
species, the petals are connate rather than coherent.
SYNOPSES
OF ANGOSTURA & CONCHOCARPUS 281
FIG. 4. Conchocarpus concinnus. A apex of stem; B flower, frontal and lateral views; C calyx, disc, and ovary in
longitudinal section, with detail of ciliate sepal margin; D opened corolla showing androecium; E staminode and
stamen, adaxial view; F disc and gynoecium; G fruit; H seed; J embryo; K young plant with the two cotyledons and
three leaves. A - G, K from Pirani & Kallunki 2743; H, J from Jardim et al. 254. Drawn by Bobbi Angell.
282 KEW BULLETIN
VOL. 53(2)
Conchocarpus cuneifolius Nees & Mart. in Nova Acta Phys.-Med. Acad. Caes.
Leop.-Carol. Nat. Cur. 11: 162, t. 21 (1823). Type: Brazil, Bahia, ad Engenho da
Ponta, Martius s.n. (holotype M). This is the only sheet of this species collected by
Martius that we have seen. The protologue states the habitat as "in sylvis prope ab
oppido Caxoeira ad Peruagua<ii fluvium, et ad S. Georgii oppidum in Comarca
dos Ilheus Provinciae Bahiensis." Because we have seen no specimen labelled with
these locality names and because Engenho da Ponta is near Cachoeira and Sao
Jorge (see Martius's itinerary in Urban 1906: 59), we take the collection cited
above to be the holotype.
Galipea
cuneifolia
(Nees & Mart.) A. St.-Hil. ex DC., Prodr. 1: 732 (1824).
Bonplandia cuneifolia
(Nees & Mart.) Spreng., Syst. Veg. 1: 781 (1825).
Cusparia cuneifolia
(Nees & Mart.) Engl. in Mart., Fl. Bras. 12(2): 116 (1874).
Angostura
cuneifolia
(Nees & Mart.) Albuq. in Acta Amazon. 11: 850 (1981).
DISTRIBUTION. Brazil
(Bahia,
Espirito
Santo).
Conchocarpus cuneifolius var. confertus Kallunki var. nov. a var. cuneifolio
inflorescentiis partialibus omnibus sessilibus, squamulosis, ad apicem pedunculi
elongati confertis, unaquaque bractea foliacea subtenta differt. Typus: Brazil, Bahia,
Sao Jos , J. A. Kallunki,
J. R. Pirani, W W Thomas & T S. dos Santos
425 (holotypus
SPF, isotypi CEPEC, K, NY and 1 to be distributed to MO).
Shrub to 0.75 m tall, unbranched, bearing congested leaves near apex of stem, the
internodes obscured. Leaves
1-foliolate; petiole none, the petiolule 1 - 1.5 cm long,
swollen at base; blade oblanceolate, long-cuneate at base, obtuse or rounded at apex,
46 - 51.5 x 10.5 - 13.8 cm, pellucid-punctate, the glands obscure on both surfaces;
midvein convex adaxially, a fimbrial vein absent. Inflorescence
a long-pedunculate,
ascending, much-modified thyrse, extra-axillary,
more than one borne on a stem, 20 -
29.5 cm long, the partial inflorescences congested, sessile, conspicuously bracteate,
clustered at apex of glabrous peduncle, the secondary axis none, with age apparently
monochasial branchlets developing as scaly short-shoots to 25 mm long; primary
bracts foliaceous, (9) 15 - 85 x (7) 13 - 50 mm, persistent; pedicels 3 - 3.5 mm long in
flower, 4 - 4.2 mm long in fruit. Calyx
lobed, c. 2 mm long, glabrous or puberulent
without, glabrous within, persistent, the lobes not overlapping at anthesis, triangular,
obtuse, 1 - 1.5 x 1 - 1.5 mm. Corolla
zygomorphic, with innermost petal in upper
position and reflexed, the other four recurved below, glabrous (except at margins of
petals at base) without, tomentulose in tube within, white; petals 12.5 - 14 x 2.2 - 3.2
mm, oblanceolate, barely coherent into a tube c. 4.5 mm long, the lobes oblong,
rounded, 8 - 9.5 mm long, flat. Androecium of 2 fertile stamens and 3 staminodes;
filaments barely adherent to the corolla, those of fertile stamens c. 5.6 mm long,
villous near apex adaxially, the staminodes 12.8 - 13 mm long, hirsutulous for whole
length adaxially, subspathulate; anthers connate initially, separated and reflexed after
dehiscence, narrow-oblong, c. 3.3 x 0.6 mm, minutely spiculate, eglandular on
connective, unappendaged at base, narrowly attached to the filament; pollen
pantoporate, spheroidal, very densely baculate (Fig. 5A, B). Disc c. 0.8 mm high,
slightly shorter than the ovary. Ovary
of 5 carpels connate at base and axially, c. 0.9
SYNOPSES OF ANGOSTURA & CONCHOCARPUS 283
FIG. 5. Pollen grains and details of exine morphology (SEM). A, B Conchocarpus cuneifolius var. confertus
(Kallunki et al. 425); A pantoporate grain; B one aperture. C C. furcatus (Sucre & Soderstrom
8921); equatorial
view of grain. D - F C. cyrtanthus (Thomas et al. 9747); D equatorial view of grain; E oblique polar view showing
four apertures; F foveolate - perforate exine. G C. inopinatus (Kallunki et al. 433); oblique polar view of grain
showing four apertures. H, J C. insignis (Pirani et al. 3052); H equatorial view of grain; J oblique polar view
showing four apertures. A, D: scale bar = 10 pm; B, F: scale bar = 2 pm; C, E, G, H: scale bar = 5 pm; J: scale
unknown. Drawn by Bobbi Angell.
mm high, umbilicate, glabrous; style c. 5.2 mm long (including stigma), glabrous;
stigma exserted from the throat at anthesis. Fruit of 3 - 5 mericarps connate at base,
each 7- 8 x 6- 7 mm, glabrous, transversely
shallowly ridged, rounded apically. Seed
tuberculate, glabrous, c. 4.2 x 3 mm, the testa crustaceous. Embryo
curved, the
cotyledons conduplicate and plicate, subequal. Seedling
phanerocotylar, the expanded
cotyledons orbiculate, truncate or emarginate at base, rounded at apex. Fig. 6.
