Nilsson1974 PDF

Pollen Morphology and Taxonomy of the Genus Quararibea S.l.
(Bombacaceae)
Author(s): S. Nilsson and A. Robyns
Source: Bulletin du Jardin botanique national de Belgique / Bulletin van de National Plantentuin
van België, Vol. 44, No. 1/2 (Jun. 30, 1974), pp. 77-99
Published by: National Botanic Garden of Belgium
Stable URL: http://www.jstor.org/stable/3667429
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Bull. Jard. Bot. Nat. Belg.
Bull. Nat. Plantentuin Belg.
44: 77-99 (30-6-1974)
Pollen morphologyand taxonomy

of the genus Quararibeas. 1. (Bombacaceae)
by
S. Nilsson & A. Robyns (*)
Abstract. - The pollen of 61 species of Quararibeas.l. (Matisia included) has

been studied by light, scanning electron and transmission electron microscopy.
Six pollen types are recognized: Q. cordata-, Q. asterolepis-, Q. bicolor-, Q. intri-
cata-, Q. stenopetala-, and Q. asymmetrica-type.Since no correlation was found
between macromorphology and pollen morphology, separation of Quararibea
Aubl. (1775) and Matisia Humb. & Bonpl. (1805) is not supported. Three new
combinations are established: Q. apaporiensis, Q. idroboi and Q. soegengii.
INTRODUCTION
The mainly South American genus Quararibea has been treated

variously by different authors. Baillon (1871) and Vischer (1920)
included Matisia Humb. & Bonpl. (P1. Aequin. 1 : 9. 1805) in the
genus Quararibea Aubl. (Hist. P1. Gui. Fr. : 691. 1775) as well as
Cuatrecasas (1948) and A. Robyns (1964). Later Cuatrecasas (1954,
1971) decided to keep the two genera separate on flower- and fruit-
morphological grounds, the fundamental differences between the
two genera being : ovary 2-celled and fruit generally monospermous
in Quararibea versus ovary 5-celled and fruit 5-seeded in Matisia;
staminal tube entire or slightly dentate at the apex in Quararibea
versus staminal tube with five long antheriferous lobes in Matisia.
Erdtman (1952) investigated the pollen morphology of about
30 species of Matisia and Quararibea indicating that the pollen
grains (p. 76) <show an array of forms which will probably be
of value for classification purposes >. Fuchs (1967), studying the
(*) S. Nilsson, PalynologiskaLaboratoriet,Wallenberglaboratoriet,Lilla Frescati,

S-104 05 Stockholm 50 (Sweden); A. Robyns, Jardin botanique national de Belgi-
que, Departement des Spermatophytes et des Pteridophytes.- Manuscript re-
ceived February 14, 1974.
- 78
pollen morphology of the Bombacaceae including 25 species of

Matisia and Quararibea, concluded (p. 119) «<that various poly-
morphic genera might be more naturally delimited on the basis
of their palynomorphology ». Tsukada (1964, 1965) described the
pollen morphology of Matisia longiflora Gleason. In connection
with descriptions of three new species of Quararibea, Q. dolicho-
siphon, Q. intricata and Q. malacocalyx, notes on the pollen mor-
phology were included by A. Robyns & S. Nilsson (1970, 1972).
In the latter studies like in the present one electron microscopy
(scanning and transmission) has contributed new and valuable
information on the fine structure of the pollen grains examined.
Six different pollen types are distinguished, and an attempt has
been made to elucidate the taxonomy by means of these pollen
morphological criteria.
MATERIALS AND METHODS
Pollen material was obtained from specimens in the following

herbaria : A, BR, COL, F, GH, MO, NY, P, S, US, and VEN.
Altogether 61 species (109 specimens) were examined. The polleni-
ferous material was acetolyzed. For light microscopical analyses
the pollen grains were embedded in unstained glycerine jelly. The
slides were sealed with paraffin. Non-acetolyzed and osmium-
stained pollen of some species was embedded in Epon and cut on
a LKB-microtome. The ultra-thin sections were post-stained with
uranyl acetate and lead citrate before examination with a trans-
mission electron microscope (Zeiss EM-9A). For scanning electron
microscopy, acetolyzed, dried pollen was used. The pollen grains
were coated with gold evaporated from a tungsten filament in a
conventional vacuum coating unit. A Stereoscan Mk II a (Cam-
bridge Scientific Instrument Co.) and a Jeol (JSM U3) instrument
were used for the examination.
Terminology follows that of Erdtman (1952).
GENERAL POLLEN DESCRIPTION
Pollen grains radially symmetrical, isopolar, medium-sized to

usually large or very large (equatorial diameter from 39 ,um to
- 79 -
128 ,/m), suboblate to spheroidal, 3-6-(zono-)porate, the pores

rounded to broadly elliptical (colpoid), bordered by a smooth or
patterned margin consisting mainly of thickened nexine; exine
3-10 /m thick, divisible into sexine and nexine subtendedby intine;
sexine reticulate, foveolate, ornate, or consisting of different types
of processes, the latter often cavernulous;nexine with granules or
stout, rod-like processes; NPC = 344, 444, 544, 644. Orbicules
(Ubisch bodies) rounded or irregularin shape, more or less smooth
or crumpled, with a structure similar to that of the muri, mural
fragments or processes, often with one or more inner cavities.
KEY TO THE POLLEN TYPES

(based on acetolyzed material)
1. Pollen grains reticulate to foveolate, infrequently ornate, the muri smooth,

the nexine (lumina) smooth, with few scattered microprocesses,or some-
times finely granular:
2. Muri supported by few long and stout bacules . . . . 1. Q. cordata-type
2. Muri supported by numerous short and slender bacules .
. . . . . . . . . . . . . . . . . . . . 2. Q. asterolepis-type
1. Pollen grains provided with processes, or reticulate, or with mural frag-
ments, or with smooth loop-like densely arranged elements; processes,
muri and mural fragments with a finely striate structure;nexine (lumina)
coarsely and densely granular (except in Q. asymmetrica-type):
3. Pollen grains with isodiametric to elongated processes, infrequently also
with mural fragments .. . . . . ....... 3 Q. bicolor-type
3. Pollen grains without processes:
4. Pollen grains reticulate, the muri occasionally broken into fragments
. ... . . . . . .. . . . . . . . . 4. Q. intricata-type
4. Pollen grains non-reticulate:
5.Pollen grains with mural fragments, densely and irregularly dis-
tributed ........ 5. Q. stenopetala-type
5. Pollen grains with thin, smooth, loop-like elements, densely distrib-
uted or sometimes arrangedin a reticulate way 6. Q. asymmetrica-type
DESCRIPTION OF THE POLLEN TYPES
1. Q. cordata-type.- Fig. 1-8.

