matisia and quararibea (bombacaceae) should be retained as separate genera

Matisia and Quararibea (Bombacaceae) Should Be Retained as Separate GeneraAuthor(s): William S. AlversonSource: Taxon, Vol. 38, No. 3 (Aug., 1989), pp. 377-388Published by: International Association for Plant Taxonomy (IAPT)Stable URL: http://www.jstor.org/stable/1222268 .

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TAXON 38(3): 377-388. AUGUST 1989

MATISIA AND QUARARIBEA (BOMBACACEAE) SHOULD BE RETAINED AS SEPARATE GENERA

William S. Alverson'

Summary The genera Quararibea Aublet and Matisia Humboldt & Bonpland were described 214 and 184

years ago, respectively, and have been variously combined and separated during their history. This study indicates that Quararibea differs completely from Matisia by its reduced ovarian locule number (2 to 4, versus 5 in Matisia), and, less definitively, by its short (to 4 mm), radially-symmetrical staminal lobes. In keeping with the original generic concepts, these genera are easily distinguished and should be maintained as separate. An examination of the variability of macromorphological and pollen characters suggests that Quararibea is a relatively uniform derivative of a Matisia-like ancestor. The presence or absence of a spicy odor in dried leaves is not a reliable indicator of generic identity, despite previous claims that the odor was a practical means by which to distinguish the two genera.

Introduction The pantropical family Bombacaceae includes about 31 genera and 250 species, among

which are the well-known baobab (Adansonia), kapok (Ceiba), durian (Durio) and balsa (Ochroma). Other, poorly-known genera occur as ecologically important components of primary lowland tropical forests of the New World. The largest of these, Matisia Humb. & Bonpl. and Quararibea Aubl., together total at least 60 species. Both genera are members of the tribe Quararibeae (Garcia-Barriga, 1952; the Matisieae of Hutchinson, 1967), which is defined primarily on the basis of its "simple" (perhaps unifoliolate) leaves and the fusion of filaments into elongate staminal columns.

At the commencement of a recent taxonomic revision of Mesoamerican and Antillean Quararibea (Alverson, 1986), the genus was treated broadly (inclusive of Matisia), its generic limits based on Robyns' treatment for the Flora of Panama (1964) and the last comprehensive monograph of the genus by Vischer (1919). During the course of these studies, I was forced to conclude that these are quite distinct taxa worthy of recognition at the generic level.

The following is an annotated history of the relationship of these two genera, with special attention given to characters used to combine or separate them. I present and discuss new morphological evidence that they are distinct entities. Finally, a key to the genera and a few generic transfers are given.

Taxonomic History of Matisia and Quararibea The original description of the genus Quararibea consisted of a plate of Quararibea

guianensis (Fig. 1) and a terse description published by Fus6e Aublet in 1775 (pp. 691- 694) based on material collected in French Guiana. Both the plate and the description illustrate characters that typify the genus and contrast it with the later-described Matisia: A long staminal tube formed by fused filaments (10 cm long in this instance) and bearing short apical teeth, and a 2-locular ovary.

OlofSwartz published a very brief description ofMyrodia turbinata, based on Caribbean material, in his Prodromus (1788: 102-103). At the same time, he placed Quararibea guianensis under the new name Myrodia longiflora, making it evident that he viewed the two genera as synonymous. This view was repeated twelve years later in his Florae indiae

Herbarium, Botany Department, University of Wisconsin, Madison, WI 53706, U.S.A.

377



378 TAXON VOLUME 38

occidentalis (1800: 1225-1228, pl. 22). The genus Myrodia is now universally recognized as a synonym of Quararibea.