DISTRIBUTION.
Brazil (Bahia); known only from the type locality.
BRAZIL.
Bahia: Mun. Una, 9 km from Sao Jos6 and BR-101 on road to Una, c.
15003'S, 39014'W, 29 Jan. 1993 (fl., fr.), Kallunki et al. 425 (holotype SPF, isotypes
284 KEW BULLETIN VOL.
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FIG. 6. Conchocarpus
cuneifolius var. confertus.
A apex of stem; B flower, frontal and lateral views, and floral bud; C
filament, recurved after loss of anther; D fertile stamens, abaxial view; E disc and gynoecium; F young plant and
seedling; G seed; H fruit, after and before dehiscence; J embryo, lateral and adaxial views. All from Kallunki et
al. 425. Drawn by Bobbi Angell.
SYNOPSES OF ANGOSTURA & CONCHOCARPUS 285
MO, NY), estrada Sdo Jos6/Una, km 9 a partir da BR, Fazenda Santa Rosa, 6 Oct
1992 (fl., fr.), Amorim et al. 833 (U).
HABITAT. Disturbed Southern Bahian wet forest on steep, rocky slope.
The flowers of this variety are larger and more zygomorphic than those of the
typical variety, but the most striking difference is found in the inflorescence. In
both, the partial inflorescences are bracteate, sessile, long-lived cymes of which
some branchlets develop as short-shoots. The partial inflorescences of the typical
variety are well spaced along the rachis and subtended by bracts that measure 8 -
34 x 4 - 14 mm, while those of var. confertus are congested at the apex of the
peduncle and subtended by usually longer and much wider bracts that measure 15
- 85 x 13 - 50 mm.
Conchocarpus cyrtanthus Kallunki sp. nov. C. insigni simillima, sed ab eo foliis
angustioribus plerumque obtusis, inflorescentia breviori, et corolla majori tubo
longiori arcuato provisa differt. Typus: Brazil, Bahia, Itacare, J. R. Pirani &J. A.
Kallunki
2748 (holotypus SPF; isotypi NY, SPF, 1 to be distributed to MO).
Treelet 1 - 6 m tall, to 2 cm dbh, unbranched or 1-branched, bearing leaves
near apex of stem, the internodes evident. Leaves 1-foliolate; petiole 1.8 - 11.5
cm long; blade oblanceolate, rarely narrow-elliptic, acute and sometimes cuneate
at base, rounded or obtuse (rarely acute) at apex, 14 - 48.5 x 3 - 9 cm, not or
inconspicuously pellucid-punctate, no glands visible on either surface; midvein
plane to shallowly canaliculate adaxially, a fimbrial vein absent. Inflorescence
an
erect, narrow thyrse, terminal, becoming lateral and leaf-opposed after
development of the bud in the axil of the opposed leaf, (3.5) 4.5 - 15.8 cm long
including the peduncle, this 1 - 3 cm long, the axes + pubescent, the partial
inflorescences dichasial at first node, briefly monochasial at higher nodes, the
secondary axis 0 - 5 mm long; primary bracts 4.5 - 6 mm long, deciduous before
anthesis; pedicels 1 - 4 mm long in flower. Calyx
of free sepals or deeply lobed to
1 mm from base, glabrous or appressed-pubescent without and within, ciliolate,
creamy white (in vivo), persistent in fruit, the sepals or lobes ? overlapping at
anthesis, the outer ones ovate, rounded, 2.5 - 6.5 x 2.2 - 4 mm, the inner ones +
elliptic, rounded, 2.2 - 5.5 x 1.5 - 3 mm. Corolla zygomorphic, with innermost
petal in upper position, sinuate-strigose on tube and glabrous (or sparsely
pubescent) on lobes without, tomentulose on throat and glabrous on lobes
within, bright green in bud, creamy white at anthesis; petals 15.5 - 18 mm long,
connate into a curved tube 10 - 11 mm long, the upper lobe folded
longitudinally, cucullate, 2.6 - 3.2 mm wide (when unfolded), ascending, the
other lobes oblong, rounded, 1.8 - 3 mm wide, recurved below, flat. Androecium
of 2 fertile stamens and 3 staminodes; filaments adnate to the corolla, those of
fertile stamens 6.5 - 9 mm long, short-villous at apex adaxially, the staminodes
9.5 - 11 mm long, villous adaxially, subterete and glabrous at apex, white; anthers
lanceolate, 3.5-4.3 x 1- 1.5 mm, glabrous, eglandular on connective,
unappendaged at base, broadly attached to the filament, white; pollen 3 - 4-
colporate, euprolate, foveolate-perforate (Fig. 5D - F). Disc 0.6- 1.3 mm high,
the margin inflexed and covering the ovary. Ovary
of 5 carpels free except in the
286 KEW BULLETIN
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style, 0.7 - 1 mm high, umbilicate, glabrous; style 4.5 - 7 mm long (including
stigma), glabrous; stigma not exserted from the throat at anthesis. Fruit of usually
2 free mericarps, each 10 - 12 x c. 10 mm, sparsely appressed-pubescent,
transversely ridged, rounded apically. Seed
unknown. Fig. 7.
DISTRIBUTION. Brazil (Bahia, Espirito Santo).
BRAZIL.
Bahia: without additional locality, Luschnath s.n. (NY); Mun. Almadina,
Rod. Almadina/Ibitupt, entrada a 7 km Serra do Sete Paus, c. 12 km da entrada,
Faz. Cruzeiro do Sul, 14044'06"S, 39041'46"W, 578 m, 28 Feb 1997 (fl.), Jardim et al.