Pollen grains71-102 X 80-110 tm, suboblateto oblate-spheroidal,
3-porate, the pores rounded, 7-18 um in diam., bordered by a
4-7 Jtm wide usually smooth margin; exine 4-5 itm thick; sexine
thicker than nexine, reticulate to foveolate, rarely ornate, hetero-
brochate,the muri 1-2(-5) [kmwide, smooth, simpli- to duplibaculate,
supported by relatively few, long and stout bacules 3-4 um long,
1-6 -tmwide, rounded to angular or of varying shape in cross-
- 80
section, the lumina or foveolae usually rounded, (0.5-)1-5 ~m in

diam.; nexine smooth or sometimes finely granular. Orbicules 3-8 atm
in diam., rounded to irregular in shape, + smooth, with one or
more inner cavities.
The following species have pollen grains of the Q. cordata-type:
Q. cordata, Q. longitubulosa, and Q. soegengii.
Inter- and intraspecific variation : One collection of Q. cordata
(Sparre 18216) has pollen with relatively thin bacules (fig. 6).
2. Q. asterolepis-type. - Fig. 9-25.
Pollen grains 35-96 X 39-99 ,um, suboblate to prolate-sphe-
roidal, 3-porate, the pores rounded to oval, 2-13 ,Lm in diam.,
bordered by a 2-7 ,am wide and usually smooth margin, commonly
bounded by ellipse-shaped muri; exine 2-4 ,m thick; sexine thicker
than nexine, foveolate to reticulate, heterobrochate, the muri
1-10 mm wide, smooth, simpli- to multibaculate, usually supported
by numerous relatively short and thin bacules 1-2 /m long, 1-2 tm
wide or less, the foveolae or lumina usually rounded to oval, less
commonly angular, (0.5-)1-16 ,m in diam.; nexine smooth or finely
granular, or provided with relatively few, short, rod-like micro-
processes. Orbicules not frequently observed, 3-6 ,tm in diam.,
rounded in shape, ± smooth, with one or more inner cavities.
The following species have pollen grains of the Q. asterolepis-
type: Q. amazonica, Q. aristeguitae, Q. asterolepis, Q. bilobata,
Q. bracteolosa, Q. ciroana, Q. duckei, Q. fieldii, Q. floribunda,
Q. foenigraeca,Q. funebris,Q. gigantiflora,Q. grandifolia,Q. gua-
temalteca, Q. guianensis, Q. idroboi, Q. leptandra, Q. lomensis,
Q. loretoyacuensis, Q. magnifica, Q. obovalifolia, Q. parviflora,
Q. parvifolia,Q. platyphylla,Q. pterocalyx, Q. schultesii, Q. spatu-
lata, Q. stenophylla, Q. steyermarkii, Q. turbinata, Q. velutina,
Q. yunckeri and Quararibea sp. (Dressler 3477).
Inter- and intraspecific variation : The size of the pollen grains
varies greatly with the largest diameter ranging from about 40 ,m
to nearly 100 ,im. Q. parviflora and Q. yunckeri have comparatively
small grains (largest diam. ca. 40 ,um) while e.g. Q. gigantiflora and
Q. magnifica provide large grains (largest diam. ca. 100 mm). The
main difference concerns shape and size of foveolae or lumina and
consequently width of muri, as well as number and size of sup-
porting bacules. The majority of the species investigated appears
-81
to have typically foveolate to reticulatepollen grains with more or

less rounded lumina (fig. 9-13), while some are reticulate with
relatively large and usually angular lumina and relatively narrow
muri, e.g. Q. bracteolosa (fig. 24), Q. duckei (fig. 23), Q. giganti-
flora, Q. idroboi (fig. 25), and Q. magnifica (fig. 21-22). This
difference turned out to be inconsistent as it also occurs within
a species, e.g. Q. guianensis. One collection (Irwin et al. 47196) has
large lumina while all the others are typically foveolate. The size
of bacules (cross-section) differs. In most cases the bacules are
relatively thin (< 1 /-m wide). In one collection of Q. funebris
(Skutch 1846) the bacules are relativelylarge, 3-4 tumwide (fig. 18),
approachingthe conditions in the Q. cordata-type.Q. velutina has
reticulate and verrucose grains on the same slide (fig. 19, 20).
3. Q. bicolor-type. - Fig. 26-38, 40, 42.

Pollen grains 53-114 X 56-128 vm, suboblate to spheroidal,
3-4-porate, the pores rounded to broadly elliptic, colpoid, 6-18 ,m
in diam., bordered by a 2-7 am wide margin, the latter beset with
numerous processes or mural fragments of various size and shape;
exine 4-10 jm thick (processes included); sexine 2-9 ~m thick,
consisting of isodiametric to elongated processes and infrequently
of mural fragments; processes or fragments 2-8 /-m wide, usually
cavernulous, abruptly constricted at their base or narrowed off
(gemmae, clavae, pila), or not constricted (verrucae),usually sup-
ported by bacules, the surface finely striate or granular; nexine
densely granular or with numerous, short and stout, rod-like pro-
cesses 1-2 ,umwide and 3-4 ,umhigh. Orbicules 2-15 ,tm in diam.,
with crenated outline and one or more inner cavities.
The following species have pollen grains of the Q. bicolor-type:
Q. bicolor,Q. bolivarii,Q. castano,
Q. amplifolia,Q. apaporiensis,
Q. cornucopiae, Q. cruceto, Q. dolichopoda, Q. dolichosiphon,
Q. dowdingii, Q. huallagensis, Q. lasiocalyx p.p., Q. lecythicarpa,
Q. longiflora, Q. malacocalyx, Q. muricata, Q. ochrocalyx p.p.,
Q. wurdackii.
Inter- and intraspecific variation : The variation between and
within the species mainly concerns shape and size of the processes.
Elongated processes occur in e.g. Q. bolivarii (Cuatrecasas22001)
and Q. castano (Cuatrecasas 14428). Remnants of low muri are
found in Q. dolichopoda (fig. 28), Q. dolichosiphon and in one
- 82
collection of Q. bicolor [Ducke s.n. (RB 11419), fig. 26] which

has large processes compared with another collection of the same
species (Lao 32, fig. 27). One collection of Q. ochrocalyx (Krukoff
4719, fig. 45) has pollen grains of Q. intricata-typewhile the other
collections examined (Ducke 1089 & 1783, fig. 36-38) are of Q. bico-
lor-type. One collection of Q. lasiocalyx (Ducke 962, fig. 40, 42) has
pollen of Q. bicolor-type while the others are of Q. intricata-type
(fig. 39, 41).
4. Q. intricata-type.- Fig. 39, 41, 43-45.
Pollen grains 62-105 X 72-116 /tm, subolate to oblate-sphe-
roidal, 3-4-porate, the pores rounded, 5-16 ,um in diam., bordered
by a 3-7 ,umwide margin; exine 3-4 ,m thick; sexine thicker than
nexine, coarsely reticulate to finely reticulate at the pore margin,
heterobrochate, the muri 1-3 um wide, consisting of elongated,
entangled elements, forming a striate structure, simpli- to dupli-
baculate, sometimes broken into fragments of different size and
shape, the lumina usually angular, (0.5-)1-20(-35) tm in diam., the
smallestlumina appearingas perforationsin the muri;nexine densely
granular or with numerous, short and stout, rod-like processes
1-2 um wide and 3-4 ,m high. Orbicules 2-9 ium in diam., of
irregular shape, with a crenated outline and one or more inner
cavities.
The following species have pollen grains of the Q. intricata-type:
Q. intricata (cf. A. Robyns & S. Nilsson 1972 : fig. 1-2), Q. lasio-
calyx p.p., Q. ochrocalyxp.p., Q. putumayensis,and Q. sclerophylla.
Inter- and intraspecific variation : One collection of Q. lasio-
calyx (Ducke 962) differs from the other specimens by having the
muri split into fragments, or provided with processes (pila), ca.
4-5
sm high (fig. 40, 42). The pollen grains thus resemble those
of the Q. bicolor-type. One collection of Q. ochrocalyx (Krukoff
4719) has pollen of the Q. intricata-type(fig. 45) (cf. also Q. bicolor-
type). Q. putumayensishas frequentlypollen grains with incomplete
muri (fragments). Q. sclerophylla has a comparatively smooth
nexine surface.
5. Q. stenopetala-type. - Fig. 46-53.