The discovery of a new bombacaceous tree in Colombia by Alexander von Humboldt and Aime Bonpland led to the publication of Matisia cordata in 1805, including two superb plates (Figs. 2, 3) dedicated to Francisco Matis, an artist with Mutis' Royal Botanical Expedition of New Granada. These authors provided a comparison with the previously known Quararibea (as Myrodia, p. 12):

Les deux especes de Myrodia connues sont des arbustes a feuilles lanceolees, presque sessiles, et disposres sur toute la longueur des rameaux. Le Matisia est un grand arbre a feuilles en coeur, port~es sur de longs petioles, et rapprochbes les unes des autres a l'extremite des jeunes rameaux. Les fleurs dans le Myrodia sont regulibres et axillaires; celles du Matisia sont irregulieres et 6parses sur les branches. Les 6tamines du premier sont reunies en un tube qui porte les antares A son sommet et qui n'est pas divise comme dans le Matisia: enfin le fruit de ce dernier est une baie a cinq loges, tandis que celui du Myrodia est un drupe sec qui ne contient jamais plus de deux graines.

The elegant description, plate and discussion include the two critical morphological characters that differentiate Matisia from Quararibea, namely, its staminal tube terminated by five long lobes (ca. '/3 the length of the staminal column in this case), and its five-locular ovaries. Humboldt and Bonpland's further characterization of generic differences, although accurate relative to the species then known, cannot be strictly applied to the two genera at present. There is a valid contrast between the tendency of species of Quararibea to have smaller, lanceolate, short-petiolate leaves distributed more or less evenly along the branches versus the tendency of Matisia species to have larger, cordate (or at least cordate-based), long-petiolate leaves clustered at the branch tips. However, the differences of stature and inflorescence position that are emphasized in their discussion are not consistent, as relatively large and small species, and ramiflorous species, exist in both genera (Alverson, 1984, 1986). Their characterization of Myrodia (i.e., Quararibea) fruits as dry drupes is somewhat inaccurate, the impression probably derived from statements to this effect by both Aublet (1775: 693) and Swartz (1788: 102). The "Myrodia" fruits discussed by these authors are comparable to many other Quararibea species (e.g., Q. asterolepis Pittier and Q. yunckeri Standley) and can be more accurately described as having firm, moist-fibrous mesocarps when immature, which become increasingly soft and fleshy when ripe. In any case, Hum- boldt and Bonpland clearly stated the criteria for differentiating Matisia and Quararibea in their Plantae aequinoctiales (1805: 10-12), and reiterated their generic concept in the Nova genera et species plantarum (Humboldt et al., 1823: 306-308).

Augustin Pyramus de Candolle recognized Myrodia (Quararibea) and Matisia as separate genera in his Prodromus (1824: 477):

[Myrodia] Calyx nudus tubulosus 4-5-dentatus lateraliter rumpens. Petala oblongo-linearia. Sta- minum columna longa. Antherae 10-15 [i.e., 20-30 thecae, in pairs]. Capsula drupacea 2-3-locularis, loculis 1-spermis. Bracteolae 2-3 in pedicello floris.

[Matisia] Calyx nudus irregulariter 2-5-dentatus rumpens, persistens. Petala ovata. Staminum columna apice 5-fida, antheris in quoque lobo 12 [i.e., 12 unpaired thecae on each of 5 lobes], ad latus externum secundis. Stigma 5-sulcatum. Drupa ovata 5-locularis, loculis 1-spermis. Albumen fari- naceum? Cotyledones corrugatae.

Candolle's generic descriptions are important in a number of respects. For the first time, Quararibea (sub Myrodia) fruits are correctly characterized as variable, with two to three locules, a fact overlooked by many later workers. Furthermore, the concept of Matisia promoted by Humboldt and Bonpland is faithfully followed. Of final note, the descriptions foreshadow the confusion of subsequent literature that variously considered the anthers of the two genera as either monothecate or bithecate.



AUGUST 1989 379

S: t 2

i

.10

12-

(/uiair ?iika GI•, •: 1 •/,'/IIt i;r

Fig. 1. Type illustration of Quararibea guianensis in Aublet (1775: pl. 278, caption p. 692). The original caption follows: "1. Gemma floris cujus pedunculus multi-glandularis [in reference to the bracteoles]. 2. Calix. 3. Corolla expansa. Stylus. Stamina. 4. Petalum segregatum. 5. Calix apertus. Pistillum. Stamina. 6. Tubus apertus. Germen. Stylus. Stigma. 7. Pars tubi ferentis stamina. 8. Stigma. 9. Stamina. 10. Glandule [thecae]. 11. Capsula a calice tecta. 12. Pars capsula cum amygdata. 13. Dua amygdala. 14. Amygdala interne visa."