1028 (NY); Mun. Buerarema, on road to Olivenca at 4.2 km from its junction with
the Buerarema-Vila Brasil road at c. 7 km from Buerarema, 30 March 1995 (bud),
Kallunki et al. 688 (CEPEC, NY); Mun. Itacare, Itacare, 9 Jan. 1967 (fl.), Belem &
Pinheiro 3026 (CEPEC, US), c. 3 km S da cidade em estrada vicinal partindo da
Rodovia Itacar&-Ubaitaba 49 - 50 km da BR101, S da foz do Rio de Contas, c.
14020'S, 39000'W, 6 Feb. 1993 (fl.), Pirani & Kallunki 2748 (holotype SPF; isotypes
MO, NY, SPF); Mun. Una, estrada Oliven-a/Vila Brasil, km 33, Maruim, Faz. 2 de
Julho, 13 May 1981 (fl.), Carvalho
et al. 681 (CEPEC), Estrada Olivenga-Buerarema,
37 km do inicio da estrada ndio-pavimentada
em Olivenga, 10 Feb. 1994 (fl.), Pirani
et al. 2921 (SPF); Mun. Uruguca, 7.3 km N of Serra Grande on road to Itacare,
14025'S, 39001'W, 5 Feb. 1993 (fl.), Thomas
et al. 9747 (CEPEC, NY, SPF). Espirito
Santo: Mun. Linhares, Reserva Florestal da Cia. Vale do Rio Doce, 27 Sept. 1978
(fr.), Martinelli 4979 (RB) & 7 Feb. 1986 (fl.), Farias 108 (NY) & 31 May 1993 (fr.),
Farias 608 (NY, SPF).
HABITAT. Southern Bahian wet forest.
Among the species with anthers broadly attached to the filaments (i.e., C.
concinnus, C. gaudichaudianus, C. hirsutus, and C. insignis), C. cyrtanthus is most
similar to the last. The two species share large oblanceolate leaves that usually dry
greyish green adaxially, terminal inflorescences, connate petals, and a disc +
inflexed at margin and concealing the ovary. It differs from C. insignis in its shorter
(3.5 - 15.8 vs. 15 - 25 cm) inflorescences, glabrous or sparsely pubescent (vs.
densely sericeous) sepals, and larger zygomorphic corollas (15.5 - 18 vs. 11 - 13 mm
long) with a curved (vs. straight) tube.
Conchocarpus dasyanthus Kallunki sp. nov. foliis magnis oblanceolatis, calyce
circumscissili, et antherae connectivo glanduloso-punctato C. pentagyno
similis, sed
ab eo infructescentia angustiore, staminibus fertilibus 3 (nec 2), ovario umbilicato
stylo unico proviso (nec acuto in stylos 5 attenuato), et fructu minore differt. Typus:
Brazil, Bahia, Mun. Itabuna, S. A. Mori, J. A. Kallunki & T. D. Pennington 9579
(holotypus NY; isotypus CEPEC).
Treelet 1.5 - 2 m tall, unbranched, the internodes obscured. Leaves 1-foliolate;
petiole 1 - 4.5 cm long; blade (narrow-) oblanceolate, long-cuneate and usually
ultimately obtuse or rounded at base, usually obtuse at apex, 44.5 - 64 x 9 - 11.2
cm, markedly pellucid-punctate, the glands visible as dark dots on both but
especially abaxial surface; midvein plane or convex adaxially, a fimbrial vein
present. Inflorescence
an ascending, narrow thyrse, extra-axillary, sometimes more
SYNOPSES OF ANGOSTURA & CONCHOCARPUS 287
FIG. 7. Conchocarpus cyrtanthus. A apex of stem; B flower, lateral and frontal views; C opened corolla showing
androecium, and disc and gynoecium with part of calyx removed; D anther, adaxial and abaxial views; E apex of
staminode; F disc and ovary in longitudinal section and intact ovary; G fruit. A, G from Pirani & Kallunki 2748;
B - F from Thomas et al. 9747. Drawn by Bobbi Angell.
288 KEW BULLETIN VOL.
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than one borne on the stem, 15.5 - 43 cm long including the peduncle, this 7.2 -
19.5 cm long, the axes densely puberulent, less so with age, the partial
inflorescences stalked or the upper ones sessile, dichasial at first node, briefly
monochasial at higher nodes, the secondary axis to 6 mm long; primary bracts 2 -
3 mm long, persistent through flowering but finally deciduous; pedicels 1.5 - 4 mm
long in flower and fruit. Calyx dentate, 1.2 - 2 mm long, sparsely appressed-
pubescent without, glabrous within, circumscissile, the lobes not overlapping at
anthesis, triangular, acute, c. 0.5 x 1.2 mm. Corolla
actinomorphic, densely sinuate-
strigose on tube and appressed-pubescent on lobes without, densely tomentulose
within, white; petals 6 - 8 x c. 1.7 mm, oblanceolate, coherent into a tube c. 4.5
mm long, the lobes obtuse, c. 3.5 mm long, recurved, flat. Androecium of 3 fertile
stamens and 2 staminodes; filaments adherent to the corolla, those of fertile
stamens c. 4.5 mm long, densely villous at apex adaxially, the staminodes c. 6 mm
long, densely villous in upper half adaxially, subterete at apex; anthers narrow-
ovate, c. 2.2 x 0.9 mm, villous, glandular-punctate and glabrous on connective,
unappendaged at base, narrowly attached to the filament; pollen 4-colporate,
euprolate, foveolate (see fig. 3D in Morton & Kallunki 1993). Disc 0.8 - 1.2 mm
high, equal to the ovary. Ovary of (4) 5 carpels free except in the style, c. 1 mm
high, umbilicate, densely strigulose; style 2 - 2.6 mm long (including stigma),
glabrous; stigma not exserted from the throat at anthesis. Fruit of 1 or 2 free
mericarps, each c. 9 x 6 - 8 mm, glabrous, transversely ridged, rounded apically.