Pollen grains 75-97 X 83-105 /m, suboblate to oblate-spheroi-
dal, 3-6-porate, the pores rounded, 10-14 pm in diam., borderedby
-83
a 3-5 ,tm wide margin; exine 3-5 /m thick, sexine thicker than
nexine, with mural fragments 1-2 tm wide, of varying size and
shape, densely and irregularly distributed, consisting of elongated
entangled elements forming a striate structure,simpli- to duplibacu-
late; nexine densely granular or with numerous, short and stout,
rod-like processes 1-2 ,m wide, 3-4 tm high. Orbicules 2-8 jm in
diam., with a crenatedoutline and one or more inner cavities.
The following species have pollen of the Q. stenopetala-type:
Q. coloradorum and Q. stenopetala.
6. Q. asymmetrica-type.- Fig. 54-58.
Pollen grains 62-68 X 64-98 jtm, suboblate to oblate-spheroi-
dal, 3-6-porate, the pores rounded to oval, 7-12 ,um in diam.,
bordered by a 6-8 tm wide margin; exine 2-4 -m thick; sexine
thicker than nexine, consisting of thin, smooth, loop-like elements
0.1-0.2 ,umwide, loosely and irregularlytied, and distributedat ran-
dom, or sometimes arranged in a reticulate way; nexine almost
smooth. Orbicules 5-15 jm in diam., with an irregular,crumpled
surface.
The following species have pollen grains of the Q. asymmetrica-
type: Q. asymmetricaand Q. uribei.
CORRELATION OF POLLEN TYPES

The distinguished pollen types may be arranged in two major
pollen groups, one with smooth muri and another one with muri,
mural fragments or processes provided with a fine substructure.
The finely striate or entangled structureof the sexine is, however,
not usually clearlyvisible in the light microscope.There is apparently
no correlation between these two groups.
The first major pollen group includes Q. cordata-type and
Q. asterolepis-type.The relation between these two types is obvious,
although less evident than between Q. bicolor-, Q. intricata- and
Q. stenopetala-types.
The second major pollen group consists of Q. bicolor-type,
Q. intricata-type, Q. stenopetala-type and Q. asymmetrica-type.
Q. intricata-typewith a complete reticulum may serve as a basic
morphological pollen type. Some species of this type have pollen
grains where the muri are split into fragmentsresemblingQ. steno-
- 84-
petala-type or modified into more or less isodiametric processes

similar to Q. bicolor-type, e.g. Q. lasiocalyx and Q. ochrocalyx.
The above three pollen types are obviously closely allied, while
Q. asymmetrica-typeis more remote. Q. asymmetrica-typeis pro-
bably most closely allied to Q. stenopetala-type.The thin, loop-like
elements may be derived from the elongated elements constituting
the muri or processes of the pollen types of the second major
pollen group.
Table 1 shows a schematical, hypothetical representationof the
correlationbetween the recognizedpollen types.
Q. CORDATA-type Q.BICOLOR-type
I I
Q.ASTEROLEPIS-type Q. INTRICATA-type
Q. STENOPETALA-type ASYMMETRICA-type
---Q
Table 1. Hypothetical relationship between the recognized pollen types

of Quararibea s.l. The dotted line indicates an uncertain relation.
DISCUSSION
WITH SPECIALREFERENCETO TAXONOMY
From a pollen morphologicalstandpointit does not seem justified

to draw a generic line between Matisia and Quararibeaas currently
done (Cuatrecasas1971). As pointed out in the introductoryremarks,
the most importantmacromorphologicalcharacterdifferentiatingthe
two genera is the 2- (Quararibea)or 5-locular (Matisia) ovary, the
stigmas being lobulate according to the number of ovular cells.
Furthermore,when the ovary is 2-celled, the staminal tube is entire
or slightly dentate at the apex, while the staminal tube is deeply
5-lobed when the ovary is 5-celled. Vischer (1920) and Cuatrecasas
(1971) report however that in Q. funebris an intermediatesituation
- 85
occurs, i.e. ovary 2-, 3- or 4-celled. The same variation was found
in Q. pterocalyx : in Duke 4843 (BR) and Duke 13086 (MO)
3-celled biovulate ovaries were frequently observed, while in Allen
4631 (MO) distinctly 4-celled ovaries were encountered.The varia-
tion of the numberof ovary cells is probablymuch more widespread,
but a lack of flowering material for most of the species prevents
a more thorough study.
The genus Matisia s. str. (type-species : M. cordata) and the
genus Quararibeas. str. (type-species : Q. guianensis) have closely
related pollen types, the main differences being only the number
and size of the bacules. With reference to the present study, the
genus Matisia (Q. cordata-type of pollen) would include only the
three following species : M. cordata, M. longitubulosa and M. soe-
gengii, which are also closely related with regard to macromorphol-
ogy. In the Q. asterolepsis-type are included the type-species of
Quararibea (Q. guianensis), several species which belong clearly to
the Quararibea s. str. complex, and also some species belonging
to the Matisia s. str. complex (Q. bracteolosa, Q. grandifolia,
Q. idroboi, Q. leptandra,Q. lomensis, Q. schultesii, and Quararibea
sp.). Even, if one believesthat Q. cordata-typeand Q. asterolepis-type
must be regarded as one pollen type, this type would still include
material both of Matisia s. str. and of Quararibeas. str.
Accordingto the pollen morphologicalresults, the species belong-
ing to the second major pollen group (Q. bicolor-, Q. intricata-,
Q. stenopetala-, and Q. asymmetrica-types)could be included in
a new genus, a suggestion also put forward by Fuchs (1967). All
species in this group have 5-celled ovaries and consequentlybelong
to Matisia s. str. As macromorphologydoes not allow separation
of the species included in this group from the species having pollen
of the Q. cordata-type,the suggestion of creating a new genus is
not supported. Furthermore, since the macromorphology is not
in accord with the pollen morphology (cf. table 2), it does not seem
advisable to keep Quararibeaand Matisia separate.
At the specific level, pollen morphology confirms the close rela-
tionship between Q. bracteolosa, Q. grandifolia, Q. idroboi,
Q. lomensis, Q. schultesii and Quararibea sp. (Dressier 3477),
six species characterizedby the 5- or 10-winged calyx. In agree-
ment with Cuatrecasas (1954: 172-173), Q. loretoyacuensis,
Q. amazonica (Q. wittii nom. nud.) and Q. velutina may be regarded
as closely allied. Relationship between Q. soegengii and Q. steno-
-86
petala was suggested by Cuatrecasas (1971: 468). Pollen morphol-

ogy does not support this relationship. A. Robyns & S. Nilsson
(1972 : 351) suggested that Q. malacocalyx and Q. dowdingii are
closely related which is supported by pollen morphology. The affinity
to Q. velutina is, however, not supported. There is a tendency to
form a verrucose sexine pattern, a quite different sexine ornamen-
tation than that of the Q. bicolor-intricata-types. A close relationship
between Q. huallagensis and Q. longiflora is confirmed, but the
affinity between Q. coloradorum and the two above mentioned
species is not supported. Finally, pollen morphology corroborates
the affinity between the two species included in the Q. asymmetrica-
type, Q. asymmetrica and Q. uribei.
Pollen types Macromorphology
First major pollen group

Q. cordata-type 3 species of Matisia s. str. including types-speciesof
Matisia
Q. asterolepis-type 26 species of Quararibea s. str. & 7 species of
Matisia s. str.
Second major pollen group
Q. bicolor-type
Q. stenopetala-type 25 species of Matisia s. str.
Q. asymmetrica-type
Table 2. Relation between pollen types and macromorphology
SPECIES INVESTIGATED
Quararibea amazonica Ulbrich, Verhandl. Bot. Ver. Brandenburg 50:

91 (1908).
Klug 2192 (MO, NY), Peru.
Pollen: Q. asterolepis-type
Quararibea amplifolia (Pittier) Cuatr., Lloydia 11: 191 (1948).