380 TAXON VOLUME 38

A . 5. ' 0i.

'44?er "'wo,

MAT ISIA (CoXdata.

Al' tLqww? ?

Fig. 2. Type illustration of Matisia cordata in Humboldt and Bonpland (1805: pl. 2a). The original caption reads "1, calice; 2, fleur dont on a 6te le calice pour montrer la disposition et la forme des p6tales; 3, style entour6 seulement par une partie du tube des 6tamines, et dont on a coup6 la partie sup6rieure'pour montrer le stigmate; 4, idem, d6pourvu des &tamines et montrant la forme de l'ovaire; 5, feuille dont on a doubl6 le p6tiole; 6, portion de branche montrant le disposition des fleurs."



AUGUST 1989 381

5,

ii a.

Fig. 3. Type illustration of Matisia cordata in Humboldt and Bonpland (1805: pl. 2b). The original caption reads "1, fruit entier; 2, idem, coup6 horizontalement pour montrer la disposition et la forme des graines et des cotyledons; 3, meme coupe d'un jeune fruit; 4, graine entiere; 5, un cotyledon vu par sa partie interne."



382 TAXON VOLUME 38

One year later, Pablo de La Llave published Lexarza, a new genus based on Mexican material that contained a single species, L. funebris, without reference to the older genera Matisia, Myrodia or Quararibea (Llave and Lexarza, 1825: 12). This distinct and valid species, Quararibea funebris (Llave) Vischer, has long been controversial in generic delim- itation of Quararibea due to its putatively four-locular flowers and fruits, a matter discussed below.

Henri Baillon (1871-1873, 1872-1873, 1875) was the first to synonymize Matisia, My- rodia and Quararibea, placing all under Quararibea, the oldest name. He justified these transfers by stating that: a. The fruits share many characteristics, based on his examination of an original drawing of Matisia cordata by Matis at the Paris Herbarium; b. Flowers of Myrodia [Quararibea] cacao Triana & Planchon represent a condition intermediate between Matisia and Quararibea; c. Leaves of many Matisia species, such as M. cornucopiae Triana & Planchon and M. castano Karsten & Triana, both then newly described, are extremely similar to those of many Myrodia and Quararibea species.

Baillon's transfer of Matisia into Quararibea is unwarranted for three reasons. First, the critical fruit character, i.e., number of locules, is always five in Matisia but variable between two and four within individuals and species of Quararibea (Table 1). The fact that there is no overlap in this character between the two genera was overlooked by his focus on shared, primitive (symplesiomorphic) characters found in fruits of other genera of Quar- aribeae as well (e.g., Patinoa Cuatr. and Phragmotheca Cuatr.). Secondly, from a personal examination of type material (Bonpland 1575 at P; Goudot 105 at G, P), Myrodia cacao is morphologically a typical species of Quararibea and cannot be considered an intermediate as it was by Baillon. Its fruits are two-locular and its flowers, very similar to those of Quararibea funebris of Central America, have staminal columns with a radially-symmetrical set of five apical lobes less than 4 mm long (Table 1). Finally, while it is true that the leaves of some species of Matisia greatly resemble those of some Quararibea species (e.g., leaves of Matisia ochrocalyx Schumann resemble the leaves of Quararibea guianensis more than they do those of Matisia cordata or M. arteagensis Cuatr.), this observation does not provide a strong argument for combining the two genera. It is, however, indicative of the great variability seen among the species of Matisia, other species of which bear leaves almost indistinguishable from Phragmotheca Cuatr.