Seed rugulose, glabrous, c. 6.6 x 5 mm wide, the testa papery. Embryo
curved, the
cotyledons conduplicate and plicate, subequal. Fig. 8A - E.
DISTRIBUTION.
Brazil (Bahia); known only from the type locality.
BRAZIL. Bahia: Mun. Itabuna, c. de 3 km a NW de Ju-ari, 8 March 1978, Mori et
al. 9572 (CEPEC, NY, RB) & 9 March 1978 (fl.), Mori et al. 9579 (holotype NY;
isotype CEPEC) & 6 May 1978 (fr.), Mor & Kallunki
10069 (CEPEC,
NY).
HABITAT. Southern Bahian wet forest; 200 m elevation.
The large, oblanceolate leaves of this new species are similar to those of C.
longifolius, with which it also shares a circumscissile calyx and glandular-punctate
anthers, but it differs from that species in its three (vs. two) fertile stamens,
umbilicate ovary with a single style (vs. carpels tapered into separate styles), and
smaller (9 vs. 13 - 15 mm long) fruit. Although their leaves are quite dissimilar, it
shares with C. diadematus not only a circumscissile calyx and a glandular-punctate
anther connective, but also only three fertile stamens and an umbilicate ovary with a
glabrous style. It differs from that species, however, in its shorter, stouter petiole (1 -
4.5 vs. 3.2 - 9.5 cm long), its longer (44 - 64 vs. 13 - 32.5 cm), oblanceolate (vs.
oblong or elliptic) leaf blade, and its longer (8 vs. 4.5 - 5.8 mm) petals.
FIG. 8. A- E Conchocarpus dasyanthus. A apex of stem; B part of inflorescence; C stamen, adaxial and abaxial
views; D staminode, lateral view; E disc and gynoecium. F - N C. diadematus. F apex of branch; G part of
inflorescence; H older partial inflorescence with short-shoots; J flower; K calyx with circumscissile abscission and
disc and gynoecium with part of calyx removed; L stamen, abaxial and adaxial views; M staminode; N seed. A
from Mori & Kallunki 10069; B - E from Mori et al. 9579; F, H, N from Kallunki et al. 685; G, J - M from Pirani et
al. 2968. Drawn by Bobbi Angell.
SYNOPSES OF ANGOSTURA & CONCHOCARPUS 289
290 KEW BULLETIN
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Conchocarpus diadematus Pirani sp. nov. calyce circumscissili, staminibus fertilibus
3, antherae connectivo glanduloso-punctato, atque ovario umbilicato C. dasyantho
similis, sed ab eo foliis oblongis vel ellipticis (nec oblanceolatis) atque petalis
minoribus differt. Typus: Brazil, Bahia, Ilheus, J. R. Pirani &J. A. Kallunki 2744
(holotypus SPF;
isotypi CEPEC, HUEFS, NY, and 7 to be distributed to CAS, K, MO,
MBM, RB, SP, U).
Shrub or treelet 1 - 4 m tall, to 4 cm dbh, branched or not, bearing leaves along
much of the stem, the internodes evident. Leaves
1-foliolate;
petiole (1.5) 3.2 - 9.5 cm
long; blade oblong to elliptic, usually acute or obtuse, seldom rounded at base, usually
obtuse to rounded or sometimes acute at apex, (8.5) 13 - 36.4 x (4) 5 - 12 cm,
pellucid-punctate, the glands visible as dark dots on only abaxial surface; midvein
canaliculate or sometimes slightly raised on distal 1/3 adaxially,
a fimbrial vein present.
Inflorescence
an erect, cylindrical thyrse, extra-axillary, produced near base of new
shoots, often more than one borne on a stem, 6.5 - 23 (33) cm long including the
peduncle, this 2.5 - 6 cm long, the axes minutely appressed-pubescent,
less so with age,
the partial inflorescences sessile cymules, sometimes with age two apparently
monochasial branchlets developing as short-shoots to 10 mm long; primary bracts 1.5 -
3 mm long, usually early deciduous, rarely persistent; pedicels 0.5 - 2 mm long. Calyx
dentate, c. 1.4 mm long, pubescent without, sparsely puberulous within, ciliate, pale
green (in vivo), circumscissile, the lobes not overlapping at anthesis, ovate, obtuse, 0.6
- 0.8 x c. 1.2 mm. Corolla
actinomorphic, with the two petals flanked by the three fertile
stamens in upper position, densely sinuate-strigose on both tube and lobes without,
densely tomentulose within, creamy-white; petals 4.5 - 5.8 x 1.3 - 1.5 mm, narrow-
oblong, coherent into a tube c. 2.8 mm long, the lobes acute, c. 3 mm long, spreading
to recurved, flat. Androecium of 3 fertile stamens and 2 staminodes; filaments adherent
to the corolla, those of fertile stamens c. 2.2 mm long, villous at apex adaxially, the
staminodes 3.8 - 4.2 mm long, villous in upper half especially at throat adaxially,
subterete at apex; anthers ovate to oblong, 1.4 - 2 x 0.9 - 1.2 mm, puberulent mainly
adaxially, conspicuously glandular-punctate on connective, unappendaged at base,
narrowly attached to the filament; pollen 4 - 6-colporate, euprolate, foveolate. Disc c.
0.7 mm high, slightly shorter than the ovary. Ovary
of 5 carpels free except in the style,
c. 0.8 mm high, umbilicate, densely strigulose; style 0.9 - 1 mm long (including
stigma), glabrous; stigma not exserted from the throat at anthesis. Fruit usually of 1
mericarp, 8 - 10 x 6 - 7 mm, sparsely
puberulent, transversely
ridged, rounded apically.