Matisia amplifolia Pittier, Bol. Soc. Venez. Cienc. Nat. 10 : 115
(1945).
Delgado 357 (type US), Venezuela.
Pollen : Q. bicolor-type
- 87 -
Quararibeaapaporiensis(Cuatr.)A. Robyns & S. Nilsson comb. nov.

Matisia apaporiensisCuatr., Bot. Mus. Leafl. Harvard Univ. 17:
82 (1955).
Schultes & Cabrera 17641 (isotype F, US), Colombia.
Quararibea aristeguietae Cuatr., Bol. Soc. Venez. Cienc. Nat. 26:

153 (1965).
Aristeguieta & Pannier 1905 (holotype VEN), Venezuela.
Pollen : Q. asterolepis-type
Quararibea asterolepis Pittier, Fedde Repert. Sp. Nov. 13: 316

(1914).
Rowley e6 Stork 782 (NY), Costa Rica; Duke 4985 (MO),
Pittier 5575 (holotype US) (fig. 17), Stern, Chambers, Dwyer
& Ebinger944 (US), Panama.
Quararibeaasymmetrica(Cuatr.)Cuatr., Lloydia 11: 191 (1948).

Matisia asymmetricaCuatr., Rev. Acad. Colomb. Cienc. 7: 49
(1946).
Cuatrecasas 15152 (isotype US) (fig. 54-57), Colombia.
Pollen : Q. asymmetrica-type
Quararibeabicolor (Ducke) Cuatr., Lloydia 11: 191 (1948).

Matisia bicolor Ducke, Arch. Jard. Bot. Rio de Janeiro 3 : 210
(1922).
Ducke s.n. (RB 11419) (isotype US) (fig. 26), Brazil; Asplund
13322 (S), Rafael Lao M. 32 (US) (fig. 27), Peru.
Quararibeabilobata A. Robyns, Ann. Missouri Bot. Gard. 55: 54

(1968).
Wurdack 2450 (isotype US), Peru.
- 88-
Quararibeabolivarii (Cuatr.)Cuatr., Lloydia 11: 191 (1948).

Matisia bolivarii Cuatr., Ciencia (Mexico) 1 : 401 (1940).
Cuatrecasas 8560 (holotype COL, isotype F), 21685 (isotype
subsp. occidentalis Cuatr., BR), 22001 (US), Colombia.
Quararibeabracteolosa (Ducke) Cuatr., Lloydia 11: 191 (1948).

Matisia bracteolosaDucke, Bol. Tecn. Inst. Agron. Norte (Belem)
4: 17 (1945).
Kirkbride& Duke 1299 (BR), Panama; Prance, Pena & Ramos
3468 (US) (fig. 24), Brazil.
Quararibea castano (Triana & Karsten) Cuatr., Lloydia 11: 191

(1948).
Matisia castano Triana & Karsten in Triana, Nuev. Jen. Esp.
PI. Fl. Neogranadina (Bogota) : 24 (1854).
Cuatrecasas 17589 (BR, COL), 14428 (US), Colombia.
Quararibea ciroana Cuatr., Trop. Woods 101: 16 (1955).

Duque-Jaramillo 553 (isotype F), Colombia.
Quararibea coloradorum (Benoist) Cuatr., Lloydia 11: 191 (1948).

Matisia coloradorum Benoist, Bull. Soc. Bot. Fr. 80 : 335 (1933).
Benoist 3007 (holotype P) (fig. 48), Cazalet & Pennington 5096
(NY) (fig. 46, 49), Little Jr. 6158 (US) (fig. 47), Ecuador.
Pollen : Q. stenopetala-type
Quararibea cordata (Humb. & Bonpl.) Vischer, Bull. Soc. Bot.

Geneve, ser. 2, 11 : 206 (1920).
Matisia cordata Humb. & Bonpl., PI. Aequin. 1 : 10 (1805).
Glaziou 9636 (P), Brazil; Black 46-27 (NY), Colombia; Little Jr.
6456 (US) (fig. 5), Sparre 18216 (S) (fig. 6), Ecuador; Killip
& Smith 27273 (F), Tessman 3087 (S) (fig. 1-4, 7-8), Peru.
Pollen: Q. cordata-type
-89-
Quararibea cornucopiae (Triana & Planchon) Vischer, Bull. Soc.

Bot. Geneve, ser. 2, 11: 206 (1920).
Matisia cornucopiae Triana & Planchon, Ann. Sc. Nat., Bot.,
ser. 4, 17 : 326 (1862).
Goudot s.n. (P), Colombia.
Quararibeacruceto (Cuatr.) Cuatr., Lloydia 11: 191 (1948).

Matisia cruceto Cuatr.,Rev. Acad. Colomb. Cienc. 6 : 550 (1946).
Cuatrecasas 12982 (isotype US), Colombia.
Quararibeadolichopoda A. Robyns, Ann. Missouri Bot. Gard. 54:

185 (1967).
Correa, Dressier & Calderon 817 (US) (fig. 28), Panama.
Quararibeadolichosiphon A. Robyns & S. Nilsson, Bull. Jard. Bot.

Nat. Belg. 40: 353 (1970).
Kirkbride& Duke 1178 (holotype BR), Panama.
Quararibea dowdingii (Sprague) Vischer, Bull. Soc. Bot. Geneve,

ser. 2, 11: 206 (1920).
Matisia dowdingii Sprague, Trans. Proc. Bot. Soc. Edinb. 22:
428 (1905).
Sprague 370 (isotype US), Colombia.
Quararibea duckei Huber, Bull. Soc. Bot. Geneve, ser. 2, 6: 186

(1915).
Lems 5245 (NY), Surinam; Nilo T. Silva 1357 (BR) (fig. 23),
Brazil.
Quararibeafieldii Millsp., Field Mus. Nat. Hist., Bot. Ser. 1: 309

(1896).
Gaumer 879 (isotype US), Mexico.
- 90
Quararibea floribunda (St.-Hil. & Naudin) K. Schum. in Mart., Fl.

Bras. 12 (3): 242 (1886).
Myrodia floribundaSt.-Hil. & Naudin, Ann. Sc. Nat., Bot., ser. 2,
18 : 212 (1842).
Claussen 900 (NY), Glaziou 1676 (NY), Brazil.
Quararibeafoenigraeca Cuatr., Phytologia 4: 473 (1954).