The most all-inclusive and perhaps most influential treatment of Quararibea is that of Wilhelm Vischer (1919). With an argument analogous to that of Baillon, this Swiss botanist presented Q. fieldii Millspaugh as an intermediate between Quararibea and Matisia, based on the depth of the staminal lobes (p. 201):

Il n'existe pas non plus une difference bien tranchee entre la forme de la colonne staminodiale des Archiquararibea [his subgenus, containing Q. guianensis and other species with very short apical teeth on the staminal column] et des Matisia. Chez les premiers, on ne voit, en general, que cinq dents obtuses; chez les derniers, cinq branches bien separees; mais pour ce caractere le Quararibea fieldii Millsp. est parfaitement intermediaire.

In his opinion, the coup de grace to the maintenance of Quararibea and Matisia as separate genera can be found in the ovarian characteristics of Quararibea (Lexarza)funebris: "I1 ne reste donc comme caractere distinctif que le nombre des loges de l'ovaire. Mais il perd aussi sa valeur generique par la constatation que dans le Quararibea funebris il y a exactement un nombre intermbdiaire [i.e., 4 locules]."

In view of the morphological information now available, Vischer's first argument is weak because the type of QuararibeaJieldii Millsp. (Gaumer 879, F!) has symmetrically-arrayed staminal lobes only 3 mm long, within the range of variation of Quararibea s. str. Note that in comparison to his illustration of the staminal column apex of Q. Jieldii, the apex of Q. fynebris indicates very little lobing (Fig. 4A, B). Vischer's illustrations are misleading; my measurements of all available dried and liquid-preserved material show lobe length to



AUGUST 1989 383

vary from 2-4 mm in the latter species. In fact, I consider these two taxa to be conspecific and barely separable at the level of subspecies (Alverson, 1986). The erroneous impression given by Vischer's illustrations may be due to the fact that the staminal column of the type of Q. fJieldii was pressed and dried in such a way that the apical lobes were spread, giving the impression that they were rather profoundly divided. Furthermore, the type illustration of Q. fJieldii includes a drawing of the apex of its staminal column, drawn as if cut longitudinally on one side so as to better illustrate its internal anatomy (Fig. 4C). It is possible that an examination of this plate led Vischer to erroneously believe that the apical lobes of the staminal tube of Q. fJieldii were divided unequally and arrayed asymmetrically, as they are in most species of Matisia (Fig. 4D).

Vischer's contention that Quararibea funebris represents a true intermediate between Quararibea and Matisia must also be rejected. The variability in locule number (two to four) within Q. funebris is in keeping with other species of Quararibea s. str. and is completely distinct from the uniformly five-locular species of Matisia (Table 1).

In the three decades following Vischer's work, many new Central and South American species of Quararibea and numerous floristic treatments were published, virtually all of which treated the species within an all-inclusive Quararibea (e.g., Cuatrecasas, 1948, and Garcia-Barriga, 1952). This view was later rejected by Cuatrecasas himself (1954a, 1954b, 1971) who, upon further consideration, decided that the two genera were distinct. Schultes (1957) followed Cuatrecasas by maintaining the two genera as distinct. In contrast, Robyns (1964) combined the two, citing Baillon (1871-1873) and Vischer (1919).

Palynological investigations by Nilsson and Robyns (1974, 1986) have been the most recent word on this subject. They maintain Matisia within Quararibea because of the lack of correlation between macromorphology and pollen morphology in the group. In 1974, they described six pollen "types"; five were composed solely of species considered here to be Matisia, and one was dominated by Quararibea s. str. but also included Matisia bracteolosa and eight closely related species of Matisia.

It is my view that these palynological data do not contradict the use of macromorphological characters to separate the two genera. The specialized pollen type shared by all species of Quararibea s. str. and the Matisia bracteolosa alliance probably indicates a close evolutionary association, i.e., the "asterolepis-type" pollen seems to represent a synapo- morphy relative to the remainder of the Bombacaceae. However, within this synapomor- phic group defined by pollen is nested another group, Quararibea s. str., that is defined by a reduction of locule number and increased staminal fusion. Overall, it is not clear that palynological data should take precedence over macromorphological data that define monophyletic groups in this family. The primacy of palynology as a basis for generic limits in the Bombacaceae is questioned elsewhere by Steyermark and Stevens (1988), who argue that pollen differences within Bombax L. s.l. are insufficient in and of themselves to justify Robyns' (1963, 1971) segregation of Rhodognaphalopsis, which they consider as synonymous with Pochota Ramirez Goyena (Bombacopsis Pittier). In a somewhat analogous manner, I argue that palynological data alone form insufficient grounds to impede the segregation of Quararibea from Matisia on macromorphological characters, unless such data were to demonstrate that Quararibea is polyphyletic, which they do not. The sound macromorphological differences between these two genera should take precedence in this case, particularly in view of the parallel and convergent evolution of pollen characters that seems to have occurred in the family (Nilsson and Robyns, 1986).