Seed
subrugulose, glabrous, 6.2 - 6.8 x 4 - 4.8 mm, the testa papery. Embryo
curved, the
cotyledons conduplicate and plicate, subequal. Fig. 8F - N.
DISTRIBUTION. Brazil
(Bahia,
Rio deJaneiro).
BRAZIL. Bahia: Mun. Cairui,
Ilha de Tinhare, Morro de Sao Paulo, 10 April 1982
(fl., fr.), Pinto et al. 24/82 (CEPEC, HRB); Mun. Cruz das Almas, Reserva Florestal
do Instituto Baiano do Fumo, 27 April 1956 (fr.) Lordelo 56-346 (ALCB) & 12 Nov.
1983 (fr.), Pinto et al. 380/83 (CEPEC, HRB); Mun. Gandu, Gandu, estrada a Ibiai,
23 Oct. 1970 (fl., fr.), T S. Santos
1190 (CEPEC, U); Mun. Ibicarai, on BR-415, 5 km
W of the gate to the city of Ibicarai, 14?52'S, 39038'W, 7 Feb. 1993 (fl.), Kallunki &
Pirani 443 (NY, SPF), a 3 km W da cidade na Rodovia Itabuna-Vit6ria da Conquista
(BA-265), 14o52'S, 39?38'W, 16 Feb. 1988 (fl.), Pirani et al. 2341 (NY, SPF), Rodovia
SYNOPSES OF ANGOSTURA & CONCHOCARPUS 291
Itabuna-Itapetinga, a 41 km da BR-101 e 5 km do portao da Cidade de Ibicarai, 14
Feb. 1994 (fl., fr.), Pirani et al. 2968 (CEPEC, NY- 2 sheets, SP, SPF); Mun. Ilh6us, 5
a 6 km SW de Olivenga na estrada que liga Olivenia ao povoado do Maruim, 29 July
1993 (fr.), Jardim et al. 247 (CEPEC, NY), 6 km from Olivenga on road to Serra das
Trempes, 14'45'S, 39'02'W, 30 March 1995 (fl., fr.), Kallunki
et al. 685 (CEPEC, NY),
estrada que liga a Esta?,ao
hidromineral de Olivenca ao povoado de Vila Brasil, 5 km
ao Sudoeste de Olivenga, 8 Feb. 1982 (fl.), Mattos Silva et al. 1488 (CEPEC,
NY), a 6
km a partir da Igreja de Oliven-a na estrada para Maruim, approx. 14'57'S,
39000'W, 6 Feb. 1993 (fl., fr.), Pirani & Kallunki 2744 (holotype SPF, isotypes CAS,
CEPEC, HUEFS, K, MBM, MO, NY, RB, SP, U), estrada vicinal de Olivenga a
Buerarema, 6 km de Olivenga, 12 Feb. 1994 (fl., fr.), Pirani et al. 2951 (NY, SPF),
road from Olivenga to Serra das Trempes, 6 km from Oliven-a, 39003'W, 14'59'S, 1
Feb. 1992 (fl.), Thomas et al. 9015A (CEPEC, NY), road from Olivenca to Maruim,
6.1 km W of Olivenca, 14'59'S, 39003'W, 1 May 1992 (fl.), Thomas et al. 9059A
(CEPEC, NY, SPF), 5.4 km west of Olivenga on road to Vila Brasil & Maruim,
14015'28"S, 39002'06"W, 5 Feb. 1994 (fl., fr.), Thomas et al. 10278 (CEPEC, NY);
Mun. Jussari, 3 km NW of Jussari on road to Palmira, c. 15010'S, 39031'W, 10 May
1995 (fl.), Kallunki
et al. 745 (CEPEC, NY); Mun. Una, 19 May 1965 (fl., fr.), Belem
&
Magalhaes
1065 (CEPEC, US 2 sheets) & 2 June 1966 (fl., fr.), Belem
& Pinheiro
2401
(CEPEC, NY, US), Pedras, 13 Nov. 1969 (fl.), T. S. Santos 495 (CEPEC). Rio de
Janeiro: Mun. Cabo Frio, Armadio dos Biizios, entre Emerencia e Tucuns, 27 Dec.
1981 (fl., fr.) Martinelli & Worcman 7818 (RB).
Without locality: 1814 - 17 (fl.), Bowie
& Cunningham
68 (BM); s.d. (fl.), Sello
547
(BM).
HABITAT. Southern Bahian wet forest, Southern Bahian moist forest, restinga forest.
This new species is most similar to C. dasyanthus
and C. bellus,
with which it shares
lateral inflorescences, a circumscissile calyx, three fertile stamens with a glandular-
punctate connective, and an umbilicate, strigulose ovary with a glabrous style. The
main differences among these taxa are discussed under C. bellus and C. dasyanthus.
In addition, in C. diadematus and C. bellus,
the partial inflorescences of the thyrse
when fully mature consist of a (sub) sessile pair of monochasia.
The specific epithet, which means small crown, refers to the appearance of the
circumscissile calyx.
Conchocarpus elegans (A. St.-Hil.) Kallunki & Pirani comb. nov.
Galipea elegans
A. St.-Hil., Fl. Bras. Merid. 1: 90 (1825). Type: Brazil, Rio de Janeiro,
Sebastianopolis, St. Hilaire s.n. (holotype P). The holotype is sterile. In the
protologue, St. Hilaire explained that the flowers were lost before he wrote the
description and, therefore, he could not describe them. Because the leaves are
not distinctive, nothing about this specimen allows us to match it with any other.
Engler, however, in Flora Brasiliensis, applied this name (as Cusparia elegans) to
several fertile specimens that may or may not be the same taxon represented by
the St. Hilaire type but that represent a distinct taxon. Therefore, following
Article 9.7 of the International Code of Botanical Nomenclature (Greuter et al.