Cuatrecasas 14890 (isotype BR), Colombia.
Quararibeafunebris(La Llave)Vischer,Bull. Soc. Bot. Geneve,ser. 2,

11: 205 (1920).
Lexarza funebris La Llave in La Llave & Lexarza, Nov. Veget.
Descript. 2: 7 (1825).
Martinez-Calderon122 (A, US), Mexico; Skutch 1846 (A) (fig. 18),
Guatemala.
QuararibeagigantifloraPittier,Bol. Soc. Venez. Cienc. Nat. 10: 114

(1945).
Delgado 356 (type US), Venezuela.
Quararibea grandifolia (Little Jr.) Cuatr., Lloydia 11: 191 (1948).

Matisia grandifolia Little Jr., Jour. Wash. Acad. Sci. 38 : 102
(1948).
Little Jr. 6146 (holotype US), Ecuador.
Quararibeaguatemalteca(J. D. Sm.) Standley& Steyerm.,Field Mus.

Nat. Hist., Bot. Ser. 23: 62 (1944).
Myrodia guatemaltecaJ. D. Sm., Bot. Gaz. 16: 2 (1891).
von Tiirckheim 1410 (syntype US), Guatemala.
- 91
Quararibeaguianensis Aubl., Hist. P1. Gui. Fr.: 692 (1775).

For. Dept. Br. Guiana 5466 (NY), Guiana; Archer 7558 (NY)
(fig. 9-13), Ducke 268 (A, MO), Irwin, Egler & Pires 47196 (MO)
(fig. 14-16), Brazil; Schunke 290 [isotype Q. machin Macbride,
Field Mus. Nat. Hist., Bot. Ser. 13(3A) : 618 (1956), US], Peru.
Quararibea huallagensis (Cuatr.) Macbride, Field Mus. Nat. Hist.,

Bot. Ser. 13 (3A): 616 (1956).
Matisia huallagensisCuatr.,Phytologia4: 476 (1954).
Ll. Williams 4572 (holotype F), Peru.
Quararibeaidroboi (Cuatr.) A. Robyns & S. Nilsson comb. nov.

Matisia idroboi Cuatr., Phytologia 4 : 476 (1954).
Idrobo & Schultes 1001 (isotype COL), 1069 (US) (fig. 25),
Colombia.
Quararibeaintricata A. Robyns & S. Nilsson, Bull. Jard. Bot. Nat.

Belg. 42: 347 (1972).
Prance, Pena & Ramos 3017 (isotype S), Brazil.
Pollen : Q. intricata-type
Quararibealasiocalyx (K. Schum.) Vischer, Bull. Soc. Bot. Geneve,

ser. 2, 11 : 206 (1920).
Matisia lasiocalyx K. Schum.in Mart., Fl. Bras. 12(3) : 239 (1886).
Cowan 38297 (MO), 38362 (NY) (fig. 39, 41), Ducke 962 [MO,
US (fig. 40, 42)], Pires & Cavalcante 52654 (MO) (fig. 43, 44),
Brazil.
Pollen : Q. intricata-type;Ducke 962 cf. Q. bicolor-type
Quararibealecythicarpa(Ducke) Cuatr., Lloydia 11: 191 (1948).

Matisia lecythicarpaDucke, Bol. Tecn. Inst. Agron. Norte (Belem)
4 : 18 (1945).
Ducke 1286 (type US), Brazil.
-92
Quararibea leptandra Cuatr., Lloydia 11: 185 (1948).

Cuatrecasas22086 (isotype US), Colombia.
Quararibea lomensis Cuatr., Lloydia 11: 187 (1948).

Klug 2039 (S), Peru.
Quararibealongiflora (Gleason) Cuatr., Lloydia 11: 191 (1948).

Matisia longiflora Gleason, Phytologia 1 : 26 (1933).
Lawrance 538 (isotype US) (fig. 32-34), Colombia.
Quararibea longitubulosa A. Robyns, Ann. Missouri Bot. Gard. 55:

56 (1968).
Wurdack 2102 (holotype US), Peru.
Pollen : Q. cordata-type
Quararibea loretoyacuensis Cuatr., Phytologia 4: 474 (1954).

Schultes & Black 8550 (holotype F), Colombia.
Quararibea magnifica Pittier, Bol. Soc. Venez. Cienc. Nat. 6: 15

(1939).
Steyermark91484 (NY), Steyermark& Steyermark95441 (NY)
(fig. 21, 22), Venezuela.
Quararibea malacocalyx A. Robyns & S. Nilsson, Bull. Jard. Bot.

Nat. Belg. 42 : 349 (1972).
Killip & Smith 27025 (isotype NY), Klug 1110 (NY) (fig. 35),
Peru.
Quararibea muricata Cuatr., Bot. Mus. Leafl. Harvard Univ. 15:

53 (1951).
Prance, Pena, Forero, Ramos & Monteiro 4713 (NY), Schultes
& Pires 9010 (holotype F), Brazil.
- 93 -
Quararibeaobovalifolia Pittier, Bol. Soc. Venez. Cienc. Nat. 6: 14

(1939).
Pittier 12203 (isotype MO), Venezuela.
Quararibeaochrocalyx (K. Schum.) Vischer, Bull. Soc. Bot. Geneve,

ser. 2, 11: 206 (1920).
Matisia ochrocalyx K. Schum. in Mart., Fl. Bras. 12(3) : 238
(1886).
Ducke 1089 (MO) (fig. 30, 31, 38), 1783 (NY) (fig. 36, 37),
Krukoff4719 [A (fig. 45), MO], Brazil.
Pollen : Q. bicolor-type; Krukoff 4719, Q. intricata-type
QuararibeaparvifloraLundell, Wrightia3: 121 (1965).

Contreras 3520 (isotype US), Guatemala.
Quararibeaparvifolia Standley, Field Mus. Nat. Hist., Bot. Ser. 4:

232 (1929).
Shank & Molina 4474 (US), Costa Rica.
QuararibeaplatyphyllaPittier& J. D. Sm., Bot. Gaz. 23 : 239 (1897).

Skutch 4851 (A), Costa Rica.
Quararibeapterocalyx Hemsl., Diagn. P1.Nov. Mex. Centr.-Amer.:

4 (1878); Biol. Centr.-Amer.1: 127 (1879).
Duke 4843 (MO), Pittier 4010 (GH), Stern, Chambers, Dwyer
& Ebinger 212 (MO), Panama.
QuararibeaputumayensisCuatr.,Lloydia 11: 186 (1948).

Klug 1881 (A, F), Colombia.
- 94-
Quararibea schultesii Cuatr., Bot. Mus. Leafl. Harvard Univ. 14:

31 (1949).
Schultes 6775 (F), Colombia.
Quararibeasclerophylla(Cuatr.)Cuatr., Lloydia 11: 191 (1948).

Matisia sclerophylla Cuatr., Rev. Acad. Colomb. Cienc. 6: 549
(1946).
Cuatrecasas14146 (isotype US), Colombia.
Quararibeasoegengii (Cuatr.) A. Robyns & S. Nilsson comb. nov.

Matisia soegengii Cuatr., Phytologia 20 : 468 (1971).
Soegeng s.n. (Sept. 1961) (holotype US), Ecuador.
Pollen : 0. cordata-type
QuararibeaspatulataDucke, Trop. Woods 76: 20 (1943).

Ducke 963 (isotype US), Brazil.
Quararibeastenopetala (Standley & Cuatr.) Macbride, Field Mus.