The palynological studies of Nilsson and Robyns corroborate a pattern seen in nearly every other morphological character of these genera: Quararibea is much more uniform, Matisia much more variable. This generalization holds for leaf, corolla, staminal column, pollen, fruit, and seedling characters, but not for locule number (Table 2). Thus, Quararibea appears to be a relatively uniform, specialized derivative of a Matisia-like ancestor. Tests of this hypothesis should be directed towards determining whether the reduced number of



384 TAXON VOLUME 38

'-.

f. f?z~ I

iir

A

~t~? v C~

Fig. 4. Staminal column apices of three Quararibea and one Matisia species. In each, the distal portion of the staminal tube and its apical lobes are shown. Thecae are visible in 4A-C, and the distal portion of the style and the stigma are visible in 4B and 4D. A. Quararibeafieldii, with remote thecae as an artifact of drying; B. Q. funebris; C. Q. fieldii, with tube cut longitudinally to illustrate internal anatomy; D. Matisia malacocalyx, showing the long, asymmetrically-arrayed staminal lobes typical of Matisia. (4A, B from Vischer, 1919: fig. 3; 4C from Millspaugh, 1896: pl. 19; 4D redrawn from Robyns and Nilsson, 1972: fig. 6.)

locules and increased fusion of staminal filaments seen in Quararibea s. str. truly define a monophyletic group. Segregation of Quararibea makes Matisia a paraphyletic group but, if deemed necessary, the situation could be resolved by further segregation of genera from this heterogeneous assemblage once molecularly-based phylogenies, and additional character state data for some of the poorly known species, become available.



AUGUST 1989 385

Table 1. Variability of ovarian locule number, and staminal lobe length and symmetry, of all Central American and selected South American species of Quararibea and Matisia. Data from type descriptions, herbarium specimens, and Alverson (1986).

Number of locules Staminal lobes

in ovary Symmetrya Length (mm)

Species 2 3 4 5 SY AS Min. Max.

Q. amazonica Ulbr. X X 0.8 1.0 Q. aristeguitae Cuatr. X X 1.0 1.5 Q. aurantiocalyx Alversonb X X 2.5 3.0 Q. bilobata Robyns X X 2.0 4.0 Q. cacao Triana & Planchon X X 2.5 3.3 Q. ciroana Cuatr. X X - ca. 1

Q. duckei Huber X X - ca. 2

Q. gomeziana Alversonb X X - 1.2

Q. guianensis Aublet X X 0.8 1.4 Q. parvifolia Standley X X 0.3 0.7 Q. pendula Alversonb X X 0.8 1.1 Q. platyphylla Pittier & J. D. Smith X X 0.7 2.3

Q. santaritensis Alversonb X X - ca. 1

Q. velutina Cuatr. X X - ca. 2

Q. yunckeri Standley X X 0.4 1.4 Breedlove 35002c X X 0.7 0.9

Q. asterolepis Pittier X X X 0.5 3.0

Q. costaricensis Alversonb X X X 0.5 2.5 Q. turbinata (Sw.) Poiret X X X 0.8 1.5

Q. pumila Alverson X X X 0.9 1.1

Q. funebris (Llave) Vischer X X X X 0.5 4.0

Q. pterocalyx Hemsley X X X X 1.0 3.1 M. alata Little X ? X 10 16 M. arteagensis Cuatr. X ? ? - 6 M. bicolor Ducke X ? X 6 8 M. bracteolosa Ducke X X 5 9 M. coloradorum Benoist X ? X 12 14 M. cordata Humb. & Bonpl. X X X 12 21 M. dolichopoda (Robyns) Cuatr. X X 4 6 M. dolichosiphon (Robyns & Nilsson) Alverson X X 11 13 M. exalata Alversonb X X - 8 M. huallagensis Cuatr. X ? X 8 10 M. intricata (Robyns & Nilsson) Alverson X ? ? 4 5 M. jefensis (Robyns & Nilsson) Alverson X X 5 7