292 KEW
BULLETIN VOL. 53(2)
1994), we designate as an epitype the following specimen identified by Engler as
Cusparia elegans: Brazil, Rio de Janerio, Corcovado, Vauthier 548 (epitype
designated here G, duplicate at P).
Cusparia
elegans
(A. St-Hil.) Engl. in Mart., Fl. Bras. 12(2): 121 (1874).
Angostura
elegans
(A. St.-Hil.) Albuq. in Acta Amazon. 11: 850 (1981).
DISTRIBUTION. Brazil (Rio deJaneiro).
Conchocarpus fanshawei (Sandwith) Kallunki & Pirani comb. nov.
Cuspariafanshawei Sandwith in Kew Bull. 3: 306 (1948). Type: Guyana, Potaro R.,
Mahdia Creek, at 108 mile on Bartica-Potaro road, Fanshawe in Forest
Dept. 3494
(holotype K; isotypes K, NY).
Angosturafanshawei
(Sandwith) Albuq. in Acta Amazon. 11: 850 (1981).
DISTRIBUTION. Guyana, Brazil (Amazonas, Parai).
Conchocarpus fissicalyx Pirani sp. nov. ab omnibus congeneribus calyce connato
fissura adaxiali instructo, antherae appendice apicali elongata, atque petalis roseis
optime distincta. Typus: Brazil, Bahia, rodovia Guaratinga p/ Tabela, R. S. Pinheiro
2063 (holotypus CEPEC).
Shrub 2 m tall, the internodes evident. Leaves 1-foliolate (or perhaps simple);
petiole 1 - 2 cm long, not swollen at apex; blade oblanceolate, obtuse or truncate at
base, obtuse or short-acuminate at apex, 31 - 35.6 x 6.4 - 10 cm, slightly bullate at
main and secondary veins, pellucid-punctate, the glands visible as dark dots on both
surfaces; midvein slightly sunken (or raised distally) adaxially,
a fimbrial vein absent.
Inflorescence
a densely flowered, pyramidal thyrse, borne near apex of branch, 6.5 -
7.5 cm long including the peduncle, this c. 1 cm long, the axes appressed-pubescent,
the partial inflorescences stalked, dichasial at first node, the secondary axis (at least
the lower ones) 2.5 - 3 mm long; primary bracts c. 6 mm long, persistent; pedicels 2
- 4 mm long in flower. Calyx
dentate, urceolate, at anthesis usually splitting adaxially
almost to the base, 5- 7 mm long, very sparsely appressed-puberulent especially on
lobes without, glabrous within, green (in vivo), glandular-tuberculate, the lobes not
overlapping at anthesis, triangular, acute, c. 0.7 mm long. Corolla
densely sinuate-
strigulose without, tomentulose especially at throat and glabrous below throat within,
pink; petals 16.5 - 20.6 x c. 2 mm, linear, coherent only at throat forming a tube 5 - 6
mm long, the lobes acute, c. 12 - 15 mm long, spreading, subtriquetrous. Androecium
of 2 fertile stamens (flanking the slit in the calyx) and 3 staminodes; filaments of
fertile stamens free from the corolla, c. 3.5 mm long, shortly villous only at apex
adaxially, the staminodes adherent to the corolla, 15.4 - 19.3 mm long, subterete and
pubescent in upper half; anthers narrow-ovate, 1.6 - 1.7 x 0.7 - 1 mm, glabrous,
eglandular on connective, unappendaged at base, narrowly attached to the filament,
with a flattened, acute apical appendage c. 4 mm long exserted from the throat at
anthesis. Disc c. 0.9 mm high, slightly shorter than the ovary. Ovary
of 5 carpels free
except in the style, c. 1.4 mm high, tapered into the style, densely strigulose; style c. 2
mm long (including stigma), strigulose; stigma not exserted from the throat at
anthesis. Fruit not known. Fig. 9.
SYNOPSES
OF ANGOSTURA & CONCHOCARPUS 293
FIG. 9. Conchocarpus fissicalyx. A apex of branch; B younger flower bud; C older flower bud with slit calyx; D
flower; E staminode, adaxial and abaxial views; F stamen, abaxial and adaxial views; G disc and gynoecium with
part of calyx removed. All from R. S. Pinheiro 2063. Drawn by Bobbi Angell.
294 KEW BULLETIN
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DISTRIBUTION.
Brazil (Bahia); known only by the type collection. In a recent visit
to the type locality, we found just a few remaining patches of forest and were unable
to trace this species there.
BRAZIL.
Bahia: km 2 da rodovia Guaratinga p/ Tabela, 29 March 1973 (fl.),
Pinheiro
2063 (holotype CEPEC).
HABITAT. Forest.
This species is very distinct in the unilaterally split calyx and the elongate,
flattened, apical appendage of the anthers that is exserted from the throat at
anthesis. The relatively short, densely flowered inflorescences are similar to those of
C. gaudichaudianus and C. odoratissimus. Its long, glabrous leaves bear no evident
pulvinus at the petiole apex. In the specimen, the filaments are not adherent to the
corolla, but the staminodes are; given the presence of pubescence on the adaxial
surface of the petals and the abaxial surface of the filaments, it is likely that in life
(before complete drying) the filaments are at least to some extent adherent to the
petals at least at their apices. Conchocarpus oppositifolius,
C. obovatus,
C. macrocarpus,
C.
adenantherus, and C. cauliflorus also bear flattened (but much smaller) apical
appendages, but all lack the connate calyx split adaxially almost to the base, peculiar
to C.
fissicalyx,
and all of them have a cream-coloured, not pink, corolla.
Conchocarpus fontanesianus (A. St.-Hil.) Kallunki & Pirani comb. nov.
Galipea fontanesiana A. St.-Hil. in Bull. Sci. Soc. Philom. Paris, Ser. 3, 10: 132 (Sept.