Nat. Hist., Bot. Ser. 13 (3A): 620 (1956).
Matisia stenopetala Standley & Cuatr., Phytologia 4 : 477 (1954).
Klug 2972 [holotype F (fig. 50-53), isotype US], Wurdack 2130
(US), Peru.
Pollen : Q. stenopetala-type
Quararibea stenophylla Pittier, Fedde Repert. Sp. Nov. 13: 317

(1914).
Pittier s.n. (Inst. phys.-geogr. 12410) (type US), Costa Rica.
QuararibeasteyermarkiiCuatr., Fieldiana: Bot. 28: 363 (1952).

Steyermark61959 (holotype F), Venezuela.
Pollen : Q. asterolepis-type.
- 95 -
Quararibea turbinata (Sw.) Poiret in Lam., Encycl. Meth., Bot.,

Suppl. 4: 636 (1816).
Myrodia turbinataSw., Nov. Gen. Sp. PI. Prodr. 102 (1788).
Britton & Britton 8097 (NY), Puerto Rico; Britton & Cowell
413 (NY), St. Kitts; Fairchild 2805 (US), Grenada.
Quararibeauribei Garcia-Barriga& Hernandez,Mutisia 2: 1 (1952).

Uribe-Uribe1746 (COL) (fig. 58), Colombia.
Pollen : Q. asymmetrica-type
Quararibeavelutina Cuatr., Phytologia 4: 475 (1954).

Schunke 235 [holotype F, isotype US (fig. 19, 20)], Peru.
Quararibea wurdackii A. Robyns, Ann. Missouri Bot. Gard. 55:

58 (1968).
Wurdack 2129 (holotype US) (fig. 29), Peru.
Quararibeayunckeri Standley, Field Mus. Nat. Hist., Bot. Ser. 9:

306 (1940).
Yunker, Koepper & Wagner 8010 (isotype US), Honduras.
Quararibea sp.
Dressier 3477 (MO), Panama.
Acknowledgements: We wish to express our sincere thanks to
the Directors and the Curatorsof the various herbariafor allowing
us to sample polleniferousmaterialand for having sent us specimens
on loan. The transmission electron micrographs were taken by
Miss Elisabeth Grafstr6m and the scanning electron micrographs.
(Jeol) by Mr. Sadao Ayoyagi.
- 96
REFERENCES
Baillon, H. (1871) Sur les Quararibea.Adansonia 10: 146-149.

Cuatrecasas,J. (1948) Studies in South Americanplants, I. Lloydia 11: 185-225.
Cuatrecasas, J. (1954) Disertaciones sobre Bombacaceas. Rev. Acad. Colombiana
Cienc. Exact. Fis. Nat. 9 (35): 170-177.
Cuatrecasas, J. (1971) Miscellaneous notes on neotropical flora. Phytologia 20:
465-481.
Erdtman, G. (1952) Pollen morphology and plant taxonomy. Angiosperms (An
introduction to palynology. I): XII + 539 p., 261 fig., 1 front. Almqvist &
Wiksell, Stockholm.
Fuchs, H. P. (1967) Pollen morphology of the family Bombacaceae. Rev. Palaeo-
botan. Palynol. 3: 119-132.
Robyns, A. (1964) Bombacaceae, in Flora of Panama, Part VI. Ann. Missouri
Bot. Card. 51: 37-68.
Robyns, A. (1968) Bombacaceae neotropicae novae II. New species of Eriotheca,
Hampea and Quararibea.Ann. Missouri Bot. Gard. 55: 51-59.
Robyns, A. & Nilsson, S. (1970) Bombacaceaeneotropicaenovae III. A new species
of Quararibea from Panama. Bull. Jard. Bot. Nat. Belg. 40 : 353-356.
Robyns, A. & Nilsson, S. (1972) Bombacaceae neotropicae novae IV. Two new
species of Quararibeafrom Amazonia. Bull. Jard. Bot. Nat. Belg. 42: 347-352.
Tsukada, M. (1964) Pollen morphology and identification III. Modern and fossil
tropical pollen with emphasis on Bombacaceae.Pollen et Spores 6: 393-462.
Tsukada, M. (1965) Taxonomic significance of Bombacaceaepollen morphology.
Amer. Journ. Bot. 52: 652-653.
Vischer, W. (1920) Sur les QuararibeaAubl. Un genre de Bombacacees a ovaire
infere. Bull. Soc. Bot. Geneve, ser. 2, 11: 199-210.
LEGENDS TO THE FIGURES
(Abbreviations: PhMG = photomicrograph;SEMG = scanning electron micro-

graph; TEMG = transmissionelectron micrograph).
Fig. 1. - Quararibea cordata (Humb. & Bonpl.) Vischer (Tessman 3087, S),
pollen grain in equatorial view (SEMG X ca 550).
Fig. 2. - Id., part of the sexine with bacules visible in oblique view in the
lumina (SEMG X ca 2500).
Fig. 3. - Id., part of the sexine with an almost smooth orbicule on the surface
(SEMG X ca 3300).
Fig. 4. - Id., fragment of sexine showing reticulum and bacules standing on
the nexine (SEMG X ca 3500).
Fig. 5. - Id. (Little Jr. 6456, US): a, reticulate sexine pattern at high focus;
b, the same, bacules at high focus; c, the same, bacules at low focus
(PhMG X 750).
Fig. 6. - Id. (Sparre 18 216, S): a, reticulate-foveolate sexine pattern at high
focus; b, the same, bacules at high focus; c, the same, bacules at low
focus (PhMG X 750).
Fig. 7. - Id. (Tessman 3087, S), thin section through the exine showing a
murus (bacule) in longitudinal section, the nexine, and a rounded
orbicule in cross-section with smooth outline (TEMG X ca 8000).
Fig. 8. - Id., thin section through the exine with sexine consisting of tectum
and bacules, and one-layered nexine (TEMG X ca 4000).
-97
Fig. 9. - Quararibea guianensis Aubl. (Archer 7558, NY), pollen grain in