M. leptandra (Cuatr.) Cuatr. X X 8 10 M. longitubulosa (Robyns) Cuatr. X ? ? - 20 M. malacocalyx (Robyns & Nilsson) Alverson X X 7 8 M. muricata (Cuatr.) Cuatr. X X 6 8 M. obliquifolia Standley X X 7 10 M. ochrocalyx Schumann X X 5 8 M. palenquiana (Robyns) Alverson X ? ? 3 4 M. sanblasensis (Robyns) Cuatr. X X - -

M. schultesii (Cuatr.) Cuatr. X X 4 5 M. soegengii Cuatr. X X 20 22 M. stenopetala Standley & Cuatr. X X 8 9

a SY = staminal lobes radially symmetrical; AS = lobes asymmetric; ? = unknown but possible symmetry. b Recently published species (Alverson, 1989, in press). c Denotes type of sp. nov. ined. in Alverson (1986).



386 TAXON VOLUME 38

In summary, morphological differences between Matisia and Quararibea that were elu- cidated 184 years ago still serve to readily distinguish them in fruit and flower. Based on the correlated characters of locule number and staminal column lobing they form two distinct groups that should be maintained as separate genera.

Key to the Genera Apical lobes of staminal column usually unequally divided and not radially symmet-

rical at anthesis, (3) 4-22 mm long; ovaries uniformly 5-locular ............... ......................................................... Matisia Humb. & Bonpl. Apical teeth or lobes of staminal column radially symmetrical, to 4 mm long; ovaries

with 2 to 4 locules ................. ...................... Quararibea Aubl.

A small number of generic transfers are necessitated by the preceding discussion. Most other transfers were made previously by Cuatrecasas (1954a, 1971).

Matisia dolichosiphon (Robyns & Nilsson) Alverson, comb. nov. Quararibea dolichosiphon Robyns & Nilsson, Bull. Jard. Bot. Nat. Belg. 40: 353-356. 1970.

Matisia intricata (Robyns & Nilsson) Alverson, comb. nov. Quararibea intricata Robyns & Nilsson, Bull. Jard. Bot. Nat. Belg. 42: 347-349. 1972.

Matisia jefensis (Robyns & Nilsson) Alverson, comb. nov. Quararibea jefensis Robyns & Nilsson, Bull. Jard. Bot. Nat. Belg. 45: 323-325. 1975.

Matisia malacocalyx (Robyns & Nilsson) Alverson, comb. nov. Quararibea malacocalyx Robyns & Nilsson, Bull. Jard. Bot. Nat. Belg. 42: 349-352. 1972.

Matisia palenquiana (Robyns) Alverson, comb. nov. Quararibea palenquiana Robyns, Bull. Jard. Bot. Nat. Belg. 46: 235-236. 1976.

In conclusion, a previously reported but erroneous criterion for distinguishing the two

genera should be mentioned. Although chemical investigations of the Quararibeae tribe of the Bombacaceae are still in their infancy, their origins stretch back more than 200 years. Aublet (1775) and other early investigators of Quararibea (Swartz, 1800; Triana and Plan- chon, 1862; Baillon, 1871-1873) noticed the extremely persistent, spicy fragrance emanated

by dried specimens. As noted by Schultes (1957), some specimens more than a century old retain this odor, which has been variously described as that of sweet clover (Melilotus spp.), slippery elm (Ulmus rubra Muhl.), curry powder, fenugreek (Trigonella foenum- graecum L.), licorice (Glycyrrhiza glabra L.), and cudweed or everlasting (Gnaphalium obtusifolium L.). The chemical principle responsible for the odor has recently been identified as a lactone (Raffauf and Zennie, 1983), one of a family of metabolites that includes the first alkaloid reported from the Bombacaceae (Raffauf et al., 1984).