1823). Type: Brazil, Rio de Janeiro, Iraruama, St. Hilaire s.n. (holotype P, n.v.;
isotype P).
Rauia racemosa Nees & Mart. in Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat.
Cur. 11: 169, t. 24 (1823). Type: Brazil, [Bahia], ad viam Felisbertiam, Wied-Neuwied
s.n. (holotype BR, isotype BR). Although "via Felisbertia" is not mentioned in
Bokermann's (1957) itinerary of Wied-Neuwied's travels in Brazil, the date of the
collection given in the protologue (Jan. 1817) and information provided by Cruden
et al. (1991) would place the locality in the vicinity of Ilheus and Itabuna, Bahia.
Bonplandia
fontanesiana (A. St.-Hil.) Spreng., Syst. Veg. 1: 780 (1825).
Cuspariafontanesiana
(A. St.-Hil.) Engl. in Mart., Fl. Bras. 12(2): 122 (1874).
Angosturafontanesiana
(A. St.-Hil.) Albuq. in Acta Amazon. 11: 850 (1981).
DISTRIBUTION. Brazil (Bahia, Rio de Janeiro, Sao Paulo).
Despite inquiries made to F. Stafleu, to The Hunt Institute of Botanical
Documentation, to the Library
of the Linnaean Society of London, and to the Library
of the Royal Botanic Gardens, Kew,
we have not been able to determine the month of
publication of the Nees and Martius article and, therefore, which of the two names
(G.
fontanesiana
and R. racemosa)
has priority.
We follow Engler's choice of the former.
Conchocarpus furcatus Kallunki sp. nov. inflorescentiae partialis cymosae ramulis
duobus secundis racemiformibus C. santosii
similis, sed corolla breviori (7.5 nec 14-
20 mm longa) praecipue differt. Typus: Brazil, Espirito Santo, Mun. Mimoso do Sul,
D. Sucre
& Ti Soderstrom
8921 (holotypus SPF,
isotypi NY, RB).
SYNOPSES OF ANGOSTURA & CONCHOCARPUS 295
Shrub 3 m tall, usually unbranched, the internodes evident. Leaves 1-foliolate;
petiole 2.2 - 6.5 cm long; blade narrow-elliptic or -oblong, acute or obtuse at base,
acute at apex, 10.5 - 33.1 x 3.1 - 6.8 cm, densely pellucid-punctate, the glands visible
as dark dots on both surfaces; midvein impressed adaxially, a fimbrial vein absent.
Inflorescence
a narrow thyrse, extra-axillary,
24 - 37.5 cm long including the peduncle,
this 2.5 - 4 cm long, the axes densely mealy-pubescent, less so with age, the partial
inflorescences subsessile, dichasial at first node, with age monochasial at higher
nodes, the secondary axis 0 - 2.5 mm long, the two monochasial branchlets of older
partial inflorescences racemiform, secund, to 17 mm long; primary bracts deciduous
(only scars seen); pedicels 1 - 1.5 mm long. Calyx
dentate, c. 1 mm long, densely
mealy-pubescent without, glabrous within, persistent, beige, the lobes not
overlapping at anthesis, triangular, obtuse, c. 0.5 x 1.2 mm. Corolla
densely sinuate-
strigose on tube without, tomentulose on lobes without and within, white; petals 7.2 -
7.5 x 1.1 - 1.3 mm, oblanceolate, coherent into a tube c. 3 mm long, the lobes acute,
4.2 - 4.5 mm long, spreading, triquetrous. Androecium of 2 fertile stamens and 3
staminodes; filaments adherent to corolla, those of fertile stamens c. 2.6 mm long,
glabrous adaxially, the staminodes 4 - 4.5 mm long, glabrous adaxially, subterete and
hirsutulous at apex; anthers narrow-oblong, 2.6 - 2.8 x c. 0.6 mm glabrous, eglandular
on connective, unappendaged at base, narrowly attached to the filament; pollen
suboblate, foveolate-perforate (Fig. 5C). Disc c. 0.5 mm high, shorter than the ovary.
Ovary of 5 carpels free except in the style, c. 2 mm high, tapered into the style,
densely strigulose; style c. 0.9 mm long (including the stigma), strigulose; stigma not
exserted from the throat at anthesis. Fruit
unknown. Fig. 10A - E.
DISTRIBUTION.
Brazil (Espirito Santo).
BRAZIL.
Espirito Santo: Mun. Mimoso do Sul, vertente suleste de morro, 24 April
1972 (fl.), Sucre & Soderstrom
8921 (holotype SPF,
isotypes NY, RB).
HABITAT.
Not known.
This species is similar to C. punctatus and C. santosii in the secund, racemiform
branchlets of the mature partial inflorescences. Conchocarpus furcatus
is distinguished
easily from the other two primarily by its smaller corolla (7.5 vs. 14 - 20 mm long).
Conchocarpus gaudichaudianus (A. St.-Hil.) Kallunki & Pirani comb. nov.
Galipea
gaudichaudiana A. St.-Hil., Fl. Bras. Merid. 1: 89 (1825). Type: Brazil, Prov.
Rio de Janeiro, pr. Mandiocca, Gaudichaud s.n. (holotype P; possible isotype
[ Gaudichaud
1830] G).
Cusparia
gaudichaudiana (A. St.-Hil.) Engl. in Mart., Fl. Bras. 12(2): 116 (1874).
Angostura gaudichaudiana (A. St.-Hil.) Albuq. in Acta Amazon. 11: 850 (1981).
DISTRIBUTION. Brazil (Rio de Janeiro, Minas Gerais, Santa Catarina).
Conchocarpus gaudichaudianus subsp. bahiensis Kallunki subsp. nov. a subsp.
gaudichaudiano inflorescentiis longioribus laxis necnon floribus fructibusque
minoribus differt. Typus: Brazil, Bahia, Mun. Itacar6, T S. dos Santos, L. A. Mattos
Silva & V HolloweUll
3729 (holotypus CEPEC, isotypus NY).