slightly oblique equatorial view with 2 spores visible and a reti-
culate-foveolate sexine pattern (SEMG X ca 550).
Fig. 10. - Id., detail of the sexine with smooth muri and rounded to oval
lumina and a finely granular nexine floor, the bacules short and
only occasionally visible (SEMG X ca 2200).
Fig. 11. - Id., pollen grain with reticulate-foveolate sexine patern, bacules at
high focus (PhMG X 750).
Fig. 12. - Id., pollen grain with foveolate-reticulate sexine pattern, bacules at
high focus (PhMG X 750).
Fig. 13. - Id., detail of the sexine with one pore with reinforced margin
(SEMG X ca 1200).
Fig. 14. - Id. (Irwin, Egler & Pires 47 196, MO), thin section through a pore,
the thickened pore margin consisting of lamellated nexine (TEMG
X ca 2600).
Fig. 15. - Id., thin section through the sporoderm showing sexine (muri and
bacules), nexine and intine (TEMG X ca 1900).
Fig. 16. - Id., thin section showing sexine with a relatively thick murus and
short bacules, and nexine subtended by intine (TEMG X ca 4000).
Fig. 17. - Quararibea asterolepis Pittier (Pittier 5575, US): a, reticulate pollen
grain showing bacules at high focus; b, the same, at low focus, the
muri supported by numerous, relatively tiny bacules (PhMG X 750).
Fig. 18. - Quararibea funebris (La Llave) Vischer (Skutch 1846, A): a, reticu-
late pollen grain with bacules at high focus; b, the same, at low focus,
the muri supported by few, relatively large bacules (PhMG X 750).
Fig. 19. - Quararibeavelutina Cuatr. (Schunke235, US), reticulate pollen grain
at low focus (PhMG X 750).
Fig. 20. - Id., reticulate-verrucosepollen grain from the same slide in optical
cross-section (PhMG X 750).
Fig. 21. - Quararibeamagnifica Pittier (Steyermark& Steyermark95 441, NY),
part of a reticulate sexine with a pore bounded by an elliptic mural
frame, the lumina (nexine floor) more or less angular-shaped and
with a few scattered granules or microprocesses (SEMG X ca 900).
Fig. 22. - Id., a single mesh at higher magnification showing rod-like processes
in the lumen (SEMG X ca 5700).
Fig. 23. - Quararibea duckei Huber (Silva 1357, BR), reticulate pollen grain
in equatorial view showing a pore with reinforcedmargin and lumina
of unequal size and shape (SEMG X ca 900).
Fig. 24. - Quararibea bracteolosa (Ducke) Cuatr. (Prance, Pena & Ramos
3468, US), part of a reticulate sexine with smooth muri and often
angular lumina of uneven size, the nexine being finely granular and
the orbicules rounded to ovoidal and smooth (SEMG X ca 2200).
Fig. 25. - Quararibea idroboi (Cuatr.) A. Robyns & S. Nilsson (Idrobo &
Schultes 1069, US), part of a reticulate sexine showing few, relatively
large and irregularlyshaped lumina surroundedby smooth, unevenly
thick muri, and few, sparsely spaced granules in the lumina on the
nexine floor (SEMG X ca 5500).
Fig. 26. ~ Quararibea bicolor (Ducke) Cuatr. [Ducke s.n. (RB 11419), US]:
a, pollen grain with relatively large verrucae or verruca-like processes
in high focus; b, the same in optical cross-section (PhMG X 750).
Fig. 27. - Id. (Rafael Lao M. 32, US): a, pollen grain with relatively small
verruca-like processes in high focus; b, the same in optical cross-
section PhMG X 750).
Fig. 28. - Quararibea dolichopoda A. Robyns (Correa, Dressier & Calderon
817, US), part of the sexine showing verrucae united by low and
narrow muri (PhMG X 750).
-98 -
Fig. 29. - Quararibea wurdackii A. Robyns (Wurdack2129, US), detail of the

sexine at the aperture region, the colpoid pore surrounded by elon-
gated mural fragments with a striate surface structure (SEMG
X ca 4100).
Fig. 30. - Quararibea ochrocalyx (K. Schum.) Vischer (Ducke 1089, MO), thin
section through the sporoderm showing sexine with luminal micro-
processes in longitudinal section and one-layerednexine subtended by
intine (double-layered?) (TEMG X ca 8000).
Fig. 31. - Id., thin section through sexine and orbicule in cross-section (TEMG
X ca 14000).
Fig. 32. - Quararibea longiflora (Gleason) Cuatr. (Lawrance 538, US), sexine
with wart-like elongated processes (mural fragments) partly inter-
connected, and nexine with a dense layer of microprocesses (SEMG
X ca 2400).
Fig. 33. - Id., part of a finely striate murus (mural fragment) and granular
microprocesses on the nexine (SEMG X ca 10500).
Fig. 34. - Id., an orbicule with a finely striate (rugulose) surface structure
(SEMG X ca 12500).
Fig. 35. - Quararibea malacocalyx A. Robyns & S. Nilsson (Klug 1110, NY),
portion of a verrucose pollen grain (SEMG X 750).
Fig. 36. - Quararibea ochrocalyx (K. Schum.) Vischer (Ducke 1783, NY), a
finely striate globular process supported by bacular stilts (SEMG
X ca 10000).
Fig. 37. - Id., finely striate mural fragment and granular microprocesses on
the nexine (SEMG X ca 10000).
Fig. 38. - Id. (Ducke 1089, MO), a clava-like process with a granular surface
texture (SEMG X ca 9000).
Fig. 39. - Quararibea lasiocalyx (K. Schum.) Vischer (Cowan 38362, NY), a
reticulate pollen grain in equatorial view with one pore visible at the
extreme right (SEMG X ca 650).
Fig. 40. - Id. (Ducke 962, US), part of the sexine at the pore; pore surrounded
by elongated fragmentswith a striate substructure,with in the center
finely striate granules (remains of an operculum?); nexine densely
beset with microprocesses (SEMG X ca 4000).
Fig. 41. - Id. (Cowan 38 362, NY), detail of a murus consisting of an entangled
structurecomposed by thin elongated elements (SEMG X ca 13 500).
Fig. 42. - Id. (Ducke 962, US), a clava-like process with a finely striate
structure (SEMG X ca 6000).
Fig. 43. - Id. (Pires & Cavalcante 52 654, MO), thin section through the spo-
roderm showing the sexine with a murus (an upper crenated tectal
part with one hollow and an infratectal bacular part) and short
luminal processes, the nexine and the intine in longitudinal section
(TEMG X ca 8000).
Fig. 44. - Id., an orbicule in cross-sectionwith one central cavity and a crenated
outline (TEMG X ca 9500).
Fig. 45. - Quararibea ochrocalyx (K. Schum.) Vischer (Krukoff 4719, A):
a, part of a reticulate pollen grain in high focus; b, the same in low
focus (PhMG X 750).
Fig. 46. - Quararibea coloradorum (Benoist) Cuatr. (Cazalet & Pennington
5096, NY), pollen grain in equatorial view, showing 2 pores and an
obscure sexine pattern (SEMG X ca 600).
Fig. 47. - Id. (Little Jr. 6158, US), part of the sexine with irregularly shaped
mural fragmentswith a striate structure,and granularnexine (SEMG
x ca 5000).
Fig. 48. - Id. (Benoist 3007, P), part of the sexine consisting of irregularstriate
fragments, and 2 finely striate orbicules (SEMG X ca 7500).
- 99 -
Fig. 49. - Id. (Cazalet & Pennington 5096, NY), an orbicule with a striate
(rugulose) surface pattern (SEMG X ca 12500).
Fig. 50. - Quararibeastenopetala (Standley& Cuatr.) Macbride (Klug 2972, F),
thin section showing mural fragments and short luminal processes
in longitudinal section, nexine and intine (TEMG X ca 8000).
Fig. 51. - Id., an orbicule with crenated outline at the upper left (TEMG
X ca 8000).
Fig. 52. - Id., pollen grain in slightly oblique polar view, with 3 pores clearly
visible (SEMG X ca 600).
Fig. 53. - Id., part of sexine at pore region (SEMG X ca 1100).
Fig. 54. - Quararibea asymmetrica (Cuatr.) Cuatr. (Cuatrecasas 15 152, US),
pollen grain in equatorial view with one pore with reinforced margin
in face view (SEMG X ca 650).
Fig. 55. - Id., part of the sexine showing fine loop-like sexine elements irre-
gularly distributed on the more or less smooth nexine (SEMG X ca
7500).
Fig. 56. - Id., thin section showing sexine elements, nexine and intine (TEMG
X ca 8000).
Fig. 57. - Id., an orbicule with one central cavity and an irregular, partly
deeply incised outline (TEMG X ca 3700).
Fig. 58. - Quararibea uribei Garcia-Barriga& Hernandez (Uribe-Uribe 1746,
COL), an orbicule with wrinkled surface (SEMG X ca 10 000).
: v . . . :
_ ....... ^X.j|,::;' . .. .*.... . .. ^