Beginning about the time of Triana and Planchon, and greatly amplified by Schultes

(1957), the idea has been promoted that Quararibea and Matisia can be separated on the basis of the presence or absence of this spicy odor in dried specimens. The evidence does not support this claim. While it is true that no species of Matisia have this odor, there are also many species of Quararibea s. str. without it. I have yet to detect this odor in fresh leaves of either genus, and it remains to be elaborated how various methods of drying affect the chemical processes by which the fragrant lactone becomes evident in dried leaves. While I expect that careful analyses of these chemicals and their biosynthetic relatives will

someday aid in the construction of a taxonomic system that more accurately reflects the phylogeny of this group, the simple presence or absence of their odor in dried specimens does not corroborate any morphologically-based generic concept proposed to date.

Acknowledgments I thank Hugh H. Iltis, Stephen Solheim, and W. D. Stevens for critical review of the manuscript,

and the members of my Ph.D. thesis committee at the University of Wisconsin, Madison, who



AUGUST 1989 387

Table 2. Comparison of morphological variability within Quararibea and Matisia. Data from type descriptions, herbarium specimens, or from Alverson (1986).

Character Quararibea Matisia

Leaves

Shape of blade Entire, elliptic or ovate to mildly Entire or rarely lobed, elliptic, obovate (strongly obovate in Q. ovate, obovate or cordate, some pumila)a, sometimes mildly species strongly asymmetric asymmetric

Number of palmate 1 to 5 3 to 14 basal veins on blade

Petiole length (mm) 3 to 29 10 to 530

Flowers Presentation Single or paired, on new growth Single, paired or in multiflorous

near branch tips (cauliflorous in panicles, on young growth near Q. pumila) branch tips, ramiflorous or trun-

ciflorous Calyx Bracteolate in flower; usually green Bracteolate or ebracteolate; green

or greenish-brown (pale orange to dark brown, orange or gold- in Q. aurantiocalyx); generally en; alate in many species; persis- unwinged (winged in 3 species); tent or deciduous persistent

Corolla Actinomorphic to slightly zygo- Slightly to strongly zygomorphic morphic

Petal color White or greenish-white White, pale orange, pink, salmon, greenish-blue, red- or dark vi- olet

Pollen Uniform -of one type ("asterole- Variable--of six types pis type" of Nilsson and Ro- byns, 1974, 1986)

Fruits Number of locules 2 to 4 5 Number of seeds 1 to 4 1 to 5 Color of exocarp Green, yellowish-green, or orange Greenish-brown or brown to

brownish-orange

Seedlings Uniformly hypogeal and cryptocot- Mostly epigeal and phanerocoty- ylar, sensu Ng (1978); cotyle- lar; some hypogeal and crypto- dons uniformly smooth, fleshy cotylar; cotyledons greenish and and achlorophyllous foliose or thick and bullate, or

as in Quararibea

a Exceptions to many generalities about Quararibea are found in Q. pumila, a highly specialized endemic in northern Costa Rica that has evolved a palmoid growth form and dwarf stature (Alverson, 1984). In contrast, the morphological variabilty shown in Matisia is partitioned more evenly across many species or groups of species.

reviewed an earlier version of this work. The Missouri Botanical Garden provided photos of Aublet's plate of Q. guianensis, Claudia Lipke lent her photographic expertise, and Kandis Elliot prepared Fig. 4. Fieldwork in Latin America during 1982, 1983, 1985 and 1986 was made possible by grants from the J. S. Noyes Foundation via the Organization for Tropical Studies, and the E. K. and 0. N. Allen Herbarium Fund, the J. J. Davis Fund, and the Wisconsin Natural History Council of the University of Wisconsin, Madison, to whom I express my gratitude.

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matisia and quararibea (bombacaceae) should be retained as separate genera

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