296 KEW BULLETIN VOL. 53(2)
SYNOPSES OF ANGOSTURA & CONCHOCARPUS 297
Shrub 0.4 - 1 m tall, the internodes obscured. Leaves
1-foliolate; petiole 2.8 - 4.3
cm long; blade oblanceolate, cuneate at base, obtuse to rounded at apex, 21.5 - 39.5
x 5.9 - 9.2 cm, not pellucid-punctate, no glands visible on either surface; midvein
slightly canaliculate adaxially, a fimbrial vein absent. Inflorescence
a loosely branched
thyrse, axillary, often clustered, 5.7 - 8 cm long including the peduncle, this 2.5 -
4.5 cm long, the axes minutely appressed-pubescent, the partial inflorescences
stalked, loosely monochasial, the secondary axis 5 - 15 mm long; primary bracts c.
0.8 mm long, deciduous; pedicels 2.5 - 5.5 mm long. Calyx
lobed, c. 2 mm long,
appressed-pubescent and somewhat mealy without, glabrous within, ciliolate,
persistent in fruit, the lobes overlapping at anthesis, very wide-ovate, rounded, c. 1.3
x 2 mm. Corolla
sparsely sinuate-strigulose in basal 1/4 and glabrous above without,
tomentulose only at throat and otherwise glabrous within, white; petals c. 6.8 x 1.1 -
1.5 mm, connate in basal 1 - 2 mm, the lobes c. 4.6 mm long, wide-spreading, the
innermost folded longitudinally, cucullate, the others rounded, flat. Androecium
of 2
fertile stamens and 3 staminodes; filaments adnate at base to (or the staminodes
free from) the corolla, adherent to it at throat, those of fertile stamens 2.6 - 3 mm
long, connate to each other for c. 0.5 mm at base, villous at apex adaxially, the
staminodes 4.2 - 5.5 mm long, villous adaxially at throat, hirsutulous in upper half,
subterete at apex; anthers narrow-oblong, 1.7 - 2.1 x c. 0.7 mm, glabrous,
eglandular on connective, unappendaged at base, broadly attached to the filament.
Disc c. 0.7 mm high, slightly taller than the ovary. Ovary
of 5 carpels free except in
the style, 0.5 - 0.6 mm high, umbilicate, glabrous; style 1.2 - 1.6 mm long
(including stigma), glabrous; stigma not exserted from the throat at anthesis. Fruit
of 2 - 5 free mericarps, each c. 14 x 7 mm, densely puberulent and mealy when very
young, perhaps glabrescent at maturity (certainly so in very old fruits), transversely
ridged, beaked apically. Seed unknown. Fig. 11A - D.
DISTRIBUTION. Brazil (Bahia).
BRAZIL. Bahia: Camaca, estrada a Pau Brasil, 19 Jan. 1971 (fl.), T S. Santos
1356
(CEPEC, U); Mun. Itacar6, Fazenda das Almas, km 18 da Rod. Ubaitaba/Itacare, 27
Jan. 1982 (fl., fr.), T. S. Santos et al. 3729 (holotype CEPEC, isotype NY).
HABITAT. Southern Bahian wet forest.
The anthers of this subspecies are broadly attached to the filaments, as they are
in typical C. gaudichaudianus and in C. concinnus, C. cyrtanthus, C. hirsutus, and C.
insignis. Among these, only C. gaudichaudianus
has axillary inflorescences. The new
subspecies is distinguished from the typical one by its longer (5.7 - 8 vs. 1 - 1.5
cm), loosely flowered inflorescences, by its flowers smaller in all parts, e.g., the
calyx lobes shorter (1.3 vs. 1.7 - 3 mm) and rounded (vs. obtuse or acute) and the
corolla shorter (6.8 vs. 10.4 - 18 mm), and by its smaller fruiting carpels (14 x 7 vs.
18 - 19 x 11 mm).
FIG. 10. A - E Conchocarpus furcatus. A apex of branch with two inflorescences, and larger lower leaf; B one
partial inflorescence; C flower; D stamen, adaxial and abaxial views; E disc and gynoecium with part of calyx
removed. F - L C. longipes. F apex of branch with inflorescences of various ages; G flower bud; H flower; J
staminode; K stamen, adaxial and abaxial views; L disc and gynoecium with part of calyx removed. A - E from
Sucre & Soderstrom
8921; F - L from Santos & Silva 3718. Drawn by Bobbi Angell.
298 KEW BULLETIN VOL. 53(2)
FIG. 11. A - D Conchocarpus gaudichaudianus subsp. bahiensis. A apex of branch; B gynoecium with longitudinal
section of disc and calyx; C stamen, adaxial and abaxial views; D fruit. E- L C. grandis. E apex of inflorescence;
F flower; G gynoecium with longitudinal section of disc and calyx; H stamen, lateral, adaxial, and abaxial views;
J apex of branch; K seed; L embryo. A - C from Santos 1356; D from Santos et al. 3729; E - H from Santos et al.
191; J - L from Plowman et al. 9381. Drawn by Bobbi Angell.
SYNOPSES
OF ANGOSTURA & CONCHOCARPUS 299
Conchocarpus grandiflorus (Engl.) Kallunki & Pirani comb. nov.
Cusparia
grandiflora
Engl. in Mart., Fl. Bras. 12(2): 112, t. 23 (1874). Type: Brazil,
Rio de Janeiro, Macah6, Riedel 476 (lectotype designated here LE, isolectotypes
LE - 3 sheets, probable isolectotypes [all s.n.] K, M, W). Of the four sheets of this
collection at LE, two are annotated by Engler. Neither of these two is chosen here
as the lectotype because the specimen on one sheet (bearing an original label) is
sterile and the flowers of the specimen on the other sheet (bearing a second-
hand label) have been severely damaged by insects. A