^,.:*: :;L*:! iF::** .: .--:
. .....! _1 B
l.i - - '<
-
... ~~ ~ ~ _ .}
.....
..
_ _ _ _::.s:
t W,
.X>>: . _
.. ,t _ X
»« :n5r :·...x '""'~' ..I
! .':':,.:...-'"::....:::,e2
t;
'
'~
..,.~. ' <,
,:· . ~~~~~~~~~:.*
'
.*
. . ,.
! S..
i,.:
"''f . '. :.
z
ir
t
.·i"'.
~~~iki ,,,.
q=..r _ .i:.: :·"*.''r
tn
:,·..··
I·..'illd:·.l'':·.
·J
*1..·
·s:
^ ^ ;:
7'
fIESf,
Fig. 1-8. - Quararibea cordata-type.
440010C."~'
."'It
~
Ati~~Oij S
~.,3re~~-;'lss V
0Pd~k
iI
Oo~t~;
14
::
V na;
·
*r
a::i LL C C
r
I· -·4; I i:·
6P
r _·.
·.
n.,
6
:r*·c·'·,..·*
jR·'::*r&
..;
I:;.
··.s.i.
15
' ".'"::',":*
>^ i?"~~
e::'*'^:
,s~..
.,,-sf
*-··;4~~
.:9.../,
.
11
Fig. 9-16.- Quararibea asterolepis-type.

Fig. 9-16. - Quararibeaasterolepis-type.
17a
,r <
.="
.. - 22
L4
Fig. 17-25. - Quararibea asterolepis-type.
26bLU.
30
31
a
Lim
Fig. 26-3 1. - Quararibea bicolor-type.
36^ *f
I
R , s ''
I ' .. -
i s
.
o'
V.1- *
B^Fs
*
'I
---..' "
a I
4
Fig. 32-38. - Quararibea bicolor-type.
,,ri~ ~~~ ~.n-7.~21
;4.4
Fig. 39, 41, 43-45. - Quararibea intricata-type.

Fig. 40, 42. - Quararibea bicolor-type.
f^^B...
Fig. 46-51. - Quararibea stenopetala-type.
Fig. 52-53. - Quararibea stenopetala-type.
Fig. 54-58. - Quararibea asymmetrica-type.

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Pollen Morphology and Taxonomy of the Genus Quararibea S.l.

Pollen morphologyand taxonomy

S. Nilsson & A. Robyns (*)

Abstract. - The pollen of 61 species of Quararibeas.l. (Matisia included) has

The mainly South American genus Quararibea has been treated

(*) S. Nilsson, PalynologiskaLaboratoriet,Wallenberglaboratoriet,Lilla Frescati,

pollen morphology of the Bombacaceae including 25 species of

MATERIALS AND METHODS

Pollen material was obtained from specimens in the following

GENERAL POLLEN DESCRIPTION

Pollen grains radially symmetrical, isopolar, medium-sized to

128 ,/m), suboblate to spheroidal, 3-6-(zono-)porate, the pores

KEY TO THE POLLEN TYPES

1. Pollen grains reticulate to foveolate, infrequently ornate, the muri smooth,

DESCRIPTION OF THE POLLEN TYPES

1. Q. cordata-type.- Fig. 1-8.

section, the lumina or foveolae usually rounded, (0.5-)1-5 ~m in

to have typically foveolate to reticulatepollen grains with more or

3. Q. bicolor-type. - Fig. 26-38, 40, 42.

collection of Q. bicolor [Ducke s.n. (RB 11419), fig. 26] which

5. Q. stenopetala-type. - Fig. 46-53.

CORRELATION OF POLLEN TYPES

petala-type or modified into more or less isodiametric processes

Table 1. Hypothetical relationship between the recognized pollen types

From a pollen morphologicalstandpointit does not seem justified

petala was suggested by Cuatrecasas (1971: 468). Pollen morphol-

Pollen types Macromorphology

First major pollen group

Table 2. Relation between pollen types and macromorphology

Quararibea amazonica Ulbrich, Verhandl. Bot. Ver. Brandenburg 50:

Quararibea amplifolia (Pittier) Cuatr., Lloydia 11: 191 (1948).

Quararibeaapaporiensis(Cuatr.)A. Robyns & S. Nilsson comb. nov.

Quararibea aristeguietae Cuatr., Bol. Soc. Venez. Cienc. Nat. 26:

Quararibea asterolepis Pittier, Fedde Repert. Sp. Nov. 13: 316

Quararibeaasymmetrica(Cuatr.)Cuatr., Lloydia 11: 191 (1948).

Quararibeabicolor (Ducke) Cuatr., Lloydia 11: 191 (1948).

Quararibeabilobata A. Robyns, Ann. Missouri Bot. Gard. 55: 54

Quararibeabolivarii (Cuatr.)Cuatr., Lloydia 11: 191 (1948).

Quararibeabracteolosa (Ducke) Cuatr., Lloydia 11: 191 (1948).

Quararibea castano (Triana & Karsten) Cuatr., Lloydia 11: 191

Quararibea ciroana Cuatr., Trop. Woods 101: 16 (1955).

Quararibea coloradorum (Benoist) Cuatr., Lloydia 11: 191 (1948).

Quararibea cordata (Humb. & Bonpl.) Vischer, Bull. Soc. Bot.

Quararibea cornucopiae (Triana & Planchon) Vischer, Bull. Soc.

Quararibeacruceto (Cuatr.) Cuatr., Lloydia 11: 191 (1948).

Quararibeadolichopoda A. Robyns, Ann. Missouri Bot. Gard. 54:

Quararibeadolichosiphon A. Robyns & S. Nilsson, Bull. Jard. Bot.

Quararibea dowdingii (Sprague) Vischer, Bull. Soc. Bot. Geneve,

Quararibea duckei Huber, Bull. Soc. Bot. Geneve, ser. 2, 6: 186

Quararibeafieldii Millsp., Field Mus. Nat. Hist., Bot. Ser. 1: 309

Quararibea floribunda (St.-Hil. & Naudin) K. Schum. in Mart., Fl.

Quararibeafoenigraeca Cuatr., Phytologia 4: 473 (1954).

Quararibeafunebris(La Llave)Vischer,Bull. Soc. Bot. Geneve,ser. 2,

QuararibeagigantifloraPittier,Bol. Soc. Venez. Cienc. Nat. 10: 114

Quararibea grandifolia (Little Jr.) Cuatr., Lloydia 11: 191 (1948).

Quararibeaguatemalteca(J. D. Sm.) Standley& Steyerm.,Field Mus.

Quararibeaguianensis Aubl., Hist. P1. Gui. Fr.: 692 (1775).

Quararibea huallagensis (Cuatr.) Macbride, Field Mus. Nat. Hist.,

Quararibeaidroboi (Cuatr.) A. Robyns & S. Nilsson comb. nov.

Quararibeaintricata A. Robyns & S. Nilsson, Bull. Jard. Bot. Nat.

Quararibealasiocalyx (K. Schum.) Vischer, Bull. Soc. Bot. Geneve,

Quararibealecythicarpa(Ducke) Cuatr., Lloydia 11: 191 (1948).

Quararibea leptandra Cuatr., Lloydia 11: 185 (1